Annals 
of the © 


Missouri 


umber 


Volume 76, Number 1 
Spring 1989 


Annals of the 
Missouri Botanical Garden 


SEU. QU S10 A A UNE ERE 
The Annals, published quarterly, contains papers, primarily in systematic botany, con. 
tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the 
Garden will also be accepted. Authors should write the Editor for information concerning 


arrangements for publishing in the ANNALS. Inst 


of the last issue of each volume. 


Editorial Committee 


George K. Rogers 
Editor, Missouri Botanical Garden 


Amy Scheuler 
Editorial Assistant, 
Missouri Botanical Garden 


Glenda Nas. 
Magdalen Lampe 
Publications Staff 


ructions to Authors are printed in the back 


Marshall R. Crosby 


Missouri Botanical Garden 


Gerrit Davidse 
Missouri Botanical Garden 


John D. Dwyer : 
Missouri Botanical Garden & 
Saint Louis University 


Peter Goldblatt - ( Um 


Missouri Botanical Garden 


Dale E. Johnson 


_ Missouri Botanical Garden 


Henk van der Werff j 


Missouri Botanical Garden - 


For subscription information contact Department 
Eleven, P.O. Box 299, St. Louis, MO 63166. Sub- 
scription price is $75 per volume U.S., $80 Canada 


i d Mexico, $90 all other countries. Airmail deliv- _ 


-ery charge, $35 per volume. Four issues per vol- 


(O Missouri Botanical Garden 1989 —— 


Eleven, P.O. Box 299, St. Louis, MO 63166. 


The ANNALS OF THE Missouri BOTANICAL GARDE 
(ISSN 0026-6493) is published quarterly by th 
Missouri Botanical Garden, 2345 Tower Grove At 
enue, St. Louis, MO 63110. Second class posté 


paid at St. Louis, MO and additional mailing office 


POSTMASTER: Send address changes to Annals f 
THE Missouri BOTANICAL GARDEN, me? 


Volume 76 
Number 1 
1989 


Annals 
of the 
Missouri 
Botanical 


Garden 


NA 


TESSMANNIANTHUS, 

AN ARBORESCENT GENUS 
OF MELASTOMATACEAE 
NEW TO PANAMA! 


Frank Almeda? 


ABSTRACT 


Recent agr o. activity in montane cloud forests of western and central Panama has led to the discovery of two 


new species o 
eru. TE im du carinatus and T. g 


trees previously recorded from Colombia, 
ordonii are described, illustrated, compared, and contrasted with similar 
vided. 


smannianthus, a little-known genus of 


cuador, and 


species. À key to the new taxa and a md history of the genus are also prov 


In a paper describing new Peruvian flowering 
plants collected by the late Günther Tessmann, 
Markgraf (1927) proposed the new genus and 
species Tessmannianthus heterostemon, to accom- 
modate one of the tallest known trees (30-45 m) 
among neotropical Melastomataceae. Markgraf's 
circumscription of Tessmannianthus emphasized 
its unique androecial characters. The connectives 
on the larger set of dimorphic stamens are modified 
dorsally into biauriculate, ventrally upturned ap- 
pendages. The smaller anther sacs are longitudi- 
nally bifurcate distally and have connectives that 
are prolonged dorsally at the base into bilobed or 
deflexed, bifid appendages. These characters, to- 


gether with the pentamerous flowers, trilocular cap- 
sular fruits, and prevailingly lepidote indument, 
define Tessmannianthus as a natural alliance wor- 
thy of generic rank. 

For nearly fifty years Tessmannianthus was 
thought to be a monotypic genus. Wurdack (1975) 
described T. cenepensis and transferred the enig- 
matic Miconia calcarata Gleason to Tessman- 
nianthus, enlarging the genus to three species and 
extending its range from Peru northward to Co- 
lombia. 

Because of its arborescent habit, rainy season 
flowering, and evidently limited number of individ- 
uals in any one population, Tessmannianthus has 


' I thank John J. Wurdack for sharing information about Tessmannianthus, Gordon McPherson for returning to 
qplecing localities to gather flowering material, and curators of the following herbaria who sent specimens as gifts 
o 8 


O, 


* Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118- 


996], U.S.A. 


ANN. Missouni Bor. Garb. 76: 1-6. 1989. 


Annals of the 
Missouri Botanical Garden 


escaped the attention of most collectors. Each of 
the species is known from fewer than six collections, 
and open flowers of T. cenepensis and mature seeds 
of T. heterostemon remain unknown. 

The discriminating collecting efforts of Gordon 
McPherson in Panama have yielded the first col- 
lections of the two new species described here. 

mong the six merianioid genera of Melastoma- 
taceae now known from Panama (Gleason, 1958; 
D'Arcy, 1987), Tessmannianthus is likely to be 
confused only with Graffenrieda. The latter is dis- 
tinguished by a calyptriform or irregularly ruptur- 
ing calyx, dorsally arcuate stamens, and anther 
connectives modified into acute dorso-basal spurs. 
The Panamanian species of Tessmannianthus do 
not appear to be closely related to one another. 
They are, nevertheless, readily separated from all 
other Central American melastomes by the follow- 
ing characters: the small (4-7.5 x 2-6 mm) clawed 
petals are retrorsely erose-ciliolate; the larger ven- 
trally arcuate anthers open by a pair of ventrally 
inclined confluent pores; and each of the bifurcate 
anther sacs in the smaller stamens opens by a 
dorsally inclined pore. 


KEY TO TESSMANNIANTHUS IN PANAMA 
la. Upper leaf surfaces sulcate, the elevated nerves 
on lower leaf surfaces moderately covered with 
caducous mixture of gare deine (pinoid) 
sis) 7- 


n 
apical tooth; petals distally puberulent abaxially; 
larger anthers 9.5- 10 mm long .. 1. T. carinatus 

lb. Upper leaf surfaces essentially fla 
nerves on giki surfaces moderat 
sparingly cov 
torus; calyx lobes beset with an inconspicuous 
blunt rdc tooth; petals glabrous abaxially; 
larger anthers 4.5 mm long 2. T. gordonii 


Tessmannianthus carinatus Almeda, sp. nov. 
TYPE: Panama. Panamá: Cerro id b aem 
ca. 9°15'N, 79*30'W, forest near s t, 
850 m, 24 Aug. 1986 (fl), Mc Planos 9080 
Ponpe CAS; isotypes, MO, PMA, not seen). 
"Rue 


n IM A lotid ` +fal; 


novella foliorum subtus venae dd semine 
dense pinoideis ca. mm longis e 
squamis lepidotis modice intermixtis. Petioli 0.5-1.4 cm 
longi; lamina 4.3-8.5 x 1.7-4.4 cm elliptica vel elliptico- 
ovata apice breviter (2-3 mm) gradatimque acuminato 


> 


basi obtusa, -nervata (pari intramarginal tenui neglecto 
nervis secundar y inter se distantibus sub- 
tus elevatis, coriacea eti ntegra. "Pan cula 3-5.5 cm longa 


multiflora; flores 5(-6)-meri, pedicellis (ad anthesim) 3- 


6 mm longis. Hypanthium (ad torum) 7-8 mm longum; 
calycis tubus 1.5 mm longus, lobis 2.5 x 3 mm late 

ovatis, dentibus exterioribus callosis acutis ca. 1 mm emi- 
nentibus. Petala -6 mm obovato-suborbic- 

ularia ca. 2-2.5 mm unguiculata extus apicaliter modice 
puberula. Stamina dimorphica glabra; antherae ad basim 
in staminibus maioribus et staminibus minoribus biauric- 
ulatae. Ovarium triloculare et ca. % inferum. Fructus 
maturus ignotus 


Trees to 11 m tall. Older cauline internodes 
terete and glabrous; the subquadrangular distal 
branchlets, inflorescences 
moderately to densely covered with a mixture of 
dark brown shaggy-dendritic and irregularly-shaped 
lepidote hairs. Leaves of a pair essentially equal in 
size; petioles 0.5-1.4 cm long; blades coriaceous, 

-8.5 cm long, 1.7-4.4 cm wide, elliptic to 
elliptic-ovate, apex gradually acuminate with a short 
acumen (2-3 mm), this obtuse to broadly rounded 
and curved downward, base obtuse, margin entire 


vegetative buds, and 


and revolute distally, 5-nerved abaxially, the out- 
ermost pair of primaries often inconspicuous and 
concealed by the revolute margins when dry, the 
transverse secondaries elevated like the primaries 
and spaced 2.5-4 mm apart at the widest portion 
of the blade, glabrous and sulcate above at ma- 
turity, covered with a mixture of dendritic and 
lepidote hairs on the elevated primaries below, oth- 
erwise moderately beset only with lepidote hairs. 
Inflorescence a terminal multiflowered panicle 3- 
5.5 cm long; bracteoles paired, early caducous, 
linear-oblong to narrowly obovate, 1.5-5 mm long, 

.5-2 mm wide, margin entire, glabrate above, 
moderately lepidote below. Pedicels 3-6 mm long, 
densely lepidote. Hypanthia (at anthesis) cylindric, 
7-8 mm long to the torus, moderately to copiously 
lepidote. Calyx tube 1.5 mm long on flowering 
hypanthia; calyx lobes reportedly white, ovate but 
appearing deltoid abaxially because of the carinate 
clawlike tooth (ca. 1 mm high) at the incurved 
summit, 2.5 mm long (excluding tooth) and 3 mm 
wide basally between sinuses, the margins entire. 
Petals 5(-6), reportedly pale pink, 7-7.5 mm long, 
5.5-6 mm wide, obovate-suborbicular, rounded 
apically, abruptly tapering to a basal claw 2-2.5 
mm long, glabrous adaxially but puberulent distally 

on the abaxial surface, the margin retrorsely erose- 
ciliolate. Stamens 10(- 12), strongly dimorphic with 
larger stamens inserted on the torus opposite the 
calyx lobes and smaller ones inserted opposite the 
petals; filaments somewhat declinate, complanate 
and glabrous; anthers yellow and 2-celled. Larger 
stamens: filaments 6.5-7 mm long; anthers genic- 
ulate at the filament insertion, 9.5-10 mm long 
and 0.5 mm wide, subulate, ventrally arcuate and 
channeled between the thecae, opening by 2 ven- 


Volume 76, Number 1 
1989 


Almeda 3 


Tessmannianthus 


Tessmannianthus carinatus. — A. Habit. 


FIGURE 1. 
surface) si lar lepidote hairs 
abaxial surface (left), adaxial Arles (right). — 


trally inclined confluent pores; connective thick- 
ened dorsally and prolonged at the minutely pu- 
berulent base into a ventrally upturned biauriculate 
appendage, with each lobe 1-1.5 mm long and 0.5 
mm wide. Smaller stamens; filaments 5.5-6 mm 
long; anthers 6 mm long and 0.5 mm wide basally, 


— B. Representative le 
—D. to (at anthesis); petals, stamens, and style removed. — E. P. 
arger stamens Ba teral 


(left), dade view (right). Drawn by Ellen del o from the holot 


af. —C. Base of leaf blade a 
etals, 


view).—G. Smaller stamens, lateral view 


linear-oblong, erect to somewhat incurved distally 
with the 2 thecae of each anther diverging from 
one another, each theca opening by a dorsally 
inclined pore; connective thickened dorsally and 
prolonged at the base just above the filament in- 


sertion into a deflexed, deeply bifid appendage 0.5 


Annals of th 
Missouri as Garden 


mm long having each lobe caudiform in dorsal and 
ventral view. Ovary (at anthesis) ca. 75 inferior, 
3-celled, oblong, glabrous at the apex. Style 

8 mm long, 0.5-1 mm wide, glabrous, somewhat 
declinate and slightly incurved distally; stigma round 
to subtruncate. Mature capsule and seeds not seen. 


Distribution. Known only from the summit 
area of Cerro Jefe in central Panama at about 850 
m, where the new species is evidently an uncom- 
mon member of the low cloud forest vegetation. It 
grows with but appears to be greatly outnumbered 
by the vegetatively similar Cerro Jefe race of Cono- 
stegia montana (Sw.) D. Don ex DC. 

Tessmannianthus carinatus differs from other 
members of the genus in characters of the foliage, 
indument, and calyx lobes. The leaves of this species 
are leathery and sulcate, with a rounded terminal 
acumen that is turned downward. The indument 
on young branches, inflorescences, and the ele- 
vated primary nerves on lower leaf surfaces con- 
sists of a mixture of appressed lepidote hairs and 
spreading dendritic or pinoid hairs. The former hair 
type is found in all other members of the genus; 
the latter is unique to 7. carinatus. The specific 
epithet draws attention to the distinctive calyx mor- 
phology. Each lobe is beset with a carinate abaxial 
keel that forms an apically incurved clawlike tooth 
(Fig. 1D). 

Tessmannianthus carinatus appears to be with- 
out close relatives. Its leathery leaves are vaguely 
reminiscent o . heterostemon, which differs 
markedly in having larger (9-20.5 x 6.4-13.5 
cm) apically rounded obovate blades, shorter hy- 
panthia (3-4 mm) with minute caducous exterior 
calyx teeth, and elliptic-oblong, papillate petals. 


Tessmannianthus gordonii Almeda, sp. nov. 

: Panama. Chiriquí: Fortuna Dam wa- 
above Rio Hornito, 08?45'N, 
82°15'W, 1,250 m, 1 July 1987 (fl), Mc- 
Pherson 11161 (holotype, CAS; isotypes, 
AAU, BM, COL, CR, DUKE, EAP, F, K, LE, 
MEXU, MO, NY, P, PMA, QCA, TEX, U, 
US, VEN, not seen). Figure 2 


Ramuli primum quadrangulati demum teretes sicut pe- 
tioli foliorum subtus venae primariae odiar na 
squamis lepidotis demum caducis ergo indut 
tioli 0.6-1.3 cm longi; lamina 4.4-6.8 21-28 cm 
elliptica apice acuminato, basi acuta E a, 3-5- 
nervata vel 3-5-plinervata (pari tenui Sie su ipis 
firme membranacea et integra. Panicula 2.5-3.5 cm lon- 
ga multiflora; flores 5-meri, pedicellis (ad anthesim) 2- 
3.5 mm longis. s (ad torum) 3(-3. E mm lon- 
gum; calycis tubus m longus, lobis 1 x 2 mm 

ovatis, dentibus serons crassis 0.5 mm Bree 


Petala glabra 4-4.5 x 2-2.5 mm elliptico-oblonga ca. 
0.5-1 mm unguiculata. Stamina dimorphica glabra; an- 
therae ad basim in staminibus maioribus et staminibus 
minoribus distincte biappendiculatae. Ovarium triloculare; 
capsula ca. 5-6 mm longa; semina numerosa 2-2.5 mm 
longa anguste pyramidata. 


Trees 9-16 m tall. Older cauline internodes 
terete and glabrous; rangular distal 
branchlets, lower leaf surfaces, and inflorescences 
beset with a moderate covering of caducous lepi- 
dote hairs. Leaves of a pair essentially equal in 


the quad 


size; petioles 0.6-1.3 cm long; blades firmly mem- 
branaceous, 4.4-6.8 cm long, 2.1-2.9 cm wide, 
elliptic, apex acuminate, base acute to obtuse, mar- 
gin entire, 3-5-nerved or 3-5-plinerved abaxially 
with the innermost pair of primary nerves elevated 
and diverging from the median nerve in opposite 
fashion about 2-4 mm above the blade base, the 
outermost pair of primaries mostly depressed and 
inconspicuous, essentially glabrous above at ma- 
turity, sparingly to moderately beset with brown 
lepidote hairs below. Inflorescence a terminal mul- 
tiflowered panicle 2.5-3.5 cm long with ultimate 
branchlets terminating in umbelliform clusters; 
bracteoles paired, sessile, persistent to tardily de- 
ciduous on the infructescence, ovate to oblong- 
ovate, rounded apically, 0.5-3 mm long, 0.5-2 
mm wide, margin entire, glabrous on the concave 
upper surface, moderately to copiously lepidote to 
glabrate on the lower surface. Pedicels 2-3.5 mm 
long, sparingly lepidote to glabrate. Hypanthia (at 
anthesis) subcylindric to narrowly campanulate, 3 
(-3.5) mm long to the torus, glabrous or sparingly 
lepidote toward the base but typically glabrous at 
maturity. Calyx tube ca. 0.5 mm long on fruiting 
hypanthia; calyx lobes broadly ovate but appearing 
bluntly deltoid because of the terminal tooth (to 
0.5 mm long) on the glabrous abaxial surface, 1 
mm long, 2 mm wide basally between sinuses, the 
margins entire and somewhat incurved apically. 
Petals 5, glabrous, reportedly pink-white, elliptic- 
oblong to narrowly obovate, rounded to shallowly 
and irregularly lobed apically, gradually tapering 
to a short (0.5-1 mm) basal claw, 4-4.5 mm long, 
2-2.5 mm wide, retrorsely erose-ciliolate. Stamens 
10, strongly dimorphic with larger stamens inserted 
on the torus opposite the calyx lobes and smaller 
ones inserted opposite the petals; filaments some- 
what declinate, complanate and glabrous; anthers 
yellow, 2-celled. Larger stamens: filaments 5 mm 
long; anthers geniculate at the filament insertion, 
4.5 mm long, 0.5 mm wide, linear-oblong, arcuate 
and channeled ventrally between the thecae, open- 

ing by 2 ventrally inclined confluent pores; con- 
nective thickened dorsally and prolonged at the 


Volume 76, Number 1 Almeda 5 
1989 Tessmannianthus 


Tessmannianthus gordonii. — A. Habit. —B. i aa leaf (abaxial surface). — C. Base of leaf 
blade asd surface) with enlarged lepidote hairs. Fi Hypanthium (in fruit). — E. irs section of capsule and 
enveloping hypanthium. — F. Petal (abaxial surface). —G. Larger stamens (lateral view). — H. Smaller stamens, im 


view (left), dorsal view (right). —I. Seeds. Drawn by Ellen del Valle. (A-C, F-H from the holotype; D, E, 
McPherson 9877.) 


minutely puberulent base into a ventrally upturned basally, linear-oblong, erect to somewhat incurved 
deeply bifid appendage, with each lobe 1 mm long distally where the 2 thecae diverge from one 
and 0.5 mm wide. Smaller stamens: filaments 4— another, each theca opening by a dorsally inclined 
5 mm long; anthers 2.5-3 mm long, 0.5 mm wide pore; connective thickened dorsally and prolonged 


Annals of the 
Missouri Botanical Garden 


basally into a deflexed biauriculate skirtlike ap- 
pendage (0.5-0.75 mm long), with each of its lobes 
tail-like in ventral and dorsal view and obdeltoid 
in lateral view. Ovary (at anthesis) ca. 12 inferior 
3-celled, narrowly elliptic, glabrous apically. Style 
5-6 mm long, 0.5 mm wide, glabrous, declinate 
and somewhat incurved distally; stigma rounded to 
subtruncate. Fruit a many-seeded loculicidal cap- 
sule ca. 5-6 mm long and 4.5-5 mm diam. Seeds 
narrowly pyramidate, cuneate and angulate, 2-2.5 
mm long, white or beige, glabrous but vaguely 
papillate on the angles. 


» 


Distribution. Known only from montane for- 
ests of the Fortuna Dam watershed above Rio Hor- 
nito at 1,100-1,250 m in Chiriqui Province, Pan- 
ama. 

Additional specimen examined. PANAMA. CHIRIQUÍ: 
vicinity of Fortuna Dam along trail near Rio Hornito, ca. 
8?45'N, 82?15'W. Forest ca. 1,100 m, 8 Aug. 1986 (fr), 
McPherson 9877 (CAS). 


In the size, shape, and indument of its leaves, 
T. gordonii most closely resembles 7. cenepensis. 
The latter differs in the following characters: anther 
sacs of the larger stamens are linear-oblong and 
conspicuously retuse or bifurcate distally for about 
0.5 mm; anther sacs of the smaller stamens are 
bifurcate for the distal two-thirds of their length 
and have connectives that are obscurely lobed but 
not prolonged into biauriculate appendages; the 


ovary is puberulent; the petals are copiously fur- 
furaceous-lepidote abaxially; and the hypanthia are 
markedly constricted below the torus into a tubular 
neck that shortens to a tight constriction on fruiting 
hypanthia. Tessmannianthus cenepensis also ap- 
pears to differ in other modal tendencies. Its inflo- 
rescences are longer (5-8 cm) and more floriferous 
than those of T. gordonii, and its petals are broadly 
ovate and appear to lack a well-developed basal 
claw. The consistency of these latter differences 
will become apparent only when better flowering 
material is available for study. 

This species is named for Gordon McPherson 
(1947- ),collector of the type and all other known 
Panamanian specimens of this genus. His many 
fine collections of trees, shrubs, and epiphytes have 
added a number of new and interesting species to 
the flora of Panama. 


LITERATURE CITED 


D’Arcy, W. G. 1987. Flora of Panama: Checklist and 
: t. Bot. Missouri Bot. Gard., Volume 17 
GLEASON, H. A. 1958. Melastomataceae. In: Flora of 

Panama. Ann. Missouri Bot. Gard. 45: 203-304. 
MankcRaF, F. . Melastomataceae. /n: J. Milbraed 
(editor), Plantae Tessmannianae peruvianae IV. No- 
tizbl. Bot. Gart. Berlin-Dahlem 9: 1139-1154. 
WuRDACK, J.J. 1975. Certamen Melastomataceis XXIV. 
Phytologia 31: 492-500. 


THE GENUS ARGOSTEMMA 
(RUBIACEAE— 
ARGOSTEMMATEAE) 

IN BORNEO! 


Birgitta Bremer? 


ABSTRACT 


Argostemma is a large paleotropical genus of the tribe Argostemmateae. From Borneo 28 species are recognized. 
All are herbs growin, gi in rainforests. i species, A. apiculatum, A. burttii, A. geesinkii, A. gaharuense, A. brookei, 
an 


d A. calcicolum, are new ere is one new 
other genus is Neuroc wis The sister 4 d to 


name, 


ait. Argostemma is the larger genus of the tribe; the 


c 
the Argostemmateae is the mainly Central American tribe Hamelieae. 


em 
sister group outside Borneo. Nomenclature, descriptions, illustrations, maps, and a key for all species are provided. 


Argostemma Wall. in Roxb. is a large, complex 
genus with about 220 described species. The re- 
vision of the genus will be published in parts dealing 
with different geographical areas. About 51 taxa 
have been described from Borneo. There is no 
treatment covering the whole island. In this study 
I have recognized 28 species, of which six are new. 
Most of the 28 species are endemic to Borneo, 

Ithough six of them were described from speci- 
mens collected outside of Borneo. The remaining 
22 are typified on Borneo specimens. 

Only synonyms typified by specimens from Bor- 
neo are listed under the species 

During the 1950s, the late Dr. Bakhuizen van 
den Brink, Jr. studied the genus for the Flora 
Malesiana project. He annotated many herbarium 
labels about typifications. I have studied his lec- 
totypifications, and in most cases I do agree with 
him, and have accepted the lectotypifications which 
are published in this paper. 

No infraspecific taxa are recognized in this study. 
Most species are stenomorphous, and a few species 
are polymorphous. Among the latter, Æ. elato- 
stemma occurs as two differing populations on Bor- 
neo, but since its whole variation outside Borneo 


is not yet known, they are not recognized as sep- 
arate taxa. The substantial variation in Æ. borra- 
gineum is not correlated with geographical distri- 
bution and does not reveal distinct evolutionary 
lines. Argostemma hameliifolium and A. moul. 
tonii form a complex without discrete infraspecific 
units. The distinction between the two species is 
more or less preliminary. Argostemma densifolium 
is dimorphic, but as the variation seems small, I 
have treated the two morphs as one taxon. 


MATERIALS AND METHODS 


All cited specimens including types have been 
studied, unless otherwise stated. I have seen ma- 
terial from the following herbaria: A, AAU, B, BM, 

: >E, G; , HBG, K, KLU, L, MO, NY, 
P, S, SAR, SING, U, UC, and US. 

I have studied Argostemma parvifolium, A. 
A. rupestre, A. ha- 
meliifolium, A. moultonii, A. gracile, A. havi- 
landii, A. calcicolum, A. psychotrioides, and A. 
bryophilum in the field. Material of most of these 
species was fixed in the field and later embedded 
and sectioned for anatomical studies of leaves, an- 


elatostemma, A. ophirense, 


' Į am grateful to Mr. Paul Chai and his staff at the Forest Department at Kuching for assistance during my 


fieldwork, to the late Dr. Bakhuizen van den Brink, Jr. 


for 
for checking my Latin pes as well as to Mrs. Gunilla Hägglun i Miss K 


for technical assistance. I a 


Methods and to the reviewers 2 many useful comments. I than 


much information about the "pe ohn Dwyer 


s Miss Mari Källersjö 


e Pee Raven, Director o 


Garden, where I finished this study. Financial support from the Swedish Institute, the Royal emy of Sciences, 
ck 


Stockholm, the Swedish Natural Science Research Council, and the University of Sto 


holm is aci acknowledged. 


* Department of Botany, University of Stockholm, S-106 91 Stockholm, Sweden. 


ANN. MISSOURI Bor. GARD. 76: 7-49. 1989. 


Annals of the 
Missouri Botanical Garden 


thers, and ovaries. For endothecial structure, whole 
anthers of all species were mounted in Hoyer's 
solution for study. 

The species are arranged roughly in phyloge- 
netic order. Species 1-12 (the 4. parvifolium 
group) belong to one evolutionary group, species 
13-26 (the A. psychotrioides group) to a second 
group, species 28 (4. neurocalyx) to a third group, 
and species 27 (4. bryophilum) assumes an un- 
certain position within the genus. 

The drawings are my own. In the figures (4— 
31), all details of plants indicated by the same 
letter (B-T) are drawn at the same magnification 
and illustrate the same plant part. Hence, if a 
particular part is not illustrated, the corresponding 
letter is not used in the figure. The branch, indi- 
cated by letter A, is drawn twice as large in Figures 
7-15, 17, 19-22, 30, and 31 as in Figures 4-6, 
16, 18, and 23-29. 


SYSTEMATIC POSITION AND 
INTERRELATIONSHIPS WITHIN 
ARGOSTEMMA 


Argostemma is the largest genus of the Argo- 
stemmateae (Verdcourt, 1958; B. Bremer, 1987; 
Argostemmatideae fide Darwin, 1975, sphalm., 
pers. comm.). The other genus of the tribe is Neu- 
rocalyx (B. Bremer, 1979), a small genus endemic 
to Sri Lanka and southern India. The sister group 
to the Argostemmateae is the mainly Central Amer- 
ican tribe Hamelieae (B. Bremer, 1987). These 
two tribes form a monophyletic group within the 
subfamily Rubioideae. 

In Borneo there are 28 species of Argostemma. 
About two-thirds of these are endemic. The Bor- 
nean species belong to at least th t mono- 
AN groups, each having its sister group outside 
Borne 

The A parvifolium group is represented by 
species 1-12, characterized by “radial” endothe- 


cium (see morphological notes), 
leaves, rotate corollas, a narrowly ovoidal anther 
cone, thin and smooth apical appendages, and a 
glabrous style with a hardly widened to slightly 
capitate stigma. Three species from this group (4. 
parvifolium, A. elatostemma, and A. ophirense) 
also occur in the Malay Peninsula and on other 
Sunda Islands. 

e A. psychotrioides group, species 13-26, 
is ad by a sturdy connective, umbelli- 
form inflorescences, in most species a corolla cleft 
to less than half its length, and usually a style much 


anisophyllous 


longer than the stamens. Most species also have a 
"polarized" endothecium (one species cannot be 


classified as "radial" or "polarized"), and most 
species have an endothecium with distinct cell mar- 
gins (see morphological notes). Argostemma ha- 
meliifolium, A. moultonii, A. psychotrioides, and 
A. borragineum also occur on other Sunda Islands 
but not in the Malay Peninsula. 

The remaining species, 4. bryophilum (27) and 
A. neurocalyx (28), are more isolated. Argostem- 
ma neurocalyx belongs to a totally different group 
within Argostemma. It is characterized by an apical 
leaf rosette and free stamens, of which the anthers 
open by pores instead of slits. The group to which 
it belongs has a very wide distribution from West 
Africa to the Philippines. The systematic position 
of A. bryophilum is not yet clear. It does not fit 
in any of the groups mentioned above and is part 
of a complex of taxa common in the Philippines 
and New Guinea. 


MORPHOLOGICAL NOTES 


Raphides. Raphides are found in most parts 
of the plants and can be seen easily in corollas or 
in young glabrous stems. 


Colleters. On living material the colleters can 
be seen easily, but on old dry specimens they are 
hardly visible. They are of the standard rubiaceous 
type (Lersten, 1975), with elongated axial cells 
surrounded by a palisade epidermal layer. The 
colleters are found singly or a few together between 
the calyx lobes. At each node they surround the 
stem in a ring and occur on the adaxial side of the 
petiole and at stipule bases. In species with few 
colleters in each ring, the colleters are generally 
round in outline, whereas in species with a dense 
ring of several hundred colleters they are narrow, 
often reduced, and almost without a palisade epi- 
dermis (Lersten, 1974). The stipules often end in 
apical colleters. In Argostemma chaii, with fringed 
stipules, each of the processes making up the fringe 
ends in a distinct colleter. 


Hairs. In Borneo, only A. apiculatum is gla- 
brous. The other species are more or less entirely 
or partially pubescent. There are no substantial 
differences between hairs on vegetative organs and 
those on flowers. Normal hairs are monoseriate and 
completely septate, with thin to very thick walls, 
and they generally do not collapse when dry. They 
may occur all over the leaf surfaces or only on (or 
rarely only between) the veins. If there is indu- 
mentum on the inside of the corolla or on the style, 
the hairs are shorter, mostly one- or few-celled and 
thinner (Verdcourt, 1958). Another hair type oc- 
curs on, e.g., 4. rupestre (Fig. 131) and A. gee- 


Volume 76, Number 1 
1989 


Bremer 9 
Argostemma in Borneo 


sinkii (Fig. 141). On the upper leaf surfaces of 
these species there are three or five rows of stiff, 
pluricellular hairs. One row occurs along the midrib 
and the others along the margins, making them 
appear more or less serrulate. 


Leaf anatomy. | have studied in detail cross 
sections of field-fixed material of 4. bryophilum, 
A. elatostemma, A. havilandii, A. moultonii, A. 
ophirense, A. psychotrioides, and A. rupestre. 
Cross sections of young leaves of the species are 
generally very similar. With the exception of A. 
bryophilum, cross sections of these species can be 
described as follows: upper epidermis of one layer 
of large, rectangular cells with rather thin walls 
and lacking plastids; palisade chlorenchyma of one 
or a few layers of small carrot-shaped cells filled 
with plastids; spongy parenchyma of a few to many 
layers of small to medium-sized cells almost without 
plastids; lower epidermis of one layer of large, 
rounded cells with thin walls and without plastids; 
stomata quite exposed. This type of leaf also occurs 
in Neurocalyx (B. Bremer, unpubl.). Argostemma 
bryophilum, aberrant in many other ways, differs 
from the others in leaf cross section. Its epidermal 
layers are thinner with smaller cells, and the chlor- 
enchyma consists of large rounded cells with few 
plastids. 

Species whose lower leaf surface appears white 
(e.g., A. rupestre and A. gracile) have more in- 
tercellular spaces in the spongy parenchyma than 
other species. 


Endothecium. The endothecium of Argo- 
stemma (Fig. 1) was studied with light microscope 
on whole mounts of anthers. The endothecium var- 
ies between groups of Argostemma species. In A. 
bryophilum and A. neurocalyx, in many ways 
aberrant species, only narrow longitudinal bands 
of cells close to the stomium have wall thickenings, 
while in all other species from Borneo, wall thick- 
enings occur in all cells of the endothecial layer. 
In the following, the terms endothecium, endothe- 
cial layer, and endothecial type will only be used 
for those cells with wall thickenings. 

The endothecial cells are generally elongated 
longitudinally or horizontally compared with the 
line of dehiscence and with the anther-length axis, 
or rarely they are isodiametric. In Argostemma 
the wall thickenings are U-shaped ribs arranged 
transversely in relation to the longest axis of the 
cell and with their openings toward the epidermal 
layer. The ribs end in knobs. 

Dormer (1962), Nordenstam (1978), Noel 
(1983), and French (1985) have shown the sys- 


tematic significance of endothecial structures. This 


study shows that different endothecium types occur 
in Argostemma and that these types are restricted 
to different systematic units within the genus. Dor- 
mer (1962) and Noel (1983) presented systems 
for classification of endothecial structures. Dormer 
(1962) classified endothecial cells in the Compos- 
itae as “polarized” or “radial.” In a polarized en- 
dothecium the thickenings are concentrated mainly 
at the upper and lower ends of the cells, and when 
ribs occur they are parallel to the line of dehiscence. 
In a radial endothecium the cells are ribbed all 
around; on elongated cells the bars are arranged 
transversely, parallel to the line of dehiscence. I 
have found Dormer's system somewhat difficult to 
use with Rubiaceae because the endothecial cells 
are different, often elongated horizontally, i.e., 
transversely to the anther-length axis. In Compos- 
itae (Nordenstam, 1978; Dormer, 1962; K. Bre- 
mer, pers. comm.), when the cells are elongated, 
the elongation is parallel to the anther-length axis. 

Noel (1983) presented a more general system 
which embraces over 500 taxa from different fam- 
ilies. He distinguished several endothecial char- 
acters, such as size and shape of the cells, orga- 
nization of the thickenings, whether a basal plate 
and/or ribs occur, and the shape and number of 
these. In general, the endothecial cells of Argo- 
stemma can be characterized following Noel’s 
scheme (1983: 837): cells medium-sized, 27-60 
um on the longest side, isodiametric, rectangular 
or fusiform; wall thickenings localized; base plate 
absent; ribs U-shaped; 5-ca. 15 ribs per cell (in 
several species the cell margins become indistinct, 
and it is difficult to see where the cell ends); spacing 
from close to wide; branching absent; tips knobbed. 


gostemma. The first two are e 
first type, in A. bryophilum, A. hameliifolium 
Fig. 1A), 4. moultonii, and A. neurocalyx, the 
cell margins become indistinct, and the endothecial 
layer can be described as consisting of long, hor- 
izontal, rather narrow bands with many close dis- 
tinct transverse ribs and knobs. This type also 
occurs in the sister genus Neurocalyx. 1 have ex- 
amined the endothecia of Hamelia patens, Hoff- 


— 


mannia vesciculifera, and Xerococcus congestus, 
belonging to Hameliieae, the sister tribe to Argo- 
stemmateae (B. Bremer, 1987), and they are of 
the same type. 

In the second type (e.g., A. enerve, A. gracile, 
A. havilandii, and A. psychotrioides (Fig. 1B)), 
the cells are horizontally elongated and fusiform 
but with distinct cell margins. The ribs are not as 
close together, and the knobs are larger than in 
the first type. 


Annals of the 
Missouri Botanical Garden 


Oo 


: AU 


Eps JRE l. Endothecial cell thickenings. Scale a = 
rgostemma hameliifolium. —B. A. 


a —C. A. parvifolium. —D. A. alae ic 


The third type could be classified as "radial." 
This type occurs in many species, e.g., Á. ophi- 
rense, A. parvifolium (Fig. 1C), and A. rupestre. 
The cells are generally rectangular, with the lon- 
gitudinal axis parallel to the dehiscence line. The 
cells are larger than in the ““polarized” group. The 
transverse ribs are indistinct to hardly visible, and 
the knobs are smaller and look like a string of 
pearls surrounding the cell margins. 

fourth group consists solely of 4. borra- 
gineum (Fig. 1 D), whose endothecium is unique at 
least among the Borneo species of Argostemma. 
The cells are generally isodiametric and large. The 
ribs are indistinct, generally arranged parallel to 
the dehiscence line, and the knobs are indistinct. 
Since the ribs are not transversely arranged in 
relation to the anther-length axis, this type cannot 
be described as **radial," and since the knobs sur- 
round the cells on ie margins, it cannot be de- 


scribed as “polarize 


Fruit. 
cles become erect. These parts and the fruit, 


After anthesis the pedicels and pedun- 


crowned with the calyx lobes, become fleshy and 
succulent. The fruit opens by an apical operculum. 
Such fleshy capsules are not uncommon in the 
tropics (Ehrendorfer, 1983: 829). Bakhuizen f. 
(1975) said about fruits of Rubiaceae: 


I am convinced that true capsules are not found in 
distinctly 


t eshy 
known as * ‘drupe’ ”) and vise (having a 
(very) thin, dry, ks leathery pe ricarp, so- iait 
capsule’ -pyrenife rous are 
conventionally lema “berry.” 


46 


The fruit of Argostemma does not fit into this 


system. Although Bakhuizen f. was an expert on 
this genus, he had never seen it in the field (pers. 
comm.), and on dry material he had probably mis- 
interpreted the fruit type. 

When the fruit is open the fibrous endocarp 
separates from the remaining pericarp and splits 
into threads, which are visible in the opening. The 
septum, with the attached placentas, is also fibrous 
and will split. Among the resulting threads the 

umerous seeds will be exposed. Perhaps the threads 
Mii sudden dispersal. It is not uncommon to 
find specimens with many seeds germinated within 
the capsules. This kind of “vivipary” or germi- 
nation in situ is described earlier from Ophiorrhiza 
tomentosa (Tan & Rao, 1981). 


Placentation.  Placentation is axile with the 

placentas (Fig. 2) attached at the top of the septum. 

he placentas are stalked, pendulous, fleshy, and 
in front view broadly ovoidal to kidney-shaped. 


Seeds. | Argostemma has numerous, small (ca. 
0.5 mm), angular to ovoidal seeds (Fig. 3). The 
testa seems to be unique to this genus. The surface 
is mostly reticulate with a mamilla or a ring in 
each cell. 


DISTRIBUTION 


The maps (1-28) of Argostemma show that 
most localities are from northwestern Borneo. This 
is not because Argostemma is more common there, 
but indicates that this part, Sarawak, is botanically 
most well known. The same pattern can be seen 
in Memecylon (K. Bremer, 1983) and other gen- 
era. The few localities in Kalimantan result from 
comparatively few botanical expeditions, and large 
areas of former rainforest have been deforested. 

Taxonomically, Argostemma in Borneo belongs 
to at least three different species groups which are 
more or less sympatric, and many species within 
the same group are sympatric. The distributions 

of different species of Argostemma are influenced 
by altitude, a fact that is hardly evident on the 
maps. Argostemma elatostemma, A. densifolium, 
A. havilandii, A. cal- 
cicolum, A. trichosanthes, A. psychotrioides, and 
A. variegatum grow only at low altitude; and 4. 


A. gaharuense, A. chaii, 


anisophyllum, A. dulitense, A. apiculatum, A. 


burttii, A. geesinkii, A. gracile, A. brachyanthe- 
rum, and A. brookei grow only at altitudes over 
1,000 m. Some species occur at both low and high 
altitudes: 4. parvifolium, A. subfalcifolium, A. 
ophirense, A. A. hameliifolium, A. 
moultonii, and A. borragineum. 

Most of the lowland species and those growing 


rupestre, 


Volume 76, Number 1 


Bremer 11 
Argostemma in Borneo 


Is fv 


FIGURE 2. Placenta in ripe fruit (diagrammatic). ls, 
longitudinal section; fv, front view; cs, cross section. 


from low to high altitude are widespread, and many 
of them are sympatric. Of the eight species known 
only from a single locality each, four are from high 
altitude, the altitudes of three are unknown, and 


only one is from low altitude (ca. 70 


TAXONOMIC TREATMENT 


Argostemma Wall. in Roxb., Flora Indica 2 
324, 1824. Pomangium Reinwardt, Syll. Pl. 
Rat. 2: 10. 1825. Argostemmella Ridley, J. 
Bot. 65: 41. 1927. TYPE: Argostemma sar- 
mentosum Wall. (Pfeiffer, Nomenclator Bo- 
tanicus, 187% 


Erect, suberect, or creeping herbs, isophyllous 
or anisophyllous, glabrous to densely pubescent, 
often succulent. Stem usually unbranched or slight- 
ly branched, often succulent; internodes very short 
to 85 mm long. Leaves opposite or verticillate, 
when anisophyllous unequal, scattered along the 
stems, or decussate, or distichous, or rosulate on 
a short stem, or apically rosulate on a well-devel- 
oped stem, herbaceous, or coriaceous, or mem- 
branaceous; stipules interpetiolar, persistent or de- 
ciduous, small to large, usually entire, rarely slightly 
cleft or fringed, with colleters near the e on the 
adaxial side or rarely at the apex, if fringed then 
with a colleter at each point; midrib distinct, the 
primary veins distinct or obscure, the veinlets gen- 
erally not visible. Anisophyllous leaves with the 
small (nanophyllous) leaf persistent or deciduous, 
similar in shape to the stipules or to the larger leaf, 
often cordiform, with or without a short petiole. 
Inflorescences terminal, subterminal or rarely ax- 
illary, solitary or a few together, 1-160-flowered, 
cymose, corymbiform to umbelliform, glabrous or 
pubescent; peduncles up to cm long or rarely 
absent; bracts often persistent and small; pedicels 


FIGURE 3. Seed 
view. From Bremer 1668 (S). Scale bar = 100 


of A. borragineum in SEM surface 
m. 


in fruiting stage succulent. Flowers (4—)5(—6)-mer- 
ous, actinomorphic or rarely zygomorphic, perfect. 
Calyx with a very short tube; lobes shorter or rarely 
as long as the corolla lobes, erect or spreading to 
reflexed, in fruiting stage succulent, glabrous or 
pubescent. Corolla with valvate aestivation, white, 
sometimes with green nectar guides at the base, 
rotate, or subcampanulate but with recurved lobes, 
or campanulate, 3-12 mm long, with the tube more 
than 25 as long as the lobes, externally glabrous 
or pubescent, internally glabrous or pubescent with 
the pubescence often only on the lobes; lobes erect- 
spreading to recurved or reflexed. Stamens 4 or 
5(6), equal in number to the corolla lobes, inserted 
near the base of the corolla, introrse, coherent into 
an anther cone or free; if coherent then fused along 
the whole anther or partially so; thecae with two 
sacs (loculi); anthers bright yellow to cream-col- 
ored, erect or slightly upcurved; filaments free, 
short or long, mostly equal, erect or rarely curved, 
smooth, glabrous; sacs opening by vertical slits, 
rarely with pores; apical appendages ca. V$ as long 
as the anther or absent, thin to coriaceous, smooth 
to papillose; connectives distinct to indistinct from 
the thecae, smooth to papillose. Ovary 2-celled; 
style filiform, glabrous to pubescent, shortly to long- 
exserted, with a capitate, clavate, or scarcely wid- 
ened stigma, shallowly to rarely distinctly bifid. 
Fruit a succulent capsule with a fibrous endocarp, 
opening by an apical operculum, smooth or rarely 
with emergences, furrows, or ribs (at least on dry 
specimens), glabrous to densely pubescent. Seeds 

own, numerous, minute, angular or ovoid, the 
testa generally reticulate with a central mamilla or 
ring in each cavity. 


Annals of the 
Missouri Botanical Garden 


KEY TO THE SPECIES OF 4RGOSTEMMA IN BORNEO 


la. Leaves more or less condensed into an apical rosette. Stamens free at least after anthesis 2 
2a. Leaves membranaceous. Corolla campanulate, not deeply cleft. Stamens opening by ie pores ......... 
A. neurocalyx 
2b. Leaves herbaceous to coriaceous. Corolla rotate, cleft to near the base. Stamens opening by longitudinal 
slits aii 
lb. Leaves scattered along the stem, not condensed into an apical rosette. Stamens connate or free cc 3 
3a. Leaves anisophyllous 4 
(See also the insufficiently known 16. 4. flavescens not included in the key 
4a. Corolla cleft to near the base, starlike; lobes spr reading, erect- SURE or rarely reflexed, never 
recurved. Stamens as high as the style with stigma only slightly exserted 
5a. Flowers slightly Bs RU Calyx lobes almost as long as the corolla lobes. Filaments 
unequal, curved . parvifolium 
5b. Flowers actinomorphic. Calyx lobes much shorter than the corolla lobes. Filaments equal, 
straight 


6a. Plant creeping. The larger of the anisophyllous leaf pair basally very unequal-sided. 


Peduncles and the outside of the corolla pubescent. Anther cone black when pressed an 
dried 2 


A. elatostemma 
. Plant erect or creeping. The larger of the anisophyllous leaf pair unequal- or equal-side 
Peduncles and the outside of the corolla generally glabrous. Anther cone not black when 
pressed and dried 
7a. Larger leaf of the anisophyllous imd nen subfalcate; — leaves lan- 
ceolate. Fruits with longitudinal fur 3. A. subfalcifolium 
7b. Larger leaf of the anisophyllous ies not subfalcate; ine pis cil cordiform, 
ovate or lanceolate. Fruits without furrows but rarely with 5 ri 
8a. The larger of the anisophyllous leaf pair pim ind -sided or oblique; 
ig ade leaves cordiform 
9a. Stems and leaf veins pubescent. Leaves herbaceous or coriaceous. Corolla 
up to ca. 6 mm lon 


10a. 


an 
c 


es a rosette or creeping with generally short internodes and 
e to oblanceolate leaves 4. A. densifolium 
10b. Plant erect with obvious internodes and lanceolate to linear leaves 


; en 
9b. Stems and leaves glabrous or glabrate. Leaves herbaceous to mem 


bra 
ceous. Corolla 8-9 mm long o lim 


The larger of the nM leaf pair basally equal-sided; anos ci leaves 
ovate, elliptic, or lance 
11 


olat 
aves without stiff placas hairs 
12a. 7 aves pubescent. Corolla ca. 4 mm long, lobes reflexed. Fruit with 


c 
c 


. gaharuense 
12b. eh glabrous. Corolla more than 9 mm long, lobes not reflexed. 
Fruit without ribs 


tems pubescent. Leaves coriaceous with pale lower surfaces. 
Apical appendage ca. Y3 of the anther length ..... . dulitense 
13b. Stems and whole plant glabrous. Leaves herbaceous to mem- 
dures both sides of the same color. id p de 
. Y4 of the anther length 
11b. aw with stiff pa € in 3 or 
oroll 


apiculatum 
14 


rows 
ong o re; P ovate, spreading. Apical 
"appendage less n Y id de anter bulb oh y . A. burttii 
l4b. Corolla ca. 7 mm long or less; lobes ovate or loricóoldth.- erect- 
spreading. Apical a ide e ca. Y3 of the anther length 
. Co rolla lobes lanceolate, not pink in press. Stipules ovate to 
10. 


lanceolat A. rupestre 
15b. Corolla lobes ovate, often pink in press. t ovate to 
11. 


A. geesinkii 
4b. Corolla generally cleft 4 its length or less; lobes recurved, erect, or spreading. Stamens shorter 


than the much-exserted stigma 
16a. Carolla cleft to less in ¥ its length; lobes erect or spreading 
. Leaves distichous. Flowers solitary, almost sessile. Stigma disc-shaped and m 
bifid . gracile 
17b. Leaves not distichous. Flowers in up to 12-flowered inflorescences, Mud solitary, 
with peduncles. Stigma capitate and indistinctly bifid ..................... l . brachyantherum 
l6b. Corolla cleft to at least 2 its length; lobes recurved 18 
18a. Corolla cleft to near the base. Stamens free at anthesis. Stigma slightly capitate ....... 
brookei 


18b. Corolla cleft to ca. 4% its length. Stamens connate at anthesis. Stigma clavate 


Volume 76, Number 1 
1989 


Bremer 
Argostemma in Borneo 


19a. el lobes 2-6 mm long, distinctly reflexed. Style generally glabrou 


19b. Calyx lobes up to 1 


a. Plant erect. Leaves basally acute to cuneate or attenuate and Aa ed 
psychotrioides 

20b. Plant creeping. Leaves basally rounded to obtuse and unequal- -sided ..... 
26. A. variegatum 
mm long, erect. Style generally pubescent 21 

la. Stems green, smooth. Nanophyllous leaves persistent, cordiform ................ 
24. A. humifusum 


21b. Stems in lower part generally pale brown to gray with elevated corky to 
papery longitudinal ridges. Nanophyllous leaves deciduous, E ce to 


line 


. A. havilandü 
22 


3b. Leaves isophyllou 
22a 


Corolla 2 to near the base, starlike; lobes spreading or reflexed 23 


a. Leaves membr 
flow 


2-flowered. Apical anther D ue almost wantin 


ranaceous. Stipules persistent, broadly ovate to cordiform. Inflorescences l- 


. bryophilum 


8 
23b. qae herbaceous. Stipules deciduous. lodotesceti t 3-40-flowered. Apical nier ap- 


pendages form open cylinders (Figs. 16, 17 
Plant 10-100 cm, erect. Stamens with short filaments and short SP. appendage 
13. A. 


24a. 


ha meliifolium 


24b. duci: 2-40 cm, more or less creeping. Stamens with long filaments e na apical 


ages 
ju its le 


appe 
. Corolla cleft to ca. 
25a. Stems generally 


ngth; lobes recurved 
pale brown to gray with elevated corky to papery longitudinal ridges, 


. A. moultonii 
25 


without scars. Anther cone with connate apical appendages. Stigma clavate. Fruit smooth 


26a. Plant wholly covered with a dense indumentum of stiff long hairs 
2 


3. A. trichosanthes 


Plant slightly preg with soft hairs 
black avy scars, without ee ridges. Anther cone xit free 
15. 


26b. 
25b. Stems generally 


apical piven aed Siete ad Fruit with emergen 


l. Argostemma parvifolium Benn., Plantae 
Javanicae Rariores. 1: 96. 1838. TYPE: Java(?), 
Sumatra: Horsfield s.n. (holotype, BM). Fig- 


ure 


Erect, anisophyllous, densely pubescent herb. 
Stem 15-50 cm long, usually unbranched, densely 
pubescent; internodes 6-15 mm. Leaves opposite, 
very unequal; stipules persistent, 4-7 mm long, 
ovate to cordiform, acute to acuminate at apex; 
larger leaves of the anisophyllous pairs with pu- 
bescent petioles 3-6 mm long, the lamina 4-13 
X 1.4-3 cm, generally oblanceolate or obovate to 
elliptic, basally acute and oblique, marginally ser- 
rulate, apically acuminate, herbaceous, pubescent 

both surfaces; midrib and primary veins (9-18 
pairs) distinct. Nanophyllous leaves persistent, 5— 
10 m long, cordiform to ovate. Inflores- 
cences solitary or a few together, 1—9-flowered, 
corymbiform, pubescent; peduncle 0.9-3.2 cm 
long, bracts to 1 cm long, broadly ovate to lan- 
ceolate; lateral branches, if present, to 1.5 cm long; 
pedicels 0.7-1.4 cm long. Flowers 5-merous, 
slightly zygomorphic; calyx lobes 4-6 mm long, 
triangular to ovate, acuminate, pubescent; corolla 
5 mm long, cleft to near the base, externally 
pubescent, internally glabrous; lobes triangular, 
spreading. Stamens 6-10 mm lon 
ther cells and apical appendages connate except 


g. coherent; an- 


calcicolum 


A. borragineum 


for the most apical part; anther cone narrowly 
ovoid, slightly upcurved; filaments short, unequal, 
curved; sacs opening completely and longitudinally; 
apical appendage ca. V5 the anther length, thin and 
smooth; connective distinct, smooth. Style 6-10 
mm long, glabrous with a slightly capitate stigma, 
shortly exserted. Fruit without furrows or ribs. 


There is a question mark after Java on the label 
of the type specimen, and on the label there is also 
written “Sumatra,” as is indicated in the proto- 
logue. Argostemma parvifolium grows on sand- 
stone, sandy banks, and cliffs near streams, from 
350 to 1,450 m. The species is based on non- 
Borneo material. The habit of the Borneo popu- 
lation is similar to the type of A. parvifolium var. 
involucratum (Hemsl.) Bakh. f. I have not studied 
the complete variation of A. parvifolium outside 
Borneo, and so for now I avoid infraspecific taxa. 
Except for the inner floral parts, the whole plant 
is densely covered by long soft hairs. As in all 
Argostemma species, the flowers are slightly pen- 
dulous at anthesis, and in this species the flowers 
are slightly zygomorphic. The rotate corolla has at 
its center distinct green nectar guides on the upper 
half. The anther cone is bent upwards and the 
stamens, in the upper half of the flower, have 
curved filaments. It is the only Borneo species with 
these characters. 


Annals of the 
Missouri Botanical Garden 


D 
0.2mm G i 
.05cm CD ; 
._ 1cm B; 2cmEF , 
|, 20cm A 
FIGURE 4. Argostemma parvifolium. —A. Branch.— 


B. Flower bud. — C. Anthers. — D. Style. — E. Stipule. — 
F. Nanophyllous leaf. — G. Hair from outer surface of the 
corolla. A, C-F from Burtt 11521 (E); B, G from Winkler 
767 (L) 


Additional specimens examined. BORNEO. SARA- 
Gunong Berumput, 3,500 ft., 1962, Burtt 


Burtt & Woods 2366 (E, SAR); 1979, Burtt 11521 (E, 
Sj; 1,200-1,500 ft., 1978, Burtt 11640 (E, S); 1,450 
m, 1976, Ilias & Neo $38349 (L, SAR); 7th Div., Hose 
Mts., 800 m, 1975, Chai et al. S37338 (K, L. MO, 
SAR). K KALIMANTAN: W Prov., Bidang Menabei, 700 m, 
1924, Winkler 767 (L). 


2. Argostemma elatostemma ee k. f., The 
Flora of British India 3(7): 45. 1880. TYPE: 
Penang, Griffith s.n. Nd K; Bakh. f. 
in herb., confirmed here). Figure 

A. motleyi Ridley, J. Bot. 65: 39. 1927. TYPE: Borneo: 
Bangarmassing, 1857-1858, Motley 1174 (holo- 
type, K). 


Creeping, anisophyllous herb. Stem 2-36 cm 
long, slightly branched or unbranched, pubescent; 
internodes 5-25 
equal; stipules persistent, 4-10 m 
broadly ovate or cordiform, acuminate to acute at 
apex; larger leaves of anisophyllous pairs with pu- 
bescent petioles 4-7 mm long; lamina 2-9.5 x 
1.3-3.5 cm, obovate to ovate and slightly subfal- 
cate, basally subcordate to auriculate with the lobes 
unequal, marginally entire, apically acute, herba- 
ceous, glabrous above and with a few hairs on the 
veins below; midrib and primary veins (5-11 pairs) 
distinct. Nanophyllous leaves persistent, 4-12 mm 


mm. Leaves opposite, very un- 
m long, ovate to 


long, cordiform. Inflorescences solitary, 1 -20-flow- 
ered, densely corymbiform or umbelliform, pubes- 
cent; peduncle 0.8-4 cm long; bracts to 0.7 cm 
long, ovate or lanceolate; lateral branches, if pres- 
ent, to 2.5 cm long; pedicels 0.4-1 cm long. Flow- 
ers 5-merous; calyx lobes 1-2 mm long, ovate to 
triangular, acute, pubescent; corolla 5-6 mm long, 
cleft to near the base, externally pubescent, inter- 
nally glabrous; lobes ovate to lanceolate, spreading. 
Stamens 5-6 mm long, coherent; anther cells and 
apical appendages connate; anther cone ovoid, 
straight; filaments short, equal, straight; sacs open- 
ing completely longitudinally; apical appendage ca. 
13 the anther length, thin and smooth; connective 
indistinctly passing over to the thecae, smoot 

tyle 5-7 mm long, glabrous with a slightly apunte 
stigma, shortly exserted. Fruit without furrows or 
ribs. 


Hooker based 4. elatostemma on two collec- 
tions, by Lobb and Griffith. Both are suitable for 
lectotypification. Bakhuizen f. (in herb.) selected 
the Griffith specimen in K, and I follow his choice. 

Argostemma elatostemma grows in wet shady 
forests, from 100 to 750 m. It occurs in the first 
division of Sarawak and on the southern tip of 
Kalimantan. The collections from these two areas 
are different. The Sarawak specimens are alike and 
similar to the type collection from the Malay Pen- 
insula, while the Kalimantan specimen, described 
as A. motleyi, agrees well with the type of A. 
elatostemma var. obovata King (1904). In this 
study I have not dealt with any infraspecific taxa 
because I have not yet studied the complete vari- 
ation outside Borneo. The specimen from Kali- 
mantan is smaller with shorter leaves, and there is 
only one flower in the collection. Argostemma ela- 
tostemma is characterized by its creeping habit, 
rooting at the nodes, silvery mottled leaves, and 
unequal leaf bases. The stems, petioles, veins of 
the lower leaf surfaces, and the inflorescences are 
densely pubescent. The flowers are rotate with a 
long pale anther cone which blackens in the press. 


Volume 76, Number 1 
89 


Bremer 
Argostemma in Borneo 


— 100 km 
S > 500 m 


Mars 1, 2, 


Additional specimens examine ORNEO. SARA- 
WAK: Ist Div., Matan 2,300 ft 
1722 (S, SAR); 500 ft., 


Burtt & Woods 1959 (E) 1,500 ft., 1929, Clemens 


0.2mm IG 
0.5cmCD  , 
.__1cmB; 2cmEF : 
cmA ` 


perea: elatostemma. —A. Habit. — 

B. Flower.—C. Anthers tyle.— E. "ans ules.— F. 

Nenophyllous leaf. —G. Hair from outer surface of the 

T —1. Stiff, parcela zs hair. A-I from Bremer 
2 (S; A, from Motley 1174 (K). 


FIGURE 5. 


Distributions in Borneo. — 1. Argostemma parvifolium —2. A. elatostemma. 


20912 (NY); 2,500-3,000 ft., Mjoberg s.n. (US); 1893, 
Ridley 12303 (BM, K, SING); 1st Div., Bau, S of Bukit 
= 100 m, 1964, Anderson et al. S20288 (L); Gu- 

ong n, 300 ft., 1975, Burtt 8175 (Ey; 1893, 
Ridley 11753 (BM, K, SING). 


ad 


Argostemma subfalcifolium Bakh. f., Blu- 
mea 7: 332. 1953. TYPE: Borneo. Sarawak: 
Mt. Dulit, Dulit trail, ca. 100 m, 1932, Rich- 
ards 20904 (holotype, K). Figure 6. 


Erect, anisophyllous herb. Stem 15-35 cm long, 

r sely pubescent; in- 
ternodes 7-23(-45) mm lo 
very unequal; stipules deciduous or persistent, 5— 
12 mm long, ovate to lanceolate or broadly ovate, 
acute at apex; larger leaves of the anisophyllous 
pairs with glabrous or sparsely pubescent petioles 
2-12 mm long; lamina 7-22 x cm, gen- 
erally subfalcate to lanceolate or ovate to obovate, 
basally acute and oblique, marginally entire or rare- 
ly serrulate, apically acuminate or rarely acute, 
herbaceous, generally glabrous on both surfaces or 
finely pubescent on the veins below; midrib and 
primary veins (6—15 pairs) distinct, rarely obscure. 
Nanophyllous leaves deciduous or persistent, 7- 
12 mm long, lanceolate. Inflorescences solitary or 
a few together, to 45-flowered, laxly corymbiform; 
peduncle 1.5-6 cm long, glabrous; bracts to 0.7 
cm long, ovate to lanceolate; lateral branches l- 
3.3 cm long, glabrous or pubescent; pedicels 0.5— 
1.2 cm long, pubescent. Flowers 5-merous; calyx 
lobes 2-4 mm long, ovate to triangular to lanceo- 


Annals of the 
Missouri Botanical Garden 


Mars 3, 4. 


late, acute to acuminate, + glabrous; corolla 5- 
6.5 mm long, cleft to near the base, glabrate on 
both sides; lobes ovate to lanceolate, spreading. 


20cmA 


FIGURE 6. Argostemma subfalcifolium. —A. Flow- 
ering branch. — B. Flower and fruit.— C. Anthers. — D. 
Style. — E. Stipule. — F. Nanophyllous leaf. — I. Stiff, plu- 
ricellular leaf hair. A, B (fruit) from Clemens 30650 (NY); 
B (flower)-I from Burtt 8228 (E). 


Distributions in Borneo. —3. Argostemma subfalcifolium. —4. A. densifolium. 


Stamens 5-7 mm long, coherent; anther cells and 
apical appendages connate except for the most 
apical part; anther cone ovoid, straight; filaments 
short, equal, straight; sacs opening completely lon- 
gitudinally; apical appendage ca. Y3 the anther 
length, thin and smooth; connective indistinctly 
passing over to the thecae, smooth. Style 5-7.5 
mm long, glabrous with a slightly capitate stigma, 
shortly exserted. Fruit with distinct longitudinal 
furrows. 


Argostemma  subfalcifolium grows in damp 
shady places in the lowland rainforest at altitudes 
from 100 to 1,500 m. It tends to have subfalcate 
leaves. The species is characterized by rather long 
calyx lobes and by furrowed fruits. The stems are 
glabrate and sometimes slightly pubescent. The 
peduncles and lateral branches of the inflorescence 
are mostly glabrous, whereas the pedicels, ovary, 
and lower part of the calyx lobes are pubescent. 
The flowers are rotate, and the buds are distinctly 
angular in cross section. The staminal cone is high. 
From the outside the thecae are completely shed 
by the connectives but can be seen as immerse 
sacs between the filaments. 

here are a few aberrant specimens. Anderson 
4090, from Bako in Sarawak, consists of very small 
plants. They were collected on fallen tree trunks 
and have a peculiar habit, but no other details 
contradict inclusion in this species. Meijer 2286, 
from Kalimantan, has nearly cordiform nanophyl- 
lous leaves, but otherwise fits. Brooke 9194, from 
Sarawak, has very narrow leaves with obscure pri- 
mary veins. 


Volume 76, Number 1 
1989 


Bremer 17 
Argostemma in Borneo 


Additional specimens examined. BORNEO. SABAH: 
W Coast Res., Kinabulu, Penthukan, 4,000 ft., 1933, 
Clemens 30650 (BM, BO, GH, K, L, j UC); Peni- 


E 4,000 ft., 1933, Clemens 31963 (BM, GH, L, 
UC); Curulau ridge, 30-40 m, 1965, Kanis & 
Kuripin 53966 (K, L). SARAWAK: E Div , Gunong Mulu, 
00 ft., 1975, Burtt 8228 (Ey; 1 


: , Ulu Sungei Melinau, e 

, 1961, p nomi 4090 (L); 7th Div., Hose 
Ulu 'Pamalad: 900 m, 1964, ipu SI 7606 (K, L); 7 
i E of Sungei Tellini, 2,800 ft., 


, Camp Mailinau, 
Nanukan, N of Tarakan, low alt., 
L). 


(K , 


1953, Meijer 2286 


4. Argostemma densifolium Ridley, J. Bot. 
65: 39. 1927. TYPE: Borneo. Puak (near Lun- 
du?), 1894, Ridley 12448 (holotype, K; iso- 
type, SING). Figure 7. 


Argostemma. densifolium var. latifolium Ridley, J. Bot. 
5: 40. 1927. TYPE: British North Borneo: Bongaya, 
1897, Ridley s.n. (holotype, SING). 


Rosette or creeping, anisophyllous herb. Stem 
5-16 cm long, slightly branched or unbranched, 
pubescent; internodes very short to 15 mm. Leaves 
opposite, very unequal; stipules persistent, 4-6 mm 
long, ovate to cordiform, obtuse to acuminate at 
apex; larger leaves of the anisophyllous pairs with 
pubescent petioles to 5 mm long; lamina 4-14 x 
1-4.5 cm, obovate to oblanceolate, basally acute 
to subcordate and oblique or with the lobes unequal, 
marginally entire or serrulate, apically obtuse or 
acute, herbaceous, glabrous above and pubescent 
on the veins below; midrib and primary veins (10- 
17 pairs) distinct. Nanophyllous leaves persistent, 
4-10 mm long, cordiform. Inflorescences solitary 
or a few together, 3-20-flowered, laxly to densely 
corymbiform; peduncle 1.5-7 cm long, pubescent 
or rarely glabrous; bracts to 0.2 cm long, ovate 
or lanceolate; lateral branches, if present, to 3 cm 
long, glabrous or pubescent; pedicels to 2 cm long, 
pubescent or rarely glabrous. Flowers 5-merous; 
calyx lobes ca. 1 mm long, stiff, narrowly trian- 
gular, acute, pubescent; corolla ca. 5 mm long, 
cleft to near the base, glabrous on both surfaces, 
the lobes lanceolate, spreading. Stamens ca. 5 mm 
long, coherent; anther cells and apical appendages 
connate; anther cone ovoid, straight; filaments short; 
sacs opening completely longitudinally(?); apical 
appendage thin and smooth; connective indistinctly 
passing over to the thecae, smooth. Style ca. 
mm long, glabrous with a slightly capitate stigma, 
shortly exserted. Fruit without furrows or ribs. 


, 0.2mm 1 
| 1cm B; 2cmEF 
|, 10cm 


FIGURE 7. Argostemma densifolium. — A. Habit. — 
B. Flower. — E. Stipule. — F. Nanophyllous leaf. — I. Stiff, 
disce leaf hair. A, E-I from Sylvester Tong 534346 
(K); B from Nielsen 538 (K). 


There are only a few collections of this species, 
and they are in bad condition. The description of 
the flowers was prepared from undissected flowers, 
and so some information is lacking. The scanty 
information on its habitat indicates that it grows 
on steep sandstone slopes and on gentle slopes on 
litter. It was collected at low altitude in south- 
western and northeastern parts of Sarawak and in 
Sabah. I have not managed to find the locality 
"Bongaya," but according to Steenis-Kruseman 
(1950), Ridley visited the Kudat, Sandakan, and 
Lubak Bay in 1897, so it must be somewhere in 
this area. Ridley (1927) described two varieties of 
this species. I agree with him that the two popu- 
lations are different. The southwestern population 
(Fig. 7A), including the type of 4. densifolium, 
has narrow leaves and glabrous inflorescences, while 
the northeastern population, including the type of 
var. latifolium, has denser foliage with larger, 
broader leaves and pubescent inflorescences. In this 
study, however, I have chosen not to deal with 
infraspecific taxa. On herbarium labels, Bakhuizen 
f. has treated these taxa as synonyms of A. ophi- 
rense. Argostemma ophirense and its allies are 
complex, but A. ophirense and A. densifolium 
from Borneo are two distinct taxa, in my opinion. 
Argostemma densifolium has a different leaf shape, 
denser foliage, and long peduncles. 


Additional specimens examined. BORNEO. SARA- 

es Ist Div., Sabal, 1974, Sylvester Tong 534346 (K, 

MO); 4th Div., Melinau gorge pathway, 1962, Burtt 

F Woods n (E); 7th Div., Gunong Mulu, Melinau 

gorge, 100 1978, Nielsen 538 (AAU, K), 200 m, 
1978, Nielsen 445 (AAU). 


18 Annals of the 
Missouri Botanical Garden 
B 

0.2mm I 

_05cm CD i 

. 1cm B; 2cmEF 

, 10cm A 

D 
Ficu Argostemma ophirense. — A. Flowering branch. — B. Flowers. — C. Anthers. — D. Style. —E. Stipule. — 


F. ton a leaf.— I. Stiff, nekis leaf hair. A, E, F from Bremer 1688 (S), B-D, I from Burtt & Woods 


2701 (E). 


5. Argostemma ophirense Maingay ex Hook. 
f., The Flora of British India 3(7): 45. 1880. 
TYPE: Malacca. Mt. Ophir, 1867-1868, 
Maingay 2578 (holotype, K). Figure 8. 

A. gout m ba s Branch Roy. Asiat. Soc. 

1912. : Mt. Matang, Ridley 11751 
eee SING: oe. BM, K). 


Erect, anisophyllous herb. Stem 7-30 cm long, 
slightly branched or unbranched, pubescent; in- 
ternodes 3-15 mm. Leaves opposite, very unequal; 
stipules persistent, 3-5 mm long, ovate to lanceo- 
late or oblanceolate, acute to obtuse; larger leaves 
of the anisophyllous pairs with pubescent petioles, 
to 5 mm long; lamina 5-16 x 0.4-3 cm, lanceo- 
late to linear, basally cuneate or acute to subcor- 
date and oblique or with the lobes unequal, mar- 
ginally entire, apically acuminate to narrowly 
acuminate, herbaceous to coriaceous, with a few 
stiff pluricellular hairs on the upper surface, pu- 
bescent on the veins below; midrib and primary 
veins (10-20 pairs) distinct, at least below. Nano- 
phyllous leaves persistent, 3-5 mm long, cordi- 
form. Inflorescences solitary or a few together, 7— 
18-flowered, laxly corymbiform, glabrous; pedun- 
cle 1-3 cm long; bracts to 0.5 cm long, ovate to 
linear; lateral branches 1-2 cm long; pedicels 0.4— 
0.7 cm long. Flowers 5-merous; calyx lobes ca. 1 
mm long, stiff, narrowly triangular, acute, glabrous; 


corolla 4-5 mm long, cleft to near the base, gla- 
brous on both surfaces; lobes ovate, acute, spread- 
ing. Stamens 4-6 mm 
and apical appendages connate except for the most 
apical part; anther cone ovoid, straight; filaments 


long, coherent; anther cells 


short, equal, straight; sacs opening completely lon- 
gitudinally; apical appendage ca. Y the anther 
length, thin and smooth; connective indistinctly 
passing over to the thecae, smooth. Style 4-7.5 
mm long, glabrous, with a slightly capitate stigma, 
shortly exserted. Fruit without furrows or ribs. 


When Ridley (1912) described A. salicifolium 
he stated that the specimens are preserved in the 
Singapore Gardens Herbarium. Bakhuizen f. indi- 
cated, on the herbarium label, the K specimen as 
the type (= holotype). The SING specimen must 
be treated as the holotype and the K specimen as 
an isotype, as Ridley (1912) said that the specimens 
are preserved in SI 

Argostemma ophirense grows on ground or on 
rocks in forest from 50 to 1,300 m; most collec- 
tions are from under 1,000 m. 

There is little morphological variation in this 
species except for variation in leaf width and dis- 
tinctness of foliar veins. It is characterized by in- 
dumentum on the stems, petioles, and veins be- 
neath. The leaves are generally lanceolate with 
oblique bases. The cordiform nanophyllous leaves 


Volume 76, Number 1 Bremer 19 
1989 Argostemma in Borneo 
en ed 
Ze 
5 g 
v—— 100 km 


¿EF > 500 m 


s 


Maps 5, 6. 


Distributions in Borneo. —5. Argostemma 


and stipules are persistent. The inflorescences are 
lax cymes. The type specimens of 4. salicifolium, 
probably collected from near the summit of Mt. 
Matang, have very narrow leaves with indistinct 
primary veins on the upper side. The specimens 
from Kalimantan have more obovate leaves. 


Additional specimens examined. EO. SARA- 


ORN 
WAK: 1st Div., Berumput, 1954, Brooke 8670 (L); 3,000 


0.2mm | 
_0.5cm CD j 
1cm B; 2cmEF 


ophirense. —6. A. anisophyllum. 


ft., 1962, Burtt & Woods 2762 (E, SAR); 1st Div., Mt. 
Poi (7 Pueh), 1929, Clemens field no. 6958 (NY); 1929, 
Clemens field no. 7083 (NY); 3,000 ft., 1955, Purse- 
glove & Shah 4811(K, L, SING); 1st Div., Sungei Lundu, 
1962, Burtt & Woods 2701 (E, SAR); 1st Div., Lundu, 
1929, Clemens s.n. (NY); 1st Div., Bako, at Serai path, 
1979, Bremer 1688 (S, SAR); Telok Delima, 200 ft., 

1956, Purseglove 4987 (K, L, SAR, SING, NY); Ist 
Div., Mt. Matang, 1,200 ft., 1954, Brooke 9517 (BM 

Ly 1,000 ft., 1955, Brooks. 9743 (L); 2,000 ft., 1962, 


J 


, 10cm A 


D Cc 


FIGURE 9. Argos 


temma anisophyllum 


—A. Flowering branch. — B. 


Flower. — C. Anthers. — D. Style. — E. Stip- 
ule. — F. Nanophyllous leaf. — I. Stiff, pluricellular leaf hair. From Winkler 897 (HBG). 


20 


Annals of the 
Missouri Botanical Garden 


IGURE 10. Argostemma dulitense. — A. Flowering branch. — B. Flo 
F. Nanophyllous leaf. A from Richards 1711 (K); B-F from Richards 2101 (K). 


Burtt & Woods 2512 (E, SAR); 2,500 ft., 1962, Burtt 
2518 (E, SAR); 1,500 ft., 1929, Clemens 
R); , 1929, Clemens 20911 
NE K, NY); 2,000-3, 000 ft., 1890, Hullett s.n. (SING); 
27-1928, jari a oc r s.n. (NY); 1,000 ft., 1924, 
Mjöberg 205 (NY, SING, UC); 1924, Mjöberg 206 (NY, 
UC); 2,500- e: 000 Pd Mems n. (UC); 1890?, Ridley 
11751 (BM ING). KALIMANTAN: E Prov., Gunong 
Beratus (Peak er Balikpapan), 800-900 m, 1952, Kos- 
termans 7437 (BO, Ly; 750 m, 1952, Meijer 687 (BO, 
L). 


6. Argostemma anisophyllum Merr., Mitt. 
Inst. Allg. Bot. Hamburg 7: 278. 1937. TYPE: 
West Borneo. Bukit Raja, 1,250 m, 1924, 
Winkler 897 (holotype, HBG; isotype, NY). 
Figure 9. 


Erect, anisophyllous herb. Stem 10-30 cm long, 
slightly branched or unbranched, glabrous or gla- 
brate; internodes 5-25 mm. Leaves opposite, very 
unequal; stipules persistent, 4-5 mm long, broadly 

ovate to lanceolate, acute at apex; larger leaves of 
the anisophyllous pairs with glabrous petioles 3-7 
mm long; lamina 2-6.2 x 1.2-2.2 cm long, ob- 
ovate to oblanceolate, basally acute to attenuate 
and oblique, marginally entire, apically acuminate, 
herbaceous to membranaceous, with a few stiff 
pluricellular hairs on the upper surface, glabrous 
below; midrib and primary veins (6—10 pairs) dis- 
tinct. Nanophyllous leaves persistent, 2-4 mm long, 
cordiform. Inflorescences solitary, 1—5-flowered, 
laxly corymbiform, glabrous; peduncle 2-2.8 cm 
long; lateral branches 1-1.5 cm long; bracts ca. 
0.1 cm long, lanceolate or triangular; pedicels 1— 
2.2 cm long. Flowers 5-merous; calyx lobes ca. 1 


er. — C. Anthers. — D. Style. E Stipule. — 


mm long, stiff, narrowly triangular, acute, glabrous; 
corolla 8-9 mm long, cleft to near the base, gla- 
brous on both surfaces; lobes ovate to broadly 
ovate, spreading. Stamens 9-11 mm long, coher- 
ent; anther cells and apical appendages connate 
except for the most apical part; anther cone nar- 
rowly ovoid or ovoid, straight; filaments short, equal, 
straight; sacs opening completely longitudinally; 
apical appendage ca. Y the anther length, thin and 
smooth; connective indistinctly passing over to the 
thecae, smooth. Style 10-12 mm long, glabrous, 
with a slightly capitate stigma, exserted. Fruit with- 
out furrows or ribs. 


Argostemma anisophyllum grows from 1,250 
to 1,400 m at Bukit Raja. 

It has been treated as a synonym of A. ophirense 
by Bakhuizen f. on herbarium labels. Argostemma 
anisophyllum has fewer hairs on the stems, much 
smaller and thinner leaves, longer peduncles and 
pedicels, and much larger flowers. 


Additional specimen examined. BORNEO. KALI- 
MANTAN: Bukit Raja, 1,400 m, 1924, Winkler 950 (HBG). 


7. Argostemma dulitense Merr., Sarawak Mus. 

: 554. 1928. TYPE: Borneo. Sarawak: Mt. 

Dulit, 1,200-1,500 m, 1923, Mjöberg 8 (ho- 
lotype, UC). Figure 10. 


Erect anisophyllous herb. Stem 10-40 cm long, 
slightly branched or unbranched, densely pubes- 
cent; internodes to 8 mm. Leaves opposite, very 
unequal; stipules persistent, 2-4 mm long, broadly 
ovate, acute at apex; larger leaves of the aniso- 


Volume 76, Number 1 
1989 


Bremer 21 
Argostemma in Borneo 


u— 100 km 
ES > 500 m 


Maps 7, 8. 


phyllous pairs with glabrous petioles ca. 3 mm long; 
lamina 2- -1.5 cm, lanceolate or narrowly 
elliptic to obovate, basally cuneate to attenuate and 
with the lobes equal, marginally entire, apically 
acute to acuminate, coriaceous, glabrous on both 
surfaces; midrib distinct, primary veins generally 
obscure. Nanophyllous leaves persistent, ca. 5 mm 
long, ovate. Inflorescences solitary or a few to- 
gether, 1-3-flowered, glabrous; peduncle 1-2.5 
cm long; bracts to 0.3 cm long, ovate to triangular 
or lanceolate; pedicels 1-2.7 cm long. Flowers 
5-merous; calyx lobes ca. 2 mm long, stiff, narrowly 
triangular, glabrous. Corolla 9-12 mm long, cleft 
to near the base, glabrous on both surfaces; lobes 
ovate to lanceolate, spreading. Stamens ca. 7 mm 
long, coherent; anther cells and apical appendages 
connate except for the most apical part; anther 
cone ovoid or narrowly so, straight; filaments short, 
equal, straight; sacs opening completely longitu- 
dinally; apical appendage ca. V5 the anther length, 
thin and smooth; connective indistinctly passing 
over to the thecae, smooth. Style ca. 8 mm long, 
glabrous with a scarcely widened p^ shortly 
exserted. Fruit without furrows or ribs. 


Argostemma dulitense grows in shady places 
on tree trunks or on rocks near waterfalls in moss 
and heath forests. 

It is characterized by its glossy dark green upper 
leaf surfaces, pale lower leaf surfaces and very firm 
leaf texture. The stems are covered by appressed 
hairs; otherwise it is glabrous. The corolla is large 
and rotate with green spots at the base, and the 


8 


—— 100 km 
¿ES > 500 m 


Distributions in Borneo. — 7. Argostemma dulitense. —8. A. apiculatum. 


anther cone is high. The specimens from Kali- 
mantan have distinct primary veins, while the Sa- 
rawak specimens have obscure veins. 


Additional specimens examined. BORNEO. SARA- 
WAK: 4th Div., Dulit Ridge 1,250 m, 1932, Richards 
1711 (K), 1,000 m, 1932, Richards 2101 (K, Ly; 4th 
Div., Gunong Dulit, at Long Atun, 1,300 m, 1965, Asah 
ak Luang S22752 (K, L); 1,200-1,500 m, 4th Div., 
Ulu Tinjar, 2,600 ft., 1974, Sylvester Tong 34847 (K, 
L). KALIMANTAN: rov., Boekit Batue Ajah, 1896- 
1897, Jaheri in exp. E diebus 1636 (BO, L). 


8. Argostemma apiculatum B. Bremer, sp. 
nov. TYPE: Borneo. Sarawak: 7th Div., summit 
of Bukit Tibang, 5,000 ft., 1969, Anderson 
& Ilias Paie 528657 (holotype, K; isotypes, 
E, L). Figure 11. 


Herba erecta, glabra, inflorescentiis floribus paucis. 
Folia opposita, ilis parium inaequalissimis. Inflorescentia 
corymbosa, floribus grandibus. Lobi calycis rigidi, anguste 
triangulares, acuti. p corollae apiculati. Antherae et 
appendices apicales connatae praeter partem ig cw 

mam. Appendix apicali l4 anthera brevior, tenuis, lae 
Connectivum indistinctum. Stylus glaber stigmate vix "di 
d 

mma d simile sed planta glabra, lobis co- 
m ID differ 


Erect, anisophyllous, glabrous herb. Stem 2-30 
cm, unbranched; internodes 6-16 mm. Leaves op- 
posite, very unequal; stipules persistent, 5-7 mm 
long, ovate to cordiform, acute at apex; larger 
leaves of the anisophyllous pairs with petioles 3— 
5 mm long, the lamina 3-8.5 0.8-1.9 cm, 
narrowly elliptic, basally cuneate to attenuate with 


Annals of the 
Missouri Botanical Garden 


D 


, 05cm CD i 

|, 1cm B; 2cmEF , 

| 10cm A 

FIGURE ll.  Argostemma y eiue — A. Flower- 
ing branch.—B. Flower.—C. Anther —D. Style. —E. 


tipule. —F. Nanophyllous leaf. From pee & Ilias 
Paie S28657 (K). 


the lobes equal, marginally entire, apically acu- 
minate, herbaceous to membranaceous; midrib dis- 
tinct, the primary veins (10-16 pairs) obscure. 
Nanophyllous leaves persistent, 5-6 mm long, el- 
liptic to lanceolate. Inflorescences 1-4-flowered, 
laxly corymbiform, solitary or a few together; pe- 
duncle 1.5-2 cm; bracts to 0.5 cm long, lanceolate; 
pedicels to 0.8 mm. Flowers 5-merous; calyx lobes 
ca. 3 mm long, stiff, narrowly triangular, acute, 
glabrous; corolla 10-12 mm long, cleft to near the 
base; lobes ovate to broadly so, pointed, spreading. 
Stamens 4.5-5 mm long, coherent; anther cells 
and apical appendages connate except for the most 
apical part; anther cone ovoid, straight; filaments 
short, equal, straight; sacs opening ul lon- 
gitudinally; apical appendage less than 1⁄4 the an- 
ther length, thin and smooth; connective ida 
ly passing over the thecae, smooth. Style ca. 5 mm 
long, glabrous, with a scarcely widened stigma, 
shortly exserted. Fruit without furrows or ribs. 
Argostemma apiculatum was collected once at 
Bukit Tibang, on igneous soil at 1,600 m in mossy 


forest. 
It is a glabrous plant with large, broad, pointed 


¿0.2mm I , 


FIGURE 12. Argostemma burttii.—A. Habit. — B. 
Flower. — C. Anthers. — D. Style. — E. Stipule. — F. Nano- 
phyllous leaf. — I. Stiff, pluricellular leaf hair. From Burtt 
& Martin 4986 (E). 


corolla lobes. The apical appendages of the anthers 
are short. It is similar to 4. burttii because of its 
large corollas and the short apical appendages. 
Argostemma apiculatum differs from that species 
by its apiculate corolla lobes and erect glabrous 


habit 


9. Argostemma burttii B. Bremer, sp. nov. 


& Martin 4986 (holotype, E). Figure 12. 


rba repens vel suberecta, inflorescentiis floribus so so 


uis, laevis. Connectivum indistinctum. Stylus glaber stig- 
mate s en tato. 

mma apiculatum = sed foliis serrulatis, 
lobis ed non cu a differ 


Creeping or suberect, anisophyllous herb. Stem 
to 20 cm, slightly branched or unbranched, gla- 
brous; internodes 2-4 mm. Leaves opposite, very 


Volume 76, Number 1 
1989 


remer 
Argostemma in Borneo 


9 


E » "i m 


Mars 9, 10. 


unequal; stipules 2-4 mm long, broadly ovate to 
cordiform, acute at apex. Larger leaves of the 
anisophyllous pairs with glabrous petioles 1-3 mm 
long; lamina 1-3.5 X 0.4-0.9 cm, narrowly el- 
liptic, basally cuneate to attenuate with the lobes 
equal, marginally serrulate, apically acuminate, 
herbaceous to membranaceous, glabrous on both 
surfaces but with 3 or 5 rows of stiff pluricellular 
hairs on the upper surface; midrib distinct, the 
primary veins obscure. Nanophyllous leaves per- 
sistent, to 4 mm long, ovate to lanceolate. Inflo- 
rescences 1-2-flowered, monochasial, solitary or a 
few together; peduncle 1-2 cm, glabrous; bracts 
to 0.2 cm long, lanceolate to linear; pedicels 0.7- 
1.5 cm, glabrous. Flowers 5-merous; calyx lobes 
ca. 1.5 mm long, stiff, triangular to narrowly tri- 
angular, acute, glabrous; corolla 9-12 mm long, 
cleft to near the base, glabrous on both surfaces; 
lobes ovate, spreading. Stamens 4-5 mm long, 
coherent; anther cells and apical appendages con- 
nate except for the most apical part; anther cone 
ovoid, straight; filaments short, equal, straight; sacs 
opening completely longitudinally; apical appen- 
dage less than 14 the anther length, thin and smooth; 
connective indistinctly passing over to the thecae, 
smooth. Style ca. 5 mm long, glabrous, with a 
scarcely widened stigma, shortly exserted. Fruit 
without furrows or ribs. 


Argostemma burttii was found growing at bases 
of trees among mosses at ca. 1,300 m. It is a local 
endemic species from the Hose Mountains. Mr. B. 
L. Burtt has collected it at three different places. 


10 
— 100 km 
$e > 500 m 


Distributions in Borneo. —9. Argostemma burtii. —10. A. rupestre. 


It is a beautiful little plant with very large flow- 
ers. The corolla is open and flat with green spots 
at the center, and the anther cone is cream-colored. 
The border between the filaments and the thecae 
is indistinct, as in, for example, 4. apiculatum, A. 
rupestre, and A. geesinkii. With the last two it 
also shares the character of three or five rows of 
stiff, pluricellular hairs on the upper leaf surface. 
Argostemma burtii is most similar to A. apicu- 
latum, both having large corollas and short apical 
appendages, but 4. burtii differs by having ser- 
rulate leaves and creeping or suberect habit. 


Additional specimens examined. BORNEO. SARA- 
WAK: 7th Div., S Hose Mts., W of Bukit Sanpandai, 
1980, Burtt 12750 (E); E of Bukit Sanpandai, 4,500 


ft., 1980, Burtt 12817 (E). 


10. Argostemma rupestre Ridley, J. Bot. 65: 

. 1927. TYPE: Borneo. On the hill near Ma- 

tang, Haviland 1031 (holotype, K; isotypes, 
SAR, SINC). Figure 13 


Creeping or suberect, anisophyllous herb. Stem 
3-14 cm, generally unbranched, glabrous or with 
a few hairs; internodes very short to 7 mm. Leaves 
opposite, very unequal; stipules persistent, 2-7 mm 
long, ovate to lanceolate, acute at apex; larger 
leaves of the anisophyllous pairs with glabrous pet- 
ioles to 5 mm long; lamina 1.3-5.5 x 0.2-1.8 
cm, narrowly elliptic to lanceolate or oblanceolate, 
rarely elliptic or linear; basally cuneate to attenuate 
with the lobes equal; marginally serrulate, apically 
acuminate; herbaceous to membranaceous, gla- 


24 


Annals of the 
Missouri Botanical Garden 


D C 
0.2mm I i 
0.5cm CD 


u_1cm B; 2cmEF 


¿10cm A n 
FIGURE 13. Argostemma rupestre. — A. Habit. — B. 
Flower. Te Anthers. — D. Style. — E. Stipule. — F. Nano- 


agp leaf. — I. Stiff, pluricellular leaf hair. From Bre- 
r 1665 (S). 


brous on both surfaces except for 3 or 5 rows of 
stiff pluricellular hairs above; midrib distinct, the 
primary veins obscure. Nanophyllous leaves per- 
sistent, 2-10 mm long, ovate to elliptic or lanceo- 
late. Inflorescences 1-3(-5)-flowered, laxly cor- 
ymbiform, solitary or a few together; peduncle 1- 
3 cm, glabrous or pubescent; bracts 0.1-0.3 cm 
long, lanceolate; pedicels 0.5-1.5 cm, glabrous or 
pubescent. Flowers 5-merous; calyx lobes ca. 1 
mm long, stiff, triangular to narrowly triangular, 
acute, glabrous; corolla 4-7 mm long, cleft to near 
the base; lobes lanceolate, acute to acuminate, erect- 
spreading. Stamens 5-7 mm long, coherent; anther 
cells and apical appendages connate except for the 
most apical part; anther cone ovoid, straight; fil- 
aments short, equal, straight; sacs opening com- 
pletely longitudinally; apical appendage ca. V5 the 
anther length, thin and smooth; connective indis- 
tinctly passing over to the thecae, smooth. Style 
6-8 mm long, glabrous, with a slightly capitate 
stigma, shortly exserted. Fruit without furrows or 


Argostemma. rupestre is a small herb growing 
on wet sandstone boulders or other wet places in 
or near streams or waterfalls, chiefly among moss- 
es. It occurs from low altitudes up to ca. 1,500 m 
in mossy forests. 

This anisophyllous herb has glabrous leaves ex- 


.2mm | 
.5cm CD 
1cm B; 2cmEF 


9 


D [e ¿10cm A N, 
FIGURE 14. sal cag iei —A. Flowering 
branch. — B. Flower. — C. Anthers.—D. Style. 


- Stip- 
ule. —F. Nan oso leaf. —1. “Stiff eid leaf 
hair. From Geesink 9272 (L). 


cept for three or five rows of stiff pluricellular hairs 
on the midrib and along the margins. The marginal 
hairs make the leaves serrulate. The lower surface 
of the leaves is pale. The stipules and nanophyllous 
leaves are similar in shape. The corolla is cleft to 
near the base into narrow and erect-spreading lobes. 
The anther cone is high with a pale yellow color 
near the base and whitish at the apex. The inflo- 
rescences are lax cymes with one to five flowers. 
The peduncles and pedicels are narrow and gen- 
erally glabrous. The specimens from Bako are all 
pubescent on the peduncles, pedicels, and ovaries 
and have somewhat broader corolla lobes. There 
are two collections from different areas (Brooke 
9131 and Richards 2455) with very narrow leaves; 
both were collected in or near streams and may 
be ecotypes. Clemens 21737 came from an un- 
certain locality: on the herbarium label is written 

at, Upper Rajang River," while on the attached 
field label both **Gat" and *Mt. Ma 
ten. This plant has pubescent pedicels, as does the 
Bako population. 


,* * 
tang are writ- 


Additional EM examined. BORNEO, SARA- 
WAK: lst Div., N slope Mt. Penrissen, 1,000-1,100 
, UC); 1st Div., 


Sahal, 350 m, 19 


L, SAR); 4th Div. "Mt. Dulit, 300 m, 1932, Richarda 


Volume 76, Number 1 
1989 


remer 
Argostemma in Borneo 


Mars 11, 12. 


2455 (K, L); 4th Div., Ulu Sungei Chipidi, Ulu Tinjar, 
2,300 ft., 1974, Chai S34797 (K, L, MO, SAR); 7th 
Div., Sungei Melinau, 450 ft., 1980, Burtt 12667 (E); 

, Hose Mts., Bukit Salong, 1982, Burtt 12734 
(E); Bukit Sanpandai, 4,500 ft., 1980, Burtt 12796 (E); 
without loc., Clemens 21737 (NY). KALIMANTAN: W Prov., 
Bukit Raja, 1,400 m, Winkler 951 (HBG). 


11. Argostemma geesinkii B. Bremer, sp. nov. 
TYPE: Borneo. Kalimantan Timur, between Pa- 
padi and Pamilau, 1,400 m, 1981, Geesink 
9272 (holotype, L). Figure 14. 


Herba pusilla, repens, inflorescentiis floribus solitariis 


Lobi corollae erecto-patentes, in exsiccatis subrosei 
therae et apponi „apicales connatae praeter pacen 


apicalissima anth 


aequans, tenuis, E Üonnectivum E EN Stylus 
glaber stigmate leviter capitato. 

stemma rupestre simile sed ae pusilla, lobis 
corollae latis erecto-patentibus differ 


Small, creeping, anisophyllous herb. Stem 2-20 
cm, slightly branched or unbranched, glabrous; 
internodes very short to 5 mm. Leaves opposite, 
very unequal; stipules 2-4 mm long, ovate to cor- 
diform, acute to obtuse 
the anisophyllous pairs with glabrous petioles 1-2 
mm long; lamina 2.5-7 x 0. 


at apex; larger leaves of 


-1.5 cm, elliptic to 
obovate, basally cuneate to attenuate with the lobes 
equal, marginally serrulate or entire, apically acute 
to shortly acuminate or rarely obtuse, membrana- 
ceous, glabrous on both surfaces but with 3 or 5 


12 
— 100 km 
S > 500 m 


Distributions in Borneo. — 11. Argostemma geesinkii. —12. A. gaharuense. 


rows of stiff pluricellular hairs on the upper surface; 
midrib distinct, the primary veins generally ob- 
scure. Nanophyllous leaves persistent, 2-5 mm 
long, ovate to elliptic, acute. Inflorescences 1-2- 
flowered, monochasial, solitary or a few together; 
peduncle 0.2-1 cm, glabrous; bracts ca. 1 mm 
long, lanceolate to triangular; pedicels 0.5-1 cm, 
glabrous. Flowers 5-merous; calyx lobes ca. 1 mm 
long, stiff; narrowly triangular, acute, glabrous; 
corolla 5-7 mm long, cleft to near the base, gla- 
brous on both surfaces; lobes ovate to broadly 
ovate, acute, erect-spreading. Stamens 4-5 mm 
long, coherent; anther cells and apical appendages 
connate except for the most apical part; anther 
cone ovoid, straight; filaments short, equal, straight; 
sacs opening completely longitudinally; apical ap- 
pendage ca. V5 the anther length, thin and smooth; 
connective ls o passing over to the thecae, 
smooth. St -5 mm long, glabrous, with a slight- 
ly capitate sigma, shortly exserted. Fruit without 
furrows or ribs. 


Argostemma geesinkii grows on tree trunks and 
on boulders among mosses, from 950 to 1,500 m. 
A few collections are from sandstone. 

Argostemma geesinkii is a very small creeping 
species with small anisophyllous leaves and cordi- 
form stipules. It is glabrous except for the stiff 
pluricellular hairs in three or five rows on the upper 
leaf surfaces. The flowers are generally solitary on 
thin pedicels. The corolla lobes are broad and often 
become pinkish in the press. All specimens except 
Endert 3916 are very similar. This aberrant spec- 


Annals of the 
Missouri Botanical Garden 


_0.5¢ i 

cL Icm E 2cmEF 

| 10cm A » 

FIGURE 15.  Argostemma | gaharuense. — A. 


Hab- 
it.—B. Flower and fruit. — C. Anthers. — D. Style. — E. 
Stipule. — F. Nanophyllous leaf. —1. Hair from upper sur- 
face of leaf. From Burtt 2655 (E). 


imen lacks stiff hairs and has thicker leaves and 
prominent primary veins, but there are no floral 
differences between it and the other collections of 
A. geesinkii. There is another specimen from the 
same area (Endert 3800) that does not deviate 
from the others. Perhaps Endert 3916 is a different 
taxon. 


Additional specimens examined. BORNEO. SABAH: 
Kimanis road, Keningau, 4,400 ft., 1979, Collenette 
4179 (E). SARAWAK: 4th Div., Gunong Mulu, 4,000 ft., 
1962, Burtt & Woods 2185 (E); 5th Div., Bakelalan, 
4,000 ft., 1955, Brooke 10411 (L); 5th Div., Kalabit 
Highlands, Apa Balu Buli, 1,700 m, 1970, Nooteboom 
& Chai 2198 (L); 7th Div., ridge NW of Sungei Tellini, 


2.900 ft., 1978, Burtt 11443 (E); 7th Div., Bukit Lamut, 
Ulu Amau, 950 m, 1964, Ashton 521265 (L). KALI- 
MANTAN: E 


ei, 
near Kemoel, 1, 1925, Endert 3800 (L); 1,500 
m, 1925, Endert 3916 (A, K, L, SING). 


12. Argostemma gaharuense B. Bremer, sp. 
nov. TYPE: Borneo. Sarawak: Ist Div., Bukit 
Gaharu, 2,000 ft., 1962, Burtt 2655 (holo- 
type, E). Figure 15. 


Herba pusilla, ramosa, inflorescentiis floribus solitariis 


w 
m 
[a] 
in 
Ld 
g 
Ss 
E 
un 


glaber, stigmate vix dilatato 


Argostemma rupestre simile sed planta pubescens, ra- 
mosa, lobis corollae reflexis, fructu costato differt. 


Small, creeping(?) herb. Stem to 20 cm, 
branched, glabrous; internodes very short to 4 mm. 
Leaves opposite, very unequal; stipules 2-3 mm 
long, ovate to cordiform, acute at apex; larger 
leaves of the anisophyllous pairs with glabrous pet- 
ioles 1-2 mm long; lamina 0.4-0.6 x 0.2-0.4 
mm, elliptic to narrowly so, basally cuneate to 
attenuate with the lobes equal, marginally entire, 
apically acute, coriaceous, pubescent above an 
glabrous below; midrib distinct but without primary 
veins. Nanophyllous leaves persistent, 2-3 mm 
long, ovate to elliptic, acute. Inflorescences 1-6- 
flowered, laxly corymbiform, many together; pe- 
duncle 1.4-2.8 cm, glabrous; bracts ca. 1 mm 
long, lanceolate; pedicels 0.5-1.4 cm, glabrous. 

owers 5-merous; calyx lobes ca. 1 mm long, 
triangular, acute, glabrous; corolla ca. 4 mm long, 
cleft to near the base, glabrous on both surfaces; 
lobes ovate, acute, reflexed. Stamens 3.5-4 mm 
long, coherent; anther cells and apical appendages 
connate except for the most apical part; anther 
cone ovoidal, straight; filaments very short, equal, 
straight; sacs opening completely longitudinally; 
apical appendage ca. ! the anther length, thin and 
smooth; connective indistinctly passing over to the 
thecae, smooth. Style ca. 4.5 mm long, glabrous, 
with a scarcely widened stigma, shortly exserted. 
Fruit with 5 distinct ribs corresponding to the mid- 
dle vascular strands of the calyx lobes. 


Argostemma gaharuense was collected by Burtt 
at 660 m at Bukit Gaharu. Its habit is not given 
on the herbarium label. 

On the label of the type specimen is written: 
"Leaves dark green above, white below. Corolla 
white, lobes strongly reflexed. Base of anther cone 
green, tips white." It is a small, easily recognized 
plant. The leaves are dark and pubescent on the 
upper surfaces, and below they are white and gla- 
brous. The stems are distinctly branched distally 
with many apical inflorescences, each with one to 
six flowers in lax cymes. The vascular strand to 
each calyx lobe becomes a distinct rib, making the 
fruit five-ribbed. The habit is similar to that of A. 
rupestre, A. geesinkii, and A. burttii, but A. 
haruense differs from these by having monoseriate 
leaf hairs, branched stems, and reflexed corolla 
lobes. The leaf and stipule shapes are the same as 
those of A. geesinkii. The specimen from Gunong 
Apeng has narrower leaves than the type, and there 
is only one inflorescence with two flowers. However, 
it has reflexed corolla lobes and the same kind of 
indumentum. 


Volume 76, Number 1 
1989 


Bremer 27 
Argostemma in Borneo 


FIGURE 16. 


Argostemma hameliifolium.— A. Flowering branch. 


—B. Flower.—C. Anthers.— D. Style. — 


Stipule.—G. Hair from outer surface of the corolla.—G,. Hair from inner rte of the corolla. A, E from Bremer 


1752 (S); B-D, G from Richards 1799 (L). 


Additional specimen examined. BORNEO. SARAWA 
2nd Div., Gunong Apeng, 3,000 ft., 1961, Colli nette 
). 


s.n. (K 


13. Argostemma hameliifolium Wernham 
in Gibbs, J. Linn. Soc. Bot. 42: 89. 1914. 
TYPE: British North Borneo. Kinabalu: before 
Lobang rock, 5,000 ft., 1910, Gibbs 4101 
(holotype, K; isotype, BM). Figure 16. 


A. hullettii Rig: Bot. 65: 35. 1927. TYPE: Borneo 


araw atang, 2,000 ft., 1884, Hullett 329 
(lectotype, SING; Bakh. f. in herb., confirmed here) 
A. isophyllum pd Mitt. Inst > Bot. Hamburg 7: 
279. 1937. TYPE: West Borneo: Bukit Raja, 1,400 


m, 1924, Winkler 949 pni HBG; isotype, 
NY). 


Large, erect, isophyllous herb. Stem 10-100 
cm, unbranched, glabrous to pubescent, rarely with 
corky or papery bark; internodes : 
Leaves opposite, equal; stipules deciduous, 5-27 
mm long, oblong to lanceolate to ligulate, entire 
or slightly bifid, acute to acuminate or obtuse at 
apex; petiole 4-30 mm long, glabrous to pubescent; 
lamina 5-19 x 1-8.7 cm, oblanceolate or obovate 
to lanceolate or ovate, basally attenuate to acute 
or cuneate with the lobes equal, marginally entire, 

pically acuminate to acute or rarely obtuse, her- 
baceous, glabrous to finely pubescent above, gla- 
brous to pubescent below; midrib and primary veins 
(6-12 pairs) distinct. Inflorescences 3-40-flow- 
ered, umbelliform and rarely with 2 or 3 scorpioid 
branches, generally solitary; peduncle 2-7.5 cm 
almost glabrous; bracts to 5(-10) mm long, lan- 


» 


ceolate to linear; lateral branches, if present, to 
3.5 cm long, almost glabrous; pedicels 6-12 mm 
long, finely pubescent. Flowers 5-merous; calyx 
lobes 1-1.5 mm long, uo oma to ovate, acute, 
finely pubescent; corolla 4- m long, cleft to 

near the base, externally Dabo or pubescent, 
internally glabrous to pubescent; lobes lanceolate, 
spreading to reflexed. Stamens 3-6 mm long, co- 
herent; anther cells and apical appendages connate 
except for the free and incurved most apical parts, 
each of these formed into an open cylinder; anther 
cone ovoid, straight; filaments generally short, equal, 
straight or bent; sacs opening completely longitu- 
dinally; apical appendage ca. 14-13 the anther 
length, thin and smooth; connective distinct, gen- 
erally smooth. Style 4.5-7.5 mm long, pubescent 
or rarely glabrous, with a capitate stigma, shortly 
exserted. Fruit without furrows or ribs. 


Ridley based 4. hullettii on three collections, 
by Haviland and by Hullett. All are suitable for 
lectotypification. Bakhuizen f. (in herb.) selected 
the single Hullett specimen in SINC, and I follow 
his choice. The elevation mentioned in the proto- 
logue is indicated only on that specimen. 

Argostemma hameliifolium is common and is 
widely collected. It grows from low altitude to 2,000 
m. Most collections are from sandstone; a few are 
from limestone. It grows on boulders or sand in 
wet places near streams or waterfalls. 

Argostemma hameliifolium is part of a big, 
poorly understood complex, which includes 4. 
moultonii (see discussion under that species). Ap- 


28 Annals of the 
Missouri Botanical Garden 
ed ea 
13 14 


—— 100 km 
¿ER > 500 m 


Maps 13, 14. 


proximately 100 specimens have been referred to 

. hameliifolium, all large plants with large, equal, 
opposite leaves and more or less erect stems. The 
leaves vary in shape and size and in distinctness 
of nervation. In a few cases the leaves are very 
broad and obtuse; these are similar in appearance 
to A. borragineum. The stipules, of which only 
the upper first to third pairs are persistent, are 
large, ovate to ligulate, and entire or slightly cleft. 
The flower buds are narrowly ovate in outline. The 
anther cone is high with short, free apical ap- 
pendages and generally short filaments. The mar- 
gins are longer than the central part and incurved 
so that each apical appendage forms an open cyl- 
inder. The style is generally hairy with a distinct 
capitate and slightly bifid, slightly exserted stigma. 
In this treatment I maintain A. hameliifolium sep- 
arate from A. moultonii. There are many distin- 
guishing characters, though not always constant, 
and many specimens are impossible to determine 
with certainty. Argostemma hameliifolium is more 
erect and has larger vegetative organs. Its anthers 
have short apical appendages and shorter filaments. 


Additional specimens examined. BORNEO. BRUNEI: 
Kuala Belalong, Temburong, 200 ft., 1957, Ashton 42 
(K) Belatai District, Labi area, 1969, van Niel 4580 
(L). SABAH: W Coast Res., Kinabalu, E shoulder, 6,500 
ft., 1961, Chew et al. 1041 (K, L, SING); Ulu Liwagu 
and Ulu Mesilau, 1961, Chew et al. 2875 (BO, K, L, 
SING); Dallas, 3,000 ft., 1931, Clemens 26033 (BM, 
NY, UC); Tenompok, 4,500 ft., 1931, 

L, NY UC); Ten- 
ompok, 5,000 ft., 1932, Clemens 29538 (BM, BO, G, 
K, L, NY); Marai Parai, 5,000 ft., 1933, Clemens 32578 


O km 
S > 500 m 


Distributions in Borneo. — 13. Argostemma hameliifolium. —14. A. moultonii. 


(A, BM, BO, L, NY, UC); Penibukan, 4,000 ft., 1933, 
Clemens 35137 (BM); Penibukan, 4,000-5,000 ft., 1933, 
Clemens s.n. (BM); Kiau, 1915, Clemens 10059 (UC); 
1958, Collenette 116 (BM); 1,650 m, 1966, Ding Hou 
; 1937, Grieswold 29 (GH); western route, 1,780 
m, /Voote- 


(Ly Sandakan Res., Bettotan near Sandakan, 1927, Kloss 
SFN19154 (K, L, NY, SING, SV Sandakan Res., San- 
dakan, 1977, Anderson 4284 (MO, UC); Tawau Res., 
Tawau Hill, 1978, Fedilis & Sumbing 5AN88281 (K, 


cie yon Bukit Lanjak, 4, end ft., 
(L, SAR); 4th Div., Mt. . 250 m, 1979, Bremer 
ET (S, SAR); 1962, [ea & Woods 2372 (E) ace 


Purseglove a (K, L, N 
pe m, 1932, native dica 1256 (K); Dulit ridge, 
1932, native collector 1 428 (L), under 300 m, 1932, 
Richards 1052 (K); 1,100 m, 1932, Richards 1799 
m, Richards 7090 (K); 4th 
"Chai $30081 (K, L); 4th 
Div., Gunong Mulu, 4,000 ft., Burtt & Woods 2182 (E, 
SAR); N of Sungei | Piet 1978, Nielsen 204 (K); 800- 
00 m, Nielsen 494 (AAU); 4th Div., Upper Baram 
River, van Niel 3558 (L); 4th Div., Kalabit Highlands, 
Bario, 1,000 m, 1970, Nooteboom. & Chai 1704 e 
Batu Lawi, 1,250 m, 1970, Nooteboom & Chai 2 
(B, L, SAR, US); 5th Div., Maputi, 1955, Brooke e 
(L); 5th Div., Lawas, 1955, Brooke 10283 (BM, L); 5th 


Volume 76, Number 1 
1989 


Bremer 29 
Argostemma in Borneo 


FIGURE 17. 


pis Bakelalan, 4, vs E 1955, Brooke 10448 E 


puer to U leh, 

7th Div., Bukit Tibang, 3,700 ft., 1969, Anderson $28735 
k Lx "th Div., Wong Pelagus, 1954, Brooke 8984 
(BM, K); 7th Div., Teneong, 1954, Brooke 9128 (BM, 
Ly 7th Div., Gilam Pasang, 1954, Brooke 9258 (BM, 
L); 7th Div., Ulu Melinau, 1980, Burtt 12896 (E); 7th 
Bukit Disa. 1978, Burtt 11338 (E, S); 7th Div., 


Liang Gagang, 1893- 1894, Hallier 2779 (Ly W Prov., 
Amai Ambit, 1893-1894, Hallier 3230 (BO, L); E Prov., 
between Long Bawan bu Pando, 1,200 m, 1891, Gee- 
sink 9036 (L); E Prov., between Papadi and Pamilau, 
700 m, 1981, Geesink 9226 (L); E Prov., Banyankara, 


Argostemma moultonii. —A. Flowering branch. 


—C. Anthers. — D. Style. 
A-E from Burtt & Martin 5326 (E); I from Chai S34832 (K); II from Modas 921 (Ly; III Es Mjoberg 82 
(K); IV from Moulton 6695 (SING). 


— B. Flo —E. Stipule. 


1982, Axelius 198 (Sy; E Prov., Nanukan Island, south- 


54, Kostermans 921 


1925, Endert 4172 
(A, BO, K, L); E Prov., Kutei, near Tabang, 50 m, 1955, 
pomis 10570 L. 


14. oe moultonii a ee Bot. 65: 
6. 1927. TYPE: Borneo. ong Semapok, 
Upper Baram, 3,000 ft., 1920, Moulton 6695 
(lectotype, K; Bakh. f. in herb., confirmed here; 
isolectotype, SINC). Figure 17. 


A. moultonii var. hirta Ridley, J. Bot. 65: 36. 1927. 
TYPE: Borneo: Gunong Semapok, Upper Baram, 
1,000 ft., 1920, Moulton 6654 (holotype, K; iso- 
type, SING). 


30 


Annals of the 
Missouri Botanical Garden 


A. parvulum Ridley, J. Bot. 65: 36. 1927. TYPE: Borneo: 
Lio Matu, Upper Baram, 500 ft., 1920, Moulton 
6726 (holotype, 

A. velutinum Ridley, J. Bot. 65: 35. 1927. TYPE: Borneo: 
Serapi, 2,800 ft., is Pa 684 (lectotype, K; Bakh. 
f. in herb., confirme 

A. mjoebergii Merr., iiobis i Mus. J. 3: 534. 1928. 
TYPE: Borneo. Sarawak: Mt. Poi, 4,500-5,500 ft., 
1924, Mjoberg s.n. e UC). 

A. murudense Merr., Sarawak Mus. J. 3: 535. 1928. 
TYPE: Borneo. Sarawak: Mt. Murud, 1,900-2,400 
m, 1922, Mjoberg 82 (holotype, UC; isotype, K). 


Creeping to erect, isophyllous herb. Stem 2-40 
cm, unbranched or slightly branched, glabrous to 
densely pubescent, rarely with corky or papery 
bark; internodes 5-60 mm. Leaves opposite, equal; 
stipules deciduous, 2-12 mm long, ovate to lan- 
ceolate, entire or slightly bifid, obtuse to acuminate 
at apex; petiole 3-11 mm long, glabrous to pu- 
bescent; lamina 1.2-6.5 x (0.2-)0.6-3.1 cm, lin- 
ear to broadly ovate, basally rounded to attenuate, 
with the lobes equal or oblique, marginally entire 
or serrulate, apically obtuse with a short point or 
acute or acuminate, herbaceous, pubescent on both 
surfaces to glabrous; midrib distinct, the primary 
veins (4-7 pairs) distinct to obscure. Inflorescences 
1 -9-flowered, umbelliform, generally solitary; pe- 
duncle (1-)3-8 cm, glabrous; bracts to 5 mm long, 
ovate to lanceolate; pedicels 0.2-1.2 cm, pubes- 
cent. Flowers 5(-6)-merous. Calyx lobes ca. 1 mm 
long, triangular to ovate, acute, pubescent at least 
in lower parts. Corolla 3-9(-18) mm long, cleft to 
near the base, generally glabrous on both surfaces; 
lobes lanceolate, spreading to reflexed. Stamens 3- 
6 mm long, coherent; anther cells and apical ap- 
pendages connate except for the most apical parts, 
these free and incurved, forming an open cylinder; 
anther cone ovoid, straight; filaments generally long, 

equal, straight or bent; sacs opening completely 
longitudinally; apical Appenduge ca. s io er "is 
anther length, thin and smoot 
generally smooth. Style 3.5- ^ 5 mm pia de. 
cent or rarely glabrous, with a capitate stigma, 
shortly exserted. Fruit without furrows or ribs. 


Bakhuizen f. (pers. comm. and on herbarium 
labels) treated 4. parvulum as a synonym of 4. 
hameliifolium; he believed it to be a dwarf form. 
However, I have found that 4. parvulum is better 
accommodated as a synonym of 4. moultonii. 

Argostemma moultonii grows in wet, damp 
places among mosses on tree trunks, on cliffs, or 
on sandy or clayey soil, and a few collections are 
from limestone. It grows from low altitude up to 
ca. 2,000 m; most specimens are from ca. 1,000 m. 

Ridley based A. moultonii on two collections by 
Moulton, in K. Both are suitable for lectotypifi- 


cation. Bakhuizen f. (in herb.) selected one of the 
specimens, Moulton 6695, and I follow his choice. 

Ridley based A. velutinum on three collections, 
in K, by Haviland, Moulton, and himself. All are 
suitable for lectotypification. Bakhuizen f. (in herb.) 
selected the Haviland specimen, and I follow his 
choice. 

Argostemma moultonii and A. hameliifolium 
form a complex in which a number of species have 
been recognized (see above). This complex is well 
defined as a group by unique characters: equal, 
opposite leaves; long, glabrous peduncles; pubes- 
cent pedicels; corolla cleft to near the base and 
with narrow, spreading or reflexed lobes; and con- 
nate anthers with distinct apical appendages which 
are free in the upper part and have margins longer 
than the central parts and incurved to form an 
open cylinder. There are at least two species (4. 
hameliifolium and A. moultonii) involved in the 
complex. | cannot recognize more in spite of the 
fact that this group is the most widely collected in 
Borneo. There are a few specimens similar in habit 
to the highly variable species 4. aequifolium Ridley 
(1901) from the Malay Peninsula and Sumatra, 
but this species does not have the unique stamen 
characters. Specimens similar to the type of A. 
hameliifolium, erect, large plants, are referred to 
A. hameliifolium, but the rest could be separated 
into several variable groups within A. moultonii. 
There would be many specimens that could not be 
placed with certainty. 

Argostemma moultonii needs further eed I 
have dealt with it in a manner similar to Dr. 
huizen f. (pers. comm.). Argostemma OIM 
generally can be characterized and separated from 
A. hameliifolium by having a smaller, creeping 
habit, generally longer internodes, smaller stipules, 
and anthers with long filaments and long apical 
appendages. Within 4. moultonii at least six local 
populations with distinct morphological characters 
can be identified. From some localities different 
populations are found without continuous variation 
between them. Characters separating these popu- 
lations are indumentum of the stems, leaves, and 
floral parts and the shape, texture, and nervation 
of the leaves. The following examples illustrate this. 
At the summit of Mt. Matang the plants (including 
the type of 4. velutinum) have ovate leaves with 
acuminate apices, distinct primary veins, and long 
petioles, and the plants are covered by long hairs. 
Close to that locality, at Lundu and Berumput, the 
collections (including the type of 4. mjoebergii) 
are similar in habit but less pubescent and have 
shorter petioles and less distinct veins. At Gunong 
Murud several different populations occur. One is 


Volume 76, Number 1 
1989 


Bremer 31 
Argostemma in Borneo 


D C 


0.2mm G , 

_05cm CD , 

ı Icm B; 2cmE j 

¿20cm A E 


FIGURE 18. 


almost glabrous (dry specimens) with very thick, 
veinless, elliptic, acuminate leaves and with very 
long internodes (Burtt & Martin 5326, Noote- 
boom & Chai 1935). Except for the very long 
internodes, they are similar to the type of A. par- 
vulum from Baram. Another population includes 
the type of A. murudense with small, broad, ovate 
to rounded, obtuse to acute, densely pubescent 
leaves. These specimens are similar to the type of 
A. moultonii from the upper Baram. Martin 38848 
represents a different population with glabrous, ob- 
ovate leaves with distinct prominent veins. From 
Ulu Tinjar in Baram, there is a collection (Chai 
34832) with glabrous linear leaves. From Mt. Kin- 
abalu, there are several different populations—one 
(Gardner 88) is glabrous with thick, veinless leaves 
but different from the glabrous specimens from 
Gunong Mulu. There are also specimens with ovate 
to obovate, slightly pubescent leaves. Plants from 
Kutei and Gunong Tapa Sia in Kalimantan (Endert 
3614, 4238 and Axelius 112B) are small rosettes 
with ovate, obtuse leaves having very long hairs. 
These collections are similar to Nooteboom 921 
from Gunong Alab in Sarawak. Perhaps these rep- 
resent something different and distinct from 4A. 
moultonii, but based on the scanty material avail- 
able, it is impossible to decide. 


Additional specimens examined. BORNEO. BRUNEI: 
Kuala Belalong, 1959, Ashton 199 (K). SABAH: W Coast 


Argostemma borragineum. — A. Flowering branch. — B. Flow 


r. — C. Anther 
le. —G. Hair from outer surface of the corolla. — C,. Hair from inner ab of the mo 


(S) 


s. — D. Style. — E. Stip- 
From Bremer 1668 


Res., Kinabalu, Silau Silau trail, 1,400 m, 1979, An- 
derson 4349 (MO, UC); Bebangan River, 5,000 ft., 1964, 
Chew & Corner RSNB4618 (K); Tenompok, 5,000 ft., 


1933, Clemens 51670 (B 
ai Parai, 6,000 ft., 1958, Collenette 60 (BM); 1937, 
Grieswold 27 (GH); Lumu-Lumu, 1957, Sinclair 9212 
E, L); W Coast Res., Gunong Alab, Crocker range, 1,600 
m, 1980, Argent 1308 (E); 1,400 m, 1969, Nooteboom 
921 (B, L); 1,600 m, 1969, i aie. t 1035 (L); San- 
dakan Res., Tawau Plateau, 50 m, 77, Gardner 88 
(E, D); Interit Res., Sipitang, Ulu de ah, 2,750 ft., 

Wood SAN16695 (L, SING). SARAWAK: lst Div., Mt. 


6,000-8,000 ft., 


, 1929, Clemens 223221 (NY); 2,500- 

3,000 ft., Mjóberg s.n. (UC); 1927, native collector s.n. 
Le 1905, Ridley 12304 (BM, K, SING); 1st Div., 
Santubong, near summit, 1909, Brooke 1034 (BM); 1st 
Div., Gunong Gading, 2,800 ft., 1962, Burtt & Woods 
2682 (E) lst Div., Mt. Poi, 5,000-6,000 ft., 1929, 


Gunong Berumput, 4,800 ft., 
L); 300 ft., 1962, Burtt & Woods 2761 (E); 3,500 ft., 
1962, Burtt & Woods 2845 (E); 2nd Div., E of Bukit 
Sadon, 1982, Axelius 112B (S); 4th Div., Mt. Dulit, Dulit 
1932, Richards 1537 (K, L); Dulit ridge, 


US); 4th Div., Mt. Murud, 1,900- 2,400 ft., 1922, Mjo- 
berg 85 (UC); 1,900-2,400 ft., 1922, Mjöberg 88 (NY, 
UC); 1914, Moulton 105 (probably cited as 165 in the 


32 


Annals of the 
Missouri Botanical Garden 


rotologue of 4. velutinum) (SING); 1,100 m, 1970, 
Monos & Chai 1856 (L, SAR, US); 1,700 m, Noote- 
boom & Chai 1935 (B, L, US); 4th Div., Sungei Melinau, 
1962, Burtt & Woods 2293 (E); 4th Din. Gunong Mulu, 
4,200 ft., Anderson S4516 (K, SAR); 1,700 m, Argent 
& Jermy 1038 (E), 1,300 m, 1978, Argent & Coppins 
1076 (Ey 1962, Burtt & Woods 2090 (E); 4,500 ft., 
1962, Burtt & Woods 2095 (E, SAR); 4,500 ft., 1962, 
Burtt & di 2152 (E), 150 m, 1978, Hansen 187 
Uh 1,300 m, 1977, Martin S38848 (L, SAR); 1,800 

, 1978, Nielsen 838 (AAU); 5th Div., 
Maputi, 2,500 f 
Bakelalan, 4,000 ft., 1955, Brooke 10408 (L); 5th Div., M 
from Bakelalan to Gunong Murud, 6,000 ft., 1967, Burtt 
& — 5326 (E); 7th Div., Bukit Bakar (= ?Bakak), 
980 m, 1975, Ilias Paie $36357 (L); 7th Div., Sungei 
Nai Punan Bah, 1973, Chai et al. S33328 (L, MO); 7th 
Div., e Punan Bah, 1979, Tong & Banyeng 833271 
(K, L, MO, SAR); 7th Div., Sungei Brearan, 1978, Burtt 
11355 n 'S, 11356 (E); Tth Div., Sungei Dema-nawai, 
1978, Burtt 11431 (E), 7th Div., Sungei Tellini, 2,800 
ft., 1978, Burtt 11401 (E); 2,850 ft., 1978, Burtt 11409 
(E, S); 7th Div., S Hose Mts., Bukit Salong, 1980, Burtt 
12709, 12908 (E), 950 m, 1976, Chai S37296 (K, L, 

AR); 7th Div., S Hose Mts., E of Bukit Sanpandai, 1980, 
Burtt 12797 (E ); 7th Div., Ulu Mujong, 200 m, 1964, 
Ashton $12132 (K, L); 900 m, 1964, Ashton 521238 
m, 1964, Ashton S21249 (A, L); 7th 
, 1969, Ilias Paie 528445 
(K, B, KALIMANTAN: W Po. 


,600 m, 1925, Endert 4238 (BO, L); E Prov., 
a Papadi a nd Pamilau, 1,200 m, 1981, Gansink 
9255 (L); E Prov., Gunong Tapa Sia, 1,800 m, 1981, 
Geesink 9112 (L). 


15. Argostemma borragineum Blume ex DC., 
rod. 4: 417. 1830. TYPE: Blume in G-DC 
(holotype, G-DC, not seen; microfiche, LD!). 
igure 1 
A. borragineum var. rotundifolium Valeton in H. Wink- 
ler, Bot. Jahrb. 44: 548. 1910. TYPE: S.O. Borneo: 
Batu Babi in Lumowia, 1908, Winkler 2853 (lec- 
totype, L; Bakh. f. in herb., confirmed here; isolec- 
totypes, BM, BO). 
A. e dns Valeton, Icones Bogorienses 4: 27 
914. TYPE: Borneo: Oeloe Tjihan, 1898-99, n 
djah 319 (lectotype, BO; Bakh. f. in herb., 
firmed here). 
A. nutans var. borneense Ridley, J. Bot. 65: 33. 1927. 
TYPE: Borneo: ys unong Semapok (**Temapok" Rid- 
ley, 1927, sphalm.), 1,000 ft., 1920, Moulton 6653 
(holotype, K; b SING). 


Large, erect, isophyllous herb. Stem to 100 cm, 
glabrous to finely pubescent, black, with distinct 
scars in upper part; internodes 5-45 mm. Leaves 
opposite, equal; stipules persistent, ca. 10 mm long, 
broadly ovate, obtuse to acute at apex; petiole 0.5— 
7 cm, pubescent to glabrous; lamina 4.5-21 x 
2.5-13 cm, broadly ovate to broadly obovate, ba- 
sally rounded or acute to attenuate with the lobes 


equal, marginally entire or serrulate, apically ob- 
tuse, then sometimes with a point, or acute, her- 
baceous, glabrous to pubescent, most prominently 
pubescent on the veins below; midrib and primary 
veins (4-12 pairs) distinct. Inflorescences 25-160- 
2 or 3 scorpioid 
branches, solitary or a few together; peduncle 2- 
8 cm, glabrous to pubescent; bracts to 1 cm long, 
oblong to ovate; branches, if present, to 5 cm, 
glabrous to pubescent; pedicels to 2 cm, pubescent. 
Flowers 5-merous; calyx lobes 1 -2 mm long, ovate 
to triangular or lanceolate, pubescent at least on 
basal parts; corolla m long, cleft to about 
middle, externally pubescent, internally glabrous 
to pubescent on the lobes; lobes ovate to triangular, 
recurved. Stamens 2-4 mm long, coherent, cori- 
aceous; anther cells and apical appendages connate 
except for the most apical part; anther cone ovoid 
or shortly so, straight; filaments very short, equal, 
straight; sacs opening completely longitudinally; 
apical appendage ca. the anther length, thin and 
smooth; connective indistinctly passing over to the 


flowered, umbelliform or wit 


thecae, smooth. 
with a capitate stigma, long-exserted. Fruit without 
furrows or ribs but covered with emergences. 


-6 mm long, pubescent, 


Winkler based A. borragineum var. rotundi- 
folium on one collection with several specimens. 
All specimens are suitable for lectotypification. 
Bakhuizen f. (in herb.) selected the specimen in L 
and I follow his choice. 

Valeton did not indicate any collection in the 
protologue of A. streblosifolium. In the Bogor her- 
barium (BO) there is only one specimen (Amdjah 
319) that fits the picture and the protologue. Bak- 
huizen f. (in herb.) has selected that specimen, and 
I follow his choice. 

Argostemma borragineum is one of the most 
common Argostemma species from low altitudes 
to ca. 2,000 m. It often grows near shady streams. 
All notes concerning its habitat place it on sandy, 
loamy soil, sandstone, or sedimentary rocks. It is 
highly variable. The majority of the plants are 
large, to 1 m high, with black and scarred stems. 
The opposite leaves are equal, large, and rounded, 
and the stipules are broad and distinctive. The 
inflorescences are densely pubescent and generally 
very large, sometimes with more than 150 flowers. 
They commonly develop two or three scorpioid 
branches, which become most prominent after an- 
thesis. The corollas are cleft for about half their 
length into recurved lobes. The stamens are unique. 
The entire stamen is coriaceous and (on dry spec- 
imens) dark, and no differences can be discerned 
between the connective and the thecae, at least 
not from the dorsal side. The apical appendages 


Volume 76, Number 1 


Bremer 33 


1989 Argostemma in Borneo 
eta ea 
A 
Ad T3 
Sy RN 
1 5 5 ^ 4 d EF g 1 6 
RC rff ees 
100 km Ra O ba d. 00 km 
¿EE > 500 m #% > 500 m 


Mars 15, 16. 


are separated in a characteristic way. The style is 
generally densely pubescent and bears a distinct 
capitate stigma strongly exserted. There are also 
plants uniformly smaller in all organs except for 
the flower, with smooth stems and more densely 
pubescent inflorescences, but there are also other 
small plants which have scarred stems. Several 
different populations have been collected from Mt. 
Kinabalu. I have not found any correlation between 
the morphological variation and any pattern of 
distribution. 


Additional specimens examined. BORNEO. SABAH: 
oast Res., Gunong Alab, 1,500 m, 1969, Nooteboom 
1031 (B, L, US); W Coast Res., Kinabalu, below Kam- 
barangan, 6,000 ft., 1962, Bogle 545 (GH); Kiau, 1915, 
10058 (BO, UC); Penibukan, 4,000-5,000 ft., 
M); ridge above Pina Taki, 4, 000- 
1933, y a. CH M, BO, GH, NY); 
Marai DO spur, 5,000 ft., Clemens 32962 (A, BO); 
Marai Parai, Gigisan Creek, 4, 500 ft., 27 Mar. 1933, 
Clemens 33962 (BM, G, L, UC); headwiter of Sadikan, 
5,000 ft., 5 May 1933, Clemens 33962 (NY, must be 


US); between 
u, M ft., 1954, ho — 
, Sungei Lundu w Gun 
Perigi, 1962, Burtt & Woods 2710 (E, S lst Div. 
Mt. Matang, near summit, 2,300 ft., 1979, Bremer 1720 


1913, Moulton's native collectors 
365 (BM); 2,000 ft., 1960, Sinclair 10355 (E); 1st Div., 
Kampong Grumbing, Padawan, 30 m, 1975, Othman 
Ismawi S37437 (L, MO, SAR); 1st Div., Mt. Penrissen, 
4,400 ft., 1924, Mjoberg 218 (NY, UC): Ist Div., Sabal, 
500 m, 1979, Bremer 1668 (S, SAR); above the lorena 


Mjoberg s.n. (UC); 


Distributions in Borneo. — 15. Argostemma borragineum. —16. A. flavescens. 


office, 1979, Bremer 1673 (S, SAR); 4th Div., Bukit 
Mersing, 3,000 ft., 1956, Purseglove 5282 (A, K, L, 
SAR, SING); 5th Div. ., route from Bakelalan to Gunong 
Murud, 3,800 ft., 1967, Burtt & Martin 5216 (E); 7th 
Div., Sungei Boda area, 1980, Burtt 13003 (E); 7th Div., 
Bukit Goram, 1975, Chai S36137 (L, MO); 7th Div., 
junction of Sungei Tekalit and Sungei Mengiong, 1979, 
Bremer 1742, 1743 (S, SAR); 7th Div., Ulu Sungei 
Sadampa, 1,500 ft., 1969, Anderson & Ilias Paie $28295 
(E, L); 7th Div., foothills of Bukit Batu Tibang, 3,100 
ft., 1969, pue 528396 (A, K, L); 7th Div., Sungei 
Balang, 1969, Anderson & p Paie S2888 1 (A, E, K, 
L, SAR). KALIMANTAN: E Prov., Balikpapan District, Men- 
tawir region, 1955, Pa oiro 10121 (L); S of Gunong 
Balikpapan, 1985, Axelius 217 (S); E Prov., W Kutei, 
near Laih Leng, 200 m, 1925, Endert 2952 (BO, L); W 
Prov., Bukit Raja, 1,250 m, 1924, Winkler 879 (HBG, 
NY) 1,400 m, 1924, Winkler 1007 (HBG, NY), W 
Prov., Sungei Malang, 70 m, 1925, Winkler 1282 (HBG, 
NY); W Prov., Bukit Obat, 150 m, Winkler 1353 (HBG); 


Prov., Nanga Era, 100 m, 1925, Winkler 1562 
(HBG); W Prov., Liang Gagang, 1893-1894, Hallier 
2686 (BO, L). 

._2cmEF 
¿10cm A 
rae 19. Argostemma flavescens. — A. Flowering 


—E. Stipule. —F. Nan 


bra erudi leaf. From Ha- 
d 84/75 (SING). 


Annals of the 
Missouri Botanical Garden 


y 
D 
_0.2mmG ] 
05cm CD j 
1cm B; 2cmEF 
| 10cm A 
FIGURE 20.  Argostemma  chaii. —A. Habit. — B. 
Flower. — C. Anthers. — D. Style. — E. Stipule. —G. Hair 


e 
from outer surface of the sella: From Brooke 10268 
(L) 


16. Argostemma flavescens Bakh. f., Blumea 
7: 331. 1953. TYPE: Borneo. Path to Matang, 
Haviland 84/75 (holotype, SING). Figure 19. 


Creeping, anisophyllous herb. Stem ca. 10 cm, 
branched, pubescent; internodes 4-8 mm. Leaves 
opposite, very unequal; stipules persistent, to 3 
mm long, ovate to cordiform, acute at apex; larger 
leaves of the anisophyllous pairs with pubescent 
petioles 3-8 mm long; lamina 1-3 x 0. -1.5 em, 
obovate to ovate or elliptic, basally obtuse to acute 
and oblique, marginally entire to serrulate, apically 
acute, herbaceous, pubescent between the veins; 
midrib and primary veins (5-7 pairs) distinct. Na- 
nophyllous leaves persistent, ca. 3 mm long, ovate. 
Inflorescences 1-3-flowered, solitary; peduncle 1- 
2 cm, pubescent; bracts to 0.3 cm, lanceolate; 
pedicels 0.5-0.8 cm, pubescent. Flowers 5-mer- 
ous; calyx lobes ca. 1 mm long, broadly ovate, 
acute, pubescent in lower part; corolla 4 mm long. 
Stamens 5. Style pubescent, with an elliptic stigma, 
long-exserted. 


Argostemma flavescens is known from only two 
very poor collections. Both were examined, but 
there is still a great deal of information missing 
about the species. It has a characteristic habit, and 
its leaves are densely pubescent, particularly be- 
tween the veins. In all other Borneo species the 
indumentum is more dense on the veins. The in- 
formation on the flowers in the description is taken 


from Bakhuizen f. (1953, and pers. comm.). He 
placed A. flavescens close to A. borragineum be- 
cause of its stamens. In my opinion it is not possible 
to determine its closest relatives within Argostem- 
ma at this time. 


Additional specimen examined. BORNEO. SARAWAK: 
1st Div., Mt. Matang, Beccari 621 (L). 


17. Argostemma chaii B. Bremer, nom. nov. 
Argostemma enerve Ridley (non Ridley 1912), 
nom. illeg., J. Bot. 65: 28. 1927. TYPE: Bor- 
neo. Sarawak: native collector 1227 (holo- 
type, K). Figure 20. 


Small, erect, isophyllous herb. Stem very short 
to 11 cm, unbranched or slightly branched, gla- 
brous; internodes very short to 10 mm long. Leaves 
opposite, subequal; stipules persistent, ca. 1 mm 
long, broadly ovate to cordiform, apically fringed 
with each point ending in a gland; petiole 3-9 mm 
long, glabrous; lamina 1.2-7 x 0.8-4.5 cm, ovate 
or elliptic or rarely lanceolate, basally acute to 
attenuate with the lobes equal, marginally entire, 
apically acute to obtuse, herbaceous to coriaceous, 
glabrous or with a few hairs above; midrib distinct, 
the primary veins obscure or absent. Inflorescences 
2-15(-25)-flowered, umbelliform, solitary or a few 
together; peduncle 2-1 glabrous; bracts to 
0.6 cm long, ovate to lanceolate; pedicels 0.3-1.2 
cm, pubescent or glabrous. Flowers 5-merous; ca- 
lyx lobes 0.5-1 mm long, ovate to triangular, acute, 
glabrous to pubescent; corolla 3-5 mm long, cleft 
to near the base, externally pubescent or glabrous, 
internally glabrous; lobes lanceolate to oblong, erect- 
spreading. Stamens 3-4 mm long, at anthesis prob- 
ably free; anther cells free, the apical appendage 
first connate then free, not forming a cone; fila- 
ments ca. 4% as long as the anthers, equal, straight; 
sacs opening completely longitudinally; apical ap- 
pendages very short, thick and smooth; connective 
distinct, smooth. Style ca. 3-6 mm long, glabrous, 
with a discoid stigma, long-exserted. Fruit without 
furrows or ribs. 


This species is named after Mr. Paul Chai of 
the Forest Department, Sarawak. Argostemma 
chaii grows on dead wood, wet rocks, or wet ground 
in the lowland rainforest, to 300 m. This species 
is a very small herb, with leaves more or less 
condensed into rosettes. Its primary veins are ob- 
scure, and the stipules are broad and fringed api- 
cally with colleters. The flowers are characteristic, 
with thick corolla lobes and free anthers. The fil- 
aments are wide, fixed at the base of the corolla, 
and adnate with it for a short distance. The stigma 
is discoid. 


Volume 76, Number 1 
1989 


Bremer 35 
Argostemma in Borneo 


Distributions in Borneo. — 17. 


Mars 17, 18. 


Argostemma chaii is a rare species collected 
from two different areas in Sarawak. The collec- 
tions from the 5th Division are hairy on the ped- 
icels, calyx, and on the outside of the corolla. The 
leaves also have scattered hairs. From the lst 
Division the plants (including the type specimen) 
are glabrous in all parts. From Mt. Matang in the 
lst Division the leaves are distinctly larger than 
from other localities. 

Bakhuizen f. (pers. comm.) has treated this taxon 
as a synonym of A. inaequale (Bennett, 1838). 
Their habits are similar, but their flowers are dif- 
ferent. Argostemma inaequale does not have the 
thick corollas nor the discoid stigma, and its anthers 
have long apical appendages, while the anther ap- 
pendages of A. chaii are very short. 


PEREA specimens examined. BORNEO. SARA- 
WAK: 1st Div., Bako, Telok Asam, 100 ft., 1956, Purse- 
b ose 5096 (SING); lst Div., Mt. Matang, 800 ft., 1954, 


Brooke 9474 (L), 9478 (BM, L); 1st Div., Krangi, 1954, 
To 8502, 8512 (BM, L); 4th Div., G. Mulu, 120 

1978, Nielsen 368 (AAU); Sth Div., Lawas, 1955, 
Boake 10542, 10268 (BM, L). 


18. Argostemma gracile Stapf, Trans. Linn. 
Soc. London Bot. 4: 168. 1894. Argostemella 
gracilis (Stapf) Ridley, J. Bot. 65: 41. 1927. 
TYPE: Borneo. Kinabalu, Penokok, 3,500 ft., 
Haviland 1325 (holotype, K; isotypes, SAR, 
SING). Figure 21. 


A. ub ERE Merrill, Mitt. Inst. Allg. Bot. Hamburg 
279. 1937. Type: West Borneo: Bukit Tilung, 


Argostemma chaii. —18. A. gracile. 


750 m, 1925, Winkler 1479 (holotype, HBG; iso- 
type, NY) 

Small, creeping, anisophyllous herb. Stem 5-20 

cm, slightly branched or unbranched, glabrous; 


( 
CA 
Wr 


_0.2mmG j 
0.5cm CD 


oa | 

v Icm B; 2cmEF 

, 10cm $: 

FIGUR Argostemma gracile. —A. Habit.—B. 


Flower, inner surface of calyx.—B,. Inside view of part 
ll 


f rolla. A from Bremer 1667 
5), B-G from Argent & Connie 1142 (E). 


36 


Annals of the 
Missouri Botanical Garden 


i CD ; 
u Icm B; 2cmEF , 


, 10cm A LL 
FIGURE 22. Lo did lea cire, —À 
Habit.—B. Flower.— C. —D. Style. — E. Stip- 


ule. — F. Nanophyllous af. 6. Hair from outer surface 
of the corolla. A from Bogle et al. 389 (GH); B-G from 
Hou 225 (L) 


internodes 2-5 mm. Leaves opposite, very un- 
equal, distichous, pale silvery underneath; stipules 
persistent, ca. 2 mm long, broadly ovate or cor- 
diform to auriculiform, ending apically in a distinct 
point; stipules and petioles + fused into an am- 
plexicaul sheath. Larger leaves of the anisophyllous 
pairs with pubescent petioles 0.5-1.5 mm long; 
lamina 0.6-1.7 x 0.3-0.7 cm, oblong to narrowly 
oblong or rarely obovate, basally rounded or acute 
with the lobes equal or oblique, marginally entire, 
apically obtuse or obtuse with a point, or acute, 
herbaceous to coriaceous, glabrous or with scat- 
tered hairs somewhat concentrated along the mid- 
rib on both sides; midrib distinct but without pri- 
mary veins. Nanophyllous leaves persistent, 4-5 
mm long, ovate. Inflorescences l-flowered, soli- 
tary, without peduncle; pedicel 0.2 cm, pubescent. 
Flowers 5(or 6)-merous; calyx lobes 2-3 mm long, 
triangular to cordiform, pubescent in lower parts; 
corolla ca. 8 mm long, cleft less than Y its length, 
externally glabrous and internally pubescent only 
along the margins of the lobes; lobes broadly tri- 
angular, spreading. Stamens ca. 2.5 mm long, free; 
anther cells and apical appendage free, not forming 
a cone; filaments ca. 14-14 as long as the anthers, 
equal, straight; sacs opening completely longitu- 
dinally; apical appendages almost absent, thick and 


slightly papillose; tive distinct, smooth. Style 
ca. 5 mm long, alabrona, with a discáid, bifid stig- 


ma, long-exserted. Fruit without furrows or ribs. 


Argostemma gracile grows on fallen rotten logs 
or on rocks among mosses in submontane or mon- 
tane forests from 1,000 to 1,600 m. One collection 
(Bremer 1667) is from low elevation, 350 m, in 
an area of lowland rainforest with big boulders 
covered by mosses. 

Argostemma gracile is easily recognized, with 
its creeping habit and distichous, anisophyllous 
leaves, the nanophyllous leaves above the larger 
leaves. The leaves are thick with a distinct midrib 
but without other visible veins. They are silvery 
white beneath. The petiole and midrib have a few 
hairs on both surfaces, but one exception is the 
type specimen of A. podochiloides with completely 
glabrous leaves. The stipules and petioles are more 
or less connate with the amplexicaul sheath. This 
character, combined with the glabrous leaves, has 
been used for recogniton of 4. podochiloides, a 
species not accepted here. Flowers in A. gracile 
are solitary on very short pedicels and without 
peduncles. The corolla is campanulate with spread- 
ing lobes. The stamens are free, and there are 
practically no apical appendages. The stigma is 
distinctly bifid and discoid. 


Additional specimens examined. BORNEO. SABAH: 
W Coast Res., SPEO Ulu Langanani, 3,800 ft., 1961, 
Chew et al. 1713 (A, K, L, SAR, SING); Tenompok, 
5,000 ft., 1932, Luis 269264 (BM, BO, K, L, NY); 
Cohunbon River, 4,500 ft., 1933, Clemens 34110 (BM, 


BO, GH, L, NY, UC); Penibukan. 4, Vis ft., 1933, Clem- 
ens 32113 (B, BO, BM, C, GH, K, N Y, UC); Penibukan, 
4,000-5,000 ft., 1932, Clemens s.n. (BM NY); Marai 


Parai spur, 1915, Clemens 11076 (UC). SARAWAK: lst 
Div., Sabal, 350 m, 1979, Bremer 1667 (S, SAR); 2nd 
Div., Lubok Antu District, near Bukit Sengkajang, 1974, 
Chai S34006 (K, L, SAR); 4th Div., Gunong Mulu, W 
ridge near cam 
1142 (E); near camp 4, 4,000 ft., 
2176 (E, SAR); 1,180 m, 1976, Martin S38185 (L, 
SAR); 4th Div., Mt. Dulit ridge, 1,200 m, 1932 Richards 
1689 (K, L); S slope, 1,800 m, Nielsen 833 (AAU); Sth 
Div., Bakelalan, 4,000 ft., 1955, Brooke 10461 (BM, 
L); 5th Div., route from Bal 
ft., 1967, Burtt & Tee 
District, 2,900 ft., 1 


(E) camp 6, 4,600 ft., 1980, Burtt 12848 (E); 7th Div., 
summit ridge of Bukit Tibang, 5, d ft., 1069 po 
& Ilias Paie S28667 (KLU, SAR). KALIMANTAN: 


Prov., W Kutei, near Tabang, 600 m, 1956, "adde 
12851 (L). 


19. Argostemma brachyantherum Stapf, 
Trans. Linn. Soc. London Bot. 4: 168. 1894. 
Argostemmella brachyanthera (Stapf) Rid- 
ley, J. Bot. 65: 41. 1927. TYPE: Borneo. Kin- 


Volume 76, Number 1 
1989 


Bremer 37 
Argostemma in Borneo 


"ta 
af 
; 


A 
19 t 
— 100 km 
Sk > 500 m 


MaPs 19, 20. 


abalu, 3,000 ft., 
K). Figure 22. 


Haviland 1326 (holotype, 


A. Rinabatuense Wernham in Gibbs, J. Linn. Soc. Bot. 
8. 1914. TYPE: British North Borneo. Kinabalu, 

near Dahobang and Kinitaki rivers, 4,000-5,000 

ft., 1910?, Gibbs 4056 (holotype, BM; isotype, K). 


Erect, anisophyllous herb. Stem 3-40(-80) cm, 
slightly branched or unbranched, densely pubes- 
cent in upper part, in lower p with corky or 
papery bark; internodes 2-20(-45) mm. Leaves 
opposite, very unequal; stipules deciduous or per- 
sistent, 4-10 mm long, oblong to lanceolate, acute 
to acuminate at apex; larger leaves of the aniso- 
phyllous pairs with pubescent petioles 2-5 mm 
long; lamina 2.5-9 x 1-3 cm, obovate or oblan- 
ceolate or elliptic to narrowly elliptic, basally acute 
to obtuse with the lobes equal or oblique, marginally 
entire or serrulate, apically acute to acuminate, 
herbaceous, glabrous or with scattered hairs above, 
pubescent on the veins below; midrib and primar 
veins (6-14 pairs) distinct. Nanophyllous leaves 
deciduous or persistent, 3-10 mm long, ovate to 
lanceolate, acute to acuminate. Inflorescences l- 

(-12)-flowered, umbelliform, solitary or a few to- 
gether; peduncle 1-3 cm, pubescent; bracts 0.2- 
0.7 cm long, ovate to lanceolate; pedicels 0.5-1.5 
cm long, pubescent. Flowers 5-merous; calyx lobes 
2-5 mm long, ovate to triangular, acute, sparsely 
pubescent; corolla campanulate, cleft less than 43 
their length, externally pubescent, internally pa- 
pillose to pubescent only on the lobes; lobes tri- 
angular, acute, erect(?). Stamens 2-3(-4) mm long, 


Distributions in Borneo. — 19. Argostemma brachyantherum. — 20. A. brookei. 


free; anther cells and apical appendages free, not 
orming a cone; filaments short, equal, straight; 
sacs opening completely longitudinally; apical ap- 
pendage /,—Y/ the anther length, thick and slightly 
papillose; connective distinct, smooth. Style 3-6 
mm long, glabrous, with a capitate stigma, long- 
exserted. Fruit without furrows or rib 


Argostemma brachyantherum grows in sub- 
montane or montane forest between 1,000 and 
2,100 m in northeastern Borneo. 

This species is generally small to medium sized 
with anisophyllous leaves and a few large, cam- 
panulate flowers with free, very short anthers. The 
indumentum, particularly on the leaves, differs 
among the collections, otherwise the species is rath- 
er uniform. The specimen Argent & Coppins 1081 
is much larger than those of other collections and 
also has many more flowers. Some characters of 
this specimen deviate from the typical range for 
the species and are enclosed in parentheses in the 
description. 


Additional specimens examined. BORNEO. SABAH: 
W Coast Res., Kinabalu, vicinity of Kambaranga, 7,040 
re 1962, Bogle et al. 389 (GH); Tenompok, 5,000 ft., 

, Clem 28179 (BM, BO, G, K, L, NY); Marai 
he 5, 000 p$ 1933, Clemens 35136 (BM, BO, L, 
NY, UC); Marai Pata spur, 1915, Clemens 10981 (UC); 
Upper Kinabalu, 6,000 ft., 1933, Clemens 51020 (BM, 
G, K, NY); Marai Parai, 5,000 ft., 1958, Collenette A77 
(BM); on ridge parallel to Eua River, 45 ft., Colle- 
nette A103 (BM); 1,650 m, 1966, Ding Hou 225 (K, 
Ly Kamarangoh, 7,000 ft., 1967, Price 194 (K) W 
Coast Res., Gunong Alab, 1,400 m, 1969, Nooteboom 


38 


Annals of the 
Missouri Botanical Garden 


p 


¿0.2mm G ' 
05cm CD i 
ı Icm B; 2cmEF 
_20cm A — 


FIGURE 23. Argostemma brookei.—A. Habit.—B. 
Flower. — C. Anthers. — D. Style. — E. Stipule. — F. Nano- 
phyllous leaf. —G. Hair from outer surface of the corol- 
la. —G,. Hair from inner surface of the corolla. A from 


Brooke 8562 (G); B-G from Clemens 20064 (NY). 


922 (L). SARAWAK: 4th Div., Gunong Mulu, west ridge, 
1,800 m, 1978, Argent & Coppins 1081 (E); ridge at 
about 5,000 ft., 1962, Burtt & Woods 2150 (E); 4,000 
ft., 1962, Burtt & Woods 2178, 2187 (E); path from 
Milinau Paku, 4,200 ft., 1961, Anderson 4516 (K, L). 
KALIMANTAN: E Prov., W Kutei, 1925, Endert 4171 (A, 
BO, K, L 

20. Argostemma brookei B. Bremer, sp. nov. 
TYPE: Borneo. Sarawak: 1st Div., Berumput, 
4,800 ft., 1954, Brooke 8562 (holotype, L; 
isotypes, BM, G, UC). Figure 23. 


Herba inflorescentiis valde pubescentibus, staminibus 
liberis. Folia opposita, illis parium inaequalissimis. Inflo- 
rescentia umbelliformis, floribus pusillis. Lobi calycis ovati 
ad triangulares, acuti, ep Corolla profunde fissa. 
Lobi corollae recurvi. Stamina libera, sed approximata. 
Appendix apicalis V4 bu die antherae aequans, cras- 
sa, laevis. Connectivum distinctum. Stylus glaber, stigmate 
leviter capitato. 

rgostemma havilandii, A. calcicolum, A. trichosan- 
thes, A. humifusum, A. psychotrioides, Á. variegatum 
simile sed lobis corollae profunde fissis, staminibus liberis 
differt 


Erect, anisophyllous herb. Stem 5-22 cm, 
slightly branched or unbranched, finely pubescent; 
internodes 3-15 mm. Leaves opposite, generally 
very unequal; stipules deciduous or persistent, 4— 
6 mm long, ovate, acuminate at apex; larger leaves 
of the anisophyllous pairs with pubescent petioles 
2-35 mm long; lamina 3-70 x 0.8-2.2 cm, ob- 


lanceolate to narrowly elliptic, basally acute with 
the lobes equal, marginally entire, apically acu- 
minate, rarely acute or obtuse, herbaceous, gla- 
brous above, pubescent on the veins below; midrib 
and primary veins (6-10 pairs) distinct. Nano- 
phyllous leaves deciduous or persistent, 4-10 mm 
long, ovate to lanceolate, acute to acuminate. In- 
florescences 5-16-flowered, umbelliform, solitary 
or a few together; peduncle 0.8-1.7 cm, pubes- 
cent; bracts ca. 0.5 cm long, lanceolate; pedicels 
0.6-1.0 cm, pubescent. Flowers 5-merous; calyx 
lobes ca. 2 mm long, ovate to triangular, acute, 
sparsely pubescent, spreading; corolla cleft to near 
the base, externally with a few hairs, internally 
pubescent only on the distal part of the lobes; lobes 
ovate to triangular, recurved. Stamens ca. 3 mm 
long, free but close together; anther cells and apical 
appendage free; no true cone; filaments short, equal, 
straight; sacs opening completely longitudinally; 
apical appendage ca. 1⁄4 the anther length, thick 
and smooth; connective distinct, smooth. Style ca. 
6 mm long, glabrous, with a slightly capitate stig- 
ma, long-exserted. Fruit without furrows or ribs. 


Argostemma brookei has been collected from 
the ground in the forest in the Poi (= Pueh) range 
at elevations of 1,000 to 2,000 m. 

It is medium sized with oblanceolate, aniso- 
phyllous leaves. The upper part of the stems, the 
peduncles, and the whole inflorescences are cov- 
ered by rather stiff hairs. The length of the calyx 
lobes is intermediate between the short ones, most 
common among the Borneo species, and the long 
lobes of A. psychotrioides. 

Most specimens of this taxon have been deter- 
mined as A. brachyantherum, another mountain 
species, which occurs mainly at Kinabalu. In habit 
they are similar, but Argostemma brookei has 
many small flowers with deeply cleft corollas having 
evidently recurved lobes. The free anthers are long- 
er and stand close together, like a cone; the apical 
appendages are much longer, and the filaments are 
shorter than in A. brachyantherum. 


e ional Vague examined. NEO. SARA- 

: 1st Div., eh), summ ae northern 

Mig 4, 400 ft., 1956, Bell 2068 (BM); 6. 000 ft., 1929, 
Pian 2006 NY); forested slopes, 1929, Cle emens 


20331 e as 500. 5,500 ft., 1924, Mjöberg s n. (UC); 
t Div., g Berumput, 3,000 1954, Brooke 
HEP (Ly 4 870 "e 1962, Burtt & Woods 2800 (E). 


21. Argostemma havilandii Ridley, J. Straits 
Branch Roy. Asiat. Soc. 61: 13. 1912. TYPE: 
Borneo: Kuching, 1893, Haviland 2958 (ho- 
lotype, SING; isotypes, BM, BO, K). Figure 
24 


Volume 76, Number 1 
1989 


Bremer 39 
Argostemma in Borneo 


|, 0. 2mm G y 
.0.5cm CD A 
._1cm B; 2cmEFH 
| 20cm A 


—" 


FicunE 24. E havilandii. —A. Habit. 
B. Flower.—C. Anthers.— D. Style. —E. Stipule. E 
Nanophyllous gs Hair from outer surface of the 
corolla. — G,. Hair from inner surface of the corolla. — 
H. Inflorescences. A, E, F from Bremer 1689 (S); C, D, 
G, H from Bremer 1662 (S). 


21 


1 
Sk > 500m 


Mars 21, 22. 


A. hallieri Valeton, m Bogorienses 4: 267. 1914. 
TYPE: Borneo: Àmai Ambit, Hallier B3228 (holo- 
e, BO; isotypes, “BO. L). 
A. lanceolatum Valeton Icones Bogorienses 4: 269. 1914. 
E: Borneo: Teysmann 11292 (lectotype, L; Bakh. 

[i in herb., confirmed here). 
A. sareta W. Smith, Notes Roy. Bot. Gard. Edin- 
burgh 8: 318. 1915. TYPE: Borneo. Sarawak: 1913, 
ative collector 85 (holotype, E). 
A. hallieri var. Babi ue Merr., Mitt. e Allg. Bot. 
279. 1937. TYPE: West Borneo: Sungei 
Bika, 50 m, 1925, Winkler 1428 Tert HBG; 

isotypes, HBG, NY). 


Erect, anisophyllous herb. Stem 10-60 cm, un- 
branched or slightly branched, glabrous or rarely 
slightly pubescent, lower and older parts pale brown 
with elevated corky to papery longitudinal ridges 
and/or with transverse bars; internodes 9-30 mm 
Leaves opposite, very unequal; stipules deciduous, 
4-7 mm long, ovate to broadly ovate, obtuse at 
apex. Larger leaves of the anisophyllous pairs with 
glabrous or finely pubescent petioles 4-8 mm long; 
lamina 4-1 —5 cm, obovate or oblanceolate 
to ovate to lanceolate, basally cuneate to acute or 
attenuate with the lobes equal, marginally entire, 
apically acuminate or rarely acute to obtuse, her- 
baceous to coriaceous, glabrous above or with a 
few hairs on the veins and margins, glabrous or 
generally pubescent on the veins below; midrib 
distinct, primary veins (6-11 pairs) aevi. dis- 
tinct. Nanophyllous leaves deciduous, 4-1 
long, lanceolate to linear. Inflorescences 4-18. 
flowered, umbelliform, solitary or a few together; 
peduncle 1.7-4.5 cm, glabrous; bracts to 0.6 cm 
long, ovate to broadly ovate; pedicels 0.7-1.8 cm, 


Distributions in Borneo. — 21. Argostemma havilandii. — 22. A. calcicolum. 


Annals of the 
Missouri Botanical Garden 


ı Icm B; 2cmE 


, 20cm A 
FIGURE 25. — Argostemma calcicolum. — A. Habit. — 
B. Flower. — C. Anthers. — D. Style. — E. Stipule. — C. Hair 


from outer surface a the AS —G,. Hair from inner 
surface of the corolla. — K. Hairs un the connectives. 

, E from Burtt & Woods 2005 (E); B-D, G, I from 
Bremer 1707 (S). 


pubescent or rarely glabrous. Flowers 5-merous; 
calyx lobes ca. 1 mm long, ovate to orbicular, 
obtuse or acute, pubescent; corolla 3-4 mm long, 
cleft to about 14, externally pubescent, internally 
papillose to pubescent only on the lobes; lobes ovate 
to triangular, recurved. Stamens 3-4 mm long, 
coherent; anther cells free, the apical appendages 
connate; anther cone ovoid to shortly so, straight; 
filaments very short, equal, straight; sacs opening 
completely longitudinally; apical appendage 1⁄4- 1% 
the anther length, thick, coriaceous, smooth; con- 
nective coriaceous, distinct, generally smooth. Style 
ca. 3-6 mm long, pubescent, with a clavate stigma, 
long-exserted. Fruit without furrows or ribs. 


Valeton based 4. lanceolatum on four collec- 
tions by Teysmann and Hallier, in BO and L. All 
are suitable for lectotypification. Bakhuizen f. (in 
herb.) has selected Teysmann 11292 in L, and I 
follow his choice. 

Bakhuizen f. selected the K specimen of A. 
havilandii as the type (= holotype). This is a 
mistake because Ridley (191 2) stated that the plants 
are preserved in the Singapore Botanical Gardens 
(note under 4. ophirense). 


Argostemma havilandii has a scattered distri- 
bution. Most, if not all, collections are from lime- 
stone. It grows from 50 to 1,100 m elevation on 
litter or more or less as an epiphyte on tree trunks, 
usually in lowland rainforest. 

It is easily distinguished. The stem is black on 
the upper part (on dry specimens), and the lower 


transverse bars. The corolla is cleft to about its 
middle into recurved lobes. The anther cone is low 
and pale yellow to creamish. The connectives are 
distinct, and the thecae are separated from each 
other at anthesis. The broad, thick apical append- 
ages are connate along their whole length. 

Argostemma havilandii is easily circumscribed. 
There is little variation among the populations ex- 
cept for width and hair cover of the leaves. From 
around Gunong Api there is a population with a 
very dense hair cover, even more distinctive than 
the densely pubescent specimens from Sungei Bika 
in Kalimantan, where they have been described as 
A. hallieri var. sparsipilum. 


Additional specimens examined. BORNEO. SARA- 
WAK: lst Div., Mt. Matang: 500 ft., 1954, Brooke 9487 
5181 (NY, 


(L); 1,400 ft., 1927-1928, native collector 
; ft., 


14, native i 2403 
near Tabang, 
Teysmann 11921 (BO, 


ng, 
lector Tode (A); 


Pro Kutei, 
termans ; 12845 (Ly without loc., 
K). 


22. Argostemma calcicolum B. Bremer, sp. 
nov. TYPE: Borneo. Sarawak: 1st Div., Bukit 
Manok, halfway between Teng Bukap and Pa- 
dawan, low alt., 1979, Bremer 1707 (holo- 
type, S; isotypes, K, L, SAR). Figure 25. 


Herba caulibus porcatis et suberosis ad papyraceis. 
Folia ien. illis parium aequalibus. Inflorescentiae um- 
belliformes aut ramis duobus scorpioides. Lobi calycis 
triangulares ad ovati, acuti. Corolla ad dimidio fissa. Lobi 
corollae recurvi. Antherae liberae. Appendices apicales 
connatae, //-'/ longitudinem antherarum aequans, cras- 


Volume 76, Number 1 
1989 


Bremer 41 
Argostemma in Borneo 


sae, coriaceae, laeves. Connectivum distinctum, papillo- 
Stylus pubescens, stigmate clavato. 
Argostemma havilandii simile, - iix plus pubes- 
cens, foliis oppositis aequalibus differ 


Erect, isophyllous herb. Stem 5-30 cm, slightly 
branched or unbranched, slightly pubescent, the 
lower and older parts pale brown with elevated 
corky to papery SOAM ridges and/or with 
transverse bars; internodes mm. Leaves op- 
posite, equal or subequal; stipules deciduous, 5- 
10 mm long, ovate to oblong, acute to obtuse at 
apex; petiole 3-14 mm long, pubescent; lamina 
5-17 
neate to attenuate with the lobes equal, marginally 
entire or serrulate, apically acuminate, herbaceous, 
pubescent on both surfaces; midrib and primary 
veins (7-9 pairs) distinct. Inflorescences 3-30- 
flowered, umbelliform or with 2 scorpioid branches, 


X 2-7 cm, obovate to elliptic, basally cu- 


solitary or a few together; peduncle 3-4.5 cm, 
pubescent, the bracts to 0.5 cm long, lanceolate; 
pedicels 0.5-0.7 cm, pubescent. Flowers 5-mer- 
ous; calyx lobes ca. 1 mm long, triangular to ovate, 
acute, pubescent; corolla ca. 4 mm long, cleft to 
about 4, externally pubescent, internally papillose 
to pubescent only on the lobes; lobes ovate to 
triangular, recurved. Stamens ca. 3.5 mm long, 
coherent; anther cells free, the apical appendages 
connate; anther cone ovoid to shortly so, straight; 
filaments very short, equal, straight; sacs opening 
completely longitudinally; apical appendage /-* 
the anther length, thick, coriaceous, smooth; con- 
nective coriaceous, distinct, papillose. Style 5-6 
mm long, pubescent, with a clavate stigma, long- 
exserted. Fruit without furrows or ribs. 


This rare species has been collected only four 
times. Haviland and Hose made the first collection, 
noting only the locality as Baram. Different col- 
lectors recorded it as growing on limestone. An- 
derson said “frequent in crevices on vertical lime- 
stone rocks." My collection is from low altitude in 
a limestone area. The earlier collections were a 
determined as A. borragineum because of the equal, 
opposite leaves. However, the stems, stipules, an- 
thers, and style are quite different. It is close to 
A. havilandii and shares various characters with 
this species: lower stems pale brown to grayish, 
these having papery to corky surfaces and elevated 
ridges; anther shape; and clavate pubescent styles. 
However, Argostemma calcicolum differs from A. 
havilandii by its denser pubescence even on the 
peduncles and stipules, isophyllous leaves, and the 
upper part of the connectives distinctly papillose 
to hairy. The hairs on the connectives are not 
homologous to those on the anthers of 4. psycho- 


L0. 2mm GK —  ., 
0.5cm CD i 
¿ lem B; 2cmE  , 
_20cm A E CE 
G, G 
FIGURE 26.  Argostemma trichosanthes. — A. Hab 
it. — B. Flower. — C. Anthers. — D. Style. — E. Stipule.— 


G. Hair from outer surface of the FREE —G,. Ha 
inner surface of the corolla. —K. Papillae or hairs from 
the connectives. From Winkler 781 (NY). 


trioides, which has smooth connectives but papil- 
lose to hairy thecae and apical appendages. In A. 
calcicolum these parts are smooth. 


Additional specimens examined. BORNEO. SARA- 

WAK: 4th Div., Gunong Subis, 1972, Anderson 531672 
(A); 4th Div., Penkalan Lobang Niah Caves, Burtt & 
Woods 2005 (E); 4th Div., Baram, 1894, Haviland & 
Hose 3409 (K). 
23. Argostemma trichosanthes Merr., Mitt. 
Inst. Allg. Bot. Hamburg 7: 280. 1937. TYPE: 
West Borneo. Bidang Menabei, 700 m, 1924, 
Winkler 781 (holotype, HBG; isotype, NY). 
Figure 26. 


Erect, densely pubescent, isophyllous herb. Stem 
ca. cm, unbranched, densely pubescent, lower 
and older parts pale brown with elevated corky to 
papery longitudinal ridges and/or with transverse 
bars; internodes mm. Leaves opposite, equal; 
stipules deciduous, 8-10 mm long, ovate, obtuse 
at apex; petiole 5—12 mm long, pubescent; lamina 


42 


Annals of the 
Missouri Botanical Garden 


Mars 23, 24. 


6-11 x 3-5 cm, obovate to elliptic, basally cu- 
neate to attenuate with the lobes equal, marginally 
serrulate, apically acuminate, herbaceous, pubes- 
cent on both sides; midrib and primary veins (ca. 
8 pairs) distinct. Inflorescences 10—20-flowered, 
umbelliform, solitary; peduncle 1.5-3.2 cm, pu- 
bescent; bracts to 0.6 cm long, lanceolate to linear; 
pedicels 0.5-0.7 cm, pubescent. Flowers 5-mer- 
ous; calyx lobes less than 1 mm long, triangular 
or ovate, acute, pubescent; corolla 3-5 mm long, 
cleft to about 14, externally pubescent, internally 
papillose to pubescent only on the lobes; lobes ovate 
to triangular, recurved. Stamens ca. 3.5 mm long, 
coherent, the anther cells free, the apical append- 
ages connate; anther cone ovoid to shortly ovoid, 
straight; filaments very short, equal, straight; sacs 
opening completely longitadiselly: apical append- 
age YY the anther length, thick, coriaceous, 
smooth; connective coriaceous, distinct, papillose 
in upper part. Style ca. 6 mm long, pubescent, 
with a clavate stigma, long-exserted. Fruit without 
furrows or ribs. 


Argostemma trichosanthes is known only from 
the type. It was collected at Bidang Menabei on 
Kalimantan at 700 m. 

Argostemma trichosanthes is close to A. havi- 
landii and A. calcicolum, with which it shares the 
stem characters pale brown to gray, papery to 
corky, and with elevated ridges on the lower stems. 
With A. calcicolum it shares opposite isophyllous 
leaves. Argostemma trichosanthes is easily distin- 


24 


— 100 km 
Ef > 500m 


D 
Kh 


Distributions in Borneo.— 23. Argostemma trichosanthes. —24. A. humifusum. 


guished by its dense indumentum. The upper part 
of the stems, particularly the younger leaves and 
most parts of the inflorescences, are densely cov- 
ered by stiff, long, thick-walled hairs. 

24. Argostemma humifusum W. Smith, Notes 
Roy. Bot. Gard. Edinburgh 8: 317. 1915 
TYPE: Borneo. Sarawak: Gunong Bayat 
(= ?Gunong Sebayat near Kuching), 1914, 
native collector D131 (holotype, E; isotypes, 
E, K, SAR). Figure 27. 


Erect, anisophyllous herb. Stem 6-16 cm, 
slightly branched or unbranched, glabrous; inter- 
nodes 4-15 mm. Leaves opposite, very unequal; 
stipules persistent, 6-8 mm long, cordiform or 

roadly ovate, acuminate at apex. Larger leaves 

of the anisophyllous pairs with finely pubescent 
petioles 5-16 mm long; lamina 4-9 x 2-4.5 cm, 
obovate, basally acute to obtuse with the lobes 
unequal, marginally serrulate, apically shortly acu- 
minate, membranaceous to herbaceous, pubescent 
on both surfaces; midrib and primary veins (6-10 
pairs) distinct. Nanophyllous leaves persistent, 5— 
10 mm long, ovate. Inflorescences 5-10-flowered, 
umbelliform, solitary or a few together; peduncle 
1.7-2.5 cm, glabrous; bracts very small, ovate to 
linear; pedicels 0.3-0.6 cm, with a few hairs. Flow- 
ers 5-merous; calyx lobes less than 1 mm long, 
ovate, acute, pubescent; corolla ca. 4 mm long, 
externally pubescent, internally glabrous(?). Sta- 
ns ca. 3 mm long, coherent; anther cells free, 


Volume 76, Number 1 
1989 


Bremer 43 
Argostemma in Borneo 


¿0.2mm G ; 
_05cm CD , 


_ 1cm B; 2cmEF 
¿20cm A - 

FIGURE 27. — Argostemma humifusum. — A. Habit. — 
B. Flower bud, and anther cone.— C. Anthers.— D. 
Style. — E. Stipule. — F. Nanophyllous leaf. —G. Hair from 


outer surface of the corolla. From native collector D131 
K). 


E, 


the apical appendages connate; anther cone ovoid 
to shortly ovoid, straight; filaments very short, equal, 
straight; sacs opening completely longitudinally; 
apical appendage /,-/ the anther length, thick, 
coriaceous, smooth; connective coriaceous, dis- 
tinct, papillose in upper part. Style ca. 5 mm long, 
glabrous to pubescent, with a clavate stigma, long- 
exserted. Fruit without furrows or ribs. 


Argostemma humifusum has been collected 
twice, but nothing was recorded about its habit or 
habitat. 

This species is characterized by its broad, ob- 
ovate leaves and large cordiform stipules. Because 
of incomplete material, I do not know the shape 
of the corolla. It has a thin corolla similar to those 
generally cleft halfway into recurved lobes. Inter- 
nally the corolla seems to be glabrous, but among 
the other species with thin corollas, the lobes are 
usually pubescent. The upper part of the connec- 
tive is probably papillose to pubescent as in 4. 
calcicolum and A. trichosanthes. 


Additional specimen examined. BORNEO. SARAWAK: 
Kutein, 1865-1867, Beccari 150 (K, L) 


25. Argostemma psychotrioides Ridley, J. 
Bot. 65: 38. 1927. TYPE: Borneo: Haviland 
689 (holotype, K; isotype, SAR). Figure 28. 


Erect, anisophyllous herb. Stem 10-100 cm, 
slightly branched or unbranched, finely pubescent 


_0.2mmG F 

_05cm CD ; 

v Icm B; 2cmEF  , 

20cm A 

FicunE 28. PI psychotrioides —A. Flow 

d branch. — B. Flower.— C. Anthers. — D. Style AE. 
pule. — F. Nanoha leaf. —G. Hai from outer sur- 

nd of the corolla. From Burtt & Martin 4991 (E). 


L 


or glabrous; internodes 7-30 mm long. Leaves 
opposite, very unequal; stipules deciduous, 6-20 
mm long, ovate to lanceolate, acuminate at apex; 
larger leaves of the anisophyllous pairs with finely 
pubescent or glabrous petioles 3-19 mm long; lam- 
ina 5-18 x 1-6 cm, obovate or oblanceolate to 
ovate or lanceolate, basally acute to attenuate with 
the lobes equal, marginally entire, apically acu- 
minate or rarely acute to obtuse, herbaceous, gla- 
brous above, glabrous to finely pubescent on the 
veins below; midrib and primary veins (7-15 pairs) 
distinct. Nanophyllous leaves deciduous, 5-13 mm 
long, ovate to lanceolate, acuminate. Inflorescences 
3-30-flowered, umbelliform, rarely with lateral 
branches to corymbiform, solitary or a few togeth- 
er; peduncle 1—4 cm, pubescent or glabrous; bracts 
0.7 cm long, ovate or lanceolate to linear; lateral 
branches, if present, to 3 cm, pubescent; pedicels 
0.5-1.5 cm, pubescent. Flowers 5-merous; calyx 


Annals of the 
Missouri Botanical Garden 


25 


00 km 
dE > 500 m 


Mars 25, 26. 


lobes 2-6 mm long, subspatulate or ovate to ob- 
long, acute, reflexed and pressed against the ovary 
concealing it even before anthesis, pubescent to 
glabrous; corolla 3-5 mm long, cleft to about Y2 
its length, externally pubescent, internally papillose 
to pubescent only on the lobes; lobes ovate to 
triangular, recurved. Stamens 2-4 mm long, co- 
herent; anther cells free, the apical appendages 
connate; anther cone ovoid to shortly ovoid, straight; 
filaments very short, equal, straight; sacs openin 

completely longitudinally; apical appendage 1⁄4- !6 
the anther length, thick, coriaceous, papillose; con- 
nective coriaceous, distinct, generally smooth. Style 
4-7 mm long, glabrous or rarely with a few hairs, 
with a clavate stigma, long-exserted. Fruit without 
furrows or ribs. 


Argostemma psychotrioides is found mostly at 
altitudes less than 250 m, although it occurs up 
A m. It grows in shady, wet places on 
exposed roots and pneumatophores in swampy for- 
ests and on boulders in streams. It also grows under 
drier conditions on limestone and at higher altitudes 
in mossy forests. 

It is easily identified by its strongly reflexed calyx 
lobes which in living material are green with white 
margins and tips. The corolla is thin and pubescent 
to densely pubescent and has recurved lobes. The 
anther cone is low, pale yellow to creamish. Only 
the apical appendages are connate, and the thecae 
are free. The thick apical appendages and the back 
of the thecae are distinctly papillose to hairy, while 
the connectives are smooth (cf. A. trichosanthes). 


26 


te 
¿Ef > 500 m 


Distributions in Borneo.— 25. Argostemma psychotrioides. — 26. A. variegatum. 


There is little variation in the characters of the 
flowers, but a few specimens (Bremer 1730, 1744, 
1748) from the 7th Division have shorter calyx 
lobes. The shape and size of the leaves as well as 
the occurrence of hairs are variable and do not 
correlate with distribution patterns. 


Additional specimens examined. ORNEO. SA- 
RAWAK: lst Div., Kuching, 1893, Haviland 2960 (BO, 
L, SAR); 1894, Haviland s.n. (BM, L); 1894, Haviland 
& Hose s.n. (A, GH); 1905, Ridley 12302 (BM, K, 
SING); lst Div., Setapok, 1966, Anderson 819690 (K, 
L, SAR); Ist Div., Mt. Matang, 1924, Mjoberg 216 (BO, 
NY, SING, UC); 2,000 ft., 1927, native collector s.n. 
(UC); 1st Div., Padawan area, Gunong Maja, 1979, Bre- 
mer 1703 (S, SAR); lst Div., Bukit Manok, halfway 
between Teng Bukap and Padawan, 1979, Bremer 1706 
(S, SAR); 1st Div., Semengoh, 1979, Bremer 1643 (S, 
SAR); Banyeng ak Nudong & Sibat ak Luang 525355 
K, L, SAR); 2nd Div., Simangang, 1955, Brooke 10716 
(BM, L); 2nd Div., Bukit Sadok, 1982, Axelius 86, 93 
(S); 2nd Div., between Entalau and Tisak, 1982, Axelius 
, Bukit Sengkajang, Lanjak- Entimau, 


— 


Nanga Pelagos, 1938, 
Daud & n SFN3567 678 (A, G, K, SAR, SING); 7th 
x Hos s., above Melinau Falls, 4,000 ft., 1967, 
tt & Martin 4991 (E); E of Bukit Sanpandai, EN 
[n Fw (Ey 7th s Sungei Melinau, Nanga 
noh, ., 1980, 2666 (E) i 
T Sins Putai, 1979, Bremer 1724 (S, SAR); mae 
tion of Sungei Tekalit and Sungei Mengiong, 1979, Bre 
mer 1730. 1744 (S, SAR); Sungei Mengiong, 1979, 
Bremer 1748 (S, SAR); 7th Div., Sungei Bene, 1980, 


£ 

= 
~ 
h 
~ 


Volume 76, Number 1 
1989 


Bremer 45 
Argostemma in Borneo 


FIGURE 29. Argostemma  variegatum. — A. Hab- 
iS —B. 2 bud and part of m PE Kis the inner 
urface. — C. Anthers. — D. . Stipule.— a- 


: d leaf. —G. Hair EE. outer surface of the co- 
rolla.—G,. Hair from inner surface of the corolla. — 
Papillae or hairs from the apical appendages. From Wi ink- 
ler 1561 (HBG, NY). 


RA 


Burtt 12935, 12951 (E); 7th Div., Gat, 1929, Clemens 
21738, 12740 da without qe native collector 294, 
4 (A, UC) Lobb s.n., 


NW of Tabang, 100-150 m, 1979, Murata et al. 1192 
). 


26. A Merr., Mitt. Inst. 
Allg. Bot. Hamburg 7: : 280. 1937. TYPE: West 
Borneo. Nanga Era, 100 m, 1925, Winkler 
1561 (holotype, HBG; isotype, NY). Figure 
29 


Creeping anisophyllous herb. Stem 10-20 cm, 
slightly branched or unbranched, pubescent; in- 
ternodes 3-17 mm long. Leaves opposite, very 
unequal; stipules persistent, 3-6 mm long, lanceo- 
ate, acuminate at apex; larger leaves of the aniso- 
phyllous pairs with pubescent petioles 2-5 mm 
long; lamina 3.5-7 x 1.4 cm, obovate to 
oblanceolate, basally rounded or obtuse with the 
lobes unequal, marginally entire or serrulate, api- 
cally obtuse or obtuse with a point, herbaceous, 
glabrous above, finely pubescent on the veins and 
margins below; midrib and primary veins (7-9 pairs) 
distinct. Nanophyllous leaves persistent, ca. 5 mm 


ESSE E 
—B. Flowe 


ay ea bryophilum. — 


Hab- 
—C. Anthers. — D. Style with pil at 
ba —E. Sti T ule. —K. Papillae from the connectives. 
Prom Burtt 12800 (E). 


long, linear, petiolelike. Inflorescences ca. 7—flow- 
ered, umbelliform, solitary; peduncle 1-3 cm, pu- 
bescent or almost glabrous; bracts ca. 0.5 cm long, 
linear; pedicels 0.5-0.9 cm, pubescent. Flowers 
5(or 6)-merous; calyx lobes 3-4 mm long, sub- 
spatulate or ovate to oblong, acute, reflexed and 
pressed against the ovary concealing it even before 
anthesis, pubescent to glabrous; corolla ca. 3 mm 
about 1%, externally pubescent, in- 


3 mm long, coherent; anther cells free, the apical 
appendages connate; anther cone ovoid to shortly 
ovoid, straight; filaments very short, equal, straight; 
sacs opening completely longitudinally; apical ap- 
pendage !4- 16 the anther length, thick, coriaceous, 

apillose; connective coriaceous, distinct, generally 
smooth. Style ca. 5 mm long, glabrous, with a 
clavate stigma, long-exserted. Fruit without fur- 
rows or ribs. 


Argostemma variegatum has been collected on 
steep soil banks in disturbed primary forest and in 
primary forest at ca. 100 m. Its habit is like that 
of A. elatostemma and A. densifolium, that is, 
creeping with anisophyllous leaves, the larger ones 
with unequal leaf-base lobes. The stems, petioles, 
and veins are distinctly pubescent. The flowers, 


46 Annals of the 
Missouri Botanical Garden 


27 
— 100 km 
ER > 500 m 


28 


100 km 
E dw m 


Mars 27, 28. Distributions in Borneo. — 27. Argostemma bryophilum. — 28. A. neurocalyx. 


however, are different from the above species; they peduncle 3-6 mm long, almost glabrous; bracts 
are similar to those of 4. psychotrioides. The calyx ca. 4 mm long, lanceolate or triangular; pedicels 
lobes are long and reflexed, the corollas are cleft ca. 7 mm long, pubescent. Flowers 5-merous; calyx 
to about half their length into pubescent, recurved lobes 2-4 mm long, triangular, acuminate; corolla 
lobes. The original collection has silvery mottled cleft to near the base, glabrous on both surfaces; 
leaves while the other does not (perhaps due to lobes 5-7 mm long, ovate to lanceolate, acuminate, 
drying). Argostemma variegatum is close to A. spreading. Stamens 4-5 mm long, coherent; anther 
psychotrioides; the similarities to 4. elatostemma cells and apical appendages connate except for the 
are superficial. most apical part; anther cone ovoid, straight; fil- 


aments short, equal, st ght; f diff tlengt h 


Additional specimen examined. BORNEO. SARAWAK: I 
7th Div., Bukit Raya, 1969, Smith S28234 (A, K, L). opening completely longitudina y; apica append- 
e very short, glabrous; connective + distinct, 


27. Argostemma bryophilum Schumann in clavate stigma, shortly exserted. Fruit without fur- 
Schumann & Lauterb., Nachträge zur Flora rows or ribs. 
der Deutschen Schutzgebirge in der Südse 
393. 1905. TYPE: from Kaiser Wilhelmsland. 
not seen. Figure 30. 


Argostemma bryophilum grows on wet rocks, 
boulders, or banks among mosses from 350 to 
1,500 m 

Small, creeping to erect, generally isophyllous It is the first species described in a large complex 
herb. Stem 2-10 cm, unbranched or slightly from New Guinea and the Philippines with more 
branched, densely pubescent; internodes 3-20 mm. than ten taxa. Bakhuizen f. (pers. comm.) has treat- 
Leaves opposite, generally equal; stipules persis- ed the complex as a single species. All details of 
tent, to 4 mm long, broadly ovate to cordiform, the complex are not known and so I have only 
acute to obtuse at apex; petiole 3-11 mm long, accepted one variable species. This is the first re- 
pubescent; lamina 1.5-4.2 x 0.6-1.9 cm, ovate port from Borneo. The collections from Borneo are 
or elliptic to lanceolate, basally cuneate to atten- all rather similar, with the exception of Axelius 
uate with the lobes equal, marginally serrulate or — 7724, which has anisophyllous leaves. The Borneo 
entire, apically acute to acuminate or obtuse with specimens form a homogenous group and may in 
a point, membranaceous, pubescent on both sur- the future be recognized as a distinct taxon separate 
faces, below most prominent on the veins; midrib from the New Guinea and Philippine members of 
and primary veins (5-8 pairs) distinct. Inflores- the complex. It is a small plant with pale green, 
cences 1-2-flowered, solitary or a few together; thin, and hairy leaves. With one exception, all 


Volume 76, Number 1 
1989 


Bremer 47 
Argostemma in Borneo 


collections have isophyllous, opposite leaves, very 
pubescent stems, and cordiform, glabrate stipules. 
The flowers are solitary, starlike and with long, 
bright yellow anther cones. The thecae distinguish 
this species from all others: the inner pollen sac of 
each theca is only ca. half as long as the outer 
one, which is almost as long as the whole anther, 
and there is no distinct apical appendage. The 
stigma is distinctly clavate with papillae covering 
the clavate area. In all flowering specimens (five 
of the seven known specimens), the stigma is totally 
covered by germinated pollen grains. The pollen 
is probably from the same because the 
stigma is partly covered by the apical anther parts. 


ower, 


I have not found more than a few pollen grains on 
the stigmas of any other species. Probably this 
species is autogamous. 


pecimens examined. BORNEO. SARAWAK: Ist Div., 
Sabal, 350 m, 1979, Bremer 1664 (S, SAR); 2nd Div., 
Bukit Sadok, 1982, Axelius 112A (S); 5th Div., Bake- 
lalan, 4,000 ft., 1955, Brooke 10406, 10464 (BM, L); 
7th Div., Hose Mts., Bukit Salong, 1980, Burtt 12733 
(E); Bukit Sanpandai, 4,500 ft., 1980, Burtt 12800 (E); 

7th Div., Bukit Dema, 1978, Burtt 11328 (E). 


28. Argostemma neurocalyx Miq., Ann. Mus. 
ot. Lugd.-Bat. 4: 229. 1869. TYPE: Suma- 
tra?: Junghuhn s.n. (holotype, U). Figure 31. 


A. deo uis Merr., Univ. Calif. Publ. Bot. 15: 275. 
1929. TYPE: British North Borneo: Tawao, 1922- 
1923, Elmer 21138 (lectotype, K; Bakh. f. in herb., 
confirmed here; isolectotypes, BO, BR, G, GH, NY, 
S, SING, U, UC, US) 


Erect anisophyllous herb with an apical rosette 
of 2-4 leaves. Stem 4-8 cm, unbranched, glabrous 
or glabrate, without distinct internodes. Leaves ver- 
ticillate, compressed in an apical rosette; stipules(?) 
persistent, 4-9 mm long, lanceolate, acuminate at 
apex; petiole absent; lamina 2-15 x 12.7-7.6 
cm, suborbicular or ovate to elliptic, basally obtuse 
or shortly attenuate and equal, marginally entire, 
apically acute to shortly acuminate, membrana- 
ceous, pubescent above, below with a few hairs on 
the veins; midrib and primary veins (7-9 pairs) 
distinct. Inflorescences 4-15-flowered, umbelli- 
form, solitary or a few together; peduncle 2.3-5.5 
cm, glabrous; bracts to 0.5 cm long, ovate; pedicels 
0.4-0.7 cm, pubescent. Flowers 4(or 5?)-merous; 
calyx lobes 1-2 mm long, ovate to broadly ovate, 
acute, glabrous; corolla ca. 5 mm long, cleft less 
than 4 its length, glabrous on both surfaces; lobes 
ovate to triangular, erect. Stamens ca. 3 mm long; 
free, not forming a cone; filaments ca. 2 as long 
as the anthers, equal, straight; sacs opening by 
pores; without apical appendages; connective dis- 


,. 05cm CD 
. 1em B; 2cmE 
| 10cm 


Pad 3l. Argostemma neurocalyx. — 


A. Habit. 
B. Flower. — C. Anthers. — D. Style. — E. Stipule? Fiom 
Elmer 21 1 38 (G, SING). 


tinct, smooth. Style ca. 4 mm long, glabrous, with 
a capitate stigma, exserted. Fruit without furrows 
or ribs. 


Merrill based 4. platyphyllum on one collec- 
i . All spec- 
imens are suitable for lectotypification. Bakhuizen 
f. (in herb.) has selected the K specimen, and 
follow his choice. 

Argostemma neurocalyx was collected just once 
on Borneo, by Elmer, near Tawau, and was de- 
scribed by Merrill as 4. platyphyllum. The pro- 
tologue says “a succulent erect herb in dense for- 
ests on moss-covered basaltic rocks over which 
water trickles." It is not only the sole collection of 
the species on Borneo, it is also the only collection 
representing this section of the genus from Borneo. 
Possibly it was erroneously labeled in the herbarium 
However, it is an easily di 


tion by Elmer; no specimen is indicate 


by Elmer or Merrill. 
tinguished species with two or four Anon: vel 
slightly unequal leaves. On dry specimens, the leaves 
are very thin and membranaceous. It is difficult to 
determine from the herbarium specimens if there 
are stipules or just smaller leaves. The umbelliform 
inflorescences have long glabrous peduncles, rather 
large, ovate bracts, and pubescent pedicels. The 
flowers are tetramerous and campanulate, and the 
corolla lobes seem to be reflexed. The stamens are 
free with long filaments and connectives ending 


each pollen sac. The pores (slits) are placed between 
the sacs of each theca. As the dorsal sacs are larger 


48 


Annals of the 
Missouri Botanical Garden 


than the ventral sacs, only the pores of the dorsal 
ones can be seen (from the inside of the flower), 
and the pores of the ventral sacs are hidden by 
the longer dorsal sacs. 


LITERATURE CITED 


BAKHUIZEN VAN DEN BRINK, R. C., JR. 1953. Florae 
Malesianae praecursores V. Notes on Malaysian Ru- 
biaceae. Blumea 7: 329- 338. 

1 A synoptical key to the genera o 

Rubiaceae of Thailand. Thai Forest Bulletin (Botany) 

9: 15-55. 


ena J J. 1838. Plantae Javanicae Rariores 1. W. 
Allen, London. 
BREMER, B. 1979. The genus Neurocalyx (Rubiaceae- 
Argostemmateae) in Ceylon. Bot. Not. 132: 399- 
407. 


1987. E sister d of the paleotropical 
tribe Argostemmateae: efined neotropical tribe 
Hamelieae (Rubiaceae, Kodesa. Cladistics 3: 35- 


BREMER, K. 83. Taxonomy of Memec pn (Melas- 
tomataceae) in Borneo. Opera Bot. 69: 
Darwin, S. P. 1976. The subfamilial, tribal and sub- 


iriliel nomenclature of the Rubiaceae. Taxon 25: 


5 : 

DORMER, K. J. 1962. The fibrous layer in the anthers 
of Compositae. New Phytol. 61: 150-153. 
aora, F. 1983. Ubersicht des ee 

ds 758-915 in E. Strasburger, Lehrbuc 
anik. Gustav Fischer Verlag, Stuttgart & New Yo rk. 
1985. Patterns of el wall thick- 


Bei J. C. 
enings in Araceae: subfamilies Pothoideae and Mon- 
steroideae J. Bot. 72: 47 

Kinc, G Argostemma. In: G. Kin 


J. S. 

Gamble, jeng for a Flora of the Malayan Pen- 
t. Soc. Bengal 72(2): 141-156. 

1974. Colleter morphology in Pavetta, 
Neorosea and Tricalysia (Rubiaceae) and its rela- 
tionship to pom leaf nodule symbiosis. Bot. J. 
Linn. Soc. 69: 125-1 
1975. Colleter types in Rubiaceae, especially 
in relation to the bacteria! leaf nodule symbiosis. Bot. 


The endothecium—a neglected 
criterion in car and phylogeny? Bothalia 14: 


833-8 
NORDENSTAM, B. 1978. Taxonomic studies in the tribe 
Senecioneae Ve aii Opera Bot. 44: 1-83. 
Riptey, H. N. 1901. The flora of Mount Ophir. J. 
Straits lira Roy. Asiat. Soc. 35: 1-28. 
New and rare Malayan he (series 
yp. J. E Branch Roy. Asiat. Soc. 61: 1-43. 
927. The genus Argostemma. J. Bot. 65: 


STEENIS- Kruseman, M. J. VAN. 
ors. In: C. G. G. J. van Steenis (editor), Flora 
Malesiana 1(1). Noordhoff- Kolff, N. V. Djakarta. 
Tan, H. & A. N. Rao. 1981. Vivipary in Ophiorrhiza 
tomentosa Jack Rubiaceae. Biotropica 13: 232-233. 
VERDCOURT, B. 1958. Remarks on the classification of 
the Rubiaceae. Bull. Jard. Bot. Bruxelles 28: 209- 
290. 


1950. Malaysian plant 


APPENDIX I 
INDEX TO COLLECTIONS 


The species number is given within parentheses after 
each collection number. Only cited specimens from Bor- 
neo are listed. 


Anderson 4090 (3), 4284 (13), 4349 (14), 4516 
(19), 8982 (21), 9106 (21), 12581 (21), 12915 (21), 
14600 (21), S4516 (14), S19690 (25), S25106 (14), 
828396 (15), 528735 (13), S30826 (21), 531672 (13), 
831672 (22); Anderson & Ilias Paie 528295 (15), 
828657 (8), 528667 (18), S28881 (15); Anderson et 
al. 520288 (2); Argent 1308 (14); Argent & Coppins 
1076 (14), 1081 (19), 1142 (18); 
1038 (14); Argent et al. 666 (21); Asah a n 
822752 (7); Ashton 42 (13), 199 (14), S12132 (14), 
317606 (3), S17691 (10), S18063 (13), S21238 (14), 


Tx 
aR 
® 
5 
- 
e 
E 
o 
= 
8 
< 


S21249 (14), S21265 (11); Axelius 86 (25), 112A (27), 
"S NM Aen a. day oe 
25); Nu at ak Luang S 


(25); Beecari( 621 (16), "150 (24), 1716 (14); Bell 2068 
(20); Bogle 545 (15); Bogle et al. 389 (19); Br 
1643 (25), d (21), 1652 (1), 1662 (21), 1664 (27), 
0), 7 (18), 1668 (15), 1673 (15), 1675 (10), 
Er (5), 1689 (21), 1703 (25), 1706 (25), 
1717 (21), 1719 (14), 1720 (15), 1722 (2), 
1725 Ae 1730 (25), 1742 (15), 1744 (25), 
1747 (13), 8 (25), 1752 (13); Brooke 1034 (14), 
8502 (17), Hoe (20), 8564 (14), 8599 (20), 8670 (5), 
8840 (25), 8984 (13), 9128 (13), 9131 (10), 9258 (13), 
9465 (2), 9474 (17), 9487 (21), 9517 (5), 9743 (5), 
9749 (15), 9770 (21), 10123 (13), 10177 (14), 10183 
(3), 10283 (13), 10406 (27), 10408 (14), 10411 (11), 
oe 10461 (18), 10464 (27), 10542 (17), 10716 
Sooty 2 (21); Burtt 2655 (12), 8175 (2), 8228 (3), 


12667 (10), 12709 (14), 


(9), 127 
12839 (25), 12848 (18) 12896 (13), 12908 (14), 12935 
rtt Sá oan 4986 (9), 4991 (25), 
6 (14); Burtt & Woods 
1959 (2), 2005 (22), 2090 a 2095 (14), 2150 (19), 
2152 (14), 2176 (18), 2178 (19), 2182 (13), 2185 (11), 
2214 (4), 2293 (14), 2366 (1), 2372 (13), 2512 (5), 
2516 (14), 2518 (5), 2682 (14), 2701 (5), 2710 (15), 
2761 (14), 2762 (5), 2800 (20), 2816 (1), 2845 (14); 
Chai 8519532 (25), S30081 (13), 530085 (21), 533836 
(13), 533980 (25), 534006 (18), 534797 (10), S536137 
(15), S36485 (13), S37296 (14); Chai et al. 533328 
(14), S37338 (1); Chew & Corner RSNB4618 (14); 
Chew et al. 1041 (13), 2875 (13), 1713 (18); Clemens 
10058 (15), 10059 (13), 10981 (19), 11076 (18), 20064 
(20) 20331 (20) 20910 (5) 20911 (5), 20912 (2), 
21737 (10), 21738 (25), 22372 (15), 26033 (13), 26990 
(13), 28179 (19), 28250 (14), 29499 (14), 29538 (13), 
30650 (3), 31077 (15), 31963 (3), 32113 (18), 32578 
(13), 32962 (15), 33962A (15), 33962 (15), 34110 
(18), 35136 (19), 35137A (13), 35137 (15), 40049 
(14), 51020 (19), 51670 (14), 223221 (14), 26926A 
(18), field n 6891 (14), field n 6920 (14), field n 6958 


Volume 76, Number 1 Bremer 49 
1989 Argostemma in Borneo 
(5), field n 6973 (14), field n 7083 (5), s.n. (5), s.n. (13, — Argostemma Wall. in Roxb. 11 
s.n. (15), s.n. (18); Collenette 60 (14), 116 (13), 4179 anisophyllum Merr 20 
(11), A77 (19), A103 (19), s.n. (12); Cox 957 (13); apiculatum B S E 21 
Darnton 568 (15); Daud & Tachun SFN35678 (25); orragineum 32 
Ding Hou 220 (13), 225 (19); Elmer 21138 (28); borragineum var. Moa Valeton in 
Endert 2952 (15), 3614 (14), 3800 (11), 3874 (14), inkler 32 
3916 (11), 4171 (19), 4172 Be 4238 (14), 4327 (3); bsc E EE 36 
Fedilis & Sumbing SAN 88281 (13); Fuchs 21031 brookei 38 
(13); Gardner 88 (14); Geesink 9036 (13), 9112 (14), bryophilum trail in Schum. & Lauterb. ...... 46 
9113 (11), 9226 (13), 9255 (14), 9272 (11); Gibbs burttii B. Bremer 22 
4056 (19), 4101 (13); Grieswold 27 (14), 29 (13); calcicolum B. Bremer 40 
Hallier 1720 (14), 2686 (15), 2779 (13), 3230 (13), chaii 34 
B3228 (21); Hansen 20 (3), 187 (14); Haviland 684 densifolium Ridl. 17 
(14), 689 (25), 1031 (10), 1325 (18), 1326 (19), 2958 Finca var. latifolium Ridl. „nn 17 
(21), 2960 (25), 8475 (16), s.n. (13), s.n. (21), s.n. (25); dulit err. 20 
Haviland & Hose 3409 (22), s.n. (25) Hullett 329 a nm Hook. f. 14 
(13), s.n. (5); Ilias Paie 828445 (14), S36357 (14); enerve Ridl. 34 
Ilias & Yeo S38349 (1); Jacobs 5065 (10); Jaheri flavescens Bakh. f. 34 
in exp. Nieuwenhuis 1636 (7); James Mamit S35046 gaharuense B. Bremer 26 
(21); Kanis & miro 53966 (3); Kloss SFN19154 geesinkii B. Bremer 25 
(135; Kostermans 7437 (5), 9216 (13), 10121 (15), gracile Stapf 35 
10570 (13), 12845 (21), 12851 (18); Lobb s.n. (25); hallieri Valeton 39 
Lo 25); Madani 90812 (13); Martin 538185 hallieri var. sparsipilum Mer 39 
(18), 538848 (14); Martin & T 6861 (13); hameliifolium Wernham in ‘Gibbs EN 27 
Meijer 687 (5), 2286 (3), 2343 (13); Mjöberg 8 (7), havilandii Ridl. 38 
82 (14), 85 (14), 88 (14), 205 (5), 206 (5), 216 (25), hullettii Ridl. 27 
218 (15), s.n. (2), s.n. (5), s.n. (13), s.n. (14), s.n. (14), humifusum M 42 
(15), s.n. (20); Motley 1174 (2); Moulton 105 isophyllum 27 
(14), 6695 (14), 6726 (14); Moulton's nativ ho kinabaluense Wernham i in Gibbs s 37 
lectors (15); Murata et al. 1 25) n lanceolatum Valeton 39 
collector 85 (21), 294 (25), 408 (25), 1227 (17), 1232 mjoebergii Merr 30 
(21), 1244 (25), 1256 (13), 1428 (13), 2403 (21), 5181 motleyi Ridl 14 
(21), D131 (24), s.n. (5), s. 1), s.n. (25); moultonii Ridl. 29 
Nielsen 204 (13), 368 (17) 494 (13), 538 (4), 833 moultonii var. hirta Ridl. 29 
(18), 838 (14); Nooteboom 921 (14), 922 (19), 1031 murudense 30 
(15), 1035 (14); Nooteboom & Aban 1552 (13), 159 neurocalyx Miq. 47 
(13); Nooteboom & Chai 1704 (13), 1744 (14), 1856 nutans var. borneense Ridl. 32 
(14), 1935 (14), 2198 (11), 2307 a Oth s- ophirense Maingay ex Hook. f. ooo 18 
mawi S37437 (15); Price 194 ( Purseglove 4987 parvifolium Be 13 
(5), 5095 (10), 5096 (17), 5129 (13), Vr EN 3), 5282 parvulum Ridl. 30 
(15), 5534 (10); Purseglove & Shah 4811 (5); Rich- platyphyllum Merr. 47 
ards 1052 (13), 1537 (14), 1689 (18), mu (7), 1790 podochiloides Merr. 35 
(14), 1799 (13), 2090A (3), 2101 (7), 2455 (10), 7090 psychotrioides Ridl. 43 
(13); Ridley 11751 (5), 11753 (2), 12302 (25), 12303 stre Ri 23 
(2), 12304 (14), 12448 (4), s.n. (4); Sanusi Tahir 8954 salicifolium Ridl 18 
(25); Sibat ak Luang S23664 (25); Sidek Kiah 2 arawakense Smi 39 
(13); Sinclair 9212 (14), 10355 (15); Sinclair et al. streblosifolium Valeton 32 
9215 (15); Smith S28234 (26); Sylvester Tong 534346 subfalcifolium Bakh. f. 15 
(4), S34847 (7); Teysmann 11292 (21), 11921 (21); trichosanthes Merr. 41 
Tong & Banyeng S33271 (14); van Niel 3558 (13), variegatum Mer 45 
4580 (13); Winkler 660 (3), 767 (1), 781 (23), 791 velutinum Ridl. 30 
(3), 879 (15), 897 (6), 949 (13), 950 (6), 951 (10), ; 
1007 (15), 1282 (15), 1353 (15), 1428 (21), 1479 (18) oe uo - Tow nc P 
1561 (26), 1562 (15), 2853 (15), Wood SAN16695 brachyant iis E j 35 
(14). gracilis (Stapf) R 


APPENDIX II 
INDEX TO TAXA 


Accepted names are in boldface, synonyms in italics. 


Pomangium Reinw. — Argostemma 


STUDIES IN NEOTROPICAL 
PALEOBOTANY. VII. 

THE LOWER MIOCENE 
COMMUNITIES OF 
PANAMA —THE 

LA BOCA FORMATION' 


Alan Graham? 


ABSTRACT 


Thirty-nine palynomorphs have been dp um from among 54 forms recovered from the lower Miocene La Boca 


Formation in the Canal region of Panama. These 


Operculodinium, Lycopodium, CMT Cyathea, Pteris (types 1 
"S e es l- p monolete fern spores (types l- ui. Gramineae, Palmae (At 


r of high altitudes: cera of 1,200 t 
fossil floras. The affini 
Crudia, all genera id in the modern vegetation of Pana 
Tertiary paleotemperature curves, 
tropical habitats in southern Central America 


y of the flora is distincidy with mu America and areas to the north. 


are ascomycete cleistothecia, the dinoflagellates si iA and 
h o 


AnS fern 
anthus 
al- 


and 2), cf. Antrop yum 
ttalea, Manicar ria , and Syn 


-5), Malvaveas. Rhizophora (nee 67-88% of five samples counted), Rubiaceae ie 1 
i i i iocene assembla 


With the sa of 


a, and consequently the paleoclimate, in agreement with 


was similar to that era today in coastal, lowland, and moderate- nnde 


The La Boca Formation is the third in a series 
of three stratigraphically and lithologically similar 
formations from the geologically complex Cana 
region of Panama yielding well-preserved fossil pol- 
len and spores. The Culebra Formation is oldest 
(basal) and is overlain by the Cucaracha Formation. 
The La Boca does not anywhere lie directly on the 
Cucaracha, but it interfingers (viz., is contempo- 
raneous) with the Pedro Miguel Formation, which 
overlies the Cucaracha elsewhere in the Canal re- 
gion. Age estimates are based primarily on mol- 
luscan (Woodring, 1957-1982) and ostracod (Van 
den Bold, 1972, 1973) data which suggest an early 
Miocene age for the three formations. 

he La Boca outcrops along both sides of the 
Panama Canal from the Pacific entrance to the 
Las Cascadas Reach. The sediments were deposited 
in an estuarine environment and include an alter- 
nating sequence of mudstones, siltstones, sand- 
stones, lignitic shales, tuffs (waterlain volcanic ash), 


— 


and limestones typical of that environmental set- 
ting. The presence of the coralliferous Emperador 
limestone member in the lower part of the for- 
mation and Rhizophora-containing lignitic shales 
indicate deposition in warm temperate to tropical, 
shallow seas and in adjacent coastal, brackish-water 
habitats. 

Fifty-four samples were collected from two ex- 
posures of the La Boca Formation along the Las 
Cascadas Reach in September 1967 (latitude 
9°04'N, longitude 79?40'W; ground elevation 52.7 
m). Locality A (samples 1-26) was at Canal marker 
1600 (1766 in the new marking system), and lo- 
cality B (samples 27-54) at markers 1622 (1788), 
1625 (1791), and 1627 (1793). Since that time 
the Canal has been widened from 90 m to 155 m, 
and compared with the present physiography, the 
1967 samples were collected about 25 m up and 
60 m out over the present Canal. The beds dipped 
back into the slope at a 20-25? angle so that now 


! The author gratefully acknowledges information provided by William Elsik (Exxon Company, U.S.A.) and William 
Evitt pied ha ity) on the fungi and dinoflagellates. Research was supported by National Science Foundation 


pent BSR-850 


epartment bs Biological Sciences, Kent State University, Kent, Ohio 44242, U.S.A. 


ANN. Missouni Bor. GARD. 76: 50-66. 1989. 


Volume 76, Number 1 
1989 


Graham 51 
La Boca Formation—Lower Miocene 
Communities 


the same lignite layers are nearly at water level. 
Other details of the collecting site and geology of 
the area are summarized in Graham et al. (1985: 
495-502). 


MATERIALS AND METHODS 


Extraction and processing techniques are de- 
scribed in Graham (1985). Slides are labeled ac- 
cording to locality, sample number, and slide num- 
ber (e.g., Pan A 5-1). Of the 54 samples collected, 
20 contained fair to well-preserved palynomorphs 
(locality A—5, 14, 16, 17, 19, 20, 21; locality 
B— 27, 28, 29, 30, 33, 34, 35, 45, 46, 47, 50, 
52, 54). Location of the specimens on the slides 
is by England Slide Finder e (e.g., ESF 
C-29). The tabulations in Table 1 are based on 
five representative samples (14, E 20, 21, 50) 
containing diverse and well-preserved palyno- 
morphs. All materials are deposited in the paly- 
nology collections at Kent State University. 


SYSTEMATICS 


Thirty-nine palynomorphs have been identified 
from the La Boca Formation (Table 1) in addition 
to 15 others that cannot presently be identified 
(unknowns 1-15). Most of the specimens have 
been recovered from other Tertiary formations in 
the Gulf/Caribbean region and discussed in pre- 
vious publications, hence the material presented 
here is synoptic. These formations and references 
are as follows: Gatuncillo (middle(?) to late Eocene, 
Panama; Graham, 1985); San Sebastian (middle 
to late Oligocene, Puerto Rico; Graham & Jarzen, 
1969); Uscari (early Miocene, Costa Rica; Gra- 
ham, 19872, b); Culebra (early Miocene, Panama; 
Graham, 19882), and Cucaracha (early Miocene, 
Panama; Graham, 1988b). Present ranges of the 
modern analogs within the Neotropics and ecolog- 
ical data are based on field observations, personal 
communication with specialists, and the literature, 
especially Croat (1978), D'Arcy (1987), Hartshorn 
(1983), Tryon & Tryon (1982), and Woodson & 
Schery (1943-1980). Terminology for vegetation 
types follows Holdridge (1947; Holdridge et al., 
1971), used by Croat (1978) and Hartshorn (1983) 
for describing the plant communities of Panama 
and Costa Rica. In instances where the modern 
analogs extend into South America, their altitudinal 
ranges and community affiliations may differ slight- 
ly from their Central American occurrences. The 
data for Central America are considered first in 
paleocommunity and paleoenvironmental recon- 
struction of the La Boca flora, since communities 


in South America are more distant and were iso- 
lated from Central America until about 3 Ma. 


FUNGI 
Ascomycete cleistothecium (Fig. 1). Flattened, 
circular, multicellular, cells cubical, 6 x 9 um, 


outer and some lateral walls of peripheral (mar- 
ginal) cells thickened, center opaque (solid?), cells 
disrupted along outer edge, forming slitlike opening 
35 um long (attachment scar?); 124 
Reproductive structures of the Plectomycetes 
group of ascomycete fungi are frequent in Gulf/ 
Caribbean Tertiary deposits but never in large num- 
bers. Although entire specimens are relatively rare, 
fragments are present in almost all samples. Another 
type with radially aligned cells and marginal spines 
was recovered from the lower Miocene Uscari se- 
quence of Costa Rica (Graham, 1987a, fig. 1). 


PYRROPHYTA 


Spiniferites (Figs. 2, 3). These dinoflagellate 
cysts, as fossils referable to Spiniferites, are com- 
mon in the Tertiary and extend back to the Early 
Cretaceous. They are produced by some species 
of the extant Gonyaulax group (Evitt, pers. comm., 

87) 


This 
is also a common Tertiary form produced by the 
modern Gonyaulax grindleyi (= Protoceratium 


Operculodinium centrocarpus (Fig. 4). 


reticulatum) and a frequent associate of Spinifer- 
ites in the Miocene. According to Evitt (pers. comm., 
1987), “These two cyst types are common con- 
stituents of Tertiary nearshore sediments. They 
occur, virtually to the exclusion of other forms, in 
the Miocene Monterrey Formation in California 
(Spiniferites much the more abundant), and to- 
gether with a very rich associated assemblage in 
the Calvert Formation along the east coast. It would 
be entirely plausible to find them in estuarine de- 
posits of Miocene age virtually anywhere. No ex- 
amples of Spiniferites, which has a very distinctive 
and easily recognized morphology (at least at the 
generic level), have been found in fully freshwater 
sediments." 


LYCOPODIACEAE 


Lycopodium (Fig. 5). 
rounded; trilete, laesurae straight, narrow, ca. 20— 
22 um long, extending to spore margin, inner mar- 
gin entire, distal surface with numerous circular 
punctae ca. | um diam., Ru face laevigate; 
wall 1-1.5 um thick; 40 u 

These spores are abis to cee of L. reflexum 


Amb triangular, apices 


52 


Annals 


of the 
Missouri Botanical Garden 


TABLE 1. 
Formation. Figures 


Identification and numerical representation of fossil palynomorphs from the lower Miocene La Boca 
are percentages based on counts of 200 specimens from four samples at locality A (14, 16, 20, 


21) and one sample at locality B (50). Minus sign (—) indicates specimens are present in the sample but were not 
encountered in counts of 200. The counts do not include clusters of 20 or more grains of Rhizophora found in 
sample 14, or Pelliciera in sample 16 


Locality A 


Locality B 


16 


20 


50 


Fungi 


Ascomycete cleistothecium 


Pyrrophyta 
Spiniferit 
Ope 


perc ulodinium centrocarpus l 


Lycopodiaceae 
Lycopodium 

Selaginellaceae 
Selaginella 

Cyatheaceae 
Cyathea 


Pteridaceae 


Pteris type 1 
Pteris type 2 


Vittariaceae 


Cf. Antrophyum = 


Trilete fern spores 


Gramineae 


Palmae 


Attalea type 


Maricaria type 


Synechanthus type = 


Aquifoliaceae 
Ilex 


Bombacaceae 


Cf. Aguiaria 


Cf. Ceiba 


Pseudobombax 


Euphorbiaceae 
Alchornea 


Juglandaceae 


Alfaroa/ Engelhardia l 


1.5 
1.5 


2.5 


Volume 76, Number 1 
1989 


Graham 53 
La Boca Formation—Lower Miocene 
Communities 


TABLE 1. Continued. 


Locality A Locality B 


16 20 21 50 


Leguminosae — Caesalpinioideae 
Crudia - 
Lentibulariaceae 
Utricularia = 
Malpighiaceae 
Type 1 
Type 2 = 
Type 3 
Type 4 = 
Type 5 
Malvaceae 
Rhizophoraceae 
Rhizophora 88 
Rubiaceae 
Type 1 "d 
Type 2 
Theaceae 
Pelliciera 2 


Unknowns 


Type 5 0.5 
Type 6 0.5 


Other unknowns 1.5 


1 0.5 Ex l 
l 0.5 = = 


Lam. and L. linifolium L. presently occurring in 
moist shaded habitats in Panama. They have been 
recovered from the Culebra, Uscari, and Paraje 
Solo formations, although the Uscari specimens 
have thicker walls (3-4 um vs. 1-1.5 um). 


SELAGINELLACEAE 


Selaginella (Fig. 6). Spherical, amb circular 
to oval-triangular; trilete, laesurae frequently ob- 
scured by dense sculpture, arms variously devel. 
oped and spores often appearing monolete, straight, 
narrow, ca. 20-24 um long, extending nearly to 


spore margin; echinate, echinae short (ca. 2-3 
um), occasionally curved, dense, bases broad; wall 
ca. 2 um thick (excluding echinae); 30-32 um. 
Selaginella is widespread in moist, shaded hab- 
itats in the Neotropics. The spores occur in low 
numbers in the Gatuncillo, San Sebastian, Uscari, 
Culebra, Cucaracha, and Paraje Solo formations. 


CYATHEACEAE 


Cyathea (Figs. 10, 13). Amb oval-triangular, 
apices rounded; trilete, laesurae straight, narrow, 
9-21 um long, extending to or nearly to spore 


54 


Annals of th 
Missouri bu m Garden 


Ven, * 


e 
sg rye dur * 


» 
A 
E 


p 5. 
= 


FIGURES 1-7. Fossil spores from the La Boca Formation, Panama. — 1. Ascomycete cisistothéenim. Pan B 27-1, 
ESF C-29, 3-4.— 2, 3. Spiniferites sp., Pan A 14-la, ESF R-46, 1-3.— 4. Operculodinium centrocarpus, Pan A 


Volume 76, Number 1 
1989 


Graham 55 
La Boca Formation—Lower Miocene 
Communities 


margin, inner margin entire, bordered by lip 2-3 
um wide with punctae 1 um diam.; distal surface 
finely punctate, proximal surface more laevigate 
near laesurae; wall 1.5-2 um thick; 32-36 um. 

Tryon & Tryon (1982: 204) noted that the 
classification of tree ferns differs among various 
authors and that the name Cyathea has been used 
to include nearly all members of the family. The 
labels on reference slides used for identification 
often reflect this confusion, especially between Cy- 
athea and Alsophila. A further complication is 
that the lip surrounding the laesurae is difficult to 
observe on some fossil specimens because of pres- 
ervation and/or orientation. According to recent 
illustrations by Gastony & Tryon (1976) and Tryon 
& Tryon (1982), trilete, micropunctate forms with 
a lip often bordered by punctae are referred to 
Cyathea, while forms with a laevigate surface be- 
neath the perine belong to Alsophila. Laevigate 
spores of similar size and morphology lacking the 
lip and bordering punctae are produced by several 
extant genera (e.g., species of Adiantum). Follow- 
ing this classification, Cyathea spores are known 
from the Culebra, Cucaracha, La Boca, and Paraje 
Solo formations (as Alsophila in the latter; Gra- 
ham, 1976, figs. 16-18). Alsophila is known from 
the Gatuncillo Formation (Graham, 1985, fig. 10, 
as trilete fern spore type 1) and from the Paraje 
Solo Formation (Graham, 1976, fig. 19, as Cy- 
athea). 

In Central America Cyathea grows primarily in 
low rain forests, montane forests, and cloud forests, 


usually between 1,500 and 2,000 m 


PTERIDACEAE 


Pteris. 
with some 55 occurring in the Neotropics from 
northern Mexico (Nuevo León) and Florida to Ar- 
gentina and Chile (Tryon & Tryon, 1982: 334). 
It usually grows at altitudes below 2,000 m in wet, 
cloud, or gallery forests. Spores are frequent but 
not abundant in the Gatuncillo, San Sebastian, Us- 
cari, Culebra, Cucaracha, and Paraje Solo for- 
mations. 


Type 1 (Figs. 8, 9). 


ces rounded, margin entire to slightly undulating; 


Pteris is a genus of about 250 species, 


Amb oval-triangular, api- 


trilete, laesurae straight, narrow, inner margin en- 
tire, 23-25 um long, extending to spore margin; 
wall with coarse irregular verrucae on distal sur- 


face, proximal surface more laevigate, hyaline mar- 


ginal flange 12-15 um wide; 62-68 um 


Type 2 (Figs. 11, 12). Smaller (40-42 um), 
the marginal flange narrower (4-5 um wide), and 
the verrucae smaller and more numerous. 


VITTARIACEAE 


Cf. Antrophyum (Fig. 14). Amb triangular, 
apices rounded; trilete, laesurae relatively small in 
relation to spore diam., 
um long, extending ca. 75 distance to spore margin, 
inner margin entire; laevigate; wall ca. 1.5 um 
thick; 50-52 um 

Antrophyum is represented by about 10 species 
in the Neotropics, growing in cloud and rain forests 
usually at elevations between 100 and 1,500 m 
from Mexico (Hidalgo) through Central America 
and the Antilles to northern Argentina and south- 
eastern Brazil (Tryon & Tryon, 1982: 360). The 
spores are similar to those of Acrostichum aureum 
L. (cf. Tryon & Tryon, 1982, figs. 49.10, 12 and 
51.8), a species expected in the La Boca and other 
estuarine formations because it grows in brackish 


straight, narrow, 12-1 


mangrove swamps. The spores of 4. aureum in 
our reference material (nine collections) have a 
granular, scabrate sculpture, while those of An- 
trophyum are more laevigate. The distinction is 
difficult among fossil specimens, and it is possible 
that both genera are represented in Gulf/ Caribbean 
Tertiary deposits. Since the fossils are laevigate, 
however, they are referred to cf. Antrophyum. 
Similar spores are known from the Gatuncillo, San 
Sebastian (not figured in Graham & Jarzen, 1969), 
Culebra, Cucaracha, and Paraje Solo (as an un- 


known in Graham, 1976, fig. 227) formations. 


OTHER TRILETE FERN SPORES 


Several trilete fern spores were recovered for 
which biological affinities could not be determined. 
Three of the more distinctive ones are described 
below. 


Type 1 (Fig. 7). Amb oval-triangular, apices 
rounded, margin entire; trilete, laesurae straight, 
narrow, inner margin entire, 12 um long, extending 
ca. % distance to spore margin; laevigate; wall 1.5 
um thick; 29-32 um. 

These spores are similar to those of several ferns, 
including Adiantum, and cannot be referred to any 


4 1, ESF L-33, 
7. Trilete fern E nm 1, Pan A 16-2, ESF X-4 


—5. Lycopodium, Pan A 21-1, ESF F-11, 3.— 6. Selaginella, Pan A 16-3, ESF P-40, 3.— 
Ld 


56 Annals of the 
Missouri Botanical Garden 


, f T a d n è / 

e: 5 1 4 ' 3 15 17 
FIGURES 8-17. Fossil spores from the La Boca Formation, Panama. 9. Pteris type 1, Pan A 16-1, ESF 

S-35, 1.—10, 13. Cyathea, Pan A 16-1, ESF X-49; Pan A 16-2, ESFL34 1L 12. Pteris type 2, Pan A 16-2, 


ESF K-34.— 14. Cf. Antrophyum, Pan B 50-2, ESF N-14, 2.—15. R fern spore type 2, Pan B 30-3, ESF Q- 
46, 1-2.— 16, 17. Trilete fern spore type 3, Pan A 21-1, ESF W.1 


Volume 76, Number 1 
1989 


Graham 57 
La Boca Formation—Lower Miocene 
Communities 


one genus. They are also known from the Uscari 


sequence of Costa Rica (Graham, 1987a, fig. 30). 
Type 2 (Fig. 15). Amb oval-triangular to cir- 


cular; trilete, laesurae straight, narrow, 17-20 um 
long, extending ca. % distance to spore margin, 
inner margin entire; finely reticulate; wall 1.5 um 


thick; 45-48 


Type 3(Figs. 16, 17). Amb triangular, trilete, 
laesurae straight, narrow, 12-14 um long, ex- 
tending to or nearly to spore margin, prominent 
lip (partially folds?), inner margin entire; distal 
surface divided into low, irregular, verrucaelike 
segments by short, sinuous furrows, proximal sur- 
face more laevigate; wall 2-3 um thick; 31-33 
um. 


MONOLETE FERN SPORES 


Reniform, monolete fern spores are common in 
all Tertiary formations studied for the Gulf/Carib- 
bean region. They are produced by members of 
the Blechnaceae, Polypodiaceae, and Pteridaceae 
and range from Paleozoic to Recent. Five types 
illustrate the range in sculpture and size. Type 1 
Fig. 18; 45 x 32 um) is laevigate; type 2 (Fig. 
19; 70 x 46 um) is finely verrucate; type 3 (Fig. 
20; 69 x 51 um) has numerous, small, densely 
arranged verrucae; type 4 (Fig. 21; 42 x 36 um) 
is coarsely verrucate; and type 5 (Fig. 22; 54 x 
36 um) has prominent, widely scattered, peglike 
verrucae. 


GRAMINEAE 


Spherical, amb circular; monoporate, pore cir- 
cular, 2 um diam., inner margin entire, annulus 
ca. 2 um wide, outer margin entire; tectate, wall 
ca. 2 um thick; psilate to faintly scabrate; ca. 3 
um (folded) (Fig. 23). 


The Gramineae ] and the spec- 


imens cannot be referred to MER single genus. Grass 
pollen continues to be rare or absent in Gulf/ 
Caribbean Tertiary deposits studied to date, with 
only a few grains previously reported from the 
Culebra and Paraje Solo formations. Specimens are 
known in the stratigraphic literature as Monopori- 
tes annulatus and range from the Paleocene (Bra- 
zil) to Recent (Muller, 1981, 1984) 


PALMAE 


Although several surveys of modern palm pollen 
are available (e.g., Ferguson, 1986; Thanikaimoni, 


1970), it is still difficult to identify Gulf/Caribbean 
Tertiary specimens. The pa 
component of the vegetation, and three types are 
recognized for the La Boca Formation. 


Attalea type (Fig. 24). Wedge-shaped, with 
greatest diam. near pole; amb ca. rectangular; 
monocolpate, colpus straight, narrow, 19-22 um 
long, inner margin entire to minutely dentate; sca- 
brate; tectate, wall ca. 1.5 um thick; 43-47 x 
25-29 um. 

Attalea is a genus of about 40 species occurring 
in South America, the West Indies, and the Old 
World tropics, with one species (4. allenii H. 
Moore) listed for Panama (D'Arcy, 1987), where 


it grows at elevations up to ca. 1,000 m 


lms were a prominent 


Manicaria type (Figs. 25-27). | Prolate, with 
greatest diam. at or near equator; monocolpate, 
colpus straight to slightly sinuous, narrow, 30-42 
um long, extending nearly entire length of grain, 
inner margin entire; finely reticulate (diam. of lu- 
men | um or slightly less); tectate-perforate, wall 
ca. 1 um thick; 36-48 x 21-30 um. 

Manicaria is a tree up to 10 m tall, with three 
species in the Antilles, Central America, and South 
America (fide Bailey, 1943), where it often occurs 
in dense groves in wet places. Similar pollen is 
known from the Gatuncillo and Culebra formations. 


Synechanthus type (Figs. 28, 29).  Prolate; 
monocolpate, colpus straight, narrow, 30-42 um 
long, extending entire length of grain, inner margin 
entire; microreticulate; tectate-perforate, wall 1.5— 
2 um thick; 36-48 x 30-38 um 

Synechanthus grows in Panama at low eleva- 
tions in the tropical moist, premontane wet, and 
premontane rain forests (Croat, 1978: 178). Sim- 
ilar pollen occurs in the Culebra Formation. 


AQUIFOLIACEAE 


Ilex (Fig. 31). Oblate-spheroidal, amb oval to 
circular, tricolporoidate, colpi straight, 18 um long, 
equatorially arranged, meridionally elongated, 
equidistant, inner margin diffuse, pores obscure, 
diam. ca. 2-3 um, circular, situated at midpoint 
of colpus; intectate, clavate, wall 3 um thick; 27 
x 18 um 

In Central America /lex commonly grows at mid 
altitudes in moist to slightly drier habitats. It is a 
frequent component of Gulf/Caribbean Tertiary 
microfossil floras in low percentages and has been 
recovered from the Gatuncillo, San Sebastian, Us- 
cari, Culebra, Cucaracha, and Paraje Solo for- 
mations. 


58 Annals 
a eae Garden 


FicunEs 18-33. Fossil mn and pollen from the La Boca Formation, Panama. — 18. Monolete fern spore type 
, Pan A 16-1, ESF D-32, — 19. Monolete fern spore type 2, Pan A 16-1, ESF T-45, 3.— 20. Mo ES igi fern 

spore type 3, Pan A 16-3, ESF J- 43, 3-4.— 21. Monolete fern LIMES type 4, Pan A 14-1a, ESF M-48, 

Monolete fern spore type 5, Pan B 50-2, ESF V-23, 1-2.—23. Gramineae, Pan A 14-1a, ESF W- n $ .— 24. 


Volume 76, Number 1 
1989 


Graha 59 
La dac Formation—Lower Miocene 
Communities 


BOMBACACEAE 


The pollen of several Bombacaceae are distinct 
(Tsukada, 1964; Nilsson & Robyns, 1986), but 
other generalized types cannot be referred to any 
one genus (see listings in Muller, 1981: 46-48). 
There is also some overlap with the closely related 
families Sterculiaceae and Tiliaceae, and in fossil 
deposits types are recovered that do not match 
exactly any extant members of the complex (Gra- 
ham, 1976, figs. 248-255). It is possible that some 
unidentified pollen represents extant species not 
yet collected or extinct taxa from early speciation 
after introduction into the Caribbean region. Fur- 
ther studies are needed on the pollen morphology 
of modern forms, including newly described taxa, 
and a complete assessment should be made of fossil 
types. 

Bombacaceae are prominent members of neo- 
tropical forests, and the pollen is frequent but not 
abundant in Tertiary sediments. Muller (1981: 46- 
48) listed the oldest occurrence of this family as 
Maestrichtian (uppermost Cretaceous) from the 
southeastern United States and cited other records 
supporting Wolfe's (1975) suggestion that the fam- 
ily originated in eastern North America and sub- 
sequently spread to South America and Africa where 
it diversified. In the Caribbean region it first ap- 
pears in the Paleocene of northern South America 
(Bombacacidites annae = Bombax ceiba type; 
Germeraad et al., 1968: 277), and in our material, 
pollen of the family occurs in the San Sebastian 
Formation and Uscari sequence. 


Cf. Aguiaria (Fig. 30; cf. modern pollen illus- 
trated by Nilsson & Robyns, 1986, fig. 7f, 
g). Oblate, amb circular; tricolpate, colpi short 
(8-10 um apex to equator), equatorially arranged, 
meridionally elongated, equidistant, inner margin 
minutely dentate, bordered by narrow margo ca. 

.5-2 um wide; finely reticulate; tectate-perforate, 
wall ca. 2 um thick; 30-34 um. 


Cf. Ceiba (Fig. 34; cf. modern pollen illustrated 
by Nilsson & Robyns, 1986, fig. 11I). Oblate, 
amb circular; tricolpate, colpi short (13-15 um 
apex to equator), equatorially arranged, meridio- 
nally elongated, equidistant, inner margin entire to 
minutely dentate, bordere 
2-3 um wide; reticulate, muri smooth, slightly bu 


by narrow margo c 


uous, narrow (ca. 0.5-1 um), lumina polygonal, 
4-5 um diam. at poles, smaller toward equator; 
tectate-perforate, wall ca. 1.5 um thick; 56-60 
um. 


Pseudobombax (Fig. 35; cf. modern pollen il- 
lustrated by Nilsson & Robyns, 1986, fig. 
61). Oblate, amb triangular, sides slightly con- 
cave, apices rounded; tricolpate, colpi short (8-10 
um apex to equator), equatorially arranged, meridi- 
onally elongated, equidistant, inner margin entire, 
bordered by faint narrow margo ca. 1 um wide; 
reticulate at poles, muri smooth, straight to oc- 
casionally curved, narrow (ca. 1 um), lumina po- 
lygonal, ca. 3 um diam., becoming minutely retic- 
ulate around apices; tectate-perforate, wall 1.5-2 
um thick; 54-56 um 

Pseudobombax is a tree to 25 m tall distributed 
from Nicaragua to Brazil and Peru. In Panama it 
typically occurs at lower elevations in the tropical 
moist forest but is also found in the premontane 
moist, tropical wet, and locally from the tropical 

). 


dry forest (Croat, 1978: 591 


EUPHORBIACEAE 


Alchornea (Fig. 32). Oblate, amb circular; tri- 
colpate, colpi straight, 6-8 um long (apex to equa- 
tor), equatorially arranged, meridionally elongated, 
equidistant, extending within 6-7 um of pole, inner 
margin entire, operculum distinct; pistei to faintly 
scabrate; tectate, wall 1.5 um thick; 1 

Pollen of Alchornea is frequent in low numbers 
in Gulf/Caribbean Tertiary deposits. It ranges from 
the lower and middle Eocene (Colombia; González 
Guzmán, 1967) to Recent (Muller, 1981). Its ecol- 
ogy and distribution have been summarized by Gra- 
ham (1987a) based on Croat (1978) and Webster 
& Burch (1967). In Central America Alchornea 
grows in the tropical moist, premontane wet, and 
premontane rain forest typically at elevations be- 
tween 300 and 2,000 m 


JUGLANDACEAE 

Alfaroa/Engelhardia (Fig. 33). Oblate, amb 
oval-triangular; triporate, pores circular, ca. 2 um 
diam., 
equidistant; psilate; tectate, wall 1.5-2 um thick; 


inner margin entire, equatorially arranged, 


E 


Attalea kip Pan A 16-2, ESF U-38, 2.— 25-27. Manicaria type, Pan A 14-1a, ESF U-34, 1-3; Pan A 20-2, 
ES 


F Y.22, 2-4; Pan A 20-2, ESF G-19, 4.— 28, 29. 


16-1, 1-3.— 30. 


ESF 1.46, Cf. Aguiaria, Pan A 163. ESF D-42. 
Alchornea, Pan A 16-3, ESF E-33, 2.— 33. Engelhardia, Pan A 20-2, ESF T-43, 


Synechanthus type, Pan A 16-1, ESF D-44, 1-3; Pan A 


— 31. Ilex, ju A 16-3, ESF K-46.— 32. 


60 Annals of the 
Missouri Botanical Garden 


FicuRES 34-46. Fossil pollen from the La Boca pius ree — 34. Cf. Ceiba, re = r^ ja ESF O- a 
2-4.—35. Pseudobombax, Pan A 20-2, ESF N-36, 36. Utricularia, Pan A 14-1 =87, 
Crudia, Pan A 16-3, ESF X-31; Pan A 16-1, ESF x. at. 1-8. —39. Malpighiaceae s E Pan A 16- 3, ESF 

39, 2.— 40. Malpighiaceae type 2, Pan B 50-2, ESF L-36, 1-2.— 41. Malpighiaceae type 3, Pan B 50-2, ESF 


Volume 76, Number 1 
1989 


Graham 61 
La Boca Formation—Lower Miocene 
Communities 


These trees are distributed from Mexico through 
Central America, usually associated with lower- to 
mid-altitude temperate forests. Pollen occurs in the 
Gatuncillo, San Sebastian, Cucaracha, and Paraje 
Solo formations. 


LEGUMINOSAE—CAESALPINIOIDEAE 


Crudia (Fig. 37, 38). Prolate; tricolporoidate, 
colpi narrow, straight, 25 um long, extending near- 
ly entire length of grain, equatorially arranged, 
meridionally elongated, equidistant, pore area faint, 
circular, situated at midpoint of colpus; distinctly 
and coarsely striate, striae generally oriented par- 
allel to long axis of grain, surface psilate, margins 
entire, occasionally appearing beaded from under- 
lying pores in foot layer /endexine; tectate but with 
occasional separation between sculpture elements, 
wall 1.5 um thick; 42-48 x 30-32 um. 

Crudia is a South American, mainly Amazonian, 
riverine tree of low altitudes (Cowan & Polhill, 
1981: 131). Pollen ranges from the Eocene in 
northern South America (as Striatocolpites ca- 
taumbus) and from the Paleocene in Africa (Mul- 
ler, 1981). It occurs in Central America in the 
Gatuncillo and Cucaracha formations. Modern pol- 
len has been studied by Graham & Barker (1981). 


LENTIBULARIACEAE 
Utricularia (Fig. 36). Oblate, amb circular; 
stephanocolpate, colpi 12-16, equatorially ar- 


ranged, meridionally elongated, equidistant, 15- 
18 um long, extending within 8-10 um of pole, 
inner margin entire; psilate to faintly scabrate; 
tectate, wall 2 um thick; 34-36 um 

Utricularia is an annual or stoloniferous peren- 
nial, insectivorous, aquatic herb growing along the 
margins of freshwater lakes and rivers, swamps, 
and marshes from Florida to Mexico and south 
through the Antilles, Central America, and South 
America. It is represented by about 12 species in 
Panama (Taylor, 1976). Pollen also occurs in the 
Paraje Solo Formation. 


MALPIGHIACEAE 


Representation of the Malpighiaceae in the Gulf/ 
Caribbean Tertiary is analogous to the Bombaca- 
ceae, wherein several types are present but many 
do not match exactly modern reference material 


or species illustrated in the literature. Generally 
similar modern forms include Banisteria, Banis- 
teriopsis, Bunchosia, Hiraea, Malpighia, and 
Mascagnia; but since the specimens are slightly 
different, they are referred to types 1- 


Type 1 (Fig. 39). Spherical, amb circular; 
periporate, pores circular, 3-4 um 
margin entire; scabrate; tectate, wall 5-6 um thick; 

2 um. 

The distinguishing features of this grain are the 
very thick wall and the absence of colpi connecting 
the pores. 


Type 2 (Fig. 40). Spherical, amb circular; 


periporate, pores circular, 3 um diam., inner mar- 


diam., inner 


gin entire, faint colpi with diffuse, granular margins 
UN the pores; scabrate; tectate, wall 2 um 
thick; 35 y 

These specimens have moderately thick walls 
and faint connecting colpi. 

Type 3 (Fig. 41). 
periporate, pores circular, 1.5- 
margin entire, colpi with diffuse, E uir margins 


E amb circular; 
iam., inner 
connecting the pores; scabrate, some scabrae point- 
ed; tectate, wall 2 um thick; 36 um 

The distinguishing features of type 3 are the 
moderately thick wall, distinct colpi, and some sca- 
brae with pointed apices. 


Type 4 (Fig. 42). 
periporate, pores circular, 3-4 um 
margin entire, colpi with diffuse granular margins 
connecting the pores; scabrate; tectate, wall 5 um 
thick; 45 um. 

These specimens differ from the thick-walled 
type 1 by having distinct colpi. 


Type 5 (Fig. 43). Spherical, amb circular; 


periporate, pores circular, 1.5 um diam., colpi with 


Spherical, amb circular; 
diam., inner 


diffuse granular margins connecting the pores, col- 
pi and pores numerous, giving scalloped appear- 
ance to outer margin of grain; psilate; tectate, wall 
5-6 um thick; 37 um 

These thick-walled specimens have numerous 
pores and colpi, and they are psilate. 


MALVACEAE 


Spherical, amb circular; apertures obscure, peri- 
porate(?), pores circular, small (ca. 1.5 um), evenly 


== 


E-27, 2-4.—42. Malpighiaceae type 4, Pan A 16-3, ESF P-43, 2-4. 
21-1, ESF R-21, 2-4. 
l 


Ade. 1.—44. Malvaceae, Pan A 
— 46. Rubiaceae type 2, Pan A 20-2, ESF L-31, 


—43. Malpighiaceae type 5, Pan A 16-3, 
— 45. Rubiaceae type 1, Pan A 20-2, ESF W-20, 1- 


62 Annals of the 
Missouri Botanical Garden 


FIGURES 47-61. Fossil pollen from the La Boca Formation, Panama. — 47. Rhizophora, Pan B-50, 2, ESF X- 
33, 2.— 48, 49. Pelliciera, Pan A 5-2, ESF N-39, 1-2; Pan A 16-3, ESF V-44, 3-4.—50. Unknown 1, Pan A 


6, Pan A 14-la, ESF Q-30. —57. Unknown 10, Pan A 14-1a, ESF 6-29, 2-4.— 58. Unknown 11, Pan B 50-2, 
ESF V-13, 3-4.— 59. Unknown 7, Pan A 16-3, ESF Q-47, 2.—60. Unknown 8, Pan B 50-2, ESF F-33, 2.— 61. 
Unknown 9, Pan A 16-1, ESF G-45. 


Volume 76, Number 1 
1989 


Graham 63 
La Boca Formation—Lower Miocene 
Communities 


distributed, inner margins entire; echinate, echinae 
hyaline, straight, broad at the base, densely ar- 
ranged, 5 um long, decreasing to 2-3 um on por- 
tions of the exine; psilate; tectate, wall 1.5-2 um 
thick; 30 um (Fig. 44). 

This unusual specimen may be an aberrant form 
and cannot be identified to genus. The Malvaceae 
are represented in the Caribbean Tertiary by Ham- 
pea/ Hibiscus (probably Hibiscus tiliaceus L.) in 
the Culebra Formation and are frequent in the 
Tertiary of northern South America (as Echiperi- 
porites estelae, late Eocene to Recent). 


RHIZOPHORACEAE 


Rhizophora (Fig. 47). 
spheroidal; tricolporate, colpi straight, 14-16 um, 
equatorially arranged, meridionally elongated, 
equidistant, inner margin entire, costae colpi ca. 
2-3 um wide, pores elongated equatorially (colpi 
transversalis), 1 X 4 um, constricted at midpoint 
of colpus; finely reticulate; tectate-perforate, wall 
2-3 um thick; 16-20 x 14-18 um 

Pollen of Rhizophora has been recovered from 
the Gatuncillo, San Sebastian, Uscari, Culebra, Cu- 
caracha, and Paraje Solo formations and comprises 
90% or more of some samples. Its fossil record 
has been discussed by Langenheim et al. (1967), 
Leopold (1969), and Muller & Caratini (1977), 
and its occurrence in the Gulf/Caribbean Tertiary 
was summarized by Graham (1985: 519, 1987a). 


Rhizophora is indicative of coastal, brackish-water 


Prolate to prolate- 


conditions in tropical to subtropical regions. It is 
known in the stratigraphic literature as Zonocos- 
tites ramonae and ranges from the late Eocene to 
Recent. In older Tertiary deposits in Latin America 
it is replaced by its presumed ecological equivalent 
Brevitricolpites of unknown biological affinity. 


RUBIACEAE 


The Rubiaceae are represented in the La Boca 
Formation by two similar pollen types typical of 
several extant genera. These tricolpate, densely 
sculptured forms are produced by Anisomeris, 
Chomelia, Guettarda, Terebraria, and others. 


Type 1 (Fig. 45). Oblate to oblate-spheroidal, 
amb circular; tricolpate, colpi equatorially ar- 
ranged, meridionally elongated, equidistant, short 
(4-5 um apex to equator), often obscured by dense 
sculpture; reticulate, muri 1.5 um wide, smooth, 
straight to occasionally curved or slightly sinuous, 
lumina 2-3 um diam.; tectate-perforate, wall 2 um 


thick; 26-36 um. 


Type 2 (Fig. 46). Similar to type 1 but is 
slightly larger (42-46 um), and the muri are thick- 
er (ca. 2-2.5 um 


THEACEAE 


Pelliciera (Figs. 48, 49). Oblate, amb circu- 
lar; tricolporate, colpi equatorially arranged, meri- 
dionally elongated, equidistant, 20-26 um, taper- 
ing to acute apex, inner margin entire, pore circular, 
3-4 um diam., situated at midpoint of colpus, inner 
margin entire; sculpture variable from finely to 
coarsely verrucate; tectate, wall 4 um diam.; size 
variable, 40-60 um 

Pelliciera is a small mangrove tree of lowland 
coastal areas from Costa Rica to northwest Colom- 
bia and Ecuador. In the Tertiary, however, it was 
widespread in the Gulf/Caribbean and is known 
from the lower middle Eocene Yellow Limestone 
Group of Jamaica, the middle(?) to late Eocene 
Gatuncillo Formation of Panama, the middle to late 
Oligocene San Sebastian Formation of Puerto Rico, 
the Oligo-Miocene Simojovel Group of Mexico, and 
from several localities in northern South America. 
Several anther-size clusters or fragments (Fig. 49) 
were recovered, indicating the plants grew in the 
immediate vicinity of the depositional basin. The 
presence of Pelliciera pollen in the La Boca For- 
pon is one of the principal a A between 


1l Culebra and C 


In the e record it is known as Psila- 
tricolporites crassus and ranges from the Eocene 
to Recent. Other details of the ecology and geologic 
record of Pelliciera have been summarized by 


Graham (1977). 


UNKNOWNS 


A number of specimens were recovered that 
could not be identified. Some of the more distinctive 
and/or abundant ones are illustrated and briefly 
described below. 


Unknown 1 (Fig. 50). Spherical, amb circu- 
lar; apertures obscure (nonaperturate?); echinate, 
echinae straight to slightly curved, dense, 2-3 um 
long; wall 1.5 um thick; 16 um 


Unknown 2 (Fig. 51). This type is similar to 
unknown 1 but is slightly larger (22 um), and the 
spines are shorter (1 um) and blunt. 


Unknown 3(Fig. 52).  Prolate, amb oval; mono- 
colpate, colpus straight, 22 um long, inner margin 
minutely dentate; reticulate, muri straight, smooth, 
1.5 um wide, lumina 1-1.5 um diam., reticulum 


Annals of the 
Missouri Botanical Garden 


becoming finer toward poles; tectate-perforate, wall 
2 um thick; 36 x 27 um. 

These monocot pollen grains may be palms but 
no exact match has been foun 


Unknown 4 (Fig. 53). Prolate, amb oval; 
monocolpate, colpi straight, 28 um long; reticulate, 
reticulum deep (columellae 3 um long), 
straight, smooth, 1 um wide, lumina 2 um diam.; 
tectate-perforate, wall 3-4 um thick; 40 x 30 
um. 


Unknown 5 (Fig. 54). 
monocolpate, colpus 42 um long, extending entire 
length of grain, inner margin entire; echinate, 
echinae straight, 1.5 um long, moderately dense; 
tectate, wall 1-1.5 um thick; 45 x 24 um 


Unknown 6 (Figs. 55, 56).  Oblate-spheroidal, 
amb circular; tricolpate, colpi often obscured by 
coarse sculpture elements, short (5-6 um long), 
equatorially arranged, meridionally elongated, 
equidistant; coarsely verrucate; tectate, wall 3 um 
thick; 22-26 um. Similar specimens occur in the 
Paraje Solo Formation. 


Unknown 7 (Fig. 59).  Oblate-spheroidal, amb 
circular; tricolpate, colpi straight, 15 um long, 
inner margin lobate; reticulate, muri straight, 
smooth, 2 um wide, lumina polygonal, 4 um diam., 
becoming finer toward colpi forming a margo, re- 
ticulum deep (columellae 5 um long); tectate-per- 
forate, wall 6 um thick; 45 um 


Unknown 8 (Fig. 60). 
lar; periporate, pores circular, 3 um diam., widely 
spaced, inner margin entire; scabrate; tectate, wall 
2 um thick; 43 um. 


Unknown 9 (Fig. 61). Oblate, amb oval.tri- 
angular; tricolpate, colpi short (4 um apex to equa- 
tor), equatorially arranged, meridionally elongated, 
equidistant, inner margin minutely dentate; echi- 
nate, echinae short (0.5 um), moderately dense, 
bases forming subreticulum; tectate, wall 1 um 
thick; 40 um 


Unknown 10 (Fig. 57).  Oblate-spheroidal, amb 
circular; stephanocolpate, colpi 6, straight, equa- 
torially arranged, meridionally elongated, equidis- 
tant, inner margin entire; scabrate; tectate, wall 
um thick; 16-21 um 

Similar specimens occur in the Culebra For- 


muri 


Prolate, amb oval; 


Spherical, amb circu- 


Unknown 11 (Fig. 58). Oblate-spheroidal, amb 
circular; tricolpate, colpi straight, tapering to acute 
apex, 6-8 um long, inner margin entire to minutely 
dentate, equatorially arranged, meridionally elon- 


gated, equidistant; finely reticulate; tectate-perfo- 
rate, wall 2 um thick; 27 um 


Unknown 12 (Fig. 62). Prolate, amb oval; 
tricolporate, colpi straight, 45 um long, equato- 
rially arranged, meridionally elongated, equidis- 
tant, bordered by costae colpi 4-5 um wide, inner 
margin entire, pores large, oval, 5 X 16 um, sit- 
uated at midpoint of colpus, inner margin entire; 
reticulate; tectate-perforate, columellae relatively 
coarse and clearly evident in median optical sec- 
tion, wall 2.5-3 um thick; 58 x 36 um. 

Unknowns 12-15 are similar to pollen of several 
Anacardiaceae and Euphorbiaceae but cannot be 
referred to any one modern genus. 


Unknown 13 (Fig. 63). This type is similar to 
unknown 12, but the columellae are shorter and 
finer; consequently, the wall is thinner (1.5-2 um) 
and the reticulum finer. 


Unknown 14 (Fig. 64). This type has an even 
thinner wall (1-1.5 um), a finer reticulum, nar- 
rower costae colpi, and smaller and slitlike pores 


(2x4 


Unknown 15 (Fig. 65). This type is similar to 
unknown 14 but is smaller (27-32 x 18-21 um). 


PALEOCOMMUNITIES AND PALEOENVIRONMENTS 


Results of the La Boca study are of interest in 
demonstrating an internal consistency in paleocom- 
munities, climates, and physiography among four 
fossil floras in two countries of Central America, 
as well as consistency with independent plate tec- 
tonic and paleotemperature data. The lower Mio- 
cene Uscari (Costa Rica), Culebra, Cucaracha, and 
La Boca (Panama) floras preserve elements of a 
mangrove swamp (Pelliciera, Rhizophora) fring- 
ing the volcanic islands constituting the isthmian 
region during most of the Tertiary. Freshwater 
marshes and swamps (palms, ferns, Utricularia) 
occupied lowland areas behind the mangrove zone. 
On the adjacent slopes were versions of the tropical 
wet, tropical moist, and premontane forests (Ly- 
copodium, Selaginella, Cyathea, Pteris, Ilex, 
Pseudobombax, Alchornea, and Crudia, with Al- 
faroa/ Engelhardia likely occupying distant, mod- 
erate highlands in the region). All available paleo- 
botanical data are consistent in showing little 
evidence of dry (including savannah) habitats. Grass 
pollen continues to be poorly represented (maxi- 
mum of 156 in one sample from the Culebra For- 
mation; less than 0.5% in two samples from the 
La Boca) to absent (Uscaria, Cucaracha forma- 
tions). If savannahs existed, as suggested by the 


Volume 76, Number 1 
1989 


Graham 65 
La Boca Formation—Lower Miocene 
Communities 


FicunEs 62-65. 
X-14, 3.— 63. Unknown 13, Pan A 
Unknown 15, Pan A 16-2, ESF W.36, 2-4 


grazing and browsing components of the Tertiary 
faunas, they were probably short-lived and devel- 
oped locally as recovery vegetation in response to 
volcanic activity documented for central Panama 
in the form of extensive ash, tuff, and basalt de- 
posits (see further discussion in Graham, 1988b). 

he affinities of the La Boca flora are clearly with 
Central America and areas to the north, reflecting 
the establishment of a land connection between 
Central America and South America only about 3 
Ma. 

The lack of any paleobotanical evidence for sig- 
nificant highlands is in agreement with plate tec- 
tonic models suggesting that elevation of the early 
volcanic islands and peninsulas gradually increased 
during the latter part of the Cenozoic, and that the 
highest elevations (3,475 m in the western moun- 
tains of Panama; 3,820 m, Cerro Chirripo, Costa 
Rica) are of recent origin. Altitudes up to about 
1,200-1,500 m would accommodate all elements 
in the four lower Miocene floras known from south- 
ern Central America. No pollen was being blown, 
washed, or transported into the depositional basin 
to suggest dry vegetation or high altitudes in the 
Gatuncillo, Uscari, Culebra, Cucaracha, or La Boca 
formations of Costa Rica and Panama. Muller 
(1959) demonstrated that pollen from a variety of 
inland and upland communities is transported by 
rivers and deposited in coastal basins. Palynological 
data from Puerto Rico and southern Mexico further 
show that many of these habitats, represented by 
both wind- and insect-pollinated elements, are in- 
cluded in the microfossil record if such habitats 
exist in the region. 


Fossil pollen from the La Boca Formation, Panama.— 62. 
16-1, ESF Q-44, 1-3.— 


Unknown 12, Pan A 21-1, ESF 
64. Unknown 14, Pan A 16-3, ESF W-33, 4.— 65. 


All taxa identified from the La Boca Formation 
except Crudia occur in the modern vegetation of 
Panama. Consequently the paleoenvironment must 
have been similar to the tropical conditions pres- 
ently prevailing in coastal, lowland, and moderate- 
altitude habitats in southern Central America. This 
is consistent with paleotemperature curves (Savin, 
19777; Savin & Douglas, 1985; Savin et al., 1975) 
that show the Uscari, Culebra, Cucaracha, and La 
Boca floras were deposited een a iol warm 
interval just before the si 
in middle and late Miocene times. 


LITERATURE CITED 


BaiLgv, L. H. 1943. Palmaceae. In: R. E. Woodson 
& R. W. Schery (editors), Flora of Panama. Ann. 
Miseouri Bot. Gard. 30: 327-3 

Cowan, R. S. & R. M. PoLniLL. . Detarieae. Pp. 
117-134 in R. M. Polhill & P. H. Raven (editors), 
Advances in Legume Systematics. Royal Botanic Gar- 


ora of cd m" Island. 


ora of Bt Checklist and 
kin dex. Monogr. Syst. Bot. Missouri Bot. Gard., Vol- 


Bici I. K. 1986. Observations on the variation 
in pollen EE of Palmae and its significance. 
anad. J. Bot. 64: 3079-3090. 

Gastony, G. J. & R. Takon. 1976. Spore morphology 

in the Cyatheaceae II. The genera Lophosoria, Me- 

taxya Sphaeropteris, Alsophila, and Nephelea. 
Amer. J. Bot. 63: 738-758. 

Nen J. H., C. A. Hoppinc & J. MULLER. 1968. 
Palynology of Tertiary sediments from tropical areas. 

alaeobot. Palynol. 6: 189-34 
GONZÁLEZ GUZMÁN, A. E. 1967. A palynological study 


66 


Annals of the 
Missouri Botanical Garden 


n the upper Los Cuervos and Mirador formations 
(lower and middle Eocene; Tibú area, Colombia). E 
J. Brill, Leiden. 
GRAHAM, A. 1976. Studies in ——Ó p 
I. The Miocene communities of Ver z, Mexi 
Ann. Missouri Bot. Gard. 63: 787- 842. 

1977. New records of Pelliceria (Theaceae/ 
Pollicerincess) 3 in the Tertiary of the Caribbean. Bio- 
tropica 9: 48-52. 

1985. Studies in Ta paleobotany. IV. 
of Panama. Ann. Missouri 


The Eocene communities 
Bot. Gard. 72: 504-534 
87a. Miocene communities and paleoen- 
vironments of southern Costa Rica. Amer. J. Bot. 
74: 1501-1518 
1987b. Tropical American floras and paleoen- 
vironments: Mexico, Costa Rica and Panama. Amer. 
J. Bot. 74: 1519-1531. 
1988a. Studies in neotropical paleobotany. V. 
The lower Miocene communities of Panama — the 
Culebra Formation. Ann. Missouri Bot. Gard. 75: 
1440-1466. 


. Studies in neotropical paisouoteny, 
VI. The lower Miocene communities of Panama 
the erario Formation. Ann. Missouri Bot. Gard. 
75: 1467-1479. 


P" BARKER. 1981. Palynology and tribal 
classification in the R panda Pp. 801-834 
. lhill & P. H. Raven (editora), Advances 
in Legume eines Royal tanic Gardens, Kew. 
& D. M. Janzen. 1969. Studies in neotropical 
paleobotany. I. The Oligocene communities of Puerto 

ico. Ann. Missouri Bot. ai 08- 

— ———, R. H. Stewart & J. L. STEWART. 1985. Stud- 
ies in ^g: seg n III. The Tertiary com- 
munities of — geology of the pollen bearing 

i : 485-503. 


sediments. a 

HARTSHORN, G. S. 1983. . 118-157 in D. 
H. Janzen (editor), Costa Rican Natural History. Univ. 

hicago Press, 

HOLDRIDGE, L. R. 1 Determination of world plant 
formations from simple climatic data. Science 105: 
367-368. 


, W. C. GRENKE, W. s HATHEWayY, T. 2 
& J. A. Tosi 1971, Forest Environments in Tro 
2 Life Zones: A Pilot Study. Pergamon Press, Ga 


ERE J. H., B. L. HACKNER & A. H. BARTLETT. 
Mangrove pollen at the depositional site of 
r from Chiapas, Mexico. Bo 


LEOPOLD, E. B. Miocene pollen and spore flora 
of Eniwetok Atoll, Marshall Islands. Profess. Pap. 
. Geol. Surv. 260-II: 1133-1185. 
MULL, J. 1959. Palynology of Recent Orinoco delta 
shelf sediments: reports of the Orinoco shelf 
expedition 5. Micropaleontology 5: 1-32 
1981. Fossil pollen records E extant angio- 
sperma. Bot. Rev. (Lancaster) 47: 1-142. 


1984. Significance of fossil pollen for angio- 
sperm history. Ann. Missouri Bot. Gard. 71: 419- 


C. CARATINI. 1977. Pollen of Rhizophora 
(Rhizophoraceae) as a guide fossil. Pollen & Spores 
61-389. 


NILSSON, S. & A. Ropyns. 1986. Bombacaceae Kunth. 
Pp. 1-59 in S. Nilsson (editor), World Pollen and 
Spore Flora 14. Almqvist & Wiksell, Stockholm. 

SAVIN, S. M. 77. The history of the Earth's surface 


; 1985. Sea level, climate, 
and the Central vd. land bridge. Pp. 303-324 


in F. G. Stehli Webb (editors), The tu 
American Biotic Interchange. Plenum, New : 
, —— —— & F. G. STEHLI. 1975. Ferry 


marine paleotemperatures. Bull. Geol. Soc. Am 


86: 1499-1510. 
STEWART, R. H. & J. L. STEWART (with the ro 
of W. P. Woodring). 1980. Geologic Map o 


Panama Canal and Vicinity, Republic of ien 

Scale 1:100,000. U.S. Geol. Surv. Misc. Invest. Map 

I-1 232. [Map also included in ee 1982, Pro- 

fess. Pap. U.S. Geol. Surv. 3 

Tivos, P. 1976. hu. In: R. E. Woodson 
& id Ar Schery (editors), Flora of Panama. Ann. 

Missouri Bot. Gard. 63: 565-580. 

A NONE 1970. Les palmiers: palynologie et 

systé matique. Pu Frang. Pondichéry, Trav. Sect 
ci. Tech. XI: 1-286. 

Tryon, R. M. AE. TRYON. 1982. Ferns and Allied 
Plants, with Special Reference to Tropical America. 
Springer-Verlag, New York. 

TsuKADA, M. 1964. Pollen morphology and identifi- 
cation III. Modern and fossil tropical pollen with 
emphasis on Bombacaceae. Pollen & Spores 6: 393- 


VAN DEN BoLD, W. A. 1972. Ostracoda of the La Boca 
Formation, Panama Canal Zone. Micropaleontology 
18: 410-442. 

La posición estratigráfica de la for- 
n La Boca, oe a del Canal. Publ. 
Ge in  ICAITI 4: 167-17 

WEBSTER, G. D. in 1967. Euphorbiaceae. 
In: R. E. Woodson 8: R. W. Schery la Flora 

of Panama. Ann. Missouri Bot. Gard. 54: ; 

Wii J. A. Some aspects of plant geography 
of the northern. hemisphere during the late Creta- 
ceous and Tertiary. Ann. Missouri Bot. Gard. 62: 

279, 

Wooprinc, W. P. 1957-1982. Geology and pom 
tology of Canal Zone and adjoining parts of Panam 
Profess. Pap. U.S. Geol. Surv. 306A-F. 

Has R. E. & R. W. ScHERY. 1945- 1980. Flora 

nama. Ann. Missouri Bot. Gard. 30 (1943) et 


A REVISION OF 
MESOAMERICAN 
PSYCHOTR [A 
SUBGENUS PSYCHOTR IA 
(RUBIACEAE), PART I: 
INTRODUCTION AND 
SPECIES 1-16! 


Clement W. Hamilton? 


ABSTRACT 


Psychotria subgenus Psychotria (Rubiaceae) comprises 61 species and eight varieties of understory shrubs and 
Cent 


enus is most diverse in Panama (40 s 


pecies) and Costa Rica 


small trees in Mexico and Central America. The subg 
(32 species), with a secondary center of diversity in southern Mexico (26 Ew Eu and "rm (26 cies). 
Endemism is also greatest in Pan (15 species). Species-rich regions in Cos Ri and EM ma are Beier c 


species complexes contain on 


nging species and several species with m 


uch dents EN Only one species- 


rich group, the P. calophylla group, lacks a continuously wide-ranging member. 


Psychotria is a pantropical and subtropical ge- 
nus of mostly shrubs but also small trees, herbs, 
and even epiphytes. I presently estimate the num- 
ber of species to be 1,650, making it the largest 
zenus in the Rubiaceae; this includes 700 species 
in the Neotropics (my estimate), 250 in Africa 
(Petit, 1964, 1966), and 700 in Asia and Oceania 
(Sohmer, pers. comm.). 

The present state of its infrageneric taxonomy 
reflects the lack of understanding of the genus as 
a whole; all that can be said with some certainty 
is that subg. Psychotria occurs in all three tropical 
regions and that each region has one or more 
groups distinguishable from subg. Psychotria. In 


Tetramerae (Hiern) Petit (1964, 1966); and in 


Asia and Oceania, the situation is much more com- 
plex (Sohmer, 1977). 

The infrageneric uncertainty extends to delim- 
itations of Psychotria and related genera; the neo- 
tropical genera Mapouria, Cephaelis, Uragoga, 
Naletonia, Nonatelia, and Notopleura have been 
variously merged with or separated from Psycho- 
tria. Most of these genera, if included in Psycho- 
tria, fall within subg. Heteropsychotria and are 
therefore not relevant here. Mapouria, named by 
Aublet in 1775, is more critical. The principal use 
of that generic name was by Mueller-Argoviensis 
(1876, 1881), who in his Rubiaceae treatment for 
Flora Brasiliensis distinguished between Ma- 
pouria and Psychotria; the former corresponded 
to what is properly considered Psychotria subg. 
Psychotria, while the latter encompassed Steyer- 


This paper is an amended portion of my 


3 pap Ph.D. 
the Missouri Botanical Garden. I thank de Division of Biolog 
tany Department of the Missouri agg Garden for Daoa and 


Biology at 
financial support during my 
E ken and to ee Krager for 


, D. Lorence, J. Steyermark, and 


ashington University and the 
study. 


Bot 
Special thanks are due 


overall id da 
C. Ta 


paa at Washington University, St. Louis, Missouri, and 
De 


cal and Biomedical Sciences and the artment of 
o A. Gentry, P. Raven, B. Schaal, and R. Foster for 
any of the ipie Bose on Psychotria 


r were very helpful. Comments on the manuscript by G. 


s: S: died and J. H. Kirkbride } have improved it mici The following persons Miré helped me significantly 
: C. & R. Ha 


at various stages of my overall study 
Wi Wes P Congdon, P. Stev im Bisnes Sirirugsa, 

r, J. Miller, P. P. Lowry, P. Hoch, W. 
J. Fishe er, D. Nicolson: and B. Hamm 
rovided material for study: the Harvard Herbaria (A, 


SLPM, TEFH, and US. 


milton, B. Tiffney, the late E. S. Bar 
W. H. Lewis, A. Templeton, G. Johnson, G. Allen, D. Neill, 
Hahn, L. Hickey, J. Beach, w D'Arcy, H. Stockwell, C. Augspurger, 
el. I appreciate also the efforts and patience of those institutions that have 


A, ECON, GH), B, CR, DUKE, ENCB, F, K, LL, MEXU, NY, 


hoorn, S. J. Gould, D. 


? Center for Urban Horticulture, GF-15, University of Washington, Seattle, Washington 98195, U.S.A. 


ANN. 


Missouni Bor. GARD. 76: 67-111. 1989. 


Annals of the 
Missouri Botanical Garden 


mark's Psychotria subg. Heteropsychotria, in- 
cluding Cephaelis. The discovery by Petit (1964), 
reaffirmed by Steyermark (1972), that the type 
species of Psychotria, P. asiatica L., falls well 
within the Mapouria generic concept makes that 
genus obsolete. 

Members of Psychotria subg. Psychotria are 
distinguished easily from all others in the genus. 
Three characters serve to enable recognition of the 
subgenus: (1) the leaves are dry red-brown or some- 
times dull green, as opposed to bright green; (2) 
the stipules are usually sheathing and caducous, 
leaving a small, usually pale ridge often with a 
fringe of red-brown hairs, as opposed to only slight- 
ly sheathing and conspicuously persistent; and (3 
the seeds, hemispherical in cross section, have two 
longitudinal furrows on the flat ventral surface and 
usually four or five (or sometimes an irregular 
number of) longitudinal furrows on the rounded 
dorsal surface. Psychotria subg. Heteropsycho- 


tria has a variety of seed cross sections, none of 


which can be confused with those of subg. Psy- 
chotria. Additional characters of whole-plant ar- 
chitecture show differentiation between the tw 

neotropical subgenera (Hamilton, 1985), as ie 
mature fruit color, which is almost always red in 
subg. Psychotria and usually blue in subg. Het- 
eropsychotria. The naturalness and monophyletic 
(or at least ““convex””; see Meacham & Duncan, 
1987) nature of subg. Psychotria is not in doubt. 

In the Neotropics, Psychotria subg. Psychotria 
consists of ca. 200 species. It accounts for the vast 
majority of Psychotria species in the West Indies, 
just under half in Mesoamerica, and perhaps a 
third in South America, where subg. Heteropsy- 
chotria is most diverse. Psychotria subg. Psycho- 
tria in Mesoamerica, the subject of this treatment, 
includes 61 species, 15 of which are recently named 
(Hamilton, 1988), and eight varieties. | have sorted 
the species into eight species-groups; their recog- 
nition as sections awaits study of the subgenus 
throughout tropical America. 

Systematic understanding of species-rich tropi- 
cal taxa such as Psychotria, Miconia, Piper, Bac- 
tris, Heliconia, and Anthurium is the critical first 
step toward understanding geographical patterns 
of diversity and processes of diversification in the 
tropics. Psychotria subg. Psychotria appears to 
be a good index taxon for describing species di- 
versity in tropical angiosperms as a whole (see 
Biogeography) and therefore provides a diversity 
data set of use to conservation planners and natural 
resource organizations. From an ecological and 
evolutionary standpoint, one hopes that results of 


the study of Psychotria can be extrapolated to 
other diversified tropical taxa. 


MORPHOLOGY AND TERMINOLOGY 


In this section, characters and character states 
are explained and illustrated only as necessary in 
order to eliminate any ambiguity regarding their 
use in descriptions and discussions. 


HABIT 


Habit states, e.g., "shrub" versus “tree,” are 
distinguished primarily by size, not by architecture 
or shape. Even l-meter shrubs in this subgenus 
are usually treelike in having a main trunk and 
lateral branch complexes. Architecture of Pana- 
manian species of Psychotria (both subgenera) has 
been studied (Hamilton, 1985); three basic types 
of branching event were found, differentiated by 
the number and orientation of new shoots to which 
the terminal node gives rise each growing season. 
In Type 1 branching, at the terminal node one 
lateral meristem gives rise to a relay axis that 
elongates the main axis by sympodial substitution. 
In Type 2 branching, two (or more) relay axes 
arise at the terminal node and remain equal, neither 
new axis assuming the same orientation as the axis 
that preceded it. Type 3 branching resembles Type 
A except m one of the new relay axes assumes 
tion as the parent axis. Most species 

of subg. Fears shows Type 3 branching in 
both the vertical and lateral directions; this serves 
as an additional field character for recognizing 
members of the subgenus. Two exceptions within 
the subgenus— P. limonensis and P. tenuifolia— 
show Type 1 and Type 2 branching, confirming 


their taxonomic affiliation based on other cha 


ac- 
teristics (see Infrasubgeneric eee Ar- 
chitecture has not been included in the descriptions 
because it has not yet been studied thoroughly for 
most species in the field 


STIPULES 


Stipules can provide valuable information as to 
interspecific relationships but prove almost worth- 
s a source of key characters, since in this 
subgenus they are usually caducous, or deciduous. 
he two stipules on opposite sides of a node are 
frequently united to some degree into a sheath. As 
the young stem grows, mechanical pressure forces 
the stipules to sever their basal connections to the 
stem and subsequently to fall off (Fig. 1). Thus 
most herbarium specimens have no stipules re- 


Volume 76, Number 1 
1989 


amilton 69 
Mesoamerican CR 
subg. Psychotr 


maining. The stipule scar consists of a ridge, usually 
pale in contrast to the stem, often with a dense 
fringe of minute red-brown hairs; the hairs often 
fall off shortly after the stipule. Exceptions to the 
above are found in the costivenia group, in which 
stipules are robust and not sheathing and thus are 
often found persisting several nodes from the ter- 
minus, and in the calophylla group, in which the 
herbaceous and suffruticose species do not grow 
sufficiently to force the stipules to break free. 

Stipule shape and size vary considerably, with 
the general shape, apex, pubescence, an 
differentiation providing critical cone be- 
tween species and species groups (Fig. 2). Taxa 
descriptions include the stipule Lr. along the 
stem followed by the maximum widt 


LEAVES 


Leaf characters are used extensively in this 
treatment in keys and as recognition characters. 
A single leaf, if examined critically, can suffice to 
identify most species in Mesoamerica. Most char- 
acter states are unambiguous; descriptive termi- 
nology is from Hickey (1979). Blade measurements 
are the length followed by the width at the widest 
point. Venation character states are observed on 
the undersides of the leaf blades. 

Secondary vein divergence is the angle formed 
at the junction of a secondary vein and the midvein, 
measured on the distal (on the leaf) side of the 
secondary vein. Most species have curved second- 
ary veins, so the tangent at the base of the sec- 
ondary vein was estimated and used. Angles were 
measured in the middle third of the leaf blade, as 
they often become more acute towards the base 
and towards the apex. 

The character of secondary venation is consid- 
ered to have basically three states: eucamptodro- 
mous, brochidodromous, and brochidodromous with 
a distinct collector vein (Fig. 3). In eucamptodro- 
mous venation, the secondary veins near the mar- 
gins do not connect one to another in robust sec- 
ondary arches or via a collector vein; they instead 
become less robust and distinct towards the margin 
and are often connected by a series of arches of 
tertiary veins (Fig. 3a). If just one connecting arch 
maintains the robustness of a secondary vein, the 
venation is said to be bod ipsun (Fig. 3b); 
consequently, the distinction between eucampto- 
dromous and brochidodromous is one of degree, 
and many species show both states. (It may be 
assumed, unless stated otherwise, that the second- 
ary arches are near the blade margin.) Much more 


| 
FIGURE 1.  Caducous stipules being forced to fall from 
a branch of Psychotria carthagenensis. Only ter 


io has fallen. Note the ridge and m 
hairs at the node where the dod base has broken free. 
(Knapp 4942.) Scale bar = 1 cm 


distinct and easily differentiated— enough so to 
serve in the first couplet in the species-identification 
key—is the third state, brochidodromous with col- 
lector vein, in which there is a distinct collector 
vein running straight or slightly undulating near 
and parallel with the margin (Fig. 3c). (Note that, 
in keys and descriptions, the second state, “‘bro- 
chidodromous," should be assumed not to include 
a distinct collector vein unless it is otherwise stated.) 

The arc shape of the secondary veins is also 
described, the states being: a) straight, b) constantly 
arcuate, and c) straight near midvein then arcuate 
near margin. 

The axils of the secondary veins often have tufts 
of hairs or domatia or sometimes both. The origin 
and function of domatia is little understood, but 
the minute holes found in domatia of mature leaves 
suggest that early instars of some insects may in- 
habit the cavities (Fig. 


Annals of the 
Missouri Botanical Garden 


NC, 
etd 
Ne 


q 


vt 


lo, 


"bu v » 
DUNT 


JT PETI IN 


NER. "n . 
MS NANA Uo Wa s Im eim 
Pe 
h, NER B 
* NS 6 
L4 


Cru 


FIGURE 2. Stipule forms found in subg. P: (ii pair — a. Sheathing, ovate, P. carthagenensis (Croat 10347). — 
iu I & Molina 31059). — c. Sheathing, mieie 


b. Ovate, the apex epa with pale center, P. co 

P. erythrocarpa (Standley 76317).—d. Sheat a lance anamensis (A. m hd 03).—e. the 
apex long-biacuminate, P. tenuifolia (Hayden 174).— e n lanceolate, the apex biaristate, P. orosiana iuis 
& Vodicka 5522).—g. Sheathing, truncate, oe from apical corners, P. Sercho idi (Lewis et al. 2103). 
Scale bars = 2 mm. 

5). This distinction is often one of degree, as the 
major difference is that in percurrent tertiary ve- 
nation the tertiaries joining the secondaries are 
more robust than the other reticulate veins, then 
considered quaternary veins. Few species, such as 


Tertiary veins in subg. Psychotria are generally 
either: a) orthogonal reticulate, joining each other 
and the secondaries, intersecting at roughly right 
angles, or b) percurrent, joining adjacent second- 
ladderlike effect between them (Fig. 


aries, making a 


Volume 76, Number 1 
1989 


mb. 


Hamilton 7 
Mesoamerican Psychotria 
subg. Psychotria 


P. micrantha, have strikingly percurrent tertiary 
venation. In several species, especially in the cos- 
tivenia group, tertiary veins known as intersecon- 
dary veins intersect the midvein between the sec 
ondary intersections and run roughly parallel with 
the secondaries. 


INFLORESCENCES 


Inflorescences in subg. Psychotria consist gen- 
erally of panicles of cymes, in which the cyme 
(dichasium) is the ultimate group of flowers and 
the panicle comprises the axes leading to the cymes 
(Fig. 6). The degree of branching of the panicle 
reflects the maximum branching found in the in- 
florescence: 2 degrees indicates that there are pri- 
mary and secondary axes; 3 degrees indicates that 
there are primary through tertiary axes, etc. (Fig. 
6a-c). Similarly, degree of branching of the cymes 
is also described (Fig. 6d-f). Cymes are often so 
contracted as to be glomerules of flowers. Bracts 
and bracteoles are present, though often incon- 
spicuous, on the axes; bracts subtend branches of 
the panicle, and bracteoles subtend the cymes. 

Inflorescence organization provides many of the 
most useful characters and so is described in great 
detail (Fig. 7). Secondary axes are borne in ranks 
along the main axis. The main axis itself is mea- 
sured; the distance between the base of the main 
axis and the point of insertion of the basal (first) 
rank of secondary axes is the peduncle (its length 
included in the measure of the main axis). Sec- 
ondary axes are usually two or four per rank; often 
when there are two or more pairs of axes per rank, 
the pairs are differentiated into a longer and a 
shorter pair (Fig. 7a). Often a specified node can 
have either one or two pairs of axes; measurements 
of a single pair are included under “the longer 
pair," while measurements of the "shorter pair" 
are relevant only when there are two pairs of axes. 
Sometimes secondary axes are reduced so that a 
cyme or flower cluster appears sessile on the main 
axis (Fig. 7b, k). The angle of divergence of the 
secondary axes (measured as is that of secondary 
veins) can also provide valuable information: species 
in the graciliflora group often have ca. 90? angles 
of divergence (Fig. 7h), and P. stockwellii shows 
obtuse angles of divergence (Fig. 7i), in striking 
contrast to the acute angles characterizing most 
species. 


FLOWERS 


The most interesting feature of the flowers of 
Psychotria, as is the case for much of Rubiaceae, 


TT 


om 


FiGURE 3. Types of leaf secondary venation. — 
PAEL Psychotria carthagenensis Y ks r 
1683).— Poem 3 x horizontalis (Spellman 


A ae 


& New s 1670).—c. Bro 
vein, P. calophylla (Dijon e et y 4556). Scale bars = 1 


is distyly: within a species many individual plants 


da 
eo 
1 
o 
a 


pin" or “long-style” morph; 
and the others have flowers with Ka Bde stamens 
and inserted style (*thrum" or *short-style" morph; 
Fig. 8d-f). This is part of an incompatibility system 
by which pin pollen can fertilize only thrum ovules 


72 Annals of the 
Missouri Botanical Garden 
E "d 
a Ml b AF 
FIGURE 4. Domatia in axils of secondary veins. —a. Psychotria remota ( 


(Correa & Dressler 1015).—c. P. viridis (Duke 10230). Scale bars = 1 cm 


and vice versa, i.e., intramorph incompatibility 
(Darwin, 1877; Ganders, 1979). From the stand- 
point of descriptive morphology, it means that flow- 
ers of both morphs must be described, with their 
different filament and style lengths. 


LL <a 

FIGURE 5. Types of leaf tertiary venation. —a. Or- 
thogonal reticulate. —b. Percurrent (with quaternary veins 
orthogonal reticulate). 


Hartshorn 1444).—b. P. marginata 


Of the 66 taxa in Mesoamerican Psychotria 
subg. Psychotria, 40 appear normally distylous, 
as judged from morphology alone. Five are “‘long- 
homostylous,”” i.e., having only one flower morph, 
in which stamens and style are both exserted. Five 
other taxa appear to have only one of the two 
distylous morphs: two are **pin-monomorphic" and 
three are ““thrum-monomorphic.”” No cases of dioe- 
cy are evident from flower morphology alone. (The 
remaining 16 taxa cannot be assigned a category 
because too few flowering specimens were available 
for analysis.) Nondistylous states of breeding bi- 
ology are not concentrated in any particular species 
group (see Infrageneric Relationships) and are 


laarzen 


iol ^ m 


E6 TY £h 


t f 7r mec 

Panicles: — a. Two degrees. — b. Three degrees. — c. Four 

degrees. Cymes: — d. One degree. — e. Two degrees. — f. 
ree degrees. 


Volume 76, Number 1 Hamilto 73 
1989 eee ten Psychotria 
subg. Psychotria 


Ae 
+ P i 
1 


h [|I i Jil ; Íi k Iš 


FIGURE 7. Inflorescences in subg. Psychotria, showing only main and secondary axes. Circles indicate reduced 
axes and sessile glomerules.—a. P. carthagenensis, with secondary axes in three ranks, two pairs (long vs. short) 
per rank.—b. P. flava, with three pairs of secondary axes in the first two ranks.—c. P. erythrocarpa, with one 
pair of secondary axes per rank.— d. P. dressleri, with a cluster of secondary axes at the apex of the primary axis. — 
e panamensis, with no peduncle. —f. P. limonensis, with secondary axes of such relative lengths as to give an 
umbellate shape to the inflorescence. —g. P. quinqueradiata, with the first rank of cad axes almost equal in 
length to the main axis. —h. P. ete MÀ with secondary axes diverging at right angles. —i. P. stockwellii, e 
secondary axes diverging at obtuse angles . remota, with all but the first rank of mue axes very short. — 
k. P. viridis, with almost all secondary axes talliosd. Scale bars = 1 cm. 


therefore hypothesized to have evolved indepen- acter, straightforwardly described. Persistent ca- 
ently of one another from primitive distyly (Ham- — lyces can also be valuable; Figure 9 indicates the 
ilton, in press). variation within the subgenus and how the relative 
asurements of corolla tubes and lobes are lengths of calyx tube and lobes can affect the aspect 
length followed by maximum width (diameter in of the fruit. 
the case of the tubes). e two seeds, often called pyrenes (cf. Stey- 
Stamen filaments are invariably fused to the — ermark, 1972; Dwyer, 1980), are hemispherical 
corolla tube; their point of separation from the tube — in cross section and thus have two surfaces to 
is often difficult to ascertain, however, so I have consider: the rounded, outside ““dorsal”” surface and 
measured the filaments from the base of the corolla — the flat, inner “ventral” surface. Longitudinal fur- 
tube. Using this measurement and that of style — rows run along both surfaces, and their number, 
length, one can determine the spatial separation depth, and regularity serve as important charac- 
between anthers and stigma. The filaments connect — ters. Figure 10 illustrates the variation within the 
to the anthers usually halfway up the introrse an- subgenus, with P. carthagenensis (Fig. 10a) rep- 


ther thecae. resenting the most common type of cross section. 
The depth of the furrows on the dorsal surface 

FRUIT ets the external appearance of the whole dried 
All fruit measurements are of dried herbarium fruit: P. remota (Fig. 10f) fruit appears deeply 


material and so will differ from those of fresh fruits mur while that of P. grandis (Fig. 101), with its 
with fleshy exterior. Fruit shape is a useful char- many shallow irregular furrows, appears smooth. 


74 


Annals of the 
Missouri Botanical Garden 


bsp: E 8. 
R. Foster Mi 
L MORÓN Scale bar = 


5k 


BIOGEOGRAPHY 


For purposes of biogeographical analysis I have 
divided Mesoamerica into nine regions (Fig. 11). 
The most species and mo f subg. 
Psychotria are found in pem (40 species; 15 
endemics); both diversity and endemism drop some- 
what in Costa Rica (32/7). There is a diversity 
"trough" in middle Mesoamerica (Honduras, El 
Salvador, and Nicaragua), with absolutely no ende- 
mism. A secondary center of diversity includes 
region 3 (eastern Veracruz, Tabasco, Oaxaca, and 


Long-style (pin) and short-style (thrum) flower morphs of Psychotria marginata. —a-c. Long-style 
f. Short-style morph (Antonio 2305). a, d, whole flowers; b, e, dissected corollas; c, 


Chiapas in Mexico) and region 5 (Guatemala and 
Belize), both areas having 26 species and a few 
endemics. Other parts of Mexico have only range 
extensions of some species found in regions 3 
and 5. 

The striking pattern of species diversity of subg. 
Psychotria within Mesoamerica appears to rep- 
resent general diversity patterns of moist- and wet- 
forest vascular plants (Fig. 11). This pattern— 
most diversity and endemism in Costa Rica and 
Panama, a secondary center in southern Mexico 


Volume 76, Number 1 
1989 


milton 
Mesoamerican Psychotria 
subg. Psychotria 


E 
ho 


t 
Y 
yA 
v m 
> 
SL 
we 
IN ay 


" é 
-— ++ 
A : 
EN t 
NM 
SP XA 
M a, e 
MSS DEC. 
NF € 
SV 3 - Ys 
d'a. $ Y 
233 5 2 26 
TEE T 
Q > = > ep 
£i 3i II 
-R = = PE 
== = = 7 
3 ~ Xï = = 
-3 2 È > 
à = E 
[t M TE 73 
EA y rd 
TEN de 
Ve DNA E 
Wi F 
SSC 


FIGURE 9. 
b. P. papantlensis (Contreras 7958). 
39552). 


(region 3) and Guatemala, and little between—is 
found in other recently monographed species-rich 
taxa of moist- and wet-forest plants (Table 1). The 
Anthurium pattern (tabulated from Croat, 1983, 
1986) is an amplified version of that of P. subg. 
Psychotria, with nearly seven times as many species 
in Panama as in Nicaragua. Cordia sect. Micran- 
thae (Miller, 1985) and Monochaetum (Almeda, 

1978), with somewhat smaller data sets, also reflect 
the general trend. Cavendishia (Luteyn, 1983) 
and Heliconia with pendent inflorescences (Kress, 
1984) also have greatest diversity in Panama and 
Costa Rica but show no secondary center in Mexico 
and Guatemala. 


All these data sets suggest strongly that the 


| 
| 


Dried fruit with p os calyces.— a. Psych ‘hotria carthagenensis (Standley & Valerio 44926).— 
. P. cocosensis (Pittier 12375).—d. 
—e. P. horizontalis cee & Witherspoon 8693). Scale bar = 1 m 


P. Posee (Lauvert & Barkley 


diversity pattern of subg. Psychotria represents 
that of many species-rich taxa and may therefore 
be of value as an index of regional diversity for 
tropical flowering plants. The primary value of this 
would be in pinpointing areas to which we should 
devote conservation effort in order to preserve the 
maximum diversity of life. Figure 12, a map of 
species numbers per one-degree square, reveals 
great differences in diversity that cannot be dis- 
missed as collecting artifact. It is also noteworthy 
that all nine squares with significant endemism are 
in Costa Rica and Panama (Table 2) 

Areas in Costa Rica and Panama with particu- 

arly high endemism are largely identical to those 
noted by Croat (1983, 1986) for Anthurium, most- 


75 


76 Annals of the 
Missouri Botanical Garden 


Cub EO ES TM fs 


a b c d e 


52s dE dh GS LB 


10. Seed cross sections. —a. Psychotria carthagenensis, the dorsal surface (d. s.) with 4 deep longitudinal 
2 


Fic 
furrows (l.f.), the ventral surface (v.s.) with 2 deep l.f.—b. P. horizontalis, the d.s. with 3-5 l.f., the v.s. wit 
Lf P. graciliflora, the d.s. with 4 l.f., the v.s. concave or with 2 broad shallow yas P. orosiana, the d.s. 
with 4-5 deep l.f. is acute ridges between, the v.s. with 2 shallow 1.f. —e. P. costivenia var. costivenia, the 


nal 


2 |.f.—g. P. olgae, the d.s. with 4 deep and numerous irregular l.f., the v.s. with 2 deep and several irregular l.f.— 
h. P. mexiae, the d.s. with 5-8 deep irregular l.f., the v.s. with 2 deep plus usually 2-4 additional irregular l.f. — 
i. P. grandis, the d.s. with 10-15 shallow irregular l.f., the v.s. pies 2 very deep 1.f.—j. P. flava, the d.s. with 6- 
10 irregular l.f., the v.s. with 1 often T-shaped l.f. Scale bars = 


ly 1,000-2,000 m elevation along the Continental Panama. The Divide in western Panama has sev- 
Divide plus isolated hilly areas. Costa Rican en- eral narrow-ranging species: P. boquetensis, P. 
demics P. chitariana, P. monteverdensis, and P. — hornitensis, P. hammelii, P. insueta, and P. cas- 
sylvivaga fit this pattern, as does P. stockwellii,  cajalensis. Just east of the Canal, the Cerro Azul- 
which extends from Alajuela to western Chiriqui, Cerro Jefe region has three endemics: P. liesneri, 


Y) | h 
1 p TC. 


AU ET J 


ws 
22/0 
440/15»; 


b 


= + 32/78 


i. L E 


105 100 95 9u 85 80 


FIGURE 11. Total and endemic species of Psychotria subg. Psychotria by region. Circled numbers are the region 
numbers; crap on the left of the slash are the total species in the area; numbers on the right are the species 


endemic to the a 


Volume 76, Number 1 


Hamilton 77 
Mesoamerican Psychotria 
subg. Psychotria 


Numbers of species/endemic species, by region (see Fig. 11), of various taxa. 


TABLE 1. 


Honduras 


Guatemala 


Total 


Panama 


and 
El Salvador Nicaragua Costa Rica 


and 
Belize 


Mexico 


species 


40/15 
148/82 
17/6 


26/1 16/0 22/0 32/7 
16/1 22/1 


4/0 


3/0 


11/0 26/3 


7/0 


Psychotria subg. Psychotria 


Anthurium' 


72/18 


37/22 


0 


7/0 


8/0 


9/0 


Micranthae? 


Cordia sect. 


3/0 


1/0 


28/16 
16/5 


3/0 
5/0 


Cavendishia* 


13/1 


2/0 


2/0 


1/0 


Heliconia with inflorescence pendent: 


! Croat, 1983, 1986. 


' Luteyn, 1983. 
^ Kress, 1984. 


. One-degree squares (see Fig. 12) with sig- 
nificant species endemism in Psychotria subg. Psycho- 
tria. 


Total 

Psychotria 
à ; subg. 

Square A ex i aS Piychokik 

Coordinates' A? B: C Species 

10°N, 84°W 3 0 2 19 
9*N, 84°W 0 2 3 8 
9*N, 83°W l l 4 20 
9*N, 82°W 1 0 2 14 
,19? 3 0 2 20 
8'N, 82?W 2 l 2 13 
8°N, 81°W 0 2 l 14 
8°N, 80°W 1 l 0 15 
eN, 77°W 2 0 2 20 


' Coordinates are of the SE corner of the square. 
Squares are highlighted in Figure 12. 
- Has been collected only i in this square. 
' Has been collected in only one additional square 
' Has been collected in only two or three additional 
squares. 


P. olgae, and P. pacorensis. Mountains near the 
Colombian border support three more: P. philacra 
(Cerro Tacarcuna), P. rosulatifolia (Cerro Mali 
and Cerro Pirre), and P. insignis (south of Puerto 
Obaldia). Three narrow-ranging species, P. lasel- 
vensis, P. sixaolensis, and P. fendleri, are found 
in areas of Caribbean lowlands, which have not 
been botanized thoroughly. It is beyond the scope 
of this treatment to speculate on the causes of 
these patterns—e.g., absolute rainfall, topograph- 
ical heterogeneity, geological disturbance— but the 
pattern suggests that Costa Rica and Panama de- 
serve the lion's share of in situ conservation focus 
in Mesoamerica. 

The biogeographies of species complexes, within 
groups, display consistent patterns that suggest 
hypotheses as to the course of evolution within the 
complexes (see Infrageneric Relationships). Many 
species complexes are comprised of one widespread 
species, for which the complex is named, plus one 
or more species with much narrower ranges: the 
Meg quinqueradiata complex (two species), 

e P. micrantha complex (two species), the P. 
ud complex (two species), the P. costive- 


plex (two species), the P 
species), the P. chiriquina complex (three species), 
and the P. panamensis complex (six species; see 
distribution maps in the introductions to each species 
group in the Systematic Treatment). Seven wide- 
spread species, such as P. carthagenensis and P. 


Annals of the 
Missouri Botanical Garden 


105 


100 95 


i. 


90 


FIGURE 12. 
(see Table 2) are shaded. 


marginata, are not obviously similar to any nar- 
row-ranging species. Group 8, the calophylla group, 
defies even tentative analysis of affinities: most 
species are narrow endemics in Costa Rica and 
Panama, with only P. calophylla and P. psycho- 
triifolia having somewhat wider ranges. The pre- 
dominant pattern of one widespread plus one or 
more narrow-ranging species accounts for 35 of 
the 61 species and suggests that in each complex 
the widespread species may be ancestral to those 
with narrow ranges. Careful phylogenetic analysis 
of the subgenus throughout the Neotropics is nec- 
essary to evaluate this hypothesis. 

Each species and variety account in the Sys- 
tematic Treatment includes a characterization of 
range in terms of habitat and climate. The habitat 
classification used is that of Holdridge (1967), par- 
ticularly from life zone maps of Costa Rica (Tosi, 
1969) and Panama (Anonymous, 1970). The ranges 
of many Mexican species are characterized ac- 
cording to a vegetation map of that country (Flores 
Mata et al., 1971). Climate types are those of 
Walter et al. (1960), discussed by Walter (1973); 
the particular climate types referred to herein are 
six: I equatorial, I(X) equatorial-mountainous, I(H) 
equatorial-tropical, II(I) tropical-equatorial, II 
tropical, and II(X) tropical-mountainous. Equato- 
rial and equatorial-mountainous climates may show 
some seasonality of rainfall, but the ratio of pre- 


Numbers of species of Psychotria subg. Psychotria per one-degree square. Squares of high endemism 


cipitation (in mm) to temperature (in degrees cen- 
tigrade) never drops below 2:1 in any month. In 
an equatorial-tropical climate, there is a mild dry 
season in which the ratio drops slightly below 2: 1 
for one or two months, usually around March. In 
a tropical-equatorial climate, the dry season is more 
extreme, lasting up to four months at the beginning 
of the year. The dry season in a tropical or tropical- 
mountainous climate usually lasts November 
through April, some of those months often having 
absolutely no precipitation. 


INFRASUBGENERIC RELATIONSHIPS 


Formal taxonomic recognition of sections awaits 
understanding of the subgenus throughout the Neo- 
tropics. For the time being, I have divided the 61 
Central American species into eight groups com- 
parable to sections. In several cases I refer to 
species complexes, which are sets of morphologi- 
cally very similar species within a group. 

The following key to species groups indicates 
major differences among the groups and is pri- 
marily didactic; it utilizes characters, such as stip- 
ules, that are useful at the species-group level but 
less so for species identification, e.g., because stip- 
ules are caducous and fruits are often not present. 
For species identification, one should use the key 
to species in the Systematic Treatment; it relies 


Volume 76, Number 1 


Hamilton 79 
Mesoamerican Psychotria 


subg. Psychotria 


largely on leaf and inflorescence characters, which 
are most consistently evident. 

Following the key are brief synopses of the eight 
species groups, including lists of species, geograph- 
ic patterns, and noteworthy character states not in 
the key. More extensive descriptions, with distinc- 
tive character states italicized, plus discussions of 
species relationships, biogeography, and breeding 
biology will be found at the beginning of each group 
in the systematic treatment. 


KEY TO SPECIES GROUPS 


la. KA papi axes almost always 4 (or 6) per 
rank in de east one rank, in usually 2 (or 3) 
and 
lb. eo axes generally 2 per rank or, if 
4 or clustered, equal in lengt 
2a. Leaves drying generally green-brown to red- 
brown to red-gray 


Group 1—carthagenensis group 
2b. E drying generally p green to yellow- 
en Gro pro group 
3a. Leaf secondary venation a poao 


w 
c 
ps 
O 
fs 
=s 
[^] 
[17 
o 
Oo 
ad 
a 
© 
A 
Ne 
< 
O 
d 
® 
m 
AS, 
d 
c 
[71 
[e] 
EB. 
= 
c- 
5 
o 
& 
SH 
a 
o 
a 
ix] 
o 
7 


eucamptodromous, the corolla lobes with apical 
extensions and/or the inflorescence bracts en- 
larged 
. Inflorescences generally pedunculate, the sec- 
ondary axes diverging usually at right angles 


os 
w 


> 
> 


. Inflorescences usually lacking peduncle, the 
secondary axes generally diverging at acute (ob- 


tuse in P. stockwellii) angles 
Sa. Leaves drying usually gray- green to green-black 
or red-black; axes elon- 


gated in all ranks; seed dorsal surface with 3-5 


ep furrows, sometimes plus several irregular 
longitudinal furrows ..... roup 3—gra ocior Bow 
5b. Leaves drying glossy red-brown; infloresc 


surface with 3 dee 
idis dd 
nts often pela cr qa or tom 
tose; po calyx spicuous, 2-5 
mm long; seed dore a usually with 4-5 
regular Cocinas furrows 


o 
p 
e= 


md group 
6b. ants omen 
ose; persiste 
ES r ied a coriaceous cup; seed dor 
surface with 4-10 irregular longitudinal furrows 
roup UU inpr pat group 
. Inflorescences umbelliform panicles of cymes, 
the bracts never enlarged; corolla lobes b dues 
apical extensions; persistent calyx inconspic- 
uous or to 0.5 mm long 


~J] 
m 


roup sns iae group 
Inflorescences panicles of c ll 
glomerules, the bracts often enlarged; carla 
lobes almost always with apical UM per- 
sistent calyx often a conspicuous 
Gro 


oup le nr group 


-~J 
P 


SPECIES GROUP SYNOPSES 


Group 1 —carthagenensis group. Species 1-7. 

The carthagenensis group comprises seven 
species: Psychotria carthagenensis, P. clivorum, 
P. lamarinensis, P. micrantha, P. neillii, P. quin- 
queradiata, and P. viridis. The Psychotria quin- 
queradiata species complex includes that wide- 
spread species plus P. which is 
endemic to northern Costa Rica. The P. micrantha 
complex includes that species plus P. neillii, whose 


lamarinensis, 


range is restricted to Caribbean southern Nicara- 
gua and northern Costa Rica. Psychotria car- 
thagenensis, P. clivorum, and P. viridis are all 
widespread without obvious close relatives. 

In addition to the key characters, members of 
this group may be recognized by their seeds having 
(3-)4-5 deep longitudinal furrows on the dorsal 
surface and 2 longitudinal furrows on the ventral 
surface. 

Group 2—costivenia group. Species 8-16. 

The costivenia group comprises nine species: 
P. balancanensis, P. costivenia, P. fendleri, P. 
flava, P. grandis, P. horizontalis, P. papantlen- 
sis, P. pleuropoda, and P. sylvivaga. The Psy- 
chotria horizontalis i^a includes the most 
widespread of the nine plus P. fendleri, which is 
endemic to Caribbean coastal Panama. The P. cos- 
tivenia complex comprises the other seven; P. 
costivenia is only slightly more widespread than 
P. grandis, with the two sharing noteworthy mor- 
phological similarity and overlapping ranges. Psy- 
chotria sylvivaga has been collected only in Costa 
Rica, and the remaining four species are restricted 
to southern Mexico, Guatemala, and Belize. 

Distinctive character states, in addition to those 
in the key, include a usually conspicuous persistent 
calyx in fruit. 


Group 3— graciliflora group. Species 17-25. 

The graciliflora v bud nine re 
in Mesoamerica: P. ba , P. graciliflora, P 
jimenezii, P. laselvensis, P. leia P. margina- 
ta, P. orosiana, P. parvifolia, and P. philacra. 
Psychotria marginata is the most widespread and 
frequently collected, and no other species closely 
resembles it morphologically. In contrast, the also 
widespread P. graciliflora is strikingly similar to 
six of the other species (the P. graciliflora com- 
plex) that occur more or less sympatrically with it: 
P. parvifolia and P. orosiana (Costa Rica and 
western Panama), P. laselvensis (Costa Rica), P 
bakeri (Los Santos, Panama), P. liesneri (eastern 
Panama province), and P. philacra (eastern Dar- 
ien, Panama, at higher elevation than P. gracili- 
flora). Psychotria jimenezii may belong to this 
complex as well. 


Annals of the 
Missouri Botanical Garden 


Members may be recognized by their key char- 
acters plus stipules often biacuminate to biaristate, 
secondary veins usually brochidodromous and di- 
verging 70°—-90°, and inflorescence secondary axes 
usually delicate. 


Group 4—remota group. Species 26. 

The remota group includes several species in 
South America but only one of those, P. remota, 
in Mesoamerica. An additional recognition char- 
acter is the winged inflorescence axes, conspicuous 
especially in the fie 


Group 5— nervosa group. Species 27-35. 

The nervosa group comprises nine species: P. 
aguilarii, P. boquetensis, P. chagrensis, P. eryth- 
rocarpa, P. fosteri, P. fruticetorum, P. jinote- 

ensis, P. mirandae, and P. nervosa. Two species 
complexes, those of P. nervosa and of P. cha- 
grensis, are discernible; three other fairly wide- 
spread species— P. fruticetorum, P. erythrocar- 
pa, and P. jinotegensis—do not closely resemble 
any others. The P. nervosa complex consists of 
widespread P. nervosa and three species of more 
restricted ranges, all of which occur allopatrically 
with or at range boundaries of P. nervosa: 
mirandae in southern Mexico, P. aguilarii in 
higher-elevation southern Guatemala, and P. bo- 


quetensis in high-elevation western Panama. The 
1 


P. chagrensis plex includes widespread P. cha- 
grensis and the probably derived P. fosteri, which 
occurs on Coiba Island, Panama, apparently allo- 
patrically with respect to P. chagrensis. 

Recognition characters not included in the key 
are ellipsoidal to narrow-ellipsoidal fruit, and inflo- 
rescences reduced to fascicles of flowers or the 
secondary axes in (2-)3 ranks. 


Group 6—panamensis group. Species 36-47. 
The panamensis group comprises 12 species in 
Mesoamerica: P. cascajalensis, P. chiriquina, P. 
cocosensis, P. dwyeri, P. hornitensis, P. lundellii, 
P. mexiae, P. olgae, P. panamensis, P. sarapi- 
quensis, P. stockwellii, and P. trichotoma. The 
largest species complex is the P. panamensis com- 
plex. It includes one widespread species, P. pan- 
amensis, whose var. ixtlanensis may owe its origin 
to introgression from P. trichotoma (see under 
Systematic Treatment), and five narrow-ranging 
species: P. dwyeri (southern Mexico, sympatric 
with P. panamensis), P. stockwellii (Costa Rica 
and Panama, sympatric), P. cocosensis (Cocos Is- 
land, Costa Rica, allopatric), P. olgae (Panama, 
+ sympatric), and P. cascajalensis (Panama, sym- 
patric). The widespread species P. trichotoma, P. 
sarapiquensis, and P. mexiae are of uncertain 


affinities; the former two may be closely related to 
. panamensis complex. The P. chiriquina 
complex includes two species whose ranges do not 
overlap (P. lundellii south to El Salvador and P. 
chiriquina south from northern Nicaragua) and 
one narrow endemic, P. hornitensis from high- 


the 


elevation western Panama at the eastern end of 
the range of P. chiriquina. 


Group 7 —tenuifolia group. Species 48, 49. 
The tenuifolia group includes only two species 
in Mesoamerica, P. limonensis and P. tenuifolia, 
both of which are widespread with extensively over- 
lapping ranges. The group is characterized by mi- 
nute flowers and secondary veins diverging usually 
70-90", in addition to the key characters. 


roup 8—calophylla group. Species 50-61. 

The calophylla group comprises 12 species: P. 
alfaroana, P. calophylla, P. chitariana, P. dress- 
leri, P. hammelii, P. insignis, P. insueta, P. mon- 
teverdensis, P. pacorensis, P. psychotrüfolia, P. 
rosulatifolia, and P. sixaolensis. Ten of the species 
have small geographic ranges; only P. calophylla 
and P. psychotriifolia are at all widespread, and 
their ranges are notably discontinuous. Psychotria 
dressleri and P. rosulatifolia are quite similar mor- 
phologically and occur in eastern Panama, and 
therefore may be supposed to be related; but be- 
yond that the affinities are very uncertain. 

Additional recognition characters are many: her- 
baceous or suffrutescent habit; usual general pu- 
bescence; sheathing stipules, these usually bifur- 
cate or biaristate with the extensions often from 
corners of the truncate sheath; and ellipsoidal fruit. 


SYSTEMATIC TREATMENT 


Psychotria L., Syst. Nat., ed. 10. 929. 1759 
(nom. cons.) subgenus Psychotria [sensu Pe- 
tit, Bull. Jard. Bot. État 34: 1-229. 1964. 
Also sensu Steyermark, Mem. New York Bot. 
Gard. 23: 406-484. 1972. | TYPE SPECIES: Psy- 


chotria asiatica L. 


Ouragoga E Am Cliff. 486. 1737. Cf. also Linnaeus, 
Gen. Pl., e . 378. 17 

Myrstiplylin P. os Civ. Nat. Hist. Jamaica. 152. 
(nom. rejic.). Sensu Hitchcock, Annual Rep. 

Mis ouri Bot. Gard. 4: 95. 1893. 
dae ars Browne, Civ. Nat. Hist. Jamaica. 160, 
. l, t. 17, fig. 
Men Aubl, Hist 
TYPE SPECIES: 


: : r- 
Argoviensis, Flora 59: 457-466, 495-498. 1876, 
and in Martius, Fl. Bras. 6(5): 384-428. 1881. Also 
sensu Bremekamp, Recueil Trav. Bot. Néerl. 31: 
248. 1934. 


Volume 76, Number 1 
1989 


Hamilton 81 
Mesoamerican Psychotria 
subg. Psychotria 


U logoga Baillon, Adansonia 12: 323. 1879. TYPE SPE- 
ragoga (nb o ee (Brot. end Sensu 
nist Revis. Gen. Pl. 1: 298-301. 


Shrubs, small trees, or less often herbs or sub- 
shrubs; stipules interpetiolar, often sheathing, usu- 
ally caducous, leaving a pale ridge with red-brown 
fringe. Leaves opposite; blades membranous to co- 
riaceous, drying red-brown or dull greenish or rare- 
ly chalky yellow- ies secondary veins eucamp- 
todromous to brochidodromous, sometimes with 
collector vein, the um sometimes with domatia or 
tufts of hairs; tertiary veins orthogonal reticulate 
to percurrent. /nflorescences terminal or pseu- 
doaxillary, panicles of cymes or of glomerules or 
rarely fascicles of flowers; secondary axes borne 
in 1-several ranks along the main axis, each rank 
with 1-3 often size-differentiated pairs of axes. 
Flowers small, (4—)5-merous, almost always disty- 
lous; corolla usually white, the tube cylindrical; 
style 2-branched, the branches bearing the stig- 
matic surfaces, the ovary 2-celled, the ovules 1 
per cell, attached basally. Fruit fleshy, maturing 
red (except reportedly purple or blackish in 
insueta and P. pacorensis); persistent calyx often 
conspicuous at distal end of fruit; seeds 2, hemi- 
spherical in cross section, the rounded dorsal sur- 
face with 3-5 regular to many irregular longitu- 
dinal furrows, usually giving the dried fruit a costate 
aspect, the flat or slightly concave dorsal surface 
with (1—)2 regular longitudinal furrows often plus 
several shallow irregular furrows. 


A pantropical subgenus of approximately 600 
species worldwide, including ca. 


enus accounts for the majority of species in 
ihe Antilles but only for 32 of 242 species treated 
by Steyermark (1972) for the Guayana Highland. 


KEY TO SPECIES 
PSYCHOTRIA SU BGENUS PSYCHOTRIA 


m- 


a. Leaf blades with collector veins distinct, 


lb. Leaf blades without collector veins ............. 13 
2a. Inflorescences reduced to fascicles of flowers 
3 
2b. Inflorescences with distinct axes 0 4 
3a. Leaf blades (6-)8-12 x (2-)3-5 cm; stipule 
eath 8-12 mm long; Coiba paar I 
) P. prem 


3b. Leaf blades (3-)4-7.5(-9.5) x (1- i 52 
(-3.5) cm; stipule sheath 4-7 mm long; w 
sprea (29) P. RUE i 
4a. Inflorescence a panicle of glomerules ................. 
4b. Inflorescence a panicle of cymes 
5a. Young stems glabrous; ite. Costa Rica . 
57) P. indatevexdensis 


5b. Young stems ferrugineous-pubescent or brown- 
puberu 
6a. Leaf blades ovate; secondary veins diverging 
over 90°; San Blas, Panama ....... . insignis 
6b. Leaf blades elliptic or oblanceolate or obovate; 
n 


mm 
long; widespread (51) P. wt. 
7b. Shrub or subshrub to 2 m tall; fruit less than 


ong 
8a. Inflorescence lacking peduncle, often with more 
than 1 pair of secondary axes per rank; Vibes 
mm long; southern Nicaragua throu 
a P psychotriifolia 
8b. Inflorescence with peduncle, with 1 pai 
secondary axes per rank; corolla tube 4 mm 
long; Limón, Costa Rica, and Bocas del Toro, 
Panama 1) P. Rad i 
9a. Young stems ferrugineous-pubescent; inflor 
cence with secondary axes usually in more 
than 1 size-differentiated pair per rank; Cos 
Rica (19) P. a 
9b. Young stems glabrous or minute-puberulent; 
inflorescence with secondary axes in 1 pair or 


bracts 4 mm long; is 
shrubs or herbs 0.5 m tall; eastern Dar 
Panama (60) P. rosulatifolia 
10b. Inflorescence with bracts to 1.2 m 
shrubs 0.4-3(-5) m tall 
lla. Leaves sessile with organic matter accumu- 
lating in leaf wl E blades 30-42 cm long; 
eraguas, Pan ) P. insueta 
llb. med petiolate, di as above; leaf blades to 


ong 
12a. Leaves broad- jac | (12-)17-29 x (6-)7.5- 
m; inflorescence secondary axes j 
s 5- 6 ranks; fruit (3.5-)4(-4. a x (3-)3. 
(-4) mm; widespread o... P. TE i 
12b. Leaves narrow-elliptic, (4-)7.5- E (1-)2.5- 
(-7.7) cm; inflorescence secondary axes in 
(2-)3(-4 1 aes fruit 4-5 x 3- Pr p mm; 
widespre 49) P. tenuifolia 
13a. Fruit shi ilios « diameter eastern Dar- 
nama a uu .. (25) P. philacra 


ien, 
13b. Fruit a di: 
14a. onde not drm (except rarely) 


14b. Inflorescence pedunculate (except rarely) ......... 37 
15a. Inflorescence reduced to fascicles of flowers 


15b. ii not reduced to fascicles of flow- 


16a. Young stems red-brown tomentose; southern 
exico .. (34) P. mirandae 
16b. Young stems glabrous 17 
7a. Leaf secondary veins 11-14, diverging 70°- 
80% Coiba ` Island, Panama 

L A 


60°; widespread ................ E quinqueradiata 
18a. Inflorescence secondary axes in 2 or more 

pairs per rank in at least 1 Fax (except for 

rare inflorescences) or in a fascicle _.. 1 
18b. Inflorescence secondary axes only in 1 pair 

per rank (except for unusual inflorescences 

having 2 pairs in one ran 
19a. Young stems glabrous 20 


82 Annals of the 
Missouri Botanical Garden 
19b. Young stems puberulent, tomentose, or fer- 2); southern Mexico through Nicaragua ...... 
rugineous-pubescent „i ce F. iaia 
20a. Leaf blades a y obovate or etn ane 3lb. Leaf blades narrow (length/ width appro 
(6-)9-15(-16) x (2-)3-7.5 cm, with s db 3); Oaxaca, Mexico s 
ondary veins 6-9(-10) pairs diverging 45- = _ (44b) P. panamensis var. ixtlanensis 
60°; corolla tubes 4-5 mm long; widespread 2a. Bark s oth s . 33 
Tun ( P quinqueradiata 32b. Bark vii fiic ridges, fissures, or fur- 
20b. Leaf blades broad-elliptic, (13-)16-20 x rows 4 
(5-)7.5-9.5 cm, with secondary veins A = 33a. Fruit spherical; southern Mexico . (39) P. dwyeri 
pairs diverging 70°-85°; corolla tubes 3 m 33b. Fruit narrow ellipsoidal; widespread ................. 
long; Alajuela, Costa Rica ..... 3) P. anda (35) P. nervosa 
2la. € je mm long; fruit red-brown to- 34a Stipules no more than 10 mm long ...................... 3 
hern Mexico ...... (34) P. mirandae 34b Stipules rarely as short as 10 mm cinco. 
21b. Stipules m ee 10 mm long; fruit glabrous 35a. Inflorescence secondary axes in 2 ranks; co 
rolla tube 3-3.5 mm long; Guatemala ......... 
22a. Fruit drying often shiny red-brown; secondary (27) P. aguilarü 
veins often drying reddish below; corolla often 35b. inflörescence secondary axes in 3 (or 4) ranks; 
dying pink; Mexico through northe "v Costa corolla tube 4-6 mm long; Nicaragua throug 
42) P. mexiae western Panama ccoo 37) P. Eum 
22b. Fruit drying dull red-brown or red- black; sec- 36a. Fruit drying often shiny red-brown; secondary 
Ried veins and corolla not as above; wide- veins often drying reddish below; corolla often 
(44) P. panamensis drying pink; Mexico through northern Costa 
23a. une stems obviously ferrugineous-pubes- Rica oo e uet Lu (42) P. mexiae 
cent or tomentose 3Ób. Fruit drying dull red- brown or red-black; sec- 
23b. Young stems iion or minutely puberulent ondary veins and corolla not as above; wide- 
28 sprea aca 
24a. Calyx lobes conspicuous, > 0.5 mm long ........ 25 38 
24b. Calyx lobes inconspicuous, = 0.3 mm long 
25a. Bark longitudinally grooved; stipules bilobed; 10 
inflorescence main axis m long; Chi- 39a. Shrub or subshrub to 1.5 m tall; corolla odd 
riqui, Panama coccion P. RR MEE with apical appendages; eastern Panama ....... 
25b. Bark ib. stipules not bilobed; up Ax (53) P. dressleri 
cence main axis (2.5-)4-6.5 cm long; 39b. Shrub or tree rarely less than 2 m tall; corolla 
temala through Nicaragua ..(33) P. p lobes without appendages .... 
26a. Leaf blades (1-)1.5-5.2 cm long, the sec- 40a. Young stems and = undersides densely =~ 
ondary veins inconspicuous to not evident be- rugineous-pubescent; fruit when dry 4 m 
low; Costa Rica and western Panama ........... long; Guatemala, N nes through Panama 
24) P Cj aug Re een oer eee 4) P. micrantha 
26b. Leaf blades at least (6-)9 cm bai ioe. c ee 
dary veins conspicuous below „n. 40b. Young stems and leaf undersides glabrous, 
27a. Stipules (10-)20-80 mm long; corolla tube Pubsirient lieu: amigas ir 
"d NBs fruit when dry not less than mm long ...... l 
diameter greater than half its length; seed y 7 
dorsal surface with 4 or 7-10 irregular lon- 4la. Inflorescence main i ae O- 3 26 cm long; 
gitudinal furrows; widespread coccion alky yellow-green above; 
P a fruit wien fae 8- -12- 13) mm long; us 
. parcens Mexico and Guatemala ............ ) P. flava 
27b. Stipules 6-11 mm long; corolla tube diameter ,,, vaca dg 
P . i i 41b ieee main axis 1.5-12 ne diee 
less than half its length; seed dorsal surface j d db b f B he 
€ 4-5 regular longitudinal furrows; wide- iib. red-brown to green-brown; fruit w 
(35) E neral dry 4.5-8 mm long; widespread „uu 
P. panamensis 
28a. Po calyx cuplike, coriaceous, 1.5 mm 42a. Bark s UN ES 43 
long; Cocos Island, Costa Rica — . 42b. Bark quee or ridged longitudinally sched 45 
8) P. cocosensis 
Ñ i ] 43a. Inflorescence with secondary axes in (second 
28b. Persistent calyx inconspicuous or a beak to to) fourth rank and above reduced so that 
mm long flowers and fruit appear clustered along main 
29a. Fruit when dry obovoid or, if ellipsoidal, at xis; widesprea (7) P. viridis 
least 7 mm long and with inflorescence axes 43b. Inflorescence not 
in 3-5 ranks 44a. Tree 10 m tall; leaf blades coriaceous; inflo- 
29b. Fruit Aer dry ellipsoidal or tigris seldom rescence main axis 6 cm long; Coclé, Panama 
over 7 l inflorescence P. cascajalensis 
secondary axes in 5-7 so 44b. Shrub to 1.5 m tall; leaf blades membranous; 
30a. Leaf secondary veins 6-9(-11); Veracruz, inflorescence main axis 1.5-3.5 cm long; Los 
Mexico, Costa Rica, and Panama . Santos, Panama 1) P. bakeri 
ba P. sar arapiquensis 45a. Leaf blades coriaceous, the margins inrolled; 
30b. Leaf secondary veins at least 12 „l... 31 fruit when dry 9-10 mm long; Panama Prov- 
3la. Leaf blades wide (length / width ge DA ince, Panama (43) P. olgae 


Volume 76, Number 1 
9 


Ham 


ilto 
a i ae Psychotria 


83 


subg. Psychotria 


on 
en 
w 


. Inflorescence axes winged; Costa Rica a 


. Inflorescence axes not winged 
. Leaf secondary veins brochidodromous with 


P 
. Dorsal seed surface with 4-5 regular 


. Leaf blades membranous to subcoriaceous, E 


es d not inrolled; fruit when dry up t 


: Stipules 8-12 x b 5-2 mm, not sheathing; - 


rescenc y axes in (2 or) 3 ranks; 

seed dorsal surface with 4 regular longitudinal 

furrows; southern Mexico, ater gn Belize 

30) P. ¿do saiia 
mm 


. Stipules (10-)20-80 x (2.5-)3.5-7 


sheathing; inflorescence secondary axes in $ 
7(-8) ranks; seed dorsal surface with 4 or 7- 
10 irregular longitudinal furrows; widespread 

44) P. panamensis 


Panama 


26) P. remota 
4.8 


secondary loops conspicously far from margin 


. Leaf secondary veins eucamptodromous or else 


brochidodromous with secondary loops near 
margin 


. Leaves membranous, drying dull green to red- 


brown; calyx lobes lanceolate, 1-2 mm long, 


. Leaves coriaceous, drying bright chalky y 


low-green above; calyx lobes triangular, 0. 5 
mm Bene to 1.5 mm long in fruit; gms Colón 


Y P. fendleri 


’ at least half of inflorescences ae a leas 


f 4 or 6 secondary axes in size- 
differentiated pairs (e.g., one longer, one short- 
er) 


. Inflorescence having ranks with either 2 sec- 


ondary axes or 4 equal axes or a cluster of 
axes, only rarely with 4 size-differentiated axes 


. Stipules with paler central triangle or keel, 
od 


often persistent at terminal 2-6 nodes 


. Stipules without paler central triangle or keel, 


. Dorsal seed surface with 10-15 irregular lon- 


M furrows, making jid er appear 
mooth; inflorescence main axis 18-33 cm 

“a 2) P. grandis 

longi- 

tudinal furrows, making dry fruit Me ridged; 

inflorescence main axis 1.5-22 c 


en widesprea 


. Inflorescence secondary axes in la or 2 ranks; 
b à 


abasco, Mexico, Guatemala, and Belize ..... 


) P. pleuropoda 
) 


. Inflorescence secondary axes in (2-)3-4(- 
ranks 


. Leaf blades thin membranous; corolla tube 4- 


5 mm long; calyx drying conspicuously pale 
in flower and fruit; southern Mexico, Guate- 
mala, Belize P. papantlensis 


. Leaf blades membranous to subcoriaceous; co- 


rolla tube 2-3.5 mm long; calyx not dryi ing 
conspicuously pale; Mexico through Nicar 
ua (9) P. iis 


. Leaf ‘gs pen dp de corolla 


tube x 1.5 mm; Coclé, 


P. cascajalensis 


6) 
. Leaf blades membranous or chartaceous or 


[e 
Ww 
a 


66a. 


66b. 


. Young 


) 
. Young stems glabrous or puberulent; dorsal 


. Mature leaf blades to 13(-16) cm 


23) 
i im spherical to slightly Ped. 3.5-4 


; x tube 2.5-3 x 


subcoriaceous; corolla tube at least 2 mm long 
or, if 1.5 mm long, the length > diameter ..... 56 
stems ferrugineous-pubescent; dorsal 
seed MES with 3 deep longitudinal furrows; 
Caribbean Nicaragua and Costa Rica ............... 


P. neillü 


seed surface not as a ove 


. Fruit when dry spherical, appearing smooth 


on outside; southern Mexico through El Sal- 
va 41) P. lundellii 


. Fruit when dry ellipsoidal or, if spherical, with 
conspicuous ee inal ridges on outside ...... 58 

. Young stems red-brown puberulent .................... 9 

. Young stems gla B rous 60 

. Inflorescence main axis 6-8.5 cm long; per- 


sistent calyx not evident or a minute beak; 
ih secl veins diverging 55?- 70*; wide- 
ead (2) P. clivorum 


; Inforescenee main axis 4.5 cm long; persis- 


calyx a cup ca. 
errs veins divergin rn 
Panamá Province, Panama ... (58) P. PN 


. Inflorescence secondary axes in 2 ranks c 


y near the apex of the main axis; 
Tabasco, Mexico (8) P. balancanensis 
ced 


. Inflorescence ege cni axes in ranks spac 


X evenly along the 


a. Leaves s green- dL or m black ............. 62 
61b. 


Leaves drying yellow-green to green-brown to 
red-brown to red-gray, never black 0. 
long; sec- 
ondary veins (8-)10-13 pairs; ^ tube 

-3 mm long; Heredia, Costa 
| P. Neben 


. Mature leaf blades to 10.5 cm long; secondar 


veins 7-9 pairs; corolla tube 2.5-5 m E g; 
Costa Rica and western Panama ............ 
P. orosiana 


m long, drying black; wides 


22) P. marginata 


(2 
. Fruit CR PUN 4-5.5(-6) mm long, drying 


red-Dro 


. Corolla Him 2 x 1.5 mm; leaf secondary veins 


(4-)5- 7(-8), the axils often with minute tufts 

of white hairs below; widespread „n 

(32) P. fruticetorum 
mm; leaf secondar 

s 7-10, the axils 5 we domatia or hairs 
in ea BEG arthagenensis 


. Inflorescence secondary axes TE E: 


main axis 


minally on the 


. Inflorescence secondary axes disposed i in reg- 


d nen main axis 

m long; Cartago, Costa Rica — LL. 

52) P. chitariana 

Leaf blades 11-29 x 4.5-10 cm, the sec- 

ondary veins (7-)11-14(-16); a ance 

main axis (1-)2-4 cm long; Costa q 
50) P. alfaroana 


. Herb or subshrub 0.1-0.3 m tall; leaf blades 
54 


bullate; western Panama P. hammelii 


. Shrub or tree at least 1 m tall; leaf blades not 


bullate 


84 Annals of the 
Missouri Botanical Garden 
68a. Leaf blades small and narrow, 2.5-3.5 x 0.5- 81b. Fruit when dry ellipsoidal, pov red-brown; 
0.7 em; rus p^ Said with up than 10 flow widespread (44) P. panamensis 
ers; Chiriquí, Panama ............. 40) P. dormiens 
68b. Leaf ruis usually inn and always wider 
Hands above; inflorescence of more than 10 GROUP 1. THE CARTHAGENENSIS GROUP 
692. Js secondary veins [4-)5-B(-9) and di- Shrub or sometimes small tree; young stems 
verging (67°-)70°-80%- 95°); leaf e glabrous or sometimes puberulent or ferrugineous- 
membranous; corola tube 2-2.5 mm pubescent; stipules usually sheathing, ovate (Fig. 
c np ie F - ES ) P. En UM a 2) or rarely lanceolate, the apex often acuminate 
v BM SOCODUATy Vems NOY ar apor Dr HON or rarely biacuminate, uniform in color (except P. 
the blades subcoriaceous with margins inrolled AAA : 
ar the corolla tbe 2.5-5 mm long in viridis with a darker central triangle), glabrous or 
70a. Fruit when n spherical 71 sometimes puberulent or ferrugineous-pubescent, 
70b. Fruit when dry ellipsoidal or obovoid ................... 75 sometimes fringed, caducous (Fig. 1). Leaf blades 
e p bie —€— 72 usually obovate to elliptic, less commonly oblan- 
. ul 1 e . . . 
12a. Young ares en ihe bark smooth fruit ceolate, drying reddish or greenish brown or gray; 
3.5-4 mm long; — Me .(22) P. marginata secondary veins 6-15 (except (18-)20-26 in P. 
72b. Young stems puberulen fissured or micrantha) pairs, diverging 45?-70*(-85?), eu- 
esie Tonga fruit only rarely less camptodromous (Fig. 3) or less commonly brochi- 
DON 73  dodromous, the axils usually lacking domatia (ex- 
73a. Persistent calyx conspicuous, 1 mm long; leaf i i E wA i i gua p "DB m t 
blades (8.5-)10-17 cm long; Costa Rica .. Sep IAE REESE TR PERRAS ME 
. (16) P. sylvivaga tertiary veins orthogonal reticulate (except per- 
73b. HD calyx inconspicuous; leaf blades current in P. micrantha and P. neillii; Fig. 5b). 
(4 RU 5-6(-6. Uh m long; eastern. Panamá Inflorescences panicles of cymes or, less common- 
vince, Panama cinc (21) P. liesneri | f el l P P adi 
y, of glomerules (P. micrantha and P. viridis), 
TAS Leaf blades (12 DIS 23 em long, drying green pedunculate except for P. lamarinensis and 
Mexico . : : 39) . dwyeri ta (Fig. 7g); secondary axes usually 
74b. Leaf blades (5- E 5-15 cm long, drying pale in 2 (or 3) size-differentiated pairs per rank 
to glossy red-brown; db veins (5 (Fig. 7a) or sometimes in 4 equal axes per rank 
8(-12); Mexico through El Sa ador M ... (P. lamarinensis and P. quinqueradiata) or some- 
Wes Tack acond uniri s divergin aa a Sai times in 1 pair per rank; bracts not conspicuously 
"ta Rica arid Panam o i D (93) -P. oaan enlarged. Corolla tubes (1.5-)2-5 mm long, the 
75b. Leaf secondary veins Mead almost always lobes without apical extensions. Fruit when dry 
« 75° 76  ellipsoidal or rarely obovoid; persistent calyx a beak 
76a. Inflorescence secondary axes diverging over ig. 9a) or not evident or rarely (P. micrantha) 
90*; Costa Rica and Chiriquí, po — . a tube ca. 0.5 mm long; seed dorsal surface with 
(46) P. stockwellii (3-) 4-5 deep longitudinal furrows, the ventral 
76b. Inflorescence secondary axes diverging less . , RU 
than 90° 77 surface with 2 deep or sometimes shallow lon- 
77a. Persistent calyx conspicuous, 1-3 mm long; gitudinal furrows (Fig. 
Guatemala through Nicaragua TE une tenis This group includes two pairs of very similar 
. 2 ed species plus the three other widespread species of 
77b. Persistent calyx inconspicuous or a beak never MR : . 
longer than: T mm unclear affinities, Psychotria carthagenensis, P. 
78a. Young stems usually SEREA stipules I clivorum, and P. viridis (Figs. 13, 14). The first 
ceolate, not sheathing; Mexico, Guatem pair, the P. quinqueradiata complex, includes its 
Belize P. erythrocarpa widespread namesake plus P. lamarinensis, which 
78b. Young ibi E es Aisa or fer- j is endemic to northern Costa Rica (Fig. 15); they 
19a arit ngth/wi Pros di NAR 0 share the following distinctive character states: in- 
79b. Leaf blade le ngth/width < 3 g] florescences lacking peduncles, and inflorescence 
80a. Fruit when dry 7-8 mm long; bark smooth; secondary axes 4 (not size-differentiated) per rank 
Mexico, Costa Rica, Panam 7 (Fig. 7g). The P. micrantha complex—including 
(45) P. sarapiquensis — the widespread P. micrantha and P. neillii, this 
80b. dis Mas diues ii bark iiid endemic to Caribbean southern Nicaragua and 
yum ongtuana y; CT) hs jc northern Costa Rica (Fig. 14)—shares a general 
8la. Fruit dry obovoid, usually black; south- ferrugineous vesture, biacuminate stipule apices, 


when 
ern Mexico through Nicaragua „uuu 
(47) P. trichotoma 


percurrent tertiary veins, and shallow longitudinal 
furrows on the seed ventral surfaces. 


Volume 76, Number 1 Hamilton 85 
1989 Mesoamerican Psychotria 
subg. Psychotria 
-+- a | T6: 
+ a. 4 
A A 9 
L + e SS 15 4 
$ ce 40/8, ay 
e 
E 
F 10 4 
300 km + Br 
e b 
105 100 95 90 85 a Ves) 
FIGURE 13. Distributions of Psychotria carthagenensis (circles) and P. viridis (triangles) in Mesoamerica. 
Of the seven species in the group, four are 


apparently normally distylous and one, Psychotria 
viridis, is homostylous (at least in Mesoamerica). 
The two remaining species are too little collected 
to determine whether they are distylous or thrum- 
monomorphic. 


LE 


. Psychotria carthagenensis Jacquin, Enum. 
Syst. Pl. Carib. 16. 1760. Cf. also Select. 
Stirp. Amer. Hist. 174, fig. 22. 1788. Ura- 
goga carthagenensis (Jacq.) O. Kuntze, Re- 
vis. Gen. Pl. 2: 959. 1891. Psychotria car- 
thaginensis auctt. TYPE: Colombia. Bolivar: 
Cartagena (fr). (Type material not seen; lo- 
cality and figure adequate to establish iden- 
tity.) Figures 1, 2a, 3a, 7a, 9a, 10a, 13. 


i des sagraeana Urban, Symb. Antill. 7: 450. 1913. 
E: Cuba. Wright 3588 (syntype, US). 


Shrub (0.5-)1-3(-6) m tall; young stems gla- 
brous, the bark smooth to irregularly furrowed; 
stipules sheathing, ovate, 6-8 x 3-5 mm (Fig. 2), 
irregularly fringed, glabrous, caducous, leaving a 
pale ridge usually with red-brown fringe (Fig. 1). 
Leaves subsessile to petiolate; petioles to 5(-10) 
mm long, glabrous, flat above; blades membranous, 
obovate or rarely elliptic, the apex acuminate, the 
base attenuate, (6-)7.5-13(-16) x (2-)2.5-5.5 
(-6) cm, glabrous above and below, drying red- 


brown to red-gray; secondary veins 7-10 pairs, 
diverging 60°-70°—75°), eucamptodromous (Fig. 
3), constantly arcuate, prominulous below, gla- 
brous, the axils lacking domatia or hairs; tertiary 
veins inconspicuous, orthogonal reticulate. /nflo- 
rescences terminal or pseudoaxillary, panicles of 
cymes (Fig. 7a); panicle branched to 3 or 4 degrees; 
main axis (2-)3.5-8 cm long, the peduncle 1.5- 
5 cm long; secondary axes in 3(—4) ranks, the first- 
rank axes 4, the longer pair 1-2.7 cm long, the 
shorter pair 0.4—1.5 cm long, the second-rank axes 
(2 or) 4, the longer pair 0.4-1.5 cm long, the 
shorter pair 0.3-0.8 cm long, the third-rank axes 
2 (or 4), the longer pair 0.3-1.0 cm long, the 
shorter pair 0.1-0.3 cm long, the fourth-rank axes 

, 0.3-0.4 cm long; cymes branched to (2-)3 
degrees; bracts and bracteoles prominent, trian- 
gular, 1 x 1 mm, the bracts sometimes enlarged 
to 8 mm long, caducous, glabrous, fringed. Flowers 
sessile to subpedicellate, the pedicels to 0.5 mm 
long; calyx cup-shaped, the tube 0.5 mm long, the 
lobes 5, triangular, minute, glabrous; corolla white, 
the tube cylindrical, 2.5-3 x mm, white pu- 
bescent in throat, the lobes linear, 1.5-2 x 1 mm; 
stamens 5, the filaments 3 mm long in pins, 3.5- 
4.5 mm long in thrums, the anthers 1 mm long; 
style 5-5.5 mm long in pins, 3-3.5 mm long in 
thrums, the branches linear. Fruit when dry ellip- 
soidal, 4.5—5.5(—6) mm long, 3.5-4.5 mm diam., 


maturing red, drying red-brown; persistent calyx 


86 


Annals of the 
Missouri Botanical Garden 


not evident or sometimes a beak (Fig. 9a); seed 
dorsal surface with 4 deep longitudinal furrows, 
the ventral surface with 2 deep longitudinal furrows 
(Fig. 10a) 

Distribution (Fig. 13). 
Mexico and Central America, occurring also in 
Florida (cultivated), Cuba, Colombia, Venezuela, 
the Guianas, Ecuador, Peru, Brazil, Bolivia, Par- 
aguay, Argentina, and Uruguay. In Central Amer- 
ica it is found mostly in Pacific coastal lowlands at 
elevations of 0—1,400 m, mostly under 400 m, in 
tropical moist to premontane moist and wet forest 
with equatorial to tropical climates. It has been 
collected in flower December-August, primarily 
February-June, and in fruit throughout the year, 
primarily September- February. 


Common throughout 


Selected specimens examined. MEXICO. CHIAPAS: SE 
of Mapastepec, Guamüchil, 7 Dec. 1950 (st), Enríquez 
6795 (MEXU); bank of Rio Salinas, 8 Feb. 1964 (fr), 
Lundell 17801 (MO); Tonala, Mojarra, nr. beach, 27 
Nov. 1947 (fr), Matuda 17144 (F, MEXU, NY); Hue- 
huetán, Islamapa, 27 June 1948 (early fr), Matuda 18002 
(MEXU — 2 sheets). oaxaca: Tuxtepec, Chiltepec, 7 June 
1965 (fl), G. Martínez C. 13 (F, ME 
PUEBLA: bosque aje Lain May 1952 
(MEXU — 2 sheets). TABASCO: Balancá 
14 May 1939 (fl), Matuda. 3025 (A, F— 2 P dani MEXU). 


Chavelas et al. ES-2442 (ENCB, MEXU); 
of Acayucan, dirt rd. nr. 0, 20 Feb. 1976 (fl), 
Croat 32755 (MO). GUATEMALA. ESCUINTLA: Cuyuta, 60 
m, Apr. wa? (A), Donnell-Smith 2074 (K, US— 2 sheets); 
Anubis, nr. Obispo, lowland Me 20 Apr. 1937 (fl), 
Muenscher 12370 (F, GH). izaBAL: Dept. Liv Er Rio 
O m, Mar. 1889 (fl), Donne aces 1821 (U 
RETALHULEU: plains between Nueva and Champeri- 
co, 120 m or less, 18 Feb. us (8. cine 87667 
(F, US). SAN MARCOS: 1-2 m of Ocós, palmetto flats, 
3-5 m, 15 e 1940 (8). Piu: 37866 (F). 
e des EZ: 7 mi. S of Tiquisate along rd. within 3 
of ocean, S A Alotenango Farm, 30-50 m, 19 June 
1942 (early fr), Steyermark 47770 (F). ZACAPA: trail 
between Rio Hondo and waterfall, 250-400 m, 10 Oct 
1939 (fr), Steyermark 29457, 29458 (F). HONDURAS. 
COMAYAGUA: u Rio Yure con Rio Humuya, 100 
agua, 200 m, 22 Nov.-31 Dec. 
1980 (fr), C. Nelson et al. 6125 (TEFH). cortés: orilla 
del Rio Humuya, 40 km N de Santa Cruz de Yojoa, 100 
m, 1-30 Nov. 1980 (fr), C. Nelson et al. 5843 (TEFH); 
8 (A), Thieme 527 
(US). OLANCHO: midway between Juticalpa and Campa- 
mento, along Rio Juticalpa, 25 Feb. 1982 (fr), Blackmore 
& Heath 1956 (TEFH); matorrales del Rio Talgua cerca 
de la Escuela Granja Demonstrativa de Catacamas, 400 
m, 26 Apr. 1957 (fl), 4. Molina R. 8337 (F, MO); Dulce 
Nombre de Culmí, 600 m, 1 May 1982 (fl), D. Molina 
109 (TEFH). YORO: nr. Coyoles, Aguan River valley, 28 
June 1938 (fl), Yuncker et al. 8040 (F, GH, K, MO, 
, US). EL SALVADOR. SAN MIGUEL: Laguna de Olomega, 
75 m, 20 Feb. 1922 (fl, Standley 21043 (GH, MO, 
US). SANTA ANA: N shore of Lago de Gúija, Hda. 
Barra, 470 m, 27 Feb. 1946 (fl), Conon 1009 (F). 


E 


— 


p 


SONSONATE: vic. Sonsonate, 220-300 m, 18-27 Mar 
1922 (A), Standley 22336 (GH, NY, US). NICARAGUA. 
CHINANDEGA: al SW del Volcán Cosigüina, 29 Sep. 1981 
(fr), Fonseca 118 (MO); vic. Chichigalpa, 90 m, 12-18 
July 1947 (A), Standley 11211 (F, US). CHONTALES: Río 
Las Vainillas, 12%01'N, 85°14’W, 150 m, 21 Jan. 1983 
(fr), Sandino 3939 (MO); vic. Juigalpa, 160 m, 4-13 
June 1947 (fl), iis rui 9303 (F); rd. from Juigalpa NE 
to La Libertad, ca. m NE of Rio Mayales, 12°12'N, 


Stevens 4087 (MO); ca. 5.3 kn W 
12%00'N, 85?12'W, 150-170 m, 7 June 1981 (fl, early 
fr), W. D. Stevens & Henrich 20521 (MO). EsrELÍ: Hwy. 
l, ca. 19.2 km N of entrance into Esteli, 13?16'N, 
86°21'W, 650-700 m, 25 Dec. 1978 (fr), W. D. Stevens 
11243 (MO); 25 Dec. 1978 (A), W. D. Stevens 11256 
(MO). GRANADA: Laguna Juan Tallo, 11?45'N, 85?*59'W, 
65-70 m, 18 Dec. 1980 (fr), P. P. Moreno 5449 (MO). 
JINOTEGA: Sta. Gertrudis, 30 km E de Jinotega, 1,000 n 
20 Jan. 1980 (fl), d & Moreno 876 (MO); c 

5 mi. E , 750 m, 6 Aug. 1977 (fl), Gon 
42883 (MO). 
12*18'N, 8642^W. 35 m, 31 May 1983 (fl), W. 

Stevens 22161 (MO); Rio Sinecapa at ford of e 


~ 


Stevens 3894 (MO). MATAGALPA: Hda. Sta. 
Ostuma, 1,300-1,600 m, 13 Dec. 1976 ay Tomlin 126 
MO). RÍO SAN JUAN: rd. to San Carlos 5 km SE of Rio 
Oyate, 11?42'N, 84°57'W, 40 m, 28 Aug. 1983 (early 
fr), Miller & Nee 1375 (MO); San Bartolo, 29 dily 1972 
(fr), Robbins 6176 (F, GH, Y). RIVAS: Punte de 
Tola, 40 m, 28 Feb. 1984 (fl), M. Castro 57 (MO); Isla 
de Ometepe, N de Volcá 
Angeles, 11?34'N, 85°37'W, 200-250 m 
(fr), didi 447 (MO). ZELAYA: “San Agustin, " al SE 
e Rama, 12%09'N, 84°12’W, 60 m, 22 May 1984 (fl), 
rro 544 (MO). Costa RICA. GUANACASTE: Nicoya, 


— 


Finca La Pacifica, 5 nas, 
1969 (fl), Davide & Pohl yes (F. MO, NY); playa de 
Sámara, al S de Nicoya, 10 m, 23 Mar. 1963 (fl), A 
iménez M. 520 (CR, F); Santa Rosa National Park, 
10?48'N, 85%40'W, 0-20 m, a Jan. 1978 (fr), Liesner 
4250 (CR, MO); Río Sapoá, 4 km N of La Cruz, 11?06'N, 
85°35" W, 2 Feb. 1978 (fr), s 4863 (MO); Comelco, 
5 km NW of Bagaces, 150 m, 30 Jan. 1973 (fl), Opler 
1683 (CR, F, MO — 2 sheets); Ballena Rd., 17 July 1918 
(early fr), Rowlee & Rowlee 166 (US); vic. Libano, 260- 
360 m, 15 Jan. 1926 (fr), Standley & Valerio 44926 
(US); banks of Rio Corubici, 17 Aug. 1967 (fr), Whitmore 
28 (DUKE, F, MO, NY). PUNTARENAS: 8 km N of Bar- 
ranca, 1 km N of Miramar turn-off, Qda. Negros, 10°02'N, 
84?45'W, 20-30 m, 30 Apr. 1983 (fl), Liesner et al. 
15125 (CR, MO). PANAMA. CANAL AREA: Barro Colorado 
Island, 23 May 1970 (fl), Croat 10347 (DUKE, F, MO, 
NY); Fort Clayton, Apr. 1965 (fl), Dwyer & Robyns 8 
MO — 2 sheets); N of Gamboa, 9°05'N, 79?45'W, 25 
m, 22 Mar. 1983 (fl), Hamilton 3677 (CR, MO); Parque 
Soberania nr. Las Cruces trail, 9%15'N, 79?45'W, 12 
Mar. 1983 (fl, fr), Hamilton et al. 3296 (MO); Mount 
Hope, 7 Apr. 1921 (fl), e 74 (US); 1 mi. from 
Gaillard Hwy. on track o a Chiva Road, 9?00'N, 
1982 i uid & Schmal- 
zel 4841 (MO) Balboa, Nov. 23-Jan. 1924 (fr), 
end 25501 en An e. Rio dp sd 
Chiriqui and Veraguas at Panam. Hwy., 8 
81°30! W, 100 m, 10 se 1983 (fr), Shain et al. p 


— 


Volume 76, Number 1 
1989 


Hamilto 87 
esos Psychotria 
subg. Psychotria 


(MO). cocLé: El Valle de Antón, 300-600 m, 2-3 Dec. 
1967 (fr), Lewis et al. 3543 (MO); Chorrerita, 21 Sep. 
1973 (fr), L. Medina 6 (DUKE); vic. Penonomé, 15- 
300 m, 23 Feb.-22 Mar. 1908 (fr), R. S. Williams 59 
(NY — 2 sheets, US). DARIÉN: nr. Yaviza, ca. O m, 8 Jan. 
1975 (fr), nd 13490 (GH, MO). HERRERa: vic. Ocú, 
aE Pasoancho, 19 Feb. 1963 (fl, fr), Stern et al. 1735 
US). Los saNTOs: Pedasi, bank of beach, 10 Aug. 
m. (early fr), a 2489 (MO, US); Rio Tonosi, vic. 
"oe 25 May 1967 (fl), dui et al. 1540 (DUKE, 
MO); along Rio Tones ca. 9 mi. N of Tonosi, 17 July 
1970 (early fr), Luteyn s Mur 1387 (DUKE, F, MO). 
PANAMÁ: Rio Majé, ca. speedboat minutes from con- 
fluence with Rio E p m, 19 Apr. 1976 (fl), Croat 
34403 (MO, NY); Cermeño, 7 Apr. 1965 (fl), Dwyer 
6588 (MO); 2 mi. W of Chepo, 6 Apr. 1972 (fl), Gentry 
497 1 (F, MO, NY); Rio Ipeti, S of Panam. Hwy., 9°03'N 
78°25'W, 100 m, 17 Sep. 1982 (st), SM eR & D'Arcy 
1336 (MO); falls of La Chorrera, 5 Apr. 1969 (fl), Lewis 
et al. 5205 (MO); Rio Crystal de Nuevo Arraiján, 4 Dec. 
1971 (fr), L. Salazar 15 (DUKE, MO); Cerro Azul, nr. 
lake, 600 m, 27 May 1966 (fl), Tyson & Blum 4066 
MO); marshy area 2 mi. S of Tocumen Airport, 25 July 
1969 (early fr), Tyson & Clewell 5875 (DUKE, MO). 
VERAGUAS: just S of Santa Fe, 450 m, 17 Nov. 1973 (fr), 
Nee 8036 (MO, US). 


— 


Modern authors have perpetuated an early mis- 
mee of Las specific epithet by spelling it “car- 
h _ instead of the correct original ** 
(D. Nicolson, pers. 
collection by Vahl, from the type region of Car- 
tagena, resides in Berlin (Willdenow 4083) and 
has been annotated as P. carthagenensis; it is, 
however, P. nervosa and is clearly not the type 
of Jacquin's species. 

Psychotria carthagenensis may be recognized 
by its obovate leaves drying dull red-brown, eu- 
camptodromous secondary venation (Fig. 3), sec- 
ondary inflorescence axes in two unequal pairs per 
rank (Fig. 7a), corolla drying red, ellipsoidal fruit 
drying reddish, and seed in cross section with the 


car- 
comm.). A 


orsal surface having four deep furrows and the 
ventral surface having two deep furrows (Fig. 10a). 
It is expected that careful analysis will reveal that 
this species exhibits more primitive character states 
than any other neotropical species in the sub 
Mexican and Guatemalan collections suggest that 
the leaf blades are larger there than elsewhere in 
Central America. 


2. Psychotria clivorum Standley & Steyer- 
mark, Publ. Field Mus. Nat. Hist., Bot. Ser 
23: 87. 1944. Based on Psychotria limonen- 
sis var. angustifolia Standley, Publ. Field Mus. 
Nat. Hist., Bot. Ser. 17: 282. 1937 (see pro- 
tologue of P. clivorum). TYPE: Guatemala. 
Suchitepéquez: Finca Mocá, 1,020 m, 7 Jan. 
1935 (fl), Skutch 2073 (holotype, F; isotype, 
A). Figure 14. 


Shrub or small tree, 2.5-6 m tall; young stems 
red-brown puberulent, the bark smooth to irregu- 
larly furrowed; stipules sheathing, ovate, the apex 
sometimes biaristate, 8-9(-14) x 4-4.5(-5) mm 
glabrous to puberulent, caducous, leaving a pale 
ridge with red-brown fringe. Leaves petiolate; pet- 
ioles 4-10 mm long, glabrous, grooved above; blades 
membranous, elliptic, the apex cuspidate, the base 
attenuate, (9-)1 2-22 x (3-)4-6.5(-8.5) cm, gla- 
brous above, glabrous below with the midvein some- 
times sparsely puberulent, drying red-gray to gray- 
green; secondary veins 9-13 pairs, diverging 55°— 
70%, eucamptodromous to brochidodromous, 
straight then arcuate near margin, prominulous 

elow, glabrous to minute puberulent below, the 
axils lacking domatia or hairs; tertiary veins evident 
to conspicuous, orthogonal reticulate to slightly 
percurrent. /nflorescences terminal or pseudoax- 
illary, elongate panicles of cymes; panicle branched 
to 4 degrees; main axis 6-8.5 cm long, the pe- 
duncle 2.5-5 cm long; secondary axes in (3-)4 
(-5) ranks, the first-rank axes 4 or rarely 2, the 
t pair 1.5-2.5 cm long, the shorter pair 0.6- 


long, the fourth-rank axes 2, 0.3-0.6 cm long, the 
fifth-rank axes 2, 0.3 cm long; cymes branched to 
1-3 degrees; bracts linear, to 2 mm long, puber- 
ulent; bracteoles minute, irregular, to 0.6 mm long, 
puberulent. Flowers sessile to subpedicellate, the 
pedicels to 1 mm long; calyx cup-shaped, the tube 
0.3-0.5 mm long, the lobes 5, triangular to barely 
evident, to 0.3 mm long, puberulent; corolla white, 
the tube cylindrical to campanulate, 2-2.5 x 1.5 
mm, white pubescent in throat, the lobes 5, linear, 
mm; stamens 5, the filaments 2.5 mm 
long in pins, 3.5-4 mm longi in thrums, the anthers 
l mm long; style 3.5-4 mm long in pins, 
mm long in thrums, the branches short, dubie: 
Fruit when dry ellipsoidal, 5-6 mm long, 4-4.5 
mm diam., maturing red, drying deep red-brown; 
persistent calyx not evident or sometimes a beak; 
seed dorsal surface with 4 deep longitudinal fur- 
rows, the ventral surface with 2 deep longitudinal 
furrows. 


Distribution (Fig. 14). Not commonly col- 
lected but widespread from southern Mexico to 
western Panama, occurring at low elevations, 0— 
200 m, in Veracruz and Oaxaca, Mexico, and 
Bocas del Toro, Panama, and higher elevations, 
600-1,400 m, in other parts of the range. It has 
b 


een found in tropical moist and premontane rain 


Annals of the 
Missouri Botanical Garden 


9 
ep 
E + + + a + 15 4 
e 
A 
a e 
j A 10 + 
300 km + "e +. 
e 
A d 
105 100 95 90 85 * “eo 
FIGURE 14. Distributions of Psychotria clivorum (triangles), P. micrantha (circles), and P. neillii (squares) in 


Mesoamerica. 


forest with equatorial to tropical climate. It has 
been collected in flower January-May, especially 
March-April, and in fruit November- March. 


Selected specimens examined. MEXICO. OAXACA: 
Juchitán, Montaña de la Pedrera, Palomares, 90 m, 14- 
17 Apr. 1970 (fl), MacDougall s.n. (NY). VERACRUZ: 
Mpio. Hidalgotitlán, del km 3 al km 5 del camino de Plan 
de Arroyos al Alvaro Obregón, 17°15'N, 94%40'W, 130- 
re m, 2 Apr. 1974 (fl), B. Dorantes 2731 (MEXU— 
2 sheets, MO); Mpio. ober ase Kreis Her- 
manos Cedillo a 2 km por el camin a Tied S dk 
m, 1 Apr. 1975 (fl), Ortíz & M e 71 (MEX 
3 sheets); Mpio. E poa brecha Hermanos Cedillo 
Agustín Melgar, 17?13'N, 94°35'W, 153 m, 26 Apr. 
1974 (fl), B. ig ae 418 (MEXU, M 
zacoalcos Rive O m, Mar. 1937 (fl), 
8677 (F). EC: QUEZALTENANGO: between Finca 
Pirineos and Finca Soledad, lower south-facing slopes of 
Volcán Santa María, between Santa Maria de Jesüs and 
Calahuaché, 1,300-1,400 m, 5 Jan. 1940 e Steyer- 
mark 33543 (F, US). san Marcos: Rio , below 
Rodeo, 600 m, 14 Mar. 1939 (fl), aller 68772, 
68783 (F); Volcán Tajumulco, Finca El Porvenir, along 
Rio Cabüs above Potrero Matasan, 1,000-1,300 m, 12 
Mar. 1940 (fl), Steyermark 37628 (A, F). NICARAGUA. 
CARAZO: 1 km E of San Marcos, ravine at edge of coffee 
plantation, 2 May 1976 (fl), Neill 285 (MO). MATAGALPA: 
Macizo de Peñas Blancas, Finca San Sebastián, 1,050 m 
24 Nov. 1981 (fr), Mid e al. 5156 s E (MEXU). 
Costa RICA. CARTAGO: Tucurrique, 650 m, Mar. 
1899 (fl), Tonduz . 13018 (US). PANAMA. BOCAS DEL TORO: 
vic. Almirante, Jan.-Mar. 1928 (fr), euge 453 (F); 
Jan.-Mar. 1928 (fl), Cooper 566 (F, GH, NY). cocLÉ 
El Valle, 800-1,000 m, 28 June 1967 (early fr), Duke 
13151 (MO—3 sheets); Cerro Pilón, to 930 m, 19 Jan. 


1968 (fr), Dwyer 8326 (MO); Cerro Pilón, 600-800 m, 
28 Mar. 1969 (fr), Dwyer et al. 4553 (MO). VERAGUAS: 
N of Santa Fe, Cerro Tute, Escuela Agricola Alto de 
Piedra, 1,000-1,200 m, 4 Feb. 1977 (fr), Folsom 1603 
AO 17 Oct. 1974 (early fr), Mori & Kallunki 2541 
MO 


onm clivorum may be recognized by its 
elliptic leaves drying red-gray to gray-green, sec- 
ondary veins well-spaced and diverging 55°-70°, 
inflorescence eea axes in two unequal pairs 
per rank, the higher ranks decreasing 
formly, corolla drying i ellipsoidal fruit, and seed 
cross section with four deep longitudinal furrows 
on the dorsal surface and two deep longitudinal 
furrows on the ventral surface. Psychotria clivo- 
rum shares many character states with P. car 
thagenensis, but the former differs in having sti- 
pules often biaristate, larger elliptic (versus obovate) 
leaf blades, more secondary veins, often brochi- 
dodromous secondary venation, and larger inflo- 


in size uni- 


rescences. 

Stipule morphology varies geographically, as 
Guatemalan plants appear more often biaristate 
while Nicaraguan plants have exceptionally large 
stipules. Secondary venation is generally eucamp- 
todromous, but in Nicaragua the secondary loops 
are sufficiently robust to appear brochidodromous. 


3. Psychotria lamarinensis C. Hamilton, Phy- 
tologia 64: 227. 1988. TYPE: Costa Rica. Ala- 


Volume 76, Number 1 
1989 


Hamilton 89 
Mesoamerican Psychotria 
subg. Psychotria 


P UM 2 a0 d T.C.4 
| H- ies a 
8 o 
e * " 
E CDU. X B "- f 


|: A 
10 
300 km + N + 
co d 
Q 
105 100 95 90 85 80 
FIGURE 15. Distributions of Psychotria lamarinensis (triangle) and P. quinqueradiata (circles) in Mesoamerica. 


juela: E of Rio San Rafael, W of La Marina, 
10%23'N, 84?23'W, 500 m, 19 May 1968 
(f), Burger & Stolze 5062 (holotype, NY; 
isotype, MO). Figure 15. 


Shrub 1.8-2 m tall; young stems glabrous, the 
bark pale, smooth; stipules ovate, 12 x 8 mm 
glabrous, caducous, leaving a pale ridge with thick 
red-brown fringe. Leaves petiolate; petioles 5-7 
mm long, glabrous, terete; blades membranous, 
elliptic, the apex acuminate, the base attenuate to 
attenuate-truncate, (13-)16-20 x (5-)7.5-9.5 
cm, glabrous above and below, drying green-brown 
above, red-brown below; secondary veins 9-11 
pairs, diverging 70°-85°, eucamptodromous, con- 
stantly arcuate, elevated below, glabrous, the axils 
lacking domatia or hairs; tertiary veins evident, 
orthogonal reticulate. Inflorescences terminal, con- 
densed globose panicles of cymes; panicle branched 
to 3 degrees; main axis 1.5-2 cm long, the pe- 
duncle lacking; secondary axes in 2-3 ranks, the 
first-rank axes 4, subequal, 0.5-1.5 cm long, the 
second-rank axes 4, subequal, 0.4-0.7 cm long, 
the third-rank axes 4, subequal, 0.2 cm long; cymes 
branched to 1-2 degrees; bracts triangular, 2 mm 
long, glabrous; bracteoles not evident. Flowers ped- 
icellate, the pedicels 0.5-1.5 mm long; calyx cup- 
shaped, 0.5 mm long, the lobes not evident to 
barely evident, glabrous; corolla white, the tube 
cylindrical, 3 x 1 mm, white pubescent in throat, 


the lobes 5, linear with 1.5 mm linear extension 
from near apex, 2 X 1 mm; stamens 5, the fila- 
ments not seen in pins, 3.5—4 mm long in thrums, 
the anthers 0.7 mm long; style not seen in pins, 
2-2.5 mm long in thrums, the branches linear. 
Fruit not seen. 


Distribution (Fig. 15). Known only from the 
type locality near La Marina, Alajuela, Costa Rica, 
at ca. 500 m elevation in a region of tropical wet 
to premontane wet forest with equatorial-moun- 
tainous climate. It was collected in flower on May 
19. 


Additional specimen examined. OSTA Rica. 
ALAJUELA: E of Rio San Rafael, W of La Marina, 10?23'N, 
84°23'W, 500 m, 19 May 1968 (fl), Burger & Stolze 
5069 (MO, NY). 


Psychotria lamarinensis may be recognized by 
its large, broad-elliptic leaves with the base nar- 
rowly subcordate, reduced globose inflorescence 
drying red-brown, and broad-ovate stipules. Psy- 
chotria lamarinensis may be a local derivative 
from P. quinqueradiata, from which the former 
differs in having much larger leaves with secondary 
veins diverging 709-85? instead of 45°-60° and 
shorter (3 mm vs. 4-5 mm) corolla tubes. 

Only a short-style flower morph has been seen, 
but only two flowering collections do not suffice to 
suggest that the species is thrum-monomorphic. 


90 


Annals of the 
Missouri Botanical Garden 


4. Psychotria micrantha Kunth in Humboldt, 
Bonpland and Kunth, Nov. Gen. Sp. 3: 
pl. 284. 1819. Uragoga micrantha (Kunth) 
Kuntze, Revis. Gen. Pl 1. 1891. Ma- 
pouria micrantha (Kunth) Wernham, Kew 
Bull. 1914: 69. 1914. TYPE: Peru(?) (fl), Bon- 
pland s.n. (holotype, P, n.v., fragment, F). 
Figure 14. 


» 


eer fh si Roemer & ga Syst. Veg. 5: 
192. 1819, non Kunth in H.B.K. (which is a syn- 
nym 

12): 364, 
Schultes: Colombia. Rio Magdalena, Humboldt . s.n. 
(holotype, B— Willdenow 4107). 


Tree 1.5-5(-8) m tall; 
ferrugineous-pubescent, the bark slightly furrowed 
longitudinally; stipules sheathing, ovate, the apex 
biacuminate, 13-18(-22) x (5-)6- 7(-8) mm, fer- 
A caducous, leaving a pale ridge 
with red-brown fringe. Leaves petiolate; petioles 
1-2.2 cm long, ferrugineous, grooved above; blades 
membranous, elliptic, the apex acuminate, the base 
cuneate to attenuate, (12-)20-32 x (5-)8-13 
cm, puberulent above, the veins ferrugineous-pu- 
bescent, ferrugineous-pubescent below, drying 
green-brown to red-brown; secondary veins 
(18-)20-26 pairs, diverging (60°-)65°-70°, eu- 
camptodromous to sometimes brochidodromous, 
constantly arcuate or straight then arcuate near 
margin, elevated the axils 
lacking domatia or hairs; tertiary veins evident to 
conspicuous, percurrent, the quaternaries orthog- 
onal reticulate. Inflorescences terminal or pseu- 
doaxillary, panicles of glomerules; panicle branched 
to 4 degrees; main axis 10.5-17 cm long, the 
peduncle 6-8.5 cm long; secondary axes in 3-4 
ranks, the first-rank axes 4 or 6, the long pair 
2.5-5.5 cm long, the medium pair 1-2.5 cm long, 
the short pair 0-0.5 cm long, the second-rank axes 
4 or 6, the long pair 0.5-2 cm long, the medium 
pair 0.2-0.8 cm long, the short pair 0-0.2 cm 
long, the third-rank axes 4, the longer pair 0.4— 
0.8 cm long, the shorter pair 0-0.4 cm long, the 
fourth-rank axes 4, the longer pair 0.4 cm long, 
the shorter pair 0.1 cm long; bracts ovate-acu- 
minate, to 6 mm long, ferrugineous-pubescent. 
Flowers sessile; calyx cup-shaped, the tube 0.2- 
0.5 mm long, the lobes 5, barely evident to tri- 
angular, to 0.3 mm long, white pubescent without; 
corolla white, the tube cylindrical, 2 x 
short white pubescent in throat, the lobes 5, ovate, 

-1.5 x 1 mm; stamens 5, the filaments 1.5 mm 
long in pins, 3 mm long in thrums, the anthers 1 


young stems densel 


elow, ferrugineous, 


] mm 


» 


mm long; style 3.5-4 mm long in pins, 2-2.5 mm 


long in thrums, the branches linear. Fruit when 
dry ellipsoidal, 4 mm long, 2.5-3(-3.5) mm diam., 
maturing red, drying black or sometimes red-brown, 
sparsely ferrugineous; persistent calyx a tube ca. 
.9 mm long, drying pale brown; seed dorsal sur- 
face with 4—5 deep longitudinal furrows, the ven- 
tral surface with 2 shallow longitudinal furrows. 


Distribution (Fig. 14). Guatemala and Nica- 
ragua through Panama, mostly on the Caribbean 
side, at elevations of 0- m in usually tropical 
moist to wet forest with equatorial to sometimes 
tropical-equatorial climate. It occurs also in Co- 
ombia, Venezuela, Peru, and Bolivia. 
Psychotria micrantha has been collected in flower 
April-September, December, and January, and in 
fruit July-March, primarily September- Decem- 


Ecuador, 


ber. 


Selected specimens examined. GUATEMALA. ALTA 
VERAPAZ: Sebol, WN he village, nr. Rio Sebol, 11 
July 1964 (fl), Contreras 5281 (LL); Chahal, Sepur Ranch, 
bordering Rio Chahal, 8 Oct. 1968 (fr), Contreras 7866 
(LL, NY). IZABAL: trail from 


Mar. 19 


R 
Steyermark 45415 (F, US). NICARAGUA. CHONTALES: Vic. 
La Libertad, 500-700 m, 29 May-1 June 1947 (fl), 
Standley 9097 (F). ZEL AYA: Cuamil o brenas a lo largo 
del Río Grande, 24 Apr. 1949 (fl), 4. Molina R. 2370 
m à 


(F, G ; Comarca del Cabo, 25 k S de Tronquera 
cerca de wa, San o, 35 m, 22 Aug. 1965 
(fl), 4. Molina R. 15088 (F, NY, US); Mpio. de Siuna, 
Danli, 100- m, 31 Sep. 1982 (fr), si 204 (MO). 
Costa RICA : ARTAGO: El Bron, 2 Tur ialba y Floren 
^ 00 m 6 July 1965 ila . Jiménez - M. 3308 (F); 

rialba, ara os del Instituto, 600 m, Oct. 1949 


3 


dan 1872 (US); Turrialba, Puente Cajón, 625 
31 May 1972 (fl), Poveda 129 (MO). LIMÓN: Golden: 
grove, drenaje de Rio Reventazón, 15 m, 23 Oct. 1951 
(fr), Shank & Molina 4399 (F, GH, US). PUNTARENAS: 
nr. Buenos Aires, between Qda. Grande and Qda. Gua 
jiniquil, 350 m, 1 Mar. 1966 (fr), 4. Molina R. et al. 
18146 (CR, F, MO, NY). PANAMA. BOCAS DEL TORO: Milla, 
26 July 1971 (fl, fr), Croat & Porter 16272 (ENCB, F, 

MO, NY); Old Bank Island, 23 Jan. 1941 (fl, fr), Wedel 
1869 (GH, MO); Little Bocas, Chiriqui Lagoon, 9 July 
1941 (A), Wedel 2508 (F, GH, MO). CANAL AREA: Barro 
Colorado Island, 20 Sep. 1968 (fr), Croat 6235 (DUKE, 
F, MO, NY); 25 June 1968 (8), Foster 641 (DUKE); rd. 
between Locks and Fort Sherman, 10 July 1971 (fl), 
Croat 15361 (MO, NY); Madden Dam, Boy Scout Rd., 
23 July 1968 (fl), Dwyer & Lallathin 8822 (F, MO, 
NY); along old Las Cruces Trail between Fort Clayton 
and Corozal, 31 Dec. 1923 (fr), Standley 29198 (US). 
COLÓN: N side of Rio Guanche, 0.5 km upstream Puerto 


Pilon—Portobelo Hwy. bridge, 5-30 m, 22 Sep. 1973 
(early fr), Nee 7104 (F, GH, MO); Palmas Bellas, 30 
May 1971 (fl), Thoms 35 (DUKE, F, MO). DARIÉN: con- 


e of Río Chucunaque and Rio Canglón, 5 July 1962 
(fl), Duke 5114 (GH, MO, US); Rio Chico across from 


Volume 76, Number 1 
1989 


Hamilton 91 
Mesoamerican Psychotria 
subg. Psychotria 


ca de Tesca, 18 July 1962 (fl), Duke 5217 (GH, MO, 
US) Cocalita, 13 Aug. 1963 (fl), Dwyer 4398 (US); Rio 
ucro, 11 June 1959 (fl), n 


de Cape, = El Real, b ew 1959 (fl), Stern et al 


P. Ww sd yA en 

mul Ep 3 (fr), 

Standley 26604 (A). SAN BLAS TM de Ho Mu- 

latupo, 17 Aug. 1967 (early d. Elias 1729 (CH, MEXU, 

MO, US); mainland opposite Ailigandi, 7-8 Dec. 1966 
FG O, NY, US 


Rio de Jesús, Rio Trinidad, 3 Aug. 1961 (A), Dwyer 1319 
(GH). 


Psychotria micrantha may be recognized by 
its densely ferrugineous vesture, large (20-32 x 
8-13 cm) elliptic leaves with many (20-26) sec- 
ondary veins and conspicuous percurrent tertiary 
veins, large panicles of glomerules, and small (4 
x 2.5-3 mm) pubescent ellipsoid fruits. 


gn 


Psychotria neillii C. Hamilton & Dwyer in 
C. Hamilton, Phytologia 64: 231. 1988. TYPE: 
Nicaragua. Río San Juan: Rio Sábalo, 2 km 
al O de Sta. Eduviges, 11?03'N, 84?29'W, 
80 m, 18 Feb. 1984 (fl, fr), P. P. Moreno 
23060 (holotype, MO). Figure 14. 


Shrub or small tree, 1-4 m tall; young stems 
ferrugineous-pubescent, the bark smooth; stipules 
sheathing, lanceolate, the apex biacuminate, 10- 

3-5 mm, ferrugineous-pubescent, ciliate, 
caducous, leaving a pale ridge with red-brown fringe. 
Leaves petiolate; petioles 4-8 mm long, ferrugin- 
eous-pubescent, terete; blades membranous, ob- 
lanceolate, the apex acuminate, the base cordate, 

8 x 4-7.5 cm, glabrous above, the midvein 
basally ferrugineous-pubescent, sparsely ferrugine- 
ous-pubescent below, drying green-brown to red- 
brown; secondary veins 12-15 pairs, diverging 
(609—)65?-75?, brochidodromous, constantly ar- 
cuate, elevated below, ferrugineous-pubescent be- 
low, the axils lacking domatia or hairs; tertiary 
veins evident to conspicuous, percurrent, the qua- 
ternaries orthogonal reticulate. /nflorescences ter- 
minal or pseudoaxillary, panicles of cymes; panicle 
branched to 3-4 degrees, the axes delicate; main 
axis (6-)9-15 cm long, the peduncle (3.5-)6-8 
cm long; secondary axes in 3 ranks, the first-rank 
axes 4, the longer pair (1.5-)3-6 cm long, the 
shorter pair (0.8-)1-3.5 cm long, the second-rank 
axes 2 or 4, the longer pair (0.8-)1.4-3.2 cm 
long, the shorter pair ca. 1.5 cm long, the third- 
rank axes 2, 0.6-1.4 cm long; cymes branched to 
2-3 degrees; bracts triangular, 4 mm long, fer- 
rugineous-pubescent; bracteoles linear, 0.5 mm 
long, ferrugineous-pubescent. Flowers sessile to 


pedicellate, the pedicels to 1 mm long; calyx cup- 
shaped, the tube 0.3 mm long, the lobes 5, tri- 
angular, 0.2 mm long, ferrugineous-pubescent; co- 
rolla cream, the tube cylindrical, 1.5-2 x 1 mm, 
white pubescent in throat, red-brown pubescent 
without, the lobes 5, triangular, 1 x 0.8 mm; 
stamens 5, the filaments not seen in pins, 2.5 mm 
long in thrums, the anthers 0.8 mm long; style not 
seen in pins, 2 mm long in thrums, the branches 
linear. Fruit when dry ellipsoidal to obovoid, 5-7 
mm long, 3-3.5 mm diam., maturing red, drying 
dark red-brown, sometimes puberulent; persistent 
calyx inconspicuous or a minute beak; seed dorsal 
surface with 3 deep longitudinal furrows, the ven- 
tral surface with 2 shallow longitudinal furrows. 


Distribution (Fig. 14). 
bean coastal Costa Rica and just north into Nic- 
aragua, at 80-400 m elevation in tropical moist 
to wet forest with equatorial climate. It has been 
collected in flower in January, February, and April 
and in fruit in February and May. 


Known from Carib- 


Additional specimens examin ned. NICARAGUA. RÍO 
SAN JUAN: nr. Cario Chontaleno, 20 km of El Castillo, 
200 m, 18-21 Apr. 1978 (fl), Neill & Vincelli 3503 
(MO). Costa RICA. HEREDIA: Magsasay, entre el cam- 
pamento Canta Rana y Rio Peje, 400 m, 14 Jan. 1983 
(A), Chacón 76 (MO). LIMÓN: 7 km of Bribri, 100- 
250 m, 4 May 1983 (early fr), L. D. Gómez et al. 2035 


MO — 2 sheets) 


Psychotria neillii may be recognized by its great 
resemblance to P. micrantha and its delicate fer- 
rugineous-pubescent inflorescence axes. Psycho- 
tria neillii differs from P. micrantha in having 
smaller (10-18 vs. 20-32 cm long) mature leaves 
with fewer (12-15 vs. 20-26) secondary veins, 
delicate inflorescence axes, and larger (5-7 mm 
vs. 4 mm long) fruit. 

All three flowering collections are thrums; ad- 
ditional collections are necessary to determine 
whether the species is distylous or thrum-mono- 
morphic. 


6. Psychotria quinqueradiata Polakowsky, 
Linnaea 41: 570. 1877. Uragoga quinque- 
radiata (Polak.) Kuntze, Revis. Gen. Pl. 2: 
962. 1891. TYPE: Costa Rica: prope UNE José 
et in silvis primaevis Carpinterae, June-Aug. 
(A), Polakowsky 135 (lectotype, B, destroyed, 
photos, GH, US). Figures 7g, 15. 


Mapouria obovata Oersted, Amér. Centr. 17, t. 14, figs. 
3, 4. 1863. Psyc pur per (Oersted) Hemsley, 
Biol. Cent.-Amer. Bot Pado non Psychotria 
obovata Ruiz & Pav E Peruv. Prodr. 2: 58. 
1799. Uragoga oer MR dia na Ku da Revis. Gen. 
Pl. 2: 957. 1891. Psychotria oerstediana (Kuntze) 


92 


Annals of the 
Missouri Botanical Garden 


Standley, Contr. U.S. Natl. Herb. 23: 1390. 1926. 
TYPE: Mexico: Liebmann s.n. (holotype, C, n.v.). 
Psychotria morae Polakowsky, Linnaea 41: 510. 1877. 
Uragoga morae (Polak.) Kuntze, Revis. Gen. Pl. 2: 
961. 1891. TYPE: Costa Rica: San Jose, June 1875 
(A), Polakowsky 171 (holotype, B, destroyed, pho- 


tos, GH, US) 


Shrub 0.5-2(-5) m tall; young stems glabrous, 
the bark pale, irregular; stipules sheathing, ovate- 
acuminate, 4— 2 10-1 mm, gla- 
brous, caducous, leaving a pale ridge with red- 
brown fringe. Leaves petiolate; petioles 0.3-1 cm 
long, glabrous, grooved above; blades membranous 
to subcoriaceous, obovate to oblanceolate or me- 
dium- to broad-elliptic, the apex acute to short 
(-long)-acuminate, the base narrow-subcordate or 
attenuate, the margin sometimes crenate, (6-)9- 
15(-16) x (2-)3-7.5 cm, glabrous above and 
below, drying red-brown to green-brown, paler be- 
low; secondary veins 6-9(-10) pairs, diverging 
45°-60°, eucamptodromous to usually brochido- 
dromous, the secondary loops usually far from mar- 
gin, increasingly arcuate toward margin, promi- 
nulous below, glabrous, the axils with domatia 
sometimes with tufts of hairs below; tertiary veins 
evident, orthogonal reticulate. Inflorescences ter- 
minal or pseudoaxillary, panicles of cymes (Fig. 
7g), sometimes reduced to appear as fascicles of 
flowers behind terminal stipule; panicle branched 
to (2-)3 degrees; main axis 0.5-3.5 cm long, the 
peduncle lacking or rarely present, ca. 1.5 mm 
long; secondary axes in (1-)2(-3) ranks, the first- 
rank axes 4, subequal to equal, 0.5-3.5 cm long, 
the second-rank axes 4, subequal, 0.1-0.6 cm 
long, the third-rank axes 4, equal, 0.1-0.2 cm 
long; cymes branched to 1-2 degrees; bracts and 
bracteoles not evident. Flowers pedicellate, the 


0.3-0.5 mm long, the | 

5, barely evident, glabrous; corolla white, the tube 
cylindrical, 4-5 x mm, white pubescent in 
throat, the lobes 5, lanceolate, 1.5- 
stamens 5, the filaments 3-3.5 mm long in pins, 
5.5-6.5 mm long in thrums, the anthers 1-1.5 
mm long; style 5.5-7.5 mm long in pins, 4-5 mm 
long in thrums, the branches linear. Fruit when 
dry ellipsoidal, 5.5-7 mm long, 4-5.5 mm diam., 
maturing red, drying red-brown; persistent calyx 
inconspicuous; seeds 2, the dorsal surface with 4— 
5 longitudinal furrows, the ventral surface with 2 
sometimes shallow longitudinal furrows. 


Distribution (Fig. 15). Widespread fom Ve- 
racruz, Mexico, through western Panama, at 0- 
1,300 m elevation in tropical moist to premontane 
wet forest with equatorial to tropical climate. This 


] mm; 


species has been collected in flower throughout the 
year, primarily January-June, and in fruit through- 
out the year 


Selected specimens examined. MEXICO. CAMPECHE: 
Santa Leonor, E of Rio San Pedro, 20 Apr. 1963 
Barlow 14/4 (GH). cHiaPas: Mpio. Ten 
River below Habenal, paraj 
13 July 1964 (early fr), Breedlove 6334 (F, NY); Mpio. 
Huixtla, 6-8 km NE of Huixtla along rd. to Motozintla, 
200 m, 30 June 1972 (fr), s 25966 (ENCB, 
MEXU — 2 sheets, MO, NY); ca 1. NE of Escuintla, 
rd. to El Triunfo, 250 m, 21 Aug. 1977 Si fr), Croat 
43826 (MO); 19 mi. E of Zapata on Z -Balancán 


; e 9428 
(NY); Huehuetán, Islamapa, 21 June 19 "rris Matuda 
17985 (MEXU). OAXACA: km 47 camino Codi. Sa das 
San Juan Guichicobi, 150 m, 7 Dec. 1974 (fr), J. Dora 
tes et al. 3779 (MEXU); ies a 1936 (B). 
Matuda 2277 (MEXU); Temazcal, 1962 (st), 
Sousa 1749 (MEXU — 2 sheets). TABASCO: “ous of 
Villahermosa, June 1964 (fl), Barlow 3/54 (MEXU); 
Balancán, Reforma, 22-26 May 1939 le Me 3146 
fl), Seler 
os Mpio. Paraiso, La U 


(A), irat 6183 (F); Estación Biologia de Los Tuxtlas, 
Feb. 1971 (fl), Calzada 157 (ENCB, F, GH, MEXU, 
US); Me Ozuluama, 8 km from Ozuluama, 8 July 1970 
(fr), Chiang 251 (F, MEXU); Hidalgotitlán, km 0-3 
camino Cedillo la Laguna, 140 m, 10 May 197 
Dorantes 3022 (MEXU); Casitas-Gutiérrez Zamora, 30 
m, 21 June 1970 (fl), Gómez-Pompa & Nevling 1158 
(GH, MEXU); Mpio. de Dos Laguna del Ostión, 0- 
4 m, 5 Jan. 1980 (fl, fr), L. Gutiérrez C. 39 (M 
Laguna Tamiahua, 50 n mi. S of Tam mpic 
(fl), LeSueur 352 (F); Santa Lucrecia, 3 Mar. 1930 (fl), 
Mell 676 (NY); Catemaco, Laguna de Sontecomapan, 
"aix 95?00'W, 5 m, 25 Mar. 1973 (fl), Menéndez 
4 (F, MEXU, MO); vic. Pánuco, 20-25 Apr. 1910 
i digni 347 Neer K, MO, NY); vic. Pueblo Viejo, 2 
10-25 Feb. 1910 (fl), Palmer 384 
(US): a Isthmus of Tehuantepec, Feb. 1895 
(fl, fr), C. Smith 590 (F, GH, MEXU—2 sheets, MO, 
NY — : 


! of Alvarado, coastal hammock 
in Mia 28 June 1970 (fl), Thorne & Lathrop 40456 
(ENCB); 2 km despues del pap 800 m, 18 June 
nate (fr), C. Vazquez n 7 (MEXU); carret. a Plan 


e las Hayas, a 21. 5 A desviación, 720 m, 2 
e 1972 (fr), C. Vázquez Y. 781 (F, MEXU); Mpio. 
de Nautla, Nautla, , 25 Mar. 1976 (fl), Ventura 


l 

12563 (ENCB, MEX a "Mpio. de Alto Lucero, Laguna 
Verde, 20 Feb. 1978 a rae vus uii MEXU). 
GUATEMALA. ALTA VERAPAZ: km E of airport, 
12 Oct. 1968 (fr), n 7906 (NY y 1.5-2 m 

of Cubilgüitz, 300-350 m, 1 Mar. 1942 (st), [nde 
44480 (F, US); along Río Sebol to junction with Río Santa 
Isabel, 125-150 m, 20 Apr. 1942 (fl), Steyermark 45802 
(US). EscUINTLA: 8 km S of Palin, 8 July 1970 (fr), 
Harmon & Dwyer 2961 (ENCB, MO). izaBAL: Petén- 
Guatemala rd., 27 May 1971 (fl), Contreras 10864 (MO). 
PETÉN: Uaxactün, nr. aguada, 24 Apr. 1931 (fl), Bartlett 


Volume 76, Number 1 
1989 


Hamilton 93 
Mesoamerican Psychotria 
subg. Psychotria 


12722 (A, F, US); Dos Lagunas, ca. 10 km W of village, 
in n 27 Dec. 1960 (fl), Contreras 1762 (NY); Do- 
lores, 300 m W of rd. at km 82, 22 May 1961 (fl), 
A 2368 (NY); La Cumbre, on Pusila River, ca. 
8 km from village, 17 May 1967 (fl), Contreras 6936 
(DUKE, F, MEXU); Tikal National Park, Aguada El Pital, 
on new Uaxactün rd., 21 Apr. 1968 (fl), Contreras 7724 
(F, MEXU); San Puls km 116 en el camino para Poptün, 
6 Dec. 1970 (fl), Ortiz 1477 (F, US); along Rio Santa 
Isabel, between mouth of Rio Sebol and El Porvenir, 100 
m, 21 Apr. 1942 (fl), Steyermark 45836 (F); along Río 
San Martin, between Cerro Ceibal and Ceibal, 50 m, 1 
May 1942 (fl), Steyermark 46172 (F). BELIZE. BELIZE: 
Burrel Boom, thicket nr. ferry, 9 June 1973 (fr), Dwyer 
11060 (F, MO, NY); Gracie Rock, 27-28 May 1974 
H4 Dwyer 1245 1 (GH, MO— 2 oW. | eiae 6 Apr. 
934 (fl), Gentle 1255 (A, F, à , US). 
CAYO: El Cayo, 2 May 1931 (fl), EDEN nuc a US); 
Teakettle, 12 May 1931 (fl), Bartlett 13143 (A, F, K); 
Vaca, 10 May 1938 (fl), Gentle 2595 (A—2 sheets, F, 
K, MEXU); 250 m W of Hummingbird Hwy. 2.5 mi. 
S of Western Hwy., 50 m, 17 July 1970 (fr), Spellman 
& Newey 1701 (MO). TOLEDO: 2 km S of Mayan village 
of San Ee ca. 8 km W of Columbia Forest Station, 12 
3 (A), Croat 24311 (F, MO); between Punta 
orda- m Antonio Rd. and Moho River, in broken co- 
hune ridge, 28 May 1949 (fl), Gentle 6754 (DUKE, F). 
HONDURAS. ATLANTIDA: vic. Tela, 0 m, 20 May 1926 (fl), 
Mitchell 87 (F, GH, US); base of hill S of San Alejo nr. 
Río San Alejo, 150-270 m, 22-27 Apr. 1947 (fl), Stand- 
ley 7728 (F); "es back of La Ceiba, bank of i hin 
River, 29 os 938 (fr), Yuncker et al. 8673 (F, GH, 
K, T OLÓN: N bank of Rio Guaimoreto 
bivan p bridge id opening of Laguna Guaimoreto, 
15%57'N, 85°54’W, 17 Oct. 1980 (fl), det 635 (F, 
MO). COMAYAGUA: San Luis, Coyocutena, 120 m, 25 May 
1932 (A), J. B. Edwards P-297 (F, GH); 3 km de La 
Libertad, cerca de Qda. La Jutera, 600 m, 20 May 1956 
(f), A. Molina R. 7048 (F, GH, US). cortés: NE de San 
Pedro Sula, Cordillera de Omoa, 200 m, 19 Apr. 1956 
(f), 4. Molina R. 6710 (F, GH, US); 50 km N Lago de 
Yojoa, Ocote Arrancado, 600 m, 1-30 Nov. 1980 (fr), 
C. Nelson et al. 6022 (TEFH). ISLAS es LA BAHÍA: Roatán 
Island, nr. town of Roatán, 50 m, Aug. 1970 (fr), 
Harmon & Dwyer 3953 (MO). SANTA BARBARA: moun- 
tains above Las Vegas, 1,050 m, 21 June 1970 (8), 
Barkley & Smith 40856 (TEFH). YORO: 19 km SE of 
Rio Viejo doy lr m, P o 1970 (fl), Davidse 
& Pohl 2204 (F, MO, NY); n yoles, ae: Rive 
valley, 28 June 1938 (fr), Yuncker o f bs 8038 (GH, K, 
MO, oh US). a SALVADOR. AHUACHAPÁN: rd. to Tacuba, 
2-3 m an Francisco Menéndez, 500 m, 28 
July 1977 es fr), Croat 42077 (MO). NICARAGUA. 
BOACO: W slope of Cerro eI 12°24'N, 85?33'W, 
900-1,000 m, 8 Oct. 197 


PER. 12%56'N, 87°31'W, 4 
983 (fl, early D Sandino & Aldubin 4379 (MO). 
CHONTALES: 2.8 km N of Cuapa, 12°17'N, 85?23'W, 
400-500 m, 30 Dec. 1983 (fr), W. D. Stevens 22696 
ES y; 1.9 km S of La Libertad, 12?12'N, 85?10'W, 530 
), W. D. Stevens & Montiel 17516 
(MO). ee upper slopes of Volcán Mombacho, 1,100 
. 1977 i Croat 39118 (MO); Isla Zapatera, 

. El Cerro, 11?41'N, 85%46'W, 400- 

E 982 (fr), Grijalva 1961 (MO), NW 
de Volcán Liner 11?49'N, 85*56'W, 19 Aug. 1981 


(fr), Sandino 1283 (MO). JINOTEGA: Sta. Gertrudis, 30 

e er an m, 20 Jan. 1980 (fl), Araquis- 
tain & Moreno 833 (MO). MATAGALPA: La Estancia “El 
Comelar,”. feat lol TE B ioc costado NE del 
982 (fr), Grijalva & 


ar P. Marine 19688 (MO) 
án epi Agir 11°32'N, 
2 (fr), em 
3450 (MO); Hda. Fátima, Mont 3 Victori 
Sapoa, 11?10'N, 85°40’W, 12 Sp 1982 (fr), Sandino 
3585 (MO). ZELAYA: Cerro Livico, 7 km of Siuna, 
500 m, 28 Apr. 1978 (fl), Neill 3651 (MO); Cerro 
Waylawas, E slope of northern range, 13?39'N, 84°49'W, 
80 m, 11 Mar. 1979 (fr), Pipoly 4377 (MO); ca. 6.3 
km S of bridge at Colonia Yolania on rd. to Colonia 
Ep 11?36'N, 84°22’W, 200-300 m, 29-31 
Oct. 1977 (fr), W. D. Stevens 4877 (MOy ca. 8.9 km 


D. Stevens 8705 (MO); Salto La Oropendola, Río Rama, 
119%57'N, 84°17'W, 15-25 m, 17 May 1978 (fl), W. D. 
Stevens 8956 (MO). Costa RICA. ALAJUELA: Colinas de 
S Ramón, 4 June 1931 (fl), Brenes 14310 

'N, 85217'W, 


Naranjo, lower slope of Volcán Miravalles, 1,000 m, 31 
Mar. 1972 (fl), Opler 662 (CR, F); above Hda. Tenorio, 
E Feb. 1956 (fr), Schubert 1052 (A, F); vic. Tilarán, 

500-650 m, 10-31 Jan. 1926 (fl), Standley & Valerio 
44986 (US). LIMÓN: La Coloma nna Farm of the United 
Fruit Co., 70 m, 6-7 Mar. 1924 (fl), Standley 36956 
US). PUNTARENAS: E of Monteverde, 10?18'N, 84°48'W, 
1,300-1,450 m, 29 Oct.-2 Nov. 1975 (fl, fr), Burger 
& Baker gel (CR, F). PANAMA. M4 y rd. to Cerro 
Hornito . from Gualaca, 1,300 m, 1 Sep. 1979 
(fr), poer "1754 (MO); i i 
from Lower Elevation Camp, 1,250 m 
(A), Antonio 4908 (ENCB, MO). cocLÉ: N rim of El Valle, 
NY, US); El 


~ 


on Salud, ca. 0 m, 7 July 1976 

MO). Los SANTOS: Los Toretos, 
10 ye 1962 (fl), DR 2445 (US). 

Psychotria quinqueradiata and P. morae were 
published simultaneously by Polakowsky and serve 
as the only instance known to me where pin and 
thrum morphs of the same species were named as 
different species. I chose to retain the former ep- 


94 


Annals of the 
Missouri Botanical Garden 


ithet for its reference to the five equal axes ra- 
diating from the base of the inflorescence, an im- 
portant recognition character (Fig. 7g). 

The glabrous, obovate leaves drying red-brown 
to green-brown, with secondary veins usually 
brochidodromous making loops far from the mar- 
gin, also aid in recognition. Psychotria quinque- 
radiata resembles P. carthagenensis in most veg- 
etative and fruit characters but differs in its 
distinctive inflorescence and brochidodromous ve- 


Stipule shape is invariable in the range of the 
species, but size varies greatly, from the typical 
4-6 x 2-3 mm in Nicaragua and Costa Rica to 
4-6 X 5 mm in Panama to 10-15 x 8 mm in 
Guatemala. 


7. Psychotria viridis Ruiz € Pavón, Fl. Peruv. 
2: 61, pl. 210, fig. b. 1799. Uragoga 
viridis (Ruiz & Pavón) Kuntze, Revis. Gen. 
Pl. 2: 963. 1891. TYPE: n.v. Figures 4c, 7k, 
13 


Psychotria glomerata d in Humboldt, Bonpland and 
Kunth, Nov. Gen. Sp. 3: 362. 1819. Uragoga glom- 
ice Revis. Gen. Pl. 2: 960. 1891. 

nth 


agoga SEES nig d Kuntze, Revis. Gen. 
Pl. 2: 961. YPE: Costa Rica: Jaris, 900 m, 
Oersted ae anes i) n.v., photo, F neg. 
22840; isotypes, K, US). 

ky trispicata Grisebach, ds M den Pars II: 

2 (cf. also Mem. Amer. Acad. Arts, n.s 

8: és 1863). Uragoga sicud (Griseb.) oa 
Revis. Gen. : 963. 1891. TYPE: Cuba: 

Andres, 1860- 1864 (fr), Wright 1280 eae 

NY; isolectotype, NY). 


Tree 2-4 m tall; young stems glabrous, the bark 
pale, smooth; stipules ovate, the apex often acu- 
minate, darker in central triangle, 8-15 x 3-6 
mm, irregularly fringed, caducous, leaving a pale 
ridge with long red-brown fringe. Leaves subsessile 
to petiolate; petioles to 8 mm long, glabrous, flat 
above; blades membranous, elliptic-obovate, the 
apex short-acuminate to acute, the base attenuate, 
(6.5-)9-15 x (3-)4-5(-6) cm, glabrous above 
and below, drying dull red-brown to green-brown; 
secondary veins 7-10 pairs, diverging 45°-55°, 
eucamptodromous, straight then arcuate 24 to mar- 
gin, elevated below, glabrous, the axils often with 
domatia below especially near apex (Fig. 4c); ter- 
tiary veins inconspicuous, orthogonal reticulate. 
Inflorescences terminal or pseudoaxillary, panicles 
of glomerules (Fig. 7k); panicle branched to 3 
degrees; main axis 6.5-12 cm long, the peduncle 


2-6 cm long; secondary axes in (4—)5 ranks, the 
first-rank axes 4, the longer pair (0.6-)1-2.5 cm 
long, the shorter pair usually reduced or to 0.3 
cm long, the second-rank axes 2 or 4, reduced and 
thus fl 
the longer pair when present 0.5-0.8 cm long, the 
third-rank axes 2 or rarely 4, reduced, the longer 
pair when present 0.4 cm long, the fourth-rank 
axes 2, reduced, the fifth-rank axes 2, reduced; 
tertiary axes reduced and thus flowers disposed in 
glomerules along secondary axes; bracts and brac- 
teoles irregular, to 1 mm long, red-brown ciliate. 
Flowers sessile; calyx cup-shaped, ca. 0.5 mm 
long, the lobes not evident or rarely 5, triangular, 
to 0.2 mm long, glabrous; corolla white, the tube 
cylindrical, 1.5 x m, thick white pubescent in 
throat, the lobes 5, lanceolate, 1 x 0.6 mm; sta- 
mens 5, the filaments 2 mm long, the anthers 0.5 
mm long; style 2.5 mm long, the branches clublike. 
Fruit when dry ellipsoidal, 4.5-5 mm long, 3-3.5 
mm diam., maturing red, drying red-brown; per- 
sistent calyx a beak to 0.5 mm long; seed dorsal 


ower dien 


d in glomerules along main axis, 


surface with 4-5 deep longitudinal furrows, the 
ventral surface with 2 deep longitudinal furrows. 


Distribution (Fig. 13). Very sparse distribu- 
tion throughout Central America at 0-1,000 m 
elevation, collected most commonly in the lowlands 
of eastern Panama, in tropical moist to premontane 
wet forest with equatorial to tropical climate. Psy- 
chotria viridis occurs also in Cuba, Hispaniola, 
Colombia, Venezuela, Ecuador, Peru (where it is 
extremely common), Brazil, and Bolivia. In Central 
America it has been collected in flower September- 
March and in fruit January-June and in Septem- 
ber. 


Selected specimens examined. MEXICO. CHIAPAS: Rio 
Salinas, in riverbank forest, above mouth of Rio Pasión, 
8 Feb. 1964 gti fr), Lundell 17811 (LL). GUATEMALA. 
PETÉN: Dolores, km 78-79 on Santo Toribio Rd., 20 Apr. 
1961 (fr), REA 2140 (DUKE); Chinchilá, Sebol rd., 
forest in corozal and zapotal, 7 Mar. 1971 (fr), Contreras 
10610 (F); Rio de la Pasión, Ceibal, in corozal on ruin 
3 Feb. 1964 (fr), Lundell 17649 (DUKE, F, LL); Can 
Ceibal (Sierra tla between mouth of Rio Sta. Monica 
and mouth of Rio San Martin, on W side of Rio rena 
75-150 m, 30 ren 1942 (fr), Steyermark 46100 (F). 


as e, 
1978 (early fr), Neill & 
Vincelli 3270 (MO). COSTA RICA. CARTAGO: 
E of Turrialba, Rio Reventazón, 9°54'N, 83°39'W, 525- 
600 m, 9 May 1983 (fr), Liesner et Mie 15301 (CR, 
MO). LiMÓN: La Palma, Sixaola, 1 Mar. 1924 (fl), it 
470 (F, K, US); drenajes de los rios Parisis na y 

tazón, 0 m, 3 Oct. 1951 (early fr), Shank jd oia 
4261 (GH, US); entre Barra Parismina y Río uero, 
9 Oct. 1951 (fl), Shank & Molina 4330 (F, En ross 


Volume 76, Number 1 
1989 


Hamilton 95 
Mesoamerican Psychotria 
subg. Psychotria 


de Rio Valle Es mo Montana Andromeda, 26-28 Oct. 
ank & Molina 4518 (F, GH, US) 


EN: ca. 2 mi. 
N of Santa Fe, ca. 25 af 15 Feb. 1967 (fr), Duke 10230 
(ECON, MO): S slope of Cerro Tacarcuna, 700-1,000 
m, A Jan. 1975 ea s ao & Mori 13902 (MO); 
Rio Chico m upstream from Nazareht, 
3°15) N, 77°25'W, ee m, a Dec. 1980 (early fr), W. 
Hahn 123 (MO); base of Cerro Sapo, 400 m, 30 Jan. 
1975 (early fr), Hammel 1136 (MO). PANAMÁ: Serrania 
e Maje, S of Ipeti, to 360 m, 17 Sep. 1982 (early fr), 
D'Ar rey 15115 (MO); 17 Sep. 1982 (fr), D'Arcy 15123 
MO); nr. Jenine, Rio Canita, 23 Sep. 1961 (fl, early uis 
Duke 3852 (GH, MO— 2 sheets); Panam. Hwy. 
ridge, 22 Jan. 1977 (fr), Folsom 1388 
a de Cañazas, ca. 15 km SW of 
Cañazas nr. Rio Torti, 8%52'N, 78%22'W, 150 m, 15 
Jan. 1983 (fr), Stein 1337 (MO). 


Psychotria viridis may be readily recognized 
by its inflorescence: all ranks of secondary axes 
except the first and second are reduced so that 
flowers and fruit appear in clusters along the main 
axis (Fig. 7k). Psychotria viridis is, in its vege- 
tative and fruit characters, almost identical to P. 
carthagenensis; the distinctive inflorescence may 
be viewed as a reduction in secondary and tertiary 
axes of that of P. carthagenensis. 

There appears to be only one flower morph, a 
homostyle, in Central American P. viridis, with 
no spatial separation between the exserted anthers 
and stigma. Leaves in Guatemala and Belize have 
fewer secondary veins than elsewhere in Central 
America. 


GROUP 2. THE COSTIVENIA GROUP 


Shrub or tree; young stems glabrous or puber- 
ulent or rarely tomentose or ferrugineous-pubes- 
cent; stipules usually not sheathing, ovate, uniform 

color or often with a pale central triangle or 
keel (Fig. 2b), glabrous or puberulent or ferrugin- 
eous-pubescent or sometimes fringed, caducous or 
often persistent at terminal 2-6 nodes. Leaf blades 
elliptic or ovate or obovate or oblanceolate, drying 
usually yellow-green or sometimes green, green- 
brown, or red-brown, often dull or pale, sometimes 
chalky above, sometimes darker below; T 
veins (6-)7-19(-21) pairs, er (45 
85°, eucamptodromous or We 5 the 
axils usually lacking domatia or duis (except some- 
times P. horizontalis and P. sylvivaga); tertiary 
veins orthogonal reticulate or less commonly per- 
current, the intersecondaries often conspicuous, 
the tertiary loops sometimes evident (P. fendleri 
and P. horizontalis). Inflorescences panicles of 
cymes or rarely of glomerules (P. flava), pedun- 
culate (except rarely in P. horizontalis); second- 


ary axes usually in 2 or 3 size-differentiated 
pairs per rank (Fig. 7b) or sometimes 2 or rarely 
4 subequal; bracts not conspicuously enlarged (to 
5 mm long in P. flava and P. pleuropoda). Corolla 
tubes 2-3.5 mm long (4-5 mm in P. papantlen- 
sis), the lobes without apical extensions. Fruit when 
dry spherical to ellipsoidal (sometimes slightly ob- 
ovoid in P. flava); persistent calyx a beak or a 
tube Fig. 9b) or of linear lobes (Fig. 9e) or not 
evident, over 1 mm long only in P. fendleri and 
P. horizontalis; dorsal seed surface with 3-5 reg- 
ular (Fig. 10b) or 6-15 irregular longitudinal fur- 
rows (Fig. 10i, j), the ventral surface with 2 often 
incompletely divided longitudinal furrows (Fig. 10e) 
or one often T-shaped longitudinal furrow (P. flava; 
Fig. 10j) rarely plus 4 irregular longitudinal furrows 
(P. sylvivaga). 


This subgroup may be subdivided into two species 
complexes. The Psychotria horizontalis complex 
includes the widespread P. horizontalis and P. 
fendleri, the latter endemic to Caribbean coastal 
Panama (Figs. 16, 17); they share the distinctive 
character states of immediately caducous stipules, 
brochidodromous secondary venation with con- 
necting loops far from the blade margin plus evident 
tertiary connecting loops, and persistent calyx at 
least 1 mm long. The P. costivenia complex, which 
includes the remaining seven species, has in com- 
mon often persistent stipules usually with a paler 
central triangle or keel, often conspicuous inter- 
secondary veins, and persistent calyx a beak or 
tube no longer than 1 mm. Two species are wide- 
spread— P. costivenia (Mexico through Nicara- 
gua; Fig. 18) and P. grandis (Guatemala through 
Panama; Fig. 19), both also in the Greater An- 
tilles—and five have small ranges: P. balanca- 

nensis (Tabasco, Mexico; Fig. 19), P. flava i 
ern Mexico and Petén, Guatemala; Fig. 17), P 
papantlensis (southern Mexico through Belize; Fig. 
16), P. pleuropoda (southern Mexico through Be- 
lize; Fig. 17), and P. sylvivaga (central Costa Rica; 
Fig. 17) 

Of the nine species in the group, six appear 
distylous; two of those, Psychotria papantlensis 
and P. pleuropoda, show some between-morph 
asymmetry in floral part lengths. Only one species, 
P. fendleri, apparently has evolved a derived state, 
thrum-monomorphy. Too few specimens of P. 
lancanensis have been seen to determine its breed- 
ing system. 


8. Psychotria balancanensis C. Hamilton, 
Phytologia 64: 219. 1988. TYPE: Mexico. Ta- 
basco: Balancán, La Palma, 1-6 June 1939 


96 Annals of the 
Missouri Botanical Garden 


—- T T T 1 : r 
M " > 4 | TC.4 
g b 20 4 

1 + + 

o. 
e > : ° 
E + © 15 4 
"e + + es. e 
[17 
eb 

[ 10 + 

300 km +- e 

e 

105 100 95 90 85 80 


FicURE 16. Distributions of Psychotria horizontalis (circles) and P. papantlensis (triangles) in Mesoamerica. 


(f), Matuda 3286 (holotype, F; isotypes, A, ules ovate-triangular, 6.5-7.5 x 3-4 mm, fringed, 
MEXU, NY). Figure 19. glabrous, caducous, leaving a pale ridge with red- 
brown fringe. Leaves petiolate; petioles 0.9-2 cm 

Shrub ca. 2.5 m tall; young stems glabrous, the long, glabrous, flat above; blades membranous, el- 
bark smooth, sometimes with shallow fissures; stip- liptic to slightly obovate, the apex acuminate, the 


LU V T T y Lu 


i TC. 7 


104 
300 km F a + 


105 100 95 90 85 ^80 


FicurE 17. Distributions of Psychotria fendleri (open circles), P. flava (solid circles), P. pleuropoda (triangles), 
and P. priis (squares). 


Volume 76, Number 1 
1989 


Hamilton 
Mesoamerican Psychotria 
subg. Psychotria 


T T T T T T T 
-l- ^ | TC.- 
q T (d 4 204 
d e 
j 13x 
o 
jj + + aL D us 154 
o% 
e 
«S 
i 10 4 
300 km + + 
b 
105 100 95 90 85 a a 
FicURE 18. Distributions of Psychotria costivenia var. altorum (triangles) and P. costivenia var. costivenia 


(circles) i in Mesoamerica. 


base attenuate, (6-)7-11(-13) x (2-)3-6 cm, 
glabrous above and below, drying dull green; sec- 


arcuate, elevated below, glabrous, the axils lacking 


ondary veins 8-10 pairs, diverging 65?-75?, eu- onal reticulate to percurrent. Inflorescences ter 
camptodromous to brochidodromous, constantly minal, panicles of cymes; panicle branched to 3- 
-l- : i | TC. 4 
&ü 20 4 
+ a 
e 9 
e 
e 
% 
>. 
10 - 
300 km + ^ 
b 
e 
105 100 95 90 85 80 
FIGURE 19. Distributions of Psychotria balancanensis (triangle) and P. grandis (circles) in Mesoamerica. 


98 


Annals of the 
Missouri Botanical Garden 


4 degrees; main axis 3.5-9 cm long, the peduncle 
2.5-7 cm long; secondary axes in 2 ranks, the 
first-rank axes 2 or 4, the longer pair 0.8-1.2 cm 
long, the shorter pair 0.2-0.8 cm long, the second- 
rank axes 4, equal, 0.3-0.7 cm long; cymes 
branched to 1-2 degrees; bracts and bracteoles 
irregular triangular, 0.7 x 0.8 mm, puberulent, 
ringed. Flowers pedicellate; pedicels 0.7-1 mm 
long; calyx cup-shaped, the tube 0.3-0.5 mm, the 
lobes 5, broadly triangular to barely evident, 0.2— 
0.3 x 0.5 mm, fringed; corolla white, the tube 
cylindrical, 2.5-3 x 1.2 mm, white pubescent in 
throat, the lobes 5(-6), triangular, spreading, 1.5 
x 1 mm; stamens 5(-6), the filaments 2.5-3 mm 
long in pins, not seen in thrums, the anthers | mm 
long; style 4.5 mm long in pins, not seen in thrums, 
the branches linear. Fruit not seen. 


Distribution (Fig. 19). 
type collection from eastern Balancan, Tabasco, in 
a region of evergreen forest and savanna with 
equatorial-tropical climate. It was collected in flow- 
er in early June. 


Known only from the 


Psychotria balancanensis may be recognized 
by its moderate-sized (7-11 cm long) broad-elliptic 
leaves drying dull green to green-brown with con- 
spicuous intersecondary veins and by its inflores- 
cence with the secondary axes in two ranks near 
the apex of the main axis. This species differs from 
P. costivenia in having broader-elliptic leaves and 
smaller inflorescences with fewer (two vs. four) 
ranks of secondary axes. 

Only the pin morph has been collected, but the 
one flowering specimen seen is not sufficient to 
suggest that the species is pin-monomorphic. 


9. Psychotria costivenia Grisebach, Pl. 
Wright., Pars II: 508. 1862. (Cf. also Mem. 
Amer. Acad. Arts, n.s. 8: 508. 1863.) Ura- 
goga costivenia (Griseb.) Kuntze, Revis. Gen. 
Pl. 2: 960. 1891. TYPE: Cuba. Oriente: prope 
villam Monte Verde dictam, Jan.-July 1859 
(fr), Wright 242 (lectotype designated herein, 
GH). Figures 2b, 10e, 18 


Shrub 0.5-5(-6) m tall; young stems glabrous 
to red-brown puberulent to sparsely tomentose, the 
bark often pale, smooth; stipules ovate, the apex 
cuspidate or aristate or biaristate, 6-10(-11) x 
3-7(-10) mm, with pale center and often evident 
central keel (Fig. 2b), glabrous to red-brown sub- 
tomentose, often fringed especially near apex, ca- 
ducous, sometimes persistent at the terminal 2-3 
nodes, leaving a pale ridge with red-brown fringe. 
Leaves sessile to petiolate; petioles to 1.5(-2) cm 


long, glabrous to puberulent, flat and sometimes 
shallowly grooved above; blades membranous to 
rarely subcoriaceous, elliptic or rarely obovate, the 
apex acute to acuminate, the base attenuate, 
(4-)4.5-17(-22) x (1-)1.5-6(-8) cm, glabrous 
above and below, drying green, yellow-green, red- 
brown, or green-brown; secondary veins 7-12 
(-18) pairs, diverging 65°-80°, eucamptodromous 
to brochidodromous, straight or constantly arcuate, 
elevated below, glabrous, the axils lacking domatia 
or hairs; tertiary veins evident, orthogonal retic- 
ulate, the intersecondaries particularly conspicu- 
ous. Inflorescences terminal or pseudoaxillary, 
panicles of cymes; panicle branched to 3-4 de- 
grees; main axis 3.5-22 cm long, the peduncle 
(0.7-)2.5-11.5 cm long; secondary axes in (2-)3- 
4(—5) ranks, the first-rank axes 2 or 4, the longer 
pair 0.5-4.7 cm long, the shorter pair 0.3-2.3 
cm long, the second-rank axes 2 or 4, the longer 
pair 0.3-2.5 cm long, the shorter pair 0.2-1.5 
cm long, the third-rank axes 2 or 4, the longer 
pair 0.1-1.5 cm long, the shorter pair 0.1-0.5 
cm long, the fourth-rank axes 2 or 4, the longer 
pair 0.1-0.8 cm long, the shorter pair 0.4 cm 
long, the fifth-rank axes 2, 0.1 cm long; cymes 
branched to 1-3 degrees; bracts linear, 2-5 mm 
long, puberulent, sometimes fringed; bracteoles ir- 
regular, to 1 mm long, red-brown pubescent. Flow- 
ers sessile or pedicellate, the pedicels to 1.5 mm 
long; calyx cup-shaped, the tube 0.3-0.7 mm long, 
the lobes 5, barely evident to triangular to linear, 
to 0.8 mm long, to 0.6 mm wide, fringed, often 
puberulent outside; corolla white, the tube cylin- 
drical, 2-3.5 x 1-2 mm, white pubescent in throat, 
the lobes 5, linear, 1.5-2 x 
5, the filaments 2-3 mm long in pins, 3.5-5 mm 


0.7-1 mm; stamens 


long in thrums, the anthers 0.8-1 mm long; style 
(3-)3.5-5 mm long in pins, 2.5-3.5 mm long in 
thrums, the branches short, clublike. Fruit when 
dry spherical to ellipsoidal, 5-7(-8.5) mm long, 
4—6(—7) mm diam., maturing red, drying red-brown 
or black; persistent calyx not evident or as a beak, 
drying green or red-brown; seed dorsal surface with 
4-5 longitudinal furrows, the ventral surface with 
2 deep longitudinal furrows, incompletely divided 


(Fig. 10e). 


Distribution (Fig. 18). Mexico through Nic- 
aragua at elevations of 20-2,800 m 


Psychotria costivenia may be recognized by its 
stipules usually persisting at the terminal two or 
three nodes and having a paler central triangle 
(Fig. 2b), its elliptic leaves drying usually green or 
yellow-green with secondary veins drying paler and 
intersecondaries particularly conspicuous, and its 


Volume 76, Number 1 
1989 


Hamilton 99 
Mesoamerican Psychotria 
subg. Psychotria 


inflorescence secondary axes in one or two unequal 
pairs per rank, the higher ranks decreasing in size 
fairly uniformly. 


KEY TO VARIETIES OF PSYCHOTRIA COSTIVENIA 

la. Leaf blades generally 4.5-10 x 1.5-3 cm, 
drying red-brown or green brown; inflorescence 
main axis 3.5-6 cm lon 


9a. P. costivenia var. altorum 
. Leaf blades generally 8-17 x 3.5-6 cm, drying 
green or yellow-green; inflorescence main axis 
8 


5.5-22 cm lon 


— 
ig" 


9b. P. costivenia var. costivenia 


9a. Psychotria costivenia Grisebach var. al- 
torum (Standley & rk wea. C. Hamilton, 
Phytologia 64: 223. 1988. Psychotria alto- 
rum Standley & vem. Publ. Field Mus. 
Nat. Hist., Bot. Ser. 23: 86. 1944. TYPE: 
Guatemala. ec SUR Montana Chi- 
charro, on lower SE-facing slopes of Volcán 
Sta. Maria, 2-4 mi. S of Sta. Maria de Jesüs, 
1,400-1,500 m, 17 Jan. 1940 (fl), Steyer- 
mark 34302 (holotype, F). Figure 18. 


Shrub: stipules ovate, the apex aristate or biaris- 
tate, 4-4. 
long, caducous, never persistent at the terminal 2- 
3 nodes. Leaves: blades narrow-elliptic, (4—)4.5— 
10(-12) x (1-)1.5-3(-4) cm, drying red-brown 
or green-brown; secondary veins 7—9(—11) pairs. 
Inflorescences few-branched panicles of cymes; 
panicle branched to 3(-4) degrees; main axis 3.5- 
6 cm long, the peduncle (0.7-)2.5-4 cm long; 
secondary axes in 3-4(-5) ranks, the first-rank 
axes 2 (or 4), 0.5-1.6 cm long, the shorter pair 
0.3-1 cm long, the second-rank axes 2 or 4, 0.3- 
0.8 cm long, the shorter pair 0.2-0.5 cm long, 
the third-rank axes 2 (or 4), 0.1-0.5 cm long, the 
shorter pair 0.2 cm long, the fourth-rank axes 2, 
0.1-0.3 cm long, the fifth-rank axes 2, 0.1 cm 
long; cymes branched to 1-2 degrees. Flowers: 
calyx lobes triangular to linear, 0.3-0.8 mm long. 
Fruit: drying red-brown; persistent calyx not evi- 
dent or sometimes as a beak, drying red-brown. 


mm, the extensions 1.5-3 mm 


Distribution (Fig. 18). Chiapas, Mexico, and 
southern Guatemala mostly at elevations of 1,000— 
2,800 m in areas of pine—oak forest with tropical 
climate. It has been collected in flower January— 
June and once in November and in fruit mostly 
September- February and rarely in May and June. 


Selected specimens examined. MEXICO. CHIAPAS: 
Chamula, Yal Ichin, 1,800 m, 3 Apr. 1965 (fl), Breedlove 


9537 (NY); Mpio. La Trinitaria, ae Tsiskaw, 30 m 
ini 1965 (fl), Breedlove 
Y); Mpio. a de inosa, 


o Mal Pus 900 m, 25 June 1972 (fl), Breedibue 


o 
1976 (fr), IRE. 1 ee Mpio. La Indepen- 
dencia, rd. from Las Margaritas to Campo Alegre, 1,600 
m, 30 Sep. 1981 (fr), (ES 53213 (MEXU); Mpio. 
La Trinitaria, Comitán 
Montebello, 42 km o 
(fr), woes & Thorne 21156 ( 
NY); Mpio. Tapachula, Soconusco, 140 m, 20 Sep. 1977 
(fr), d 368 (ME ; Volcán Tacaná, 2,800 m, 30 
MEXU, NY); 


Mar. 1939 (fl), Matuda 2926 (F, GH, K, 
¡a Paderon, 14-20 Jan. 1946 (fl), Matuda 16302 
(MEX U ATEMALA ALTA VERAPAZ: Chamal, margenes 


R. & A. Molina 12183 (F— 2 sheets, NY, US— 2 

along Rio Carchá, between Cobán and San Pedro Carchá, 

1,360 m, 26-27 Mar. 1941 (f), Standley 90065 (F); 

10-15 km W of San Cristobal, “El Derrumbe,” canyon 

of Rio Chixoy CLE 15°25'N, 90%27' W. 1,200-1,600 
b. 1969 Le 


m, 2-4 Fe r), L. O. Williams et de 4058] (F). 
BAJA VERAPAZ: E n Barrios, E of km 155/156 on Cobán 

, 13 Apr. 1975 (8). Lundell & Contreras n VR 
Sierra de yn Minas ca. mSo d a ; 


Jan. 1973 (fl), L. O. Williams et al. 958 (F) c CHI- 
MALTENANGO: lower and middle SW Eod “ Volcan Fue- 
go, above Finca Montevideo, along Barranco Fs 
and tributary of Rio Pantaleón, 1,200-1,600 m, 20 Sep. 
194 2 (fr), Steyermark 52059 (F, US). EL PROGRESO: 

ita Pg d Finca San Miguel and summit, 


r. upper limits of Finca Ca cael 1,600-2,300 m, 10 
Feb. 942 (fr), Sirera 43776 (F, 
NANGO: Barillas, Finca San Isidro, ' 300 m 
(fl), Holdridge 2331 (F, US). o 
Ep Martin Chile Verde and Colomba, 

941 (fl), Standley 85583 (F); lower 
S- facing slopes of ard Santa Maria, between Santa 
María de Jesús and Calahuaché, along great barranco 


Feb. 1940 (fr), Steyermark 36714 (F); above Finca El 
Porvenir, up Cerro de Mono, S-facing slopes of Volcán 
Tajumulco, 1,400-1,700 m, 9 Mar. 1940 (fl), Steyer- 
mark 37388 (F). soLoLÁ: Volcán Atitlán, S-facing slopes, 
1,700-3,800 m, 11 June 1942 (fr), Steyermark 47337 
(F, GH). S is Volcàn Atitlán, S slope, 1,600 
m, 22 Oct. 1934 (fr), Skutch ded (F, GH); Volcán 
s Clara, S E Finca El Naranj alid upper slopes, 

250-2,650 m, 23 May 1942 (fr), m daas 46615 
ay sheets, US) 


Psychotria altorum has been reduced to a va- 
riety within P. costivenia because the distinguish- 
ing features (see varietal key) are not sufficient to 
distinguish a species, though varietal recognition is 
useful. 

Leaf blades of this variety appear to have more 
secondary veins in Mexico than in Guatemala; per- 
sistent calyces are beaklike in Mexico and not 
evident in Guatemala. 


100 


Annals of the 
Missouri Botanical Garden 


9b. Psychotria costivenia Grisebach var. cos- 
tivenia. Figure 18. 


Mapouria miradorensis Oersted, Amér. Centr. 17, t. 14, 
9. 186 de ie miradorensis (Oerst.) 
Hemsley, Biol. Cent.-Amer. Bot. 50. 1881. Ura- 
goga miradorensis (erst. ) Kuntze, Revis. Gen. Pl. 

2: 961. 1891. TYPE: Mexico. Veracruz: Mirador 
900 m, 1841-1843 (fl), Liebmann 11662 (lecto- 
type, C, photo F neg. no. 22839; isolectotype, US). 


Shrub: stipules ovate, the apex cuspidate, 6- 
10(-11) x 3.5-7(-10) mm, caducous, often per- 
sistent at the terminal 2-3 nodes. Leaves: blades 
elliptic or rarely obovate, (7-)8-17(-22) x 
(3-)3.5-6(-8) cm, drying green, yellow-green, or 
less often green-brown; secondary veins (8-)9- 
12(-18) pairs. Inflorescences many-branched pan- 
icles of cymes; panicle branched to 3-4 degrees, 
the axes often winged; main axis 5.5-22 cm long, 
the peduncle 2.5-11.5 cm long; secondary axes 
in (2-)4 ranks, the first-rank axes (2 or) 4, the 
longer pair (0.6-)1-4.7 cm long, the shorter pair 
(0.3-)0.5-2.3 cm long, the second-rank axes 2 or 
4, the longer pair 0.5-2.5 cm long, the shorter 
pair 0.2-1.5 cm long, the third-rank axes 2 or 4, 
the longer pair (0.2—)0.4—1.5 cm long, the shorter 
pair 0.1-0.5 cm long, the fourth-rank axes 2 or 
4, the longer pair 0.1-0.8 cm long, the shorter 
pair 0.4 cm long; cymes branched to 1-3 degrees. 
Flowers: calyx lobes barely evident to triangular, 
o 0.3 mm long. Fruit: drying black; persistent 
calyx not evident or often as a beak, to 1 mm long, 
drying green. 


Distribution (Fig. 18). Widespread in Mexico 
through Nicaragua, at elevations of 20-1,500 m, 
usually at 1,000 m or lower, in evergreen to sub- 
evergreen forest with equatorial to tropical climate. 
It has been collected in flower January-June, prin- 
cipally March-May, and in fruit throughout the 
year. 


Re ECL Vo examined. MEXICO. CAM- 


PECHE: Campo E mental Forestal Tropical “El Tor- 

mento,” a carr eres a Candelaria, 20 Apr 
1966 (fl, fr), Chav E. rnández X 94 
(MEXU). CHIAPAS: ie Jitotol, Rio Hondo 4 mi. N of 
Jitotol on rd. to Pue vo Solistahuacan, 1,650 m, 


e 
20 Aug. 1965 (49. Pada 12034 (NY); Mpio. Rayón, 
do above Rayón Mezcalapa alon uo to Jitotol, 1,700 
, 12 Dec. 1 971 (fr), Breedlove 23196 (F, MO); Mpio. 
Berriozábal, 13 km N of Berriozábal nr. Pozo Turipache 
and Finca El Suspiro, 1,000 m, 24 July 1972 (fl), Breed- 
love 26314 (ENCB, MO, NY); 900 m, 25 Dec. 1972 
(fr), Breedlove & Thorne 30828 (MO, NY); Mpio. Ocos- 
ingo, 6-8 km N of Ocosingo along rd. to Bachajón, 900 
m, 24 Sep. 1972 (fr), Breedlove 27877 (MEXU, NY); 
m N of Ocozo- 
. to Mal Paso, 750 m, 19 Oct. 1965 (fr), 
Breedlove & Raven 13568 (DUKE, ENCB, NY); Mpio. 


Cintalapa, 16 km NW of Rizo de Oro along a logging 

road to Colonia Figueroa, 1,600 m, 3 Nov. 1971 (fr), 
Breedlove & Smith 21713 (ENCB, MEXU, MO, NY); 
La Grandeza, 2,016 m, 19 May 1945 (fl), Matuda 5573 
(F, LL, MEXU); Mpio. Ocosingo, Centro Arqueológico 
Bonampak, 350 m, 24 Dec. 198 
B-127 (MEXU); Mpio. Tenajapa, barrio of Tih Ha', 
raje of Mahben Chauk, 1,080 m, 28 Oct. 1966 (fr), Ton 


(fr), Ton 3525 (ENCB); Mpio. Yajalon, 67 km al S de 
alenque, sobre carret. a Ocosingo, 17*10'N, 92°09’W, 
450 m, 3 Dec. 1979 (fr), Wendt et al. 2345 pera 
GUERRERO: Mpio. Atoyac, 19 km NE n- 
iago La Unión, 770 m, 3 Nov. 1979 (fr), Koch et al. 
79288 (MEXU). OAXACA: Mpio. Tuxtepec, Isla de de 
saga en la Presa Miguel Alemán, 24 Oct. 1964 (fr), L 
González Q. 1789 agers MEXU); Putla, Arroyo Li- 
16 Dec. 1970 (fl), MacDougall H153 
(ENCB Mo Tuxt tepec, Che epec and vic. 8 
, G. Martínez-Calderón 147 (GH, MEXU. 
7.3 de ‘la carret. México-Tu uxpan, 
May 1962 (st), sabes et al. 2119 (MEXU); Mpio. 
Hueyta eic Paxta, m, 31 Jan. 1978 (fr), Ventura 
1495 1 (ENCB, MEXU). pics ROO: 8 km N of Un 
110 km SW of C 
(fr), Davidse et al. 20178 (MO). SAN LUIS POTOSÍ: A 
ladera margosa, 250 m, 17 Apr. 1956 (fl), Rzedowski 
7529 (ENCB, MEXU). rABASCO: Mpio. Macuspana, rd. 
along Rio Chinal (Río Macuspana) between Macuspana 
and El page d Sep. 1944 (fr), Gilly & E. Hernández 
X. 378 (GH, M 


arr a . 
nánges ger (ENCB, MEXU); a 11 km de Villaher- 
m 2 km ES pa rd 28 Dec. 1965 (fl), Gon- 


z & Pérez 3896 (ENCB); Balancán, La Palma, 1- 

6 Fin 1930 (fr), pads 3264 (A, F “2 sheets; MEXU). 
VERACRUZ: Teocelo, Trapiche, 1,220 m, 2 May 1980 (fl), 
arrera et al. 228 (MEXU); Papantla, v 1947 (fl), 


Kelly 183 (DUKE); 5 km limite Puebla- Veracruz, carret. 
Tlapac dn Teziutlán, 900 m, 13 July tds Nevling 
& Gómez-Pompa 1667 (GH, MEXU); banks of Arroyo 
dtt Mar. 1930 (fl, fr), Purpus 14398 (A — 2 sheets, 
F — 2 sheets); Hidalgotitlán, Brecha Hnos. Cedillo-La Es- 
cuadra, 200 m, 7 Mar. 1974 (fl B. Vázquez 123 
(MEXU — 2 sheets); Mpio. de Nautla, La Martinica, 50 
m, 25 Mar. 1971 (fl, fr), Ventura 3338 (ENCB); Mpio. 
de Totutla, El Mirador, 1,000 m, 17 Apr. 1978 (fl), 
Ventura 15208 (MEXU, MO); Mpio. Minatitlán, 4.5 km 
al E del Rio Grande, 17?17'N, 94*30'W, 170 m, 27 Feb. 
1981 (fl), Wendt et al. 2939 (ENCB, MEXU). Guar- 
EMALA. ALTA VERAPAZ: Sebol, ca. 800 m W of village, 
pg arroyo, 15 Apr. 1964 (fl), Contreras 4319 
(MO): C ] : 


N from Hw 
NE of Panzós, 800 m, 20 July 1977 (fr), Croat 41692 
(MO); Rubelsanto, Rio Salinas, 15 July 1975 (early fr), 
Lundell & Contreras 19517 (MO); 1.5-2 mi. S of 
ilgüitz, 300-350 m, 1 Mar. 1942 (fl), Steyermark 44444 
(F). CHIQUIMULA: Montana Castilla, nr. ebollas, 
along Rio Lucia Saso, 5 km SE of Que 

1,500 m, 6 Nov. 1 

HU EHUETENANGO: between Ixcan and Finca San Rafael, 
Sierra de los Cuchu ao 200- 

fr), aci 49405 (F, 

on Rio Dulce Rd., km 10, 10 Sep. 1970 (fr), Contreras 


em 


Volume 76, Number 1 
1989 


Hamilton 101 
Mesoamerican Psychotria 


subg. Psychotria 


10245 (MO); 18 km E of Los Amates, 13 June 1970 
(8), Harmon 2569 (F, GH, MO— 2 sheets); hills above 
Eximbal mining area W of El Estor, 9 Apr. 1970 (fr), 
Harmon & Dwyer 4332 (ENCB, F, GH, MO); between 
Milla 49.5 and Cristina, 65-70 m, 30 Mar. 1940 (fl), 
Steyermark 38476 (F — 2 sheets). PETÉN: Tikal National 
Park, on Aguada Aurora, in zapotal, 15 June 1960 (fl), 
Contreras 1098 (MO); Lacandón, El Caribal, low forest, 
16 Mar. 1962 (fl), Contreras 3540 (MO); Dos Lagunas, 
on Ixcanrio Rd., in zapotal, 20 Apr. 1969 (fl), Contreras 
8364 (MO); Chinchilá, Sebol Rd., 10 Mar. 1971 (fl), 
Contreras 10706 (F); Carmelita, in thicket bordering 
aviation field, 29 June 1942 (early fr), Egler 42-264 (F); 
m S of Poptün, 1,500 m, 10 June 1970 (fl), epe 
2513 (ENCB, F, MO); 13 km S of Flores, 200 m, 
June 1970 (early fr), Harmon & Dwyer 2786 (MO); t 
Cumbre, Cerro la Cueva, m E, in zapotal, 22 Mar. 
1977 (A), Lundell & Contreras 20637 (MO); entre La 
Libertad y Subin, 50 km al S de Sta. Elena, 100 m, 10 
Nov. 1965 (fr), 4. Molina R. 15498 (F— 2 sheets, NY); 
Tikal National Park, orilla del camino a El Remate, 
en el km 58, 7 Sep. 1970 (fr), Ortíz 1296 (ENCB, F, 
MO, NY); ca. 200 m de Dolores, en orillando el camino 
para el arroyo Dolores, 16 Feb. 1971 (fr), Ortíz 1586 
(ENCB, F, MO, NY); en el camino que conduce al cam- 
pamento chiclero Mushanal, a 1.5 km de la aldea de 
Uaxactün, 16 May 1973 (fl), Ortíz 2561 (ENCB, F, MO, 
NY); low forest along Río Chinajá, N of Chinajá on trail 
towards Zacatal, 50-70 m, 28 Mar. 1942 (fr), Steyer- 
a 45452 (F). SOLOLA: pine woods bordering Rio Bravo, 
nr. Finca Moca, S-facing slopes of Volcán Atitlán, 1,000- 
1,100 m, 21 June 1942 (fr), “eeu? 48002 (F, GH). 
SUCHITEPEQUEZ: Finca Moca, 1,260 8 Jan. 1935 (fl), 
Skutch 2087 (A, F). BELIZE. BELIZE: i. 20- n yg 
(8), Dwyer 12595 (DUKE, F, GH, 
NY, US); Maskall, Nov. 1933 (fr), de 928 (A, 
( 


195 
12907 (F, NY, US); Benque Viejo, borderi 
r) 


3 (fl), adi 10828 (F, 

-9 "Mar 938 (fl), B 2307 (A, F, K). 
COROZAL: sis km w of Northern Hwy., 1.5 km N of 
Buena Vista, 23 June 1973 (fr), Dwyer 11373b (GH); 
high ridge, 1931-1932 (fr), Gentle 532 (F, MEXU). 
ORANGE WALK: along Northern Hwy., mi. 58, 23 June 
1973 (fr), Dwyer 11391a (F —2 sheets, MO, NY); Honey 
Camp, Nov. 1928 (fr), Lundell 107 (F, K, US). TOLEDO: 
Golden Stream River, upper reach, Bolo Camp, 16 Apr. 
1944 (fl), Gentle 4539 (LL); between Orange Point and 
Pablow, Broken Cohune ridge, 3 Sep. 1955 (fr), Gentle 
7737 (K, MO). ME a ae uA: Villa de Taulabé, 
-13 June 1976 (early fr), 


M Santa Rita, Qda. Jaral, 700 m, 28 Aug. 1975 (fr), A. 
Molina R. & A ead 30835 (F). CORTES: La Cumbre 
desprendimiento de Sierra de Omoa, 190 m, 30 Nov. 

1950 (fr), 4. Molina R “3466 (GH). EL PARAISO: Qda. 
Tapahuasca, 1,300 m, 14 Aug. 1964 (fr), 4. Molina R. 
14623 (F, NY, US). DISTRITO CENTRAL: Barranco y Qda. 


de Zambrano, entre Zambrano y 


to Sabana Grande, 1,300 m, 9 Nov. 1966 (fr), a Molina 


R. 18652 (F, GH, NY). FRANCISCO MORAZÁN: Qda. Que- 
mada, km 21 vic. Cerro de Hule, 1,300 m, 26 Feb. 1970 
(fr), A. Molina R. 25412 (F, MO, NY). LEMPIRA: Qda. 
Bañaderos cerca de Lepaera, 1,200 m, 27 Sep. 1963 
fr), 4. Molina R. 12991 (F—2 sheets, NY). OCOTEPE- 
QUE: Lempa River between Sta. Anita and Sta. Fe, 700 
m, 3 Sep. 1975 (fr), 4. Molina R. & A. Molina 31059 
ENCB, F, MO). sANTA BARBARA: Montana Sta. Barbara, 
above Sauce nr. Lake Yojoa, 1,000 m, 7 Aug. 1948 (fr), 
L. O. Williams & A. Molina R. 14513 (GH, MO, US). 
EL SALVADOR. AHUACHAPÁN: sin. loc., 1922 (fr), Padilla 
303 (US); Sierra de Apane in TAA region of Finca 
Colima, 17-19 Jan. 1922 (fn), P, 20062 (US). 
NICARAGUA. JINOTEGA: N slope of Volcán Yali, 13°15'N, 
86?10'W, 1,200-1,400 m, 25 Oct. 1979 (fr) W. D. 
Stevens & Grijalva 15136 da NUEVA SEGOVIA: Lom 
Fria ridge W of Rio Las Manos, 20 ong 
1,200 m, 16 June 1977 (fl), Neill 2201 (MO). ZELAYA: 
Rio Punta Gorda, Atlanta, P ra del Cano el 
Guineo, bu 4? m, 11 Nov. 1981 EN 
P.P no & Sandino aso (MO). 


— 


— 


Psychotria miradorensis, described from Mex- 
ico, is indistinguishable from P. costivenia, de- 
scribed from Cuba; the latter name was published 
only a few months prior to the former. 

Leaf blades from Zelaya, Nicaragua, at the 
southern end of the geographical range, have 15- 
18 secondary veins, numbers rarely reached in the 
rest of the range. Inflorescences tend to be smaller 
in Honduras, El Salvador, and Nicaragua than 
farther north. The corolla appears generally wider 
(2 mm) in Honduras than in the rest of the range. 


10. Mega cr wa d Contr. U.S. 
Natl. Herb. 29. 1916. TYPE: Panama. 
Canal Area: i ds Isthmus p Panama, 25 
Feb. 1850 (fl), Fendler 59 (holotype, US; 
isotypes, GH, MO). Figure 17 


Shrub 1-2 m tall; young stems glabrous to 
minute puberulent, the bark smooth to longitudi- 
nally striate; stipules ovate, 5-9 x 3-4 mm, gla- 
brous, caducous, leaving a pale ridge with red- 
brown fringe. Leaves subsessile; petioles to 3 mm 
long, glabrous, flat and furrowed above; blades 
coriaceous, stiff, obovate, the apex acute to acu- 
minate, the base attenuate, the margins inrolled, 
(5-)7-9(-10) x (3-)3.5-5(-5.5) cm, glabrous 
above and below, drying bright chalky yellow-green 
above, dull green below, often partly red-brown; 
secondary veins (6-)8-10 pairs, diverging 50°- 
70%, brochidodromous, straight, elevated below, 
glabrous, the secondary loops far from margin, the 
axils lacking domatia or hairs; tertiary veins in- 
conspicuous, orthogonal reticulate, the tertiary loops 
between secondary loops and margin evident. /n- 
florescences terminal, panicles of cymes; panicle 
branched to 3(-4) degrees; main axis 5.5-9 cm 


102 


Annals of the 
Missouri Botanical Garden 


long, the peduncle 3-7 cm long; secondary axes 

in 2(-3) ranks, the first-rank axes (2 or) 4, the 
Dus pair 0.5-1.5 cm long, the shorter pair 0.2- 
0.8 cm long, the second-rank axes 2 (or 4), sub- 
equal, 0.2-1 cm long, the third-rank axes 2, 0.2 
cm long; cymes branched to 1-2 degrees; bracts 
broad, irregular, ca. 2 mm long, glabrous to pu- 
berulent; bracteoles triangular, 0.5 mm long, pu- 
berulent. Flowers pedicellate, the pedicels 1 mm 
long; calyx cup-shaped, the tube 1.5 mm long, the 
lobes 5, triangular, 0.5 x 0.7 mm, glabrous; co- 
rolla white, the tube cylindrical, 2.5 x 1.5 mm, 
white pubescent in throat, the lobes 5, 1.5 x 1 
mm; stamens 5, the filaments not seen in pins, 3— 
4 mm long in thrums, the anthers 1 mm long; style 
not seen in pins, 2-2.5 mm long in thrums, the 
branches linear. Fruit when dry spherical to ellip- 
soidal, 4-4.5 mm long, 3-4 mm diam., maturing 
red, drying black; calyx persistent, to 1.5 mm long, 
drying green; seed dorsal surface with 4 longitu- 
dinal furrows, the ventral surface with 2 longitu- 
dinal furrows. 

Distribution (Fig. 17). Known from the Ca- 
ribbean coast of Panama, just east and west of the 
Canal, collected near sea level in tropical moist to 
tropical wet forest with equatorial-tropical climate. 
It has been collected in flower February, April, 
May, and August; fruiting specimens date from 
April and August. 


Additional sti examined. PANAMA. CANAL 
AREA: Fort Sherman, from Piña to 3 mi. NE of Piña, 2 
Apr. 1973 (fl), iener 1386 (F, MO). COLON: Miguel de 
la Borda, along steep clay sea coast, 24 Apr. 1970 (fl, 
fr), Croat 10023 (F, MO, NY); Isla Grande, 11 Apr. 
1970 (fl), D'Arcy 4020 (MO); María Chiquita, nr. beach, 
10 Aug. 1967 (fl, fr), Dwyer & Kirkbride 7783 (MO 
3 sheets, NY); 4.5 km SW of Piña, Qda. Sta. Marta, 0 
5 m, 17 May 1974 (fl), Nee 11682 (MO); Nee 11688 
(MO). SAN BLAS: between Puerto Obaldia and Puerto Ar 
mila, coastal rocks, 0-100 m, 29 Apr. 1980 (fl), D'Arcy 
13676 

Psychotria fendleri may be recognized by its 
coriaceous, moderate-sized (7-9 cm long) leaves 
usually drying chalky yellow-green and with broch- 
idodromous secondary venation suggestive of that 
of P. horizontalis. Psychotria fendleri is distin- 
guished from P. horizontalis, from which it may 
be a localized derivative, by its coriaceous leaves 
and chalky color when dry. 

The eight flowering specimens examined are all 
thrums, suggesting that the species is thrum-mono- 
morphic. 


11. Psychotria flava Oersted ex Standley, J. 
Wash. Acad. Sci. 17: 341. 1927. TYPE: Mex- 
ico. Veracruz: Misantla (fl), Liebmann 11605 


(holotype, C, n.v., peu F neg. no. 22830). 
Figures 7b, 10j, 1 


Shrub or small tree, 2-10 m tall; young stems 
glabrous to puberulent, the bark pale, smooth; stip- 
ules ovate, the apex biaristate, 7-22 x 4-8 mm, 
the extensions 3-5 mm long, puberulent, fringed, 
caducous, leaving a pale ridge with red-brown fringe. 
Leaves petiolate; petioles 1.5-3.5 cm long, gla- 

rous, grooved above; blades membranous, elliptic, 
the apex acute to acuminate, the base attenuate, 
16-40 x 6-13 cm, glabrous above, puberulent 
to sometimes glabrous below, drying chalky yellow- 
green to green-brown above, darker green-brown 
below; secondary veins 14-21 pairs, diverging 60°- 
70%, eucamptodromous to brochidodromous, 
straight to constantly arcuate, elevated below, pu- 
berulent or sometimes glabrous below, the axils 
lacking domatia or hairs; tertiary veins evident to 
conspicuous, orthogonal reticulate. /nflorescences 
terminal or pseudoaxillary, panicles of glomerules 
(Fig. 7b); panicle branched to 4 degrees; main axis 
(9-)13-26 cm long, the peduncle (5-)7-15 cm 
long; secondary axes in (3-)4 ranks, the first-rank 
axes 4 or 6, the longer pair 2.5-9.5 cm long, the 
medium pair 0.7-4 cm long, the shorter pair re- 
duced or to 2 cm long, the second-rank axes 4 or 
6, the longer pair (0.3-)1.5-5 cm long, the me- 
dium pair (0-)0.7-2 cm long, the shorter pair 0.4— 
0.5 cm long, the third-rank axes (2 or) 4 or 6, the 
longer pair (0-)0.3-2 cm long, the medium pair 
(0—)0.1—0.5 cm long, the shorter pair reduced, the 
fourth-rank axes 2 (or 4), the longer pair 0.2-0.3 
cm long, the shorter pair reduced; bracts triangular 
to linear, 3-5 mm long, red-brown pubescent, 
fringed. Flowers sessile; calyx cup-shaped, the tube 
1 mm long, the lobes 5, triangular to barely evident, 
to 0.3 x 0.8 mm, puberulent, fringed; corolla 
white, the tube campanulate, 2.5-3 X ca. 1.5 
mm, white pubescent in throat, the lobes 5, linear, 
2 x 1-1.2 mm; stamens 5, the filaments 2.5-3 
mm long in pins, 4-5 mm long in thrums, the 
anthers 1 mm long; style 4.5-5 mm long in pins, 
2.5-3.5 mm long in thrums, the branches linear. 
Fruit when dry ellipsoidal to slightly obovoid, 8— 
12(-13) mm long, 6-8(- m diam., maturing 
red, drying black; calyx sometimes persistent as 
beak drying pale brown; seed dorsal surface with 
6-10 irregular longitudinal furrows, the ventral 
surface with 1 often T-shaped longitudinal furrow 
(Fig. 10j). 

Distribution (Fig. 17). 
Mexico, primarily the Gulf coast, and Petén, Gua- 
temala, occurring at elevations of 20-1,000 m, 
mostly under 200 m, in evergreen forest with equa- 


Known from southern 


Volume 76, Number 1 
1989 


Hamilton 
Mesoamerican o th 
subg. Psychotr 


103 


torial-tropical to tropical-mountainous climate. 
Psychotria flava has been collected in flower al- 

most exclusively February-May and in fruit 
throughout the year, primarily August- December. 


Selected specimens examined. MEXICO. CHIAPAS: 
Mpio. Ocozocoautla de Espinosa, 46 km N of Ocozo- 
coautla on rd. to Mal Paso, 700 m, 8 Oct. 1974 (£l, fr), 


de Lázaro Cárdenas, camino Juchitán a Sta. Maria Chi- 
malapa, 330 m, 11 Mar. 1982 (fl), Cedillo & Torres 
1143 (MEXU, MO); Dto. Choapam (= Santiago Choa- 
pan), Yaveo, trail del Chorro, 460 m, 27 Mar. 1938 (fl), 
Mexia 9221 (B, F, CH, K, MO, NY, US); Mpio. Matias 
Romero, 3-5 km al S del Aserradero La Floresta, 16- 
18 km al S de Esmeralda, 17%02'N, 94°47'W, 200 m, 
8 Apr. = (A), dee et al. 3163 (ENCB, MEXU). 
PUEBLA: Mpio. de zalan, Cuauhtapanaloyan, 21 Nov. 
1980 (fr), ie Rafael 246 (MEXU). TABASCO: km 7 
camino peor -Tapijulapa, Cerro del Madrigal, 150 
O (fl), uen. 2890 (ENCB); 32 km al SE 


. 1983 (fr), R. Fernández N. & 
Guadarrama-Zamudio 1379 (MO—2 sheets); Mpio. 
Huimanguillo, 3 km W de Chontalpa, 20 m, 2 Mar. 1972 


2 Aug. 1972 (fr), Villegas 30 (F, M 
NY); Mpio. Hidalgotitlán, km 0-3 t Pi Cedillo-La 
Laguna, 17?20'N, 94*35'W, 140 m, 10 May 1974 (fr), 
i Dorantes 3028 (ENCB, F, MEXU); Mpio. Pajapán, 
amino a Jicacal a Pajapán, 0-5 m, 20 Apr. 1980 (fr), 
onse. & J. Dorantes L. 234 (MEXU); Mpio. Cate- 
royo basuras, 100 m, 25 Sep. 
50 (ENCB, F, GH, 
MEXU); Mpio. Jesús Carranza, carret. de Palomares a 
Uxpanapa, Río Alegre, 150 m, 9 Dec. 1975 (fr), Mon- 
salvo 11 (MEXU); Mpio. Soteapan, cerca de San Fer- 
nando, Santa Martha, 18°22'N, 94°54’W, 1,123 m, 22 
Dec. 1978 (fr), Ortega et al. 1225 (ENCB); Mpio. Mis- 
m 500 m, 15 Dec. 1975 E, gs 12258 (ENCB); 
o. Tlapacoyán, El Limón, 27 Jan. 1981 de 
| mia 18139 (MEXU); Mpio Minatitlán, l km a 
de Poblada 13 en el ue a Hem th (Pob. p. 
17?16'N, 94?09'W, 13 y i (fl), Wendt 
et al. 4090 (MEXU). M 'ATEMALA. S : Remate, on 
Tikal Rd., ca. 8.5 km 
(fl), ciis 930 (M 
Machaqu . 1.8 jte S, 
treras 5304 (Li 


16 May 1961 (fl), Con- 


Psychotria flava may be recognized by its large 
elliptic leaves drying chalky yellow-green to green- 
brown and by its large panicles of glomerules with 
three unequal pairs of secondary axes per rank 
(Fig. 7b). It differs from P. costivenia in having 
much larger leaves, glomerules instead of cymes 
in the inflorescence, and 6-10 irregular (vs. 4-5 


regular) longitudinal furrows on the seed dorsal 
surface and one T-shaped (vs. two) longitudinal 
furrows on the ventral surface (Fig. 10j). 


12. Psychotria grandis Swartz, Prodr., 


maicae interioris occidentalis (fl), Swartz s.n. 
(holotype, S, n.v., photo, A). Cf. also Swartz, 
Fl. Ind. Occid. 417. 1797. Figures 101i, 19. 


Tree or shrub, (1—)4—10 m tall; young branches 
red-brown puberulent, the bark smooth; stipules 
ovate, the apex acuminate, often with central keel, 
usually diverging from stem, the margins usually 
reflexed, the center paler, 10-20(-25) x 6-14 
(-20) mm, glabrous or sometimes red-brown pu- 
berulent in center, sometimes fringed, caducous, 
leaving a pale ridge with red-brown fringe, often 
persistent at terminal 3-6 nodes. Leaves petiolate; 
petioles 0.3-2 cm long, glabrous, flat above; blades 
membranous, obovate to sometimes oblanceolate, 
the apex acuminate, the base attenuate, (10—)17— 
32(-39) x (3.5-)7.5-11(-16) cm, glabrous above, 
glabrous or densely puberulent below, drying green- 
brown to red-brown; secondary veins 13-19 pairs, 
diverging (65?-)70?-85?, eucamptodromous to 
brochidodromous, constantly arcuate, slightly el- 
evated, glabrous or densely pubescent below, the 
axils lacking domatia or hairs; tertiary veins evi- 
dent, orthogonal reticulate to slightly percurrent, 
the intersecondaries often conspicuous. /nflores- 
cences terminal or pseudoaxillary, robust panicles 
of cymes; panicle branched to 5 degrees; main axis 
18-33 cm long, the peduncle 11-18 cm long; 
secondary axes in 4—5 ranks, the first-rank axes 
(2 or) 4 or 6, the longer pair 5-10 cm long, the 
medium pair 3.5-6 cm long, the shorter pair 2.5 
cm long, the second-rank axes 2 or 4, the longer 
pair 2.5-6 cm long, the shorter pair 0.7-4 cm 
long, the third-rank axes 2 or 4, subequal, 0.7-2 
cm long, the fourth-rank axes 4, equal, 0.3-1 cm 
long, the fifth-rank axes 2, 0.3-0.5 cm long; cymes 
branched to 2-3 degrees; bracts triangular, ca. 2 
mm long, often tomentose within; bracteoles lan- 
ceolate, 0.5-1.5 mm long, red-brown puberulent. 
Flowers sessile to pedicellate, the pedicels to 1.5 
mm long; calyx cup-shaped, the tube 0.5 mm long, 
the lobes 5, triangular to barely evident, to 0.3 x 
0.5 mm, puberulent; corolla white, the tube cylin- 
drical to campanulate, 2.5 X 1.5 mm, white pu- 
bescent in throat, the lobes 5, ovate, 1.5-2 x 1 
mm; stamens 5, the filaments 2 mm long in pins, 
3.5-4 mm long in thrums, the anthers 0.8 mm 
long; style 4 mm long in pins, 2.5 mm long in 


104 


Annals of the 
Missouri Botanical Garden 


thrums, the branches linear. Fruit when dry spher- 
ical to slightly ellipsoidal, 5-6.5(-8) mm long, 4.5- 

- iam., maturing red, drying black or 
sometimes red-brown; persistent calyx inconspic- 
uous or a minute beak, drying pale brown; seed 
dorsal surface with 10-15 irregular longitudinal 
furrows, the ventral surface with 2 very deep lon- 


gitudinal furrows (Fig. 10i). 


Distribution (Fig. 19). 
from Belize, Guatemala, and adjacent Honduras 
near the Caribbean coast and also from Nicaragua— 
Panama, where it reaches the Pacific side. It occurs 
also in the Greater Antilles—Cuba, Jamaica, His- 
pañola, and Puerto Rico—and Colombia, Vene- 
zuela, and Ecuador. In Central America this species 
ranges in elevation 0-2,000 m, reaching over 500 
m only occasionally, and is found in tropical moist 
to premontane rain forest with equatorial to trop- 
ical-equatorial climate. Psychotria grandis has 
been collected in flower almost exclusively March- 
July and in fruit throughout the year, especially 
August-January. 


Commonly collected 


Selected specimens examined. GUATEMALA. ALTA 
VERAPAZ: region of Cocolá, NE of Carchá, 1,200 m, 
Apr. 1939 (fl), Standley 70305 (F); woods between Finca 
Cubilgüitz and Hda. Yaxcabanal, 300 m, 7 Mar. 1942 
(fr), Steyermark 44833 (F, US); along Río Sebol, down- 
stream from Carrizal, 150-200 m, 19 Apr. 1942 (fr), 
Steyermark 45800 (F — 2 sheets, NY); Cobán, 1,350 m, 
Apr. 1908 (fl), Türckheim 11-1828 (F, GH, MO, NY, 
US). HUEHUETENANGO: ca. 27 km N of Barillas, vic. Max- 
bal, Sierra da los Cuchumatanes, 1,500 m, 15-16 July 
1942 (early fr), Steyermark 48850 (F-2 abontd around 
Ixcan at ‘‘Patcushin,”’ 
m, 22 July 1942 (early fr), Steyermark 49197 (F). 
7 éndez, 5 Aug. 1966 aes os al ag 5907 
5 (fl), Pittier 


to 


May 1966 (fl), rat D-1 c. Quiriguá, 
75-225 m, 15-31 922 7 Standley. 23866 s 
NY, US); between es and ** resa" in Montan 


de Mico, 40-300 m 


8 


orillando el camino para La Cumbre, a km 1 3 De 
1970 (fr), Ortíz 1447. (DUKE, F, GH, VN Parque 
Nacional de Tikal, en orillando la aguada del Hotel Posada 
la Selva, lado SE, 1 June 1971 (fr), Ortíz 1827 (F, NY, 
US); San Luis, 9 May 1976 (fl), Ventur 267 (F). BELIZE. 
TOLEDO: Columbia Forest Station, nr. entrance, 11 June 
1973 (fl), Dwyer 11091 (GH, MO); Rio Temash, 26 July 
1979 (fr), Dwyer 14793 (MO); vic. Columbia Forestry 
Station, N of San Antonio, 90-150 m, 11-12 June 1973 
A E 8076 (F, MO, NY); Sand Hill, Punta Gorda, 
, 1 Sep. 1932 (fr), Schipp 1013 (A, F, GH, K, 
MOT NY). HONDURAS. ATLANTIDA: Cuyamel, 2 Jan. 1923 
(fr), Carleton 429a (US); Corozal, orilla del Rio Juana 
Leandra, 100 m, 1-3 May 1980 (fl), C. Nelson 5534 
(TEFH); E of Tela, nr. Yoro trail, 30 m, 9 Aug. 1934 
(fr), Yuncker 4975 (A, F, MO); 23 km E of La Ceiba, 


plains nr. Roma siding of S. F. Co. RR, 21 July 1938 
(fr), Yuncker et al. 8567 (F, GH, K, MO, NY, US). 
OLANCHO: Mata de Maiz, 30 km NE Culmi, 700 m, 1- 


NICARAGUA. CHONTALES: 4 km al N 
12°17'N, 85?06'W, 280 m, 12-13 May 1984 (fl), Gri- 
jalva 3793 (MO). RÍO SAN JUAN: along Rio San Juan 
between San Juan del Norte and Delta de San Juan, 0- 
50 m, 24-25 Mar. 1961 (fl), Bunting & Licht 855 (F). 
RIVAS: Isla Ometepe, Volcán Maderas, Balgüe, 11?26'N, 
85°30'W, 1,200-1,260 m, 1 May 1984 (fl), Robleto 
524 (MO). ZELAYA: Comarca del Cabo, San Mateo, 16 
mi. al S de Tronquera cerca de Rio Wawa, 35 m, 22 
Aug. 1965 (fr), 4. Molina R. 15083 (F, NY); Río Punta 
Gorda, Atlanta, “La Richard," 11%32'N, 84?05'W, 20 
m, 13 Nov. 1981 (fr), P. P. Moreno & Sandino 13027b 
(MO); 12 km NE of La Cruz de Rio Grande, 10 m, 10 
June 1978 (fl), Neill 4375 (MO); Cano El Ocote, 13°38'N, 
85°04'W, 340 m, Apr. 1983 (fl), Ortíz 1388 (MO); Caño 
Serrano, 40 km E of Nueva RAS l Apr. ea (fl), 
Sandino 4952 (MO); vic. El Recreo, on Rio Mico, 30 
m, 23 Apr.-14 May 1949 (bud), Sila. 19362 (F); 
new rd. to Mina Nueva America, more or less westward 
from ca. 14.3 km N of El Empalme on main rd. to Rosita, 
ca. 8.6 km from main rd., 27 Apr. 1978 (fl), W. D. 
Stevens 8399 (MO); Monkey Point, 11?35'N, 83°39'W, 

-20 m, 7 Apr. 1981 (bud), W. D. Stevens 20025 (MO). 
Costa RICA. ALAJUELA: rd. between Cañas and Upala, 10 
km N of Bijagua, 200 m, 26 June 1976 (early 2 Croat 
36465 (MO, US); 4 km SE of Fortuna, 10°29'N, 8 ?43'W, 
29 Apr. 1983 (fl), Liesner et al. 15222 (MO). CARTAGO: 
Rio Reventazón, nr. Inter-American Institute of Agricul- 
tural Sciences, 3 km SE of Turrialba, 9%55'N, 83?41'W, 
500-600 m, 20 June 1949 (early fr), Holm & Iltis 22 
(B, F, NY). HEREDIA: Finca La Selva, OTS field station 
on Río Puerto Viejo, 100 m, 4 Aug. 1980 (fr), Hammel 
9447 (DUKE); along Rio Vueltas from old rd. to Carillo, 
10%05'N, 84?05'W, 2,00 
er 2030 (CR 


2-16 Aug. 1938 (fr), Woodson 
8 (A, F, MO). CANAL AREA: Pipeline Rd. 5.6 
mi. from gate, 6 Dec. 1970 (fr), Croat 12757 (DUKE, 

, MO, NY); Fort Sherman, 30 Nov. 1966 (fr), Dwyer 
7179 (GH, MO); Barro Colorado Island, 25 Feb. 1932 
(fl), Woodworth & Vestal 700 (A, F, MO). CHIRIQUÍ: 
Progreso, July-Aug. diui po Cooper & Slater 186 (F, 
US); Puerto Armuelles, , 15 Aug. 1938 (fr), M. 
Davidson 1120 (F); vic. E ub. Peninsula de 


1938 (fl, fr), Allen 951 (GH, MO, NY, US); Rio Chu- 
cunaque, between Rio Membrillo and Rio Subcutí, 22 
Aug. 1966 (fr), Duke 8597 (MO, US); Rio Piña, 2 Mar. 
1967 (f), Duke 10568 (MO, US); Serrania del Sapo, 
7°40'N, 78°10'W, 450 m, 2 Jan. 1981 (fl, fr), W. Hahn 


Volume 76, Number 1 
1989 


Hamilton 105 
Mesoamerican Psychotria 
subg. Psychotria 


296 (MO); Manené to mouth of Rio Peur 28 Apr. 1968 
(fl), Kirkbride € Bristan 1460 (MO, NY); trail from 
Paya to Pucro, 12 June 1959 (early i». Stern et al. 426 
(GH, MO, US); vic. Campamento Buena Vista, Rio Chu- 
cunaque above confluence with Rio PEN 5 July 1959 
(early fr), Stern et al. 935 (GH, MO, US). Los SANTOS: 
Loma Prieta, 800-900 m, 8 y 1967 (fl), Duke 11897 
(MO); on rd. cut towards Veraguas from El Cortezo, 300- 
600 m, 26 Oct. 1978 (fr), Hammel 5321 (MO). PANAMÁ: 
vic. Pacora, 35 m, 5 Nov. 1939 (fl), Allen 2035 (GH, 
MO, NY, US); La Chorrera, frente a la fábrica Maribel, 
20 Nov. 1971 (fr), Atencio 11 (DUKE, MO); Rio Pita, 
1-3 mi. above confluence with Rio La Maestra, 14 Oct. 
1961 (early fr), Duke 4759 (GH, MO); El Llano-Carti 
rd., 1.4 km N of Panam. Hwy., 150 m, 8 Jan. 1974 
(fr), Nee & Dwyer 9235 (F, MO). SAN BLAS: headwaters 
of Río Cuadi, 100 m, 18 Dec. 1967 (fl), Duke et al. 
3652 (MO); mainland opposite Playón Chico, 0-3 mi. 
from Caribbean, 0-200 m, 4 Oct. 1972 (fr), ae 
6417 (F, MO); nr. Puerto Obaldia, 8?40'N, 77°2 

m, 16 Apr. 1982 (fl, fr), Knapp & Mallet 4641 (MO — 
2 sheets). Veraguas: Isla de Coiba, 16 Aug. 1961 (early 
fr), Dwyer 1540 (MO) 


~ 


Psychotria grandis may be recognized by its 
large size (to 10 m tall), robust keeled stipules with 
margins reflexed and usually persistent at terminal 
3-6 nodes, large (17-32 cm long) leaves with 
conspicuous intersecondary veins, large robust in- 
florescences, and fruit drying spherical and smooth, 
the latter due to the seed dorsal surface having 
many shallow irregular furrows instead of few deep 
furrows (Fig. 10i). The distinction between this 
species and P. costivenia 1s somewhat problematic, 
as it Is based largely on quantitative characters, 
such as leaf and stipule and inflorescence size. 
Differences in seed cross section (10-15 irregular 
vs. 4-5 deep regular furrows on dorsal surface) 
support the distinction. The two species occur sym- 
patrically only in northern Guatemala and southern 

elize. 

General puberulence is common only in Belize 
and Honduras. Material from Belize has the longest 
stipules; Guatemalan material has the largest in- 
florescences; and Nicaraguan fruit is generally larg- 
er than elsewhere. 


13. Psychotria horizontalis Swartz, Prodr., 
44. 1788. Uragoga horizontalis (Sw.) Kuntze, 
Revis. Gen. Pl. 1: 300. 1891. Myrstiphyllum 
horizontalis ao Millsp., Publ. Field Mus. 
Nat. Hist., Bot. Ser. 2: 102. 1900. TYPE: 
Hills (fr), Swartz s.n. (holot ype, 5, n.v., 
photo, A; isotype, B— Willdenow 4087). Cf 
also Swartz, Fl. Ind. Occid., 410. 1797. Fig- 
ures 3b, 9e, 10b, 

Psychotria dece Kunth in Humboldt, Bonpland 
& 


Kunth, Nov. Gen. Sp. 3: 358. 1819. Uragoga 
glaucescens (Kunth) Kuntze, on Gen. Pl. 2: 960 


1891. Psychotria horizontalis Sw. var. glauces- 
eyerm., Mem. Ne 
n ripa Orinoci 


me (8, fr), Bonpland s.n. (holotype, P, 


n.v.). 
Psychotria longicollis Bentham in Oersted, Vidensk. 
Dansk Naturhist. Foren. Kjebenhavn 1852: 


i ca. 
Cartago: nr. Turrialba, ca. 900 m, May (fr), Oersted 

.n. (holotype, C, n.v., photo, MO). 
Psychotria bimea L. Riley Kow Bull. 1927: dr 1927. 
Panama. Piin : between Panamá and Sa- 


T 
banas, 14 June 1924 (8). Riley "2 Uribe K, 


n otype, MO). 
Psychotria ibn ru Swartz EUM basicordata Dwyer, 
Ann. 


ouri Bot. Gard. 55: 4 


1962 (early fr), Dwyer 2384a (holotype, MO— 
2165270; isotype, MO 
ipiis horizontalis Swartz var. psilophylla Stey- 
r m. New York Bot. Gard. 23: 472. 1972. 
TYPE: Colombia. Magdalena: nr. Masinga Vieja, Sta. 
Marta, 300-450 m, May 1898-1899 (8), H. H. 
Smith 393 (holotype, NY; isotypes, F, GH). 
Shrub 1-3 m tall; young stems glabrous or 
puberulent or ferrugineous-pubescent, the bark pale, 
smooth; stipules sheathing, ovate-acuminate, 
(2.5-)4-7 x 2-3.5 mm, puberulent to ferrugin- 
eous-pubescent, caducous, leaving a pale ridge 
with red-brown fringe. Leaves petiolate; petioles 
(1-)2-13 mm long, glabrous to puberulent to fer- 
rugineous-pubescent, flat or grooved above; blades 
membranous, elliptic or rarely ovate, the apex acute 
to long-acuminate, the base cuneate to sometimes 
cordate, the margins often crenate, (6-)7.5-15 
(717) x 3-6.5(-8) cm, glabrous above, glabrous to 
puberulent on midvein to puberulent below, drying 
dull green or sometimes red-brown; secondary veins 
(6-)8-11(-14) pairs, diverging (45°-)60°-75°, 
brochidodromous, the secondary loops far from 
margin (Fig. 3b), increasingly arcuate toward mar- 
gin, prominulous below, glabrous or sometimes pu- 
berulent below, the axils sometimes with minute 
domatia below; tertiary veins evident to conspic- 
uous, orthogonal reticulate, the loops between sec- 
ondary loops and margins evident. /nflorescences 
terminal or pseudoaxillary, ie panicles of 
s panicle branched to 3 degrees; main axis 
.9-)4-10 cm long, the peduncle rarely lacking 
or (2-)3-7 cm long; secondary axes in (2-)3-4 
ranks, the first-rank axes 4 or 6, the longer pair 
0.7-)1-2 cm long, the medium pair 0.4-1.4 cm 
long, the shorter pair 0.3-0.6 cm long, the second- 
rank axes 2 or 4, the longer pair (0.2-)0.4-0.9 
cm long, the shorter pair 0.3-0.6 cm long, the 
third-rank axes 2, 0.1-0.3 cm long, the fourth- 
rank axes 2, 0.1-0.2 cm long; cymes branched to 
1(-2) degrees; bracts inconspicuous, linear to tri- 


TO 


— 


106 


Annals of the 
Missouri Botanical Garden 


angular, to 5 mm long, puberulent to ferrugineous- 
pubescent; bracteoles linear, to 1 mm long, pu- 
berulent to ferrugineous-pubescent. Flowers ped- 
icellate, the pedicels 0.5-1.5 mm long; calyx 
campanulate, the tube 0.5 mm long, the lobes 5, 
lanceolate, (0.5-)1-2 x 0.6 mm, puberulent; co- 
rolla white, the tube cylindrical, 2.5-3.5 x 1.3- 

.5 mm, white pubescent in throat, the lobes 5, 
lanceolate, 1.5- m; stamens 5, the fil- 
aments (2.5-)3-3.5 mm long in pins, 5-6 mm 
long in thrums, the anthers 0.8-1 mm long; style 
(4.5-)5-6 mm long in pins, 3-3.5 mm long in 
thrums, the branches linear, often recurved. Fruit 
when dry ellipsoidal, 3.5-4.5(-5) mm long, 2.5- 
3.5(-4) mm diam., maturing red, drying red-brown, 
sometimes sparsely puberulent; persistent calyx of 
conspicuous lanceolate lobes, 1-3(-5) mm long 
(Fig. 9e); seed dorsal surface with 3-5 longitudinal 
furrows, the ventral surface with 2 longitudinal 
furrows (Fig. 10b 


Distribution (Fig. 16). 
primarily the Pacific coast from Sinaloa eastward, 
through Panama, ubiquitous throughout its range 
at elevations of 0—1,300 m, mostly below 600 m, 
in tropical moist to premontane wet to sometimes 


Known from Mexico, 


tropical dry forest (deciduous forest in western 
Mexico) with equatorial or usually tropical or some- 
times subtropical-tropical climate. Psychotria 
horizontalis occurs also in Cuba, Hispaniola, Co- 
lombia, Venezuela, the Guianas, Ecuador, and Bra- 
zil. It has been collected in flower June-August in 
Mexico and primarily March-July throughout the 
rest of its range; fruiting collections come from 
throughout the year, primarily August- March. 


Selected specimens examined. | MEXICO. CHIAPAS: 
Mpio. Frontera Comalapa, 6-8 km E of Frontera Comala- 
pa along rd. to Ciudad Cuauhtemoc, 1,000 m, 15 Aug. 
1972 (fr), Breedlove 26989 (MEXU); Mpio. Arriaga, 6 

m N of Arriaga, 250 m, 23 Aug. 1972 (fr), Breedlove 
27286 (ENCB); Mpio. Ocozocoautla de Espinosa, canyon 
at head of Rio de la Venta at the chorreadero nr. Derna, 
800-1,000 m, 1 Sep. 
(MEXU); Mpio. Ocosingo, ruins o chilan anks 

of Río Usumacinta, 300 m, 20-21 Dec. 1976 (fr), Breed 
E 42841 (MEXU, MO); 4 mi. N of Tapachula a 
. to Nueva Alemania, 250 m, 20 Aug. 1977 (fr), e 


Vallecitos, 500 m, 26 HR ae 7 (8, Hinton et al. 10368 
Y, US); Mpio. Atoyac, 19 km NE of Atoyac, 

iÓ 0 m, 3 Nov. 1979 (fr), Koch 
ag Rincón Vie ejo, nr. Agua de Obispo, 

, 11 June 0 (fl), 13 Aug. 1960 (fr), Kruse 277 
ALISCO: prida Guadalajara- -Colima, a la altura 
de Pihuamo, cerca del río, 750 m, 24 July 1966 (fl), 
Puga 569 (ENCB); Mpio. La Huerta, Estación de Biologia 
Chamela, selva baja caducifolia perturbada, 19?30'N, 
105°03'W, 25 Sep. 1981 (fr), Lott 575 (MEXU, MO); 


(ENCE B). J 


Mpio. La Huerta, Puerto Vallarta-Barra de Navidad, ca. 
7 km al SE de la Estación de Biologia Chamela, 27 Oct. 
1981 (fr), Lott & Magallanes 625 (MEXU). MICHOACÁN: 
Coalcomán, Villa Victoria, 700 m, 11 July 1939 (fl), 
Hinton 13912 (F, GH, MO, NY, US). NAYARIT: Old San 
Blas, on hill, m Oct. 1925 (fr) Ferris 5417 n 2 


m 
less, 21 July 1959 (early sl King 1955 (US); Mpio. 
Juquila, 1 km al S de Charco Redondo en el Parque 
Nacional “Lagunas de Chacahua,” 3-5 m, 4 Aug. 1979 
(fr), Loera et al. C-3 (MEXU); Jamiltepec, 1.5 km al N 
del Rosario y 13 km al N de Jamiltepec, 22 Oct. 1982 
(fr), Torres et al. 1646 (ENCB, MEXU, MO). SINALOA: 
Mazatlán, San Ignacio, San Juan, 200 m (fl), J. González 
Ortega 4013 (MEXU, US); Mazatlán, San Ignacio, Ar- 
roya del Palmarito, 220 m, 25 Sep. 1918 (fr), Montes 
& Salazar 614 (US). TABASCO: Mpio. Tenosique, a ca. 
15 km arriba de La Palma por río, a 0.5 km del rancho 
Punta de Montaña del Sr. Angel Zubieta, 4 July 1981 
(A), Cowan & Niño 3385 (MEXU, MO); Balancán, ejido 
el Soberano a 7 km del ejido El Palmar, 10 Dec. 1975 
(fr), Novelo et al. 204 (MO). GUATEMALA. CHIQUIMULA: 
moist thickets of Esquipulas River, 1,100 m, 26 Sep. 
1971 (fr), 4. Molina R. & A. Molina 26735 (ENCB, 
F). ESCUINTLA: jct. of CA-2 and Río Coyolate, 300 m, 26 
May 1970 (fl), Harmon & Fuentes 2375 (ENCB, MO— 
2 sheets). HUEHUETENANGO: canyon as to Rio Trapi- 
chillo, between Democracia and can of Chamusht, 
000-1,100 m, 24 Aug. 1942 or fr) Steyermark 
51230 (A, F). IZABAL: Quiriguá, 75-225 m, 15-31 May 
1922 (fl), Standley 24701 (GH, US). PETÉN: La Libertad 
and vicinity, 3 June 1934 (fl), M. Aguilar H: 260 (F, 
H, K, 1 . 1.5 km S of Lacandón, 5 Mar 
1962 (fr), fess 3466 (MO); bordering Lake Petén 
Itza, between San José and Chachaclum, low forest, 25 
Jan. 1971 (fr), Contreras 10396 (MO); Santa Elena, en 
orillando el camino para La Libertad, a km 20, 13 
1970 (fr), Ortíz 1500 a F, MO, NY); en el ia 
que conduce a Arroyo Pucte, km 40 S saliente de Saya 
ché, 15 June 1973 (fl), Ortíz 2698 (F, US); Cerro Ceibal 
cn Mojada), between mouth of = m Mónica an 
uth of Río San Martin, on W s rie 
75- 150 m, 30 Apr. 1942 (st), ad 4610 1 (F). 
SANTA ROSA: Santa Rosa, 900 m, May 1903 (fl), Heyde 
& Lux 4494 (F, US— 2 sheets); 
21 Dec. e AN Standley 60657 (F); 


= 


0- 

SUC iet 6 
of government exper 
16 May "1970 (A), mus 2340 (MO); vic. Tiquisate, 
100 m, 17 June 1942 (fr), Steyermark 47694 (F, GH). 
BELIZE. BELIZE: Gracie Rock, 1.5 mi. S of mile 22 on 
Western Hwy., 4-5 June 1973 (fl), Croat 23838 (F, 
MO, NY). cayo: Cocquericot, 16 Mar. 1931 (fl), Bartlett 
12043a (F); Spanish Lookout Crossing E of Belize River 
along rd., 17 Mar. 1967 (st), Dwyer et al. 167 (MO); 
vic. Blaneaneaux Lodge 5 
Pine ridge, 600 m, 24 Jan. 1974 (fr), Liesner & Dwyer 
1598 (GH, MO); Caves Branch, Mountain Cow hill, 25 
July 1976 (fl), these il 1119 (MO). ORANGE WALK: 
Indian Church, 27 June 1976 (A, fr), Arnason & Lambert 
17081 (MO). HONDURAS. COMAYAGUA: Qda. Las Mercedes, 
Valle Comayagua, 650 m, 28 June 1964 (fl), A. Molina 
R. 14378 (F, NY); Río Hondo, 5 km to La Libertad, 


Volume 76, Number 1 
1989 


Hamilton 107 
Mesoamerican Psychotria 
tri 


subg. Psychotria 


500 m, 17 Oct. 1971 (fr), A. Molina R. & A. Molina 
26857 (F). coPÁN: Copan Ruins on rd. to Sta. Rita, Qda. 
Seca, 500 m, 19 Nov. 1969 (fr), 4. Molina R. & A. 
Molina 24627 (F, NY). conTÉs: Ocote Arrancado, 50 
km N Lago de Yojoa, 600 m, Nov. 1980 (fr), C. Nelson 
et al. 5574 (TEFH). EL PARAISO: matorrales del Rio 
dro cerca del pueblo de Teupasenti, 600 m, 26- 
27 Apr. 1963 (fl), 4. Molina R. 11910 (F); Las Mesas 
region nr. Yuscarán, Aug. 1960 (early fr), Pfeifer 1511 
(US). FRANCISCO MORAZÁN: 5 km O de Cedros, orillas de 
riachuelo Chimbo, 900-1,000 m, 28-30 May 1976 (fl), 
C. Nelson & Vargas 3473 (MO); Sabana Grande, 1,100 
m, 26 Aug. 1945 (early fr), J. V. Rodríguez 3271 (F); 
drainage of Rio Yeguare, between Las Mesas and Sta. 
Clara, 14°N, 87°W, 900 m, 3 Sep. 1949 (early fr), L. 
O. Williams 15953 (F, GH). ISLAS DE LA BAHIA: Island 
of Roatán, nr. town of Roatán, 16 Aug. 1970 (early fr), 
n & Dwyer 3945 (ENCB, F, MO, NY, US 
EE o nr. San Antonio, forest El Cerro, 1,300 m, 
30 Aug. 1968 (fr), 4. Molina R. 22447 (F, NY—2 
sheets). SANTA BARBARA: W side Lago de Yojoa, 850 m, 
16 May 1972 (fl), Burch 6091 (MO, NY); Los m 
on Río Chamelecón SW of Quimistán, 265-360 m, 16- 
17 Apr. 1947 (st), Standley & Lindelie 7454 (F) y V 'ALLE: 
Planada del Puerto de San Lorenzo, 0 m, 7 955 
(early fr), A. Molina R. 5404 (F) EL SALVADOR. 
200-450 m, 28 July 1977 (fr), Eqs 42072 (MO). 
CHALATENANGO: rd. to Chalatenango, 3 wy. 
200 m, 10 June 1970 (f), Davidse & "Pohl 2070 (F, 
SAN SALVADOR: camino al Cerro > De Ag 
Panchimalco, 22 May 1968 (fl), A. 
Pérez & J. González 39 (MEXU). e Mk UA. BOACO: 


N slope of Cerro Mombachito, between Cerro and main 
o 


N). CHINANDEGA: 3-4 km al S 
Er Grande, “Los Laureles,” 13?15'N, 86°53'W, 340- 
m, 28 Sep. 1981 (fr), P. P. Moreno 11692 (MO). 
CHONTALES: bridge over Qda. Niscala along rd. v biis 
coyapa and Rio Oyate, 11?47'N, 85%01'W, 50 m, 
w 1981 (fl), Henrich & Stevens 167 (MO); eee 
km 118 carret. al Rama, “Cerro Grande," 12?09'N, 
85°31'W, 200-230 m, 20 Aug. 1982 (fr), P. P. Moreno 
16953 (MO); vic. La Libertad, 500-700 m, 29 May-1 
June 1947 (fl), Standley 9124 (F); 2.8 km N of ERES 
12°17'N, 85°23'W, 400-500 m, 30 Dec. 1983 (fr), W. 
D. Stevens 22698 (MO). ESTELÍ: Salto de Estanzuela, Río 
Estanzuela ca. 6 km S of Esteli, 13%01'N, 86?20'W, 
920-1,020 m, 1 Oct. 1979 (fr), W. D. Stevens et al. 
E de (MO). LEÓN: carret. a León, Hda. “El Chanal," 
2 km hacia León del empalme León-La Paz Centro, 
del Sep. 1982 (fr), pea & Grijalva 1282 (MO). 
MANAGUA: km 24 o 12 (carret. vieja a León), 7 
km WSW of Minen. of e de Managua, 12?04'N, 
86°26'W, 200 m, 7 July 1977 (fl), W. D. Stevens 2653 
(MO). masaYa: Parque Nacional Volcán Masaya, 
6 Aug. 1982 (fr), ie 999 (MO). MATAGALPA: Río 
uma of Matagalpa, 400 m, 25 
d p (fl), Neill 1994 (MO); N de ee ls carret. 
ón-Pancasán, 12%55'N, 85°49'W, 7 800 m 
16 qe 1982 (fl), uns & Sáenz 3074 y^. RÍO 
SAN JUAN: along rd. to San Carlos 5 km SE of Rio Oyate, 
11?42'N, 84°57'W, 40 m, 28 Aug. 1983 (fr), J. Miller 
& Nee 1370 (MO); Archipelago Solentiname, Isla San 
Fernando, 11?11'N, 84*59'W, 30 m, 18 Sep. 1982 (fr), 


Sandino 3629 (MO). ZELAYA: Caño Calcamo, ca. 5 km 
al SE de Siuna, carret. Siuna-Empalme, 13%40'N, 
84°45'W, 26 Oct. 1982 (fr), Grijalva & Burgos 1522 
(MO); nr. Río Yaoya, 4 km S of crossing of Siuna- Rosita 
hwy., 100 m, 2 May 1978 (fl), Neill 3791 (MO); Cerro 
Waylawás, E slope of northern range, 13*39'N, 84°49'W, 
80 m, 11 Mar. 1979 (fr), Pipoly 4396 (MO); Ibo Tingni, 


N of rd. between Puerto 


a , 
Oct. 1978 (fr), W. D. Stevens 10634 (MO). Costa Rica. 
ALAJUELA: Río Jesús, San Ramón, 700 m, 20 May 1982 
(fl), Carvajal 260 (MO). GUANACASTE: Bebedero, 40-50 
m, 19 June 1930 (fl, Brenes 12603 (CR, F, NY); nr. 
27 Abril, along rd. to Playa Tamarindo, 10*16'N, 
8546'W, 20-80 m, 9 Nov. 1975 (fr), Burger & Baker 
9894 (CR, F, MO); 5 mi. N of Bagaces, 14 July 1965 
(A), Croat 632 (MO); Santa Rosa National Park, 30 km 
NW of Liberia, 10%50'N, 85?35'W, 0-300 m, 1 Dec 

1976 (fr), Janzen 10426 (MO); 19 June 1978 (fl), pes 
zen 10979 (MO); 10-20 km NE of Liberia on Camino 
Sta. María, 300-650 m, 20 Sep. 1975 (fr), Utley & 
ud 3134 (DUKE). i^ CE gis Cerro dap Gorda, 

m NW of Punta de Burica, 200 m, 5 Mar. 1973 (fl), 
Pi 770 (CR, MO); ^ c ce EE Sirena, 
8*29'N, 83°36'W, 0-5 m, 6 1977 (fr), Liesner 
2975 (CR, ea i m, June 


— 
` 


Island, 5 Nov. 19 
1983 (fl), o 3 


early fr), Hamilton & Stockwell 
3706 (MO). cocLÉ: El Valle, 26 May 1970 (fl), I. Aguilar 
21 (F, MO); nr. El Cope, 27 Oct. 1967 (fr), Garner 40 
KE, MO). nos path from sea to ridge a 
Garrote, 4 Nov. 1975 (fr), D'Arcy ole (MO, pie 
Peluca, km no Transisthmia w rd. to 
Nombre de D 25 Feb. 1973 (fl), Kennedy 267 1 (GH, 
O, NY). DARIÉN: Rio Morti, ca. 250 m, 18 Sep. 1967 
(fr), ike 14178 (F, MO); S of El Real on ia of Cerro 
Pirre, 500-1,000 m, 26 Sep. 1969 (fr), Foster & Ken- 
nedy 1274 (DUKE); Ensenada del Guayabo, 18 km SE 
of Jaqué, 1-12 Apr. 1978 (fl), Garwood 772 (F); Can- 
glon, N of Panamerican Hwy., 'N, 77°45'W, 100 
m, 17 July 1982 (early fr), Hamilton et al. 513 (MO); 


su 
m, 10 May 1979 hy Hammel 7323 


along beach, June 1914 (fl), Pittier 6969 (US) 
Yaviza, along Río Chucunaque, nr. El Punteadero, 7 Ju 
1959 (early fr), Stern et al. 164 (GH, MO, US). HERRERA 


Ocú, 27 Aug. 1960 D fr), Ebinger 1058 (ENCB, F, 
MO). LOS SANTOS: 10 ortezo, 810 m, 27 
E D 1 aa i aae. 5371 (MO); N of Guaniquito, 
onosi along Río Tonosi, 100-2 
17 july 1970 (eariy fr), anda & Foster 1382 Tux 
a Ermita-San Carlos, 5 Nov. 1972 
(fr), Banal 11 [CHI Isla del Rey, trail to bis 
un: 20 July 1967 (early fr), M. Correa A. 
DUKE, MO); Rio Pita, 1-3 mi. abov e P ERA with 
| Maca, 14 Oct. 1961 (fr), Dile 4764 (GH—2 
,M ; San José Island, 28 May 1945 (f), 
posa 236 (GH, NY, US); Majé, 5 mi. up Río Majé, 
O m, 18 Nov. 1970 (fr), Foster & Kennedy 2013 
(DUKE); Taboga ‘Talend, 15 Aug. 1972 (fr), Gentry 5736 


N 
© 


108 


Annals of the 
Missouri Botanical Garden 


(MO); between Chepo and El Llano, 1 Sep. 1971 (fr), 
Gentry & Tyson 1693 (MO, n Alcalde Diaz, 11 Nov. 
1974 (fr), J. Gómez 22 (DUKE, MO); € just S d 
Panamerican Hwy., 9?10'N, 78°52 W, 

Sep. 1982 (fr), Hamilton & D'Arcy 1411 as Eoo 
Paitilla, Nov. 1921 (fr), Heriberto 207 (GH, NY, US); 
. 1971 (fr), 


i. E of Cañazas checkpoint, 8°52'N, 78°15'W, 0-50 
m, 27 Feb. 1982 (fl), Knapp 3882 (MO); Pearl Islands, 
Trapiche Island, 15 Mar. 1937 (fl), E 2d 1900 (US); 
970 e Sandoval 

am id ume 

1923 (fr), Standley 26586 (US). SAN BLAS: - Ailigandi, 
Aug. 1965 (fr), Dwyer 6820 (MO). VERAGUAS: Islas Con- 
treras, Isla Uva, 8°48'N, 81?45'W, 50 m, 18 July 1984 
(fr), Churchill 5688 (MO — 2 sheets); Cerro Tute, 8 Aug. 
1963 (early fr), Dwyer 4286 (MO); vic. Ponuga, 31 July 
1967 (early fr), Dwyer & Kirkbride 7440 (DUKE, MO); 
2 mi. W of Santiago on Panamerican Hwy., 6 Aug. 1967 
(early fr), Dwyer et al. 7558 (GH, MO, US); Coiba Island, 
Ensenada Sta. Cruz, 27 Aug. 1970 (fr), Foster ie 
ud F, MO); 18 km W of Las Minas, N slope 

to Higo, cutover areas, 720-900 m, 8 Aug. 1978 (fr) 
lene 4332 (MO). 


S, 


All the species in synonymy have been properly 
synonymized before (cf. Dwyer, 1980). Infraspe- 
cific categories are not recognized herein because 
they have been defined in terms of variation seen 
within almost any population, such as the basicor- 
date leaf blades on which P. horizontalis subsp. 
basicordata was based. 

Psychotria horizontalis may be recognized by 
its leaves drying usually dull pale green, with sec- 
ondary veins brochidodromous and making con- 
necting arches far from the margin (Fig. 3b), and 
by its fruit with conspicuous persistent calyx of 
long, linear lobes (Fig. 9e) 

Material from Guatemala, Belize, and Nicaragua 
is generally more pubescent than elsewhere in Cen- 
tral America. The calyx lobes, prominently persis- 
tent in fruit, are often much shorter in central 
Panama than elsewhere. 


14. Psychotria —€— a Hems- 
ley, Biol. Cent. Amer. Bot. 50. 1881. Based 
on Md Bui nies "inen Amér. 
Centr. p. 17, t. 14, fig. 5. 1863. Uragoga 
papantlensis (Oerst.) Kuntze, Revis. Gen. Pl. 
2: 962. 1891. TYPE: Mexico. Veracruz: Pa- 
pantla, Liebmann s.n. (holotype, C, n.v.). Fig- 
ures 9b, 16 


Shrub or tree, 1-4 m tall; young stems glabrous, 
the bark pale, smooth; stipules sheathing, ovate, 
the apex usually biaristate, 4-9 x 2.5-6 
often paler in the central triangle, the margin often 
ciliate, often persistent at terminal 2-4 nodes, leav- 
ing a pale ridge with short red-brown fringe. Leaves 


mm, 


petiolate; petioles (0.5-)1-3(-4) cm long, often 
diverging at right angles to stem, glabrous, flat 
above; blades very thin membranous, elliptic to 
oblanceolate, the apex acuminate, the base atten- 
uate, (6.5-)9-14(-17) x (1.5-)2.5-5(-6) cm, 
glabrous above and below, drying usually pale green 
to sometimes pale red-brown; secondary veins 9— 
11 pairs, diverging 65?-75?, eucamptodromous to 
brochidodromous, constantly arcuate, prominulous 
below, glabrous, the axils lacking domatia or hairs; 
tertiary veins evident, orthogonal reticulate. /nflo- 
rescences terminal or pseudoaxillary, panicles of 
cymes; panicle branched to 3 degrees; main axis 
3.5-14.5 cm long, the peduncle 2.5-11 cm long; 
secondary axes in 3(-4) ranks, the first-rank axes 
4, the longer pair 0.6-1.7 cm long, the shorter 
pair 0.4-1 cm long, the second-rank axes 4, sub- 
equal, 0.3-0.8 cm long, the third-rank axes 2 or 
4, subequal, 0.2—0.4 cm long, the fourth-rank axes 
2, 0.1 cm long; cymes branched to 1-2(-3) de- 
grees; bracts and bracteoles lanceolate, ca. 1 mm 
long, fringed, caducous. Flowers sessile to pedi- 
cellate, the pedicels to 0.7 mm long; calyx drying 
pale brown against darker petiole and receptacle, 
cup-shaped, the tube to 0.5 mm long, the lobes 5, 
triangular, 0.5 x 0.5 mm, glabrous to minutely 
ciliate; corolla white, the tube cylindrical, 4-5 x 
l mm, sparsely white pubescent in throat, the lobes 
5, lanceolate, 1.5-2.5 x m; stamens 5, 
the filaments 3 mm long in pins, 6-7.5 mm long 
in thrums, the anthers 1.3-2 mm long; style 6 mm 
long in pins, 4-5 mm long in thrums, the branches 
linear. Fruit when dry ellipsoidal, (6-)7-7.5 mm 
long, (4-)4.5-5 mm diam., maturing red, drying 
black; persistent calyx a tubular beak, drying pale 
green, 0.5-1 mm long (Fig. 9b); seed dorsal surface 
with 4 shallow longitudinal furrows, the ventral 
surface with 2 deep incompletely divided longitu- 
dinal furrows. 


Distribution (Fig. 16). 
Mexico through Petén, Guatemala, 
western Belize, at 0-400 m elevation in evergreen 
forest with usually equatorial-tropical climate. Psy- 
chotria papantlensis has been collected in flower 
March-June and in fruit in March and July—De- 


cember. 


Common in southern 
into south- 


Selected specimens examined. MEXICO. CHIAPAS: 60 
mi. SE of Palenque, gravel rd. from Palenque to Bonam- 
pak, 400 m, 5 del 1977 (fr), Croat 40180 (MO); La 
E 29 Apr. 1952 (fl), pipas: 7584 (MEXU); 41 

of Ocozocoautla, r al Paso, 17%12' 
93°40’ W, 350 m, 4-5 Aug 1965 (fr), Roe et al. 903 
(ENCB); ruinas de Palenque, 24 Mar. 191 
Seler 5502 ). TABASCO: Río Zan napa, o 
Huimanguillo, 6 Apr. 1963 (fl), Barlow 10/5 ( (GH); June 


Volume 76, Number 1 
1989 


Hamilto 109 
VERUM Psychotria 


subg. Psychotria 


1964 (f), Barlow v (MEXU); Mpio. Heroica Car- 
denas, km 21 de la carret. Cárdenas-Coatzacoalcos, 30 
Apr. 1981 d Magaña E Curiel 189 (ENCB, MEXU). 
VERACRUZ: Estacion de Biologia Tropical Los Tuxtlas, 400 
m, 4 Apr. 1972 (fl), Cedillo T ie 2 sheets, K, 
» XU); camino Catemaco-M , llegando a Mon- 
tepio, 50 m, 27 July 1977 n pom 206 (MEXU — 
2 sheets, NY); Estación de Biologia Tro opic cal E Tuxtlas, 
68 (fr), Martínez C. 1790 (A, 


lomas al S del Poblado 11 y al S d 
500 m, 2 Oct. 1980 (fr), Wendt et al. 2821 (M 

acoalcos River, 30-50 m, Mar. 1937 (fl, 
fr), L. O. E s 8798 (F, US). GUATEMALA. ALTA 
VERAPAZ: Chahal, bordering Cluiju iud in zapotal, 28 
Sep. 1968 (fr), p 7749 (MO). IZABAL: Cadenas/ 
Puerto Méndez, New Cadenas Rd., 3.5 hs from village, 
19 Aug. 1969 (fr), Pr. 8988 


MO); along Rio 
Bonita, 30-150 m, 21 941 d Steyermark 41678 
(F). PETÉN: on Sebol r km from San Luis Rd., 20 


Nov. 1966 (fr), Contreras 6618 Mut forest E iuam 
n Diego and San Di 
m, 25 Mar 1942 o RU ner 
45346a (F, NY); mue fo ore tW € 

between Río Chinajá an of Rio 
50-70 m, 29 Mar. 1942 (8). Steyermark 45492 (F. 
BELIZE. TOLEDO: San José Maya Indian village, 10 km N 
of Columbia Forest Station, 13 June 1973 (early fr), Croat 
24378 (F, GH, MO) Dwyer 11152a (DUKE, F, MO, 

US). 


o 
o 
3 


Psychotria papantlensis may be recognized by 
its elliptic long-petiolate leaves drying pale green 
and by its calyx drying conspicuously pale in flower 
and in fruit, when it contrasts markedly with the 
black-drying ellipsoidal fruit (Fig. 9b). 

As in several species, the corolla tube of the 
thrum flower morph is longer than that of the pin 
morph (5 mm vs. 4 mm); but the species appears 
otherwise normally distylous. 


15. Psychotria pleuropoda Donnell-Smith, 
Bot. Gaz. (Crawfordsville) 40: 5. 1905. TYPE: 
Guatemala. Alta Verapaz: Cubilgüitz, 350 m, 
Aug. 1903 (fr), H. von Tuerckheim 8529 
(lectotype designated herein, GH; isolecto- 
types, US— 2 sheets, fragments, F, US). Fig- 


ure 17 


Shrub 0.5-5 m tall; young stems glabrous, the 
bark pale, furrowed longitudinally; stipules ovate, 
the apex biacuminate, the central triangle drying 
pale green, 10-17 x 4-6 mm, fringed, persistent 
at terminal 3-5 nodes, leaving a pale ridge with 
red-brown fringe. Leaves petiolate; petioles 
(0.2-)0.5-1 cm long, glabrous, terete; blades 
membranous, narrow-oblanceolate, the apex long- 
acuminate, the base attenuate, (7-)12-21 x (1-) 
1.3-2.7 cm, glabrous above and below, drying 


dull pale green-brown to sometimes red-brown; sec- 
ondary veins 15-18 pairs, diverging 60?-75*, eu- 
camptodromous to brochidodromous, constantly 
arcuate, prominulous below, glabrous, the axils 
lacking domatia or hairs; tertiary veins evident, 
percurrent, the intersecondaries especially evident. 
Inflorescences terminal or pseudoaxillary, panicles 
of cymes; panicle branched to 3 degrees; main axis 
(1.5-)2.5-12 cm long, the peduncle (1-)2-9.5 
cm long; secondary axes in 1-2 ranks, the first- 
rank axes 4, the longer pair 0.3-1.3 cm long, the 
shorter pair 0.2-1 cm long, the second-rank axes 
2 or 4, subequal, 0.2-0.5 cm long; cymes branched 
to 1-2 degrees; bracts linear, to 5 mm long, ciliate; 
bracteoles irregular, ca. 1 mm long, ciliate. Flowers 
sessile to pedicellate, the pedicels to 1 mm long; 
calyx drying pale brown against darker petiole and 
receptacle, cylindrical, the tube 1 mm long, the 
lobes 5, triangular, barely evident, minutely ciliate; 
corolla white, the tube cylindrical, 2.5-3 x 1.5 
mm, white pubescent in throat, the lobes 5, lan- 
ceolate, 2.5 X 1 mm; stamens 5, the filaments 3 
mm long in pins, 3.5-4 mm long in thrums, the 
anthers 0.8-1 mm long; style 5 mm long in pins, 

.5-3 mm long in thrums, the branches clublike 
in pins and linear in thrums. Fruit when dry ellip- 
soidal, (5.5-)6 mm long, (4.5-)5 mm diam., ma- 
turing red, drying black or green-brown; persistent 
calyx a tubular beak ca. 1 mm long; seed dorsal 
surface with 4-5 deep longitudinal Ow Me 
ventral surface with 2 deep sometimes i 


divided longitudinal furrows. 


Distribution (Fig. 17). Known from Tabasco, 
Mexico, northern Guatemala, and southern Belize, 
at 50- m elevation in evergreen forest with 
equatorial-tropical to equatorial-mountainous cli- 
mate. This species has been collected in flower 
February—June and in fruit August- February. 


cted specimens AE. MEXICO. TABASCO: 

Mpio. dumis 3kma del ejido Lázaro Cárdenas, 
50 m, 10 May 197 79 (fl), runs 2069 (NY). GUATEMALA. 
ALTA VERAPAZ: Montana Yxocubvain, 4 km W of Cubil- 
guitz, 300-500 m, 12 Mar. 1942 (fl), Steyermark 44990 
(F— 2 sheets, GH, NY). izaBAL: damp forested slopes and 
barrancos, 300-900 m, 25 Dec. 1941 (fr), Steyermark 
( =m sheets, NY). PETEN: Sayaxché, Laguna 
Petex-Batun, 1 km S of wharf, 30 Mar. 1964 (fl), Con- 
treras 4130 (MEXU, MO); La Cumbre, Las Canas, E of 
of the Petén-Izabal rd., 20 Sep. 1966 (fr), 
ae 6166 (MO); 6 Mar. 1975 (fl), Lundell & 


hinaja, d Rio Chinajá and 6 mi. W of Rio San 
A O m, 29 Mar. 1942 (fl), Steyermark 45488 
(F, NY). Bod TOLEDO: San Benito Poite, nr. Otoxha, 

200 m, Feb. 1973 (st), Boster s.n. (ECON); trail fein 
Columbia Forest Station to Esperanza, 3-6 km W of San 


José rd., 180-330 m, 13 June 1973 (fl), Gentry 8172a 


110 


Annals of the 
Missouri Botanical Garden 


F, NY); Deep River Forest Reserve, limestone-derived 


(F, 
soil, Feb. 1945 (fl), Lamb 54, 62 (F) 


Psychotria pleuropoda may be recognized 
readily by its narrow-oblanceolate (length/ width — 
8-10) leaves unique in subg. Psychotria in Me- 
soamerica. It differs from P. costivenia by having 
very narrow leaf blades and inflorescences with 
relatively short secondary axes in only 1-2 (vs. 
ca. 4) ranks. The persistent calyx tube in fruit is 
also distinctive. 


16. Psychotria sylvivaga Standley, J. Wash. 
A ci. 18: 274. 1928. TYPE: Costa Rica. 
Heredia: Yerba Buena, NE of San Isidro, 2,000 
m, 28 Feb. 1926 (fr), Standley & Valerio 
49989 (holotype, US). Figure 17. 


Shrub 2-3 m tall; young stems sparsely red- 
brown puberulent, the bark pale, furrowed longi- 
tudinally; stipules broadly ovate, 9-12 x 5-6 mm, 
glabrous, sometimes ciliate near base, caducous, 
leaving a pale ridge with red-brown fringe. Leaves 
petiolate; petioles 1-2 cm long, glabrous, flat above; 

lades membranous to sometimes subcoriaceous, 
elliptic to oblanceolate, the apex acuminate to cau- 
date, the base attenuate, (8.5-)10-17 x (2.5-)3- 
4.5(-5) em, glabrous above and below, drying green- 
brown to red-brown; secondary veins 11-14 pairs, 
diverging (60?—)65?—75*(-80?), brochidodromous, 
constantly arcuate, elevated below, glabrous, the 
axils sometimes with minute tufts of red-brown 
hairs; tertiary veins evident to inconspicuous, or- 
thogonal reticulate, the intersecondaries often ev- 
ident. Inflorescences terminal or dps iet 
sparse panicles of cymes; panicle branche 1 
degrees; main axis 9.5-14 cm long, the nda 
6-7.5 cm long; secondary axes in (3-)4 ranks, the 
first-rank axes 2, 2-3.2 cm long, the second-rank 
axes 2, 0.8-1.2 cm long, the third-rank axes 2, 
0.4—0.8 cm long, the fourth-rank axes 2, 0.2-0.5 
cm long; cymes branched to 1 degree; bracts lan- 
ceolate, 3 mm long, glabrous; bracteoles triangular, 
to 1 mm long, red-brown ciliate. Flowers pedicel- 
late, the pedicels to 1 mm long; calyx cup-shaped, 
the tube 0.5 mm long, the lobes 5, triangular, 0.5 
mm long, puberulent; mature corolla, stamens, and 
style not seen. Fruit when dry spherical, 4.5-5 
mm long, 4-5 mm diam., maturing red, drying 
red-black; persistent calyx 1 mm long, drying pale 
brown; seed dorsal surface with 6-8 often irregular 
longitudinal furrows, the ventral surface with 
medium-deep often plus ca. 4 irregular longitudinal 
furrows. 


Distribution (Fig. 17). 


Known from southern 


Alajuela and San José, Costa Rica, at 2,000-2,200 
m elevation in a region of premontane to low mon- 
tane rain forest with equatorial-mountainous cli- 
mate. Psychotria sylvivaga has been collected in 
flower in December and in fruit in December and 
February. 


Additional specimens examined. COSTA RICA. SAN 
JOSÉ: nr. Laguna de la Escuadra, NE of El Copey, 2,000- 
2,200 m, 16 Dec. 1925 (fr), Piani 41924 (A, US), 
41974 (F, US); Laguna de la Chon E of Sta. Maria 
de Dota, 2,000-2,100 m, 18 Dec. "los (fl, early fr), 
Standley 42212 (K, US). 


Psychotria sylvivaga may be recognized by its 
elliptic to oblanceolate leaves drying green-brown 
with brochidodromous secondary veins 
and evident intersecondaries and by its fruit with 


to red-brown 


persistent calyx a conspicuous tube. 

tandley described a mature flower with corolla 
tube 5 X 1.2 mm, the lobes triangular-ovate, 1.5 
mm long, in his original description, but I was 
unable to find a mature flower. 


LITERATURE CITED 


ALMEDA, F. 1978 
m (Melastomataceae) in 
America. Univ. Calif. Publ. Bot. 

ANONYMOUS. 1970. Mapa ecológi d Panamá. El 
Departamento de Cartografi la del Catastro Rural, 
Reforma Agraria, Pana 

AUBLET, M. F. 1775. Histoire des Plantes de la Guiane 
Francaise, Volumes 1, 2. P.-F. Didot jeune, Paris. 

Croat, T. B. 1983. A revision of the genus Anthurium 
(Araceae) of Mexico and Central America. Part I: 
Mexico and Middle America. Ann. Missouri Bot. 

Gard. 70: 211-420. 

19 A revision of the genus Anthurium 

(Araceae) of Mexico and Central America. Part II: 

P : 


Systematics of the genus Mono- 
a and Central 


-5 
fs 


1971. Mapa de tipos de vegetación de la República 
aai Secretaria de Recursos Hidráulicos, Mex- 


a F.R. 1979. The Ar of heterostyly. New 
Zealand J. Bot. 17: 607- 

HaMiLTON, C. W. 1985. idu in neotropical 
Psychotria L. (Rubiaceae): dynamics of branc hing 
and its taxonomic significance. read J. Bot. 
881-888. 

: . New species and combinations in Me- 

soamerican Psychotria subgenus Psychotria (Ru- 

dip Phytologia 64: 219-237. 

ariations on a distylous theme in Meso 

a Ao cade subgenus Psychotria (Rubi- 

aceae). In: G. Prance & G. Gottsberger (editors), 

Reproductive Biology of Plants (working title). New 

York Bot. Gard., Bronx, New York (in press). 


Volume 76, Number 1 
1989 


Hamilton 111 
Mesoamerican Psychotria 


subg. Psychotria 


Hickey, L. J. 1979. A revised classification of the 
architecture of dicotyledonous leaves. /n: C. et- 
calfe & L. Chalk (editors), Anatomy of the Dicoty- 
ledons, 2nd edition, Volume 1. Clarendon Press, Ox- 
ord 


HOLDRIDGE, 


L. R. 1967. Life Zone E x Centro 
Científico Tropical, San José, Costa Ric 
Kress, W. J. 198 Systematics of ARR American 


Heliconia (Heliconiaceae) with pendent inflores- 
cences. J. Arnold Arbor. 65: 429-532. 

LUTEYN, J. L. . Ericaceae — Part I. Cavendishia. 
In: Flora Neotrop. 35: 1-290. New York Botanical 
Garden, New Yor 

MEACHAM, C. & T. DUN 987. The necessity of 
convex groups in biological classification. Syst. Bot. 

2: 78-90. 

MILLER, J. S. s Systematics of the Genus Cordia 
(Boraginaceae) i in Mexico and Central America. Ph.D. 
Thesis. St. Louis University, St. Louis, Missouri. 

MUELLER(-ARGOVIENSIS), J. Rubiaceae brasilien- 

ses novae. Flora 59: 457-466, 495-498 

. 1881. . In: C. F. P. von Martius 

(editor), Flora Brasiliensis 6(5): 1-470. 


Petit, E. 1964. Les espéces africaines du genre Psy- 
chotria L. (Rubiaceae)— I. Bull. Jard. Bot. État 34: 
9. 


Les espéces africaines du genr 


1966. e Psy 
chotria L. (Rubiaceae)— II. Bull. Jard. Bot. État 36: 
6 


5-19 
SOHMER, S. 1977. a I: (Ripiaceae) in the 
Hawaiian Islands. Lyonia 1: 
STEYERMARK, J. A. 1972. nos. Ds: The Botany 
of the Guayana vu. rerom Part IX. Mem. New York 
Bot. Gar 7-832 
Tost, J. A. QU Mapa ecológico de Republica de Costa 


Rica, segun clasificación de zonas de vida del mundo 


de L. R. Ho EIN Centro Cientifico Tropical, San 
Jose, Costa Ric 
WALTER, H. 1973. Vegetation of the Earth in Relation 


to Climate and the Eco-physiological Conditions. J. 
Wieser (translator). The English Universities Press, 


i. Liera & E. HaRNICKELL. 1960. Mittel- 
amerika. In: H. Walter & H. Lieth, poc 
Weltatlas. Gustav Fischer Verlag, Jen 


DIANTHOVEUS: A NEW 
GENUS OF CYCLANTHACEAE! 


Barry E. Hammel? and 
George J. Wilder’ 


ABSTRACT 


Dianthoveus cremnophilus, a terrestrial Cyclanthaceae from southwestern Colombia to northern Ecuador, is 


described as a new 


t shares 


With Schultesiophytum it 


ecies. The cupules at the ba 


na 
. The new genus differs eta enta 
staminate perianth and in seed shape and seed coat structure. 
with Evodianthus (e.g., separate fruits, terminal inflorescences, ote eh 
of these uncommon charact 
appendages. Although Dianthoveus and Dicranopygium sub. 


Diant 


ers but e shares only laminar anther 
genus Gleasonianthus uniquely share staminate flowers 
h 


with eglandular tepals, no other characters suggest a close relationship between them. 


Exploration of tropical wet forests in southern 
Central America and northwestern South America 
in recent years has greatly increased the number 
of collections and species known from these areas. 
This was especially predictable for the Cyclantha- 
ceae, which are restricted to the wettest, often 
least-explored habitats (Grayum € Hammel, 1982). 
Since the time of Harling's (1958) monograph, 
which included 178 species in 11 genera, 29 new 
species have been added to this predominantly epi- 
phytic family. More than 30 additional new species 
are now known from Central America and the 
Chocó region of South America alone (Hammel, 
unpubl.). However, only one new subgenus (Wilder, 
1978) and no other supraspecific taxa have been 
described. In fact, in recent years only 10 genera 
have been recognized; Pseudoludovia is thought 
to be a mixed collection: a spadix, perhaps of a 
Sphaeradenia, and leaves of Ludovia (Wilder, 
1978). Harling suggested that on further study it 
might be possible and logical to subdivide the three 
largest genera into additional genera, and for 
Sphaeradenia this reclassification is imminent (R. 
Eriksson, in press). Dianthoveus, the name an 
anagram of “Evodianthus,” is a totally novel, 
monotypic genus. 

This plant was first described and illustrated, as 
a short-stemmed terrestrial, in The Flora of the 


Río Palenque Science Center (Dodson & Gentry, 
1978), where it was identified as Asplundia va- 
gans Harl., a common root-climbing epiphyte of 
Central and South America. Examination of spec- 
imens of Cyclanthaceae in the herbarium at the 
Marie Selby Botanical Gardens, where vouchers 
for the Rio Palenque florula are deposited, revealed 
that the plant was neither poorly illustrated nor a 
terrestrial form of A. vagans, but rather something 
problematic, exhibiting unusual characters of sev- 
eral genera. This stimulated two field trips to Ec- 
uador, which, along with laboratory studies, con- 
vinced us that the plant is an undescribed genus. 


MATERIALS AND METHODS 


In total, well over 200 individuals in seven pop- 
ulations were examined in the field. Plants at var- 
ious stages—from early anthesis through ripe in- 
fructescences to peduncle remains—were fixed in 

or anatomical and morphological studies. 
Twenty-eight fluid-preserved collections at these 
stages were made, and numerous herbarium vouch- 
ers and live plants were collected. In the field, live 
specimens were examined to determine inflores- 
cence position, presence/absence of latex, drying 
qualities of wounded and air-dried leaves, and fea- 
tures of leaf color and texture. 


' Supported by grants from the National Science Foundation (BSR-8508463) and the Jessie Noyes Smith Foundation 


to B. Hammel 


W. John Kress bse helpful discussion and criticis 


a 
earlier draft of XR paper and Gerrit Davidse and Lucinda McDade for critical review of the submitted manuscript. 


issouri Botanical Garden, P.O. Box 299, St. 
* Cleveland State Ede 1983 East 24th St., 


Louis, Missouri 63166, U.S.A. 
Cleveland, Ohio 44115, U.S.A. 


ANN. MISSOURI Bor. GARD. 76: 112-123. 1989. 


Volume 76, Number 1 
1989 


Hammel & Wilder 113 


Dianthoveus 


Plants preserved in the field in FAA were drained 
of fluid and transported in sealed plastic bags within 
a large plastic container. Subsequently they were 
washed in water for several days and then stored 
in an aqueous solution of 5% glycerin and 50% 
ethanol. All measurements were taken from live or 
fluid-preserved plants at populations along the Río 
Pilatón (Hammel & Wilder 16058, 16059, 16066, 
16067 & 16074). Leaf measurements were taken 
from a total of 17 adult leaves from three individ- 
uals. Inflorescence measurements were taken from 
four individuals, and flower measurements come 
from ten flowers in each inflorescence. Infructes- 
cence measurements were from ten individuals, and 
fruit measurements from ten fruits in each of four 
individuals. For anatomical studies, material of lam- 
inae, roots, and stems was prepared according to 
the methods of Wilder (1985). Seeds were taken 
from mature preserved infructescences embedded 
in Tissue Tek OTC polysaccharide compound, fro- 
zen, and sectioned in a cryostat. Sections were 
mounted, stained with toluidine blue, and viewed 
with a bright field microscope. Pollen was prepared 
with and without acetolysis and with and without 
critical point drying, coated with gold, and viewed 
with a Hitachi S45D scanning electron microscope. 
Detailed studies on the morphology and develop- 
ment of vegetative parts and inflorescences, and 
the anatomy of laminae, first-order roots, and rhi- 
zomes are in progress (Wilder, unpubl.). 

In order to test our initial hypothesis of rela- 
tionship of the new genus to Evodianthus we se- 
lected 22 characters for the 11 genera of Cyclan- 
thaceae (Table 1), developed a character-by-taxon 
matrix (Table 2), and analyzed the data using PAUP 
for microcomputer, a maximum parsimony tech- 
nique (Swofford, 1985). All characters were un- 
weighted, but the four character states of “habit” 
were restricted to a specific order (Table 1 and 
discussion below). Since no logical basis was ap- 
parent for restricting the order of character trans- 
formations of other multistate characters, all other 
characters were treated as unordered. Characters 
with variable states for a particular genus were 
coded as missing (Pimentel & Riggins, 1987). Aut- 
apomorphies for each of the genera were not in- 
cluded in the analysis. 

Collections made in December 1988 were added 
in proof and not considered in the counts and 
measurements. 


‘TAXONOMY 


Dianthoveus cremnophilus Hammel & Wil- 
der, gen. et sp. nov. TYPE: Ecuador. Pichincha: 
Quito to Santo Domingo de los Colorados, Rio 


ABLE 1. Characters and character states for the 
genera of Cyclanthaceae. Characters found only in one 
genus (autapomorphies) are not included. 


A = Placentation: 0 = parietal, 1 = apical 

B = Fruit fusion: 0 = some, 1 = none 

C = Fruit type: 0 = other, 1 = succulent pixis 

D = Seed shape: 0 = terete, 1 = moderately flattened, 

E = Orientation of cells of testa: 0 = horizontal, 1 = 
vertical 

F = Outer layer of testa with enlarged, dark cells: 0 = 
no, 1 = yes 

G = Spinules on wall of inner layer of testa integument: 
0 = no, l = yes 

H = Inner tangential and radial walls of inner layer of 
integument equally much thickened: 0 = yes, 1 = 


no 
I = Inflorescence position: 0 = terminal, 1 = lateral 
J = Anther appendages: 0 = none, 1 = laminar-apicu- 
late, 2 = globular 
K = Phyllotaxis: 0 = distichous, 1 = spiral 
L = Glands on staminate tepals: 0 = absent, 1 = present 
M = Staminate flower symmetrical: 0 = yes, 
N = Staminate flower funnel-shaped: 0 = yes, 1 = no 
O = Pollen: 0 = aperturate, 1 = inaperturate 
P = Leaf styloids: 0 = unidirectional, 1 = omni, 2 = 
S 


— no 


Q = Birefringent cells in leaves: 0 = absent, 1 = present 
R = Anthers: 0 = not sessile, 1 = sessile (no basal bulb 
) 


or filament 
S= Habit: 
a: 1 = epiphyte 
O = terrestrial b: l = on rocks 
c: 1 = climber 


T = Root styloids: O = absent, 1 = pre 

U = Circumstelar sclerenchymatous ring; 37 — absent, 1 
— U-shaped, 2 = evenly thickened 

V = Root stele nonperipheral phloem: O = in fascicles, 


ispersed 
W= Spathe arrangement: 0 = dispersed, 1 = clustered 


Pilatón valley, 44 km E of Santo Domingo, 
on steep wet slope along highway, 0?23'S, 
78*50'W, 1,200 m, 18 June 1987 (infr), B. 
Hammel & G. Wilder 16058 (holotype, MO; 
isotypes, COL, DUKE, F, GB, NY, QCA, US). 


Evodianthi funiferi affinis sed habitu terrestre, statura 
majore, periantho floris masculini  uniseriato, tepalis acu- 
, connectivo an- 
therarum laminare api iculat toque, seminibus modice ap- 
planatis vel quasi teretis, perlongiori m latioribus 
d 


es 
Tor staminorum cupulas, inter omnes Cyclanthaceas 
peculiari 


114 Annals of the 
Missouri Botanical Garden 
TaBLE 2. Data matrix for the 10 genera of subfamily Carludovicoideae with Cyclanthus as hypothetical ancestor. 


Characters coded ? were inapplicable or variable characters. 


un 


ABCDEFGH JKLMNOPQRabcTUVW 
Cyclanthus 0?000000000?0?00000000000 
Asplundia 0012 70011011 7? 700000 0 7 01 0 0 
Carludovica 00011011001 1010000000010 I 
Dicranopygium 00000000101 ?1 1000001 001 0 1 
Evodianthus 0102 1001 001] 1001 101 001 1 21 I 
Ludovia 100000001001 01 02 10 7? 0 ? 021 0 0 
Sphaeradenia 1100? 1001201 11001010001 1 0 
Stelestylis 1010? 1001 201 1 1001.01 00 0 1 1 0 
Thoracocarpus 00120011101 100020000 1 0 2 0 0 
Dianthoveus 0 10 1 10 1 10 1 1001 1 101 0001 10 I 
Schultesiophytum 0 1 O 1 110011 110102000000 0 0 1 


Terrestrial herbaceous plant, often forming 
clumps by vegetative branching; the rhizomes 7- 
53 cm long, to about 6 cm diam. in mature spec- 
imens, often exposed and held erect by numerous 
stout, monomorphic, adventitious roots up to 79 
cm long and 0.85 cm diam. Foliage leaves of adult 
plants 145-271.5 cm long, with spiral (dispersed, 
sensu Harling) phyllotaxy, crowded on the rhizome 
at internodes mostly less than 1 cm apart, even- 
tually detaching and leaving the naked rhizome 
with well-defined leaf scars; sheaths 52-101 cm 
long, dull green and remaining intact; petioles 21- 
77 cm long; lamina unicostate or rarely and 
conspicuously subtricostate, bifid to 42 or slightly 
less, 72-107 cm long, 16.5-30 cm wide at the 
base of the median sinus, the segments 9-16 cm 
wide; each half of the lamina with 19-26 folds, 
lanceolate with a short-acuminate tip, dark satiny 
green above, dull and paler below, scabrous when 


in- 


dry. Inflorescences solitary and terminal but be- 
coming laterally displaced by the subtending, pre- 
cociously expanding renewal bud. Peduncles 25- 
34 cm long at anthesis, becoming 41-61 cm long 
and curving down in mature fruiting stage, 1. 

1.8 cm diam. Spathes of two kinds, both iue 
until very late flowering stage: 3 conspicuous 
spathes, thick, green and fleshy, differentiated into 
sheath, lamina, and sometimes petiole, 9.5-25.8 
cm long, ca. 5-6 cm wide, + clustered and inserted 
no more than 3 cm below the spadix, collectively 


surrounding the spadix, the lowermost much larger 
than the upper 2 and unusual in bearing petiolate 
and often bifid lamina; up to 6 inconspicuous 
spathes, comprising membranous greenish white 
scale leaves, up to 4.7 cm long and 1 cm wide, 
inserted above the conspicuous spathes and all + 
at one level, usually subtending flowers and often 
inserted above the base of the spadix, arising onl 

from one side of the inflorescence. Spadices cylin- 
drical, at anthesis 7.8-9.5 cm long, 2.6-3.1 cm 
diam., sordid white, in ripe fruiting stage becoming 
11-14.8 cm long, 4-6 cm diam., green. Staminate 
flowers + obpyramidal, 3.3-5.5 mm long; recep- 
tacle approximately triangular in top view, ca. 2- 
3.3 mm wide, adorned with numerous multicellular 
papillae forming a cupule around the base of each 
anther; perianth lobes 10-18, thin and narrowly 
triangular-acuminate, (1-)1.3-2.7 mm long, 
eglandular, distributed + evenly on the receptacle 
margin; stamens 19-32, the anthers + elliptic but 
often widest above the middle, 0.7-1.1 mm long, 
0.4-0.8 mm wide, the thecae occupying the lower 
,— of the anther and separated by the broad 
upper portion of connective, this terminating in a 
laminar apiculate tip; filaments minute, 0.1-0.2 
mm long or lacking; basal bulbs lacking. Pollen 
grains mostly ovoid but varying to nearly globose, 
15-21.5 um long, 14-15.5 um wide, psilate, in- 
aperturate. Pistillate flowers free to base, narrowly 
turbinate, at anthesis 3-3.5 mm long (from base 


FiGURE 1. 


and all staminodia remove 


16058. —E. Mature fruit, top view. Bar = 5 mm; Ham 
Hammel & Wilder 16067. 
16067 


Flowers and fruits of pony cremnophilus. 
mm; Hammel & Wilder 16066. 

parallel to axis of spadix, D al 1 aide saisie a to axis of spadix. Bar — 5 mm; 
] mel & Wilder 16058. —F. Staminate flower. Bar 

— 6G, H. Anthers, front view O and side view (H). Bar = 


> 


—A, B. Young Pig flowers, B with one tepal 
—C, D. Mature fruits, C viewed from side 
el 


soa 


0.5 mm; Hammel & ue 


Volume 76, Number 1 
1989 


Hammel & Wilder 
Dianthoveus 


115 


116 


Annals of the 
Missouri Botanical Garden 


km | 


FIGURE 2. 


to top of stigma), 3.8-4.1 mm wide, E ripe fruiting 
stage becoming 2.1-2.4 cm long, 1.5-1.7 cm wide 
green at maturity; tepals free to pee (partially 
connate in age), arising from the top of the inferior 
ovary, long-acuminate, the tips folding over the 
stigmas and then curving out, at anthesis 2-4.3 
2-4.3 mm wide, in fruiting stages the 
tips often breaking off, becoming acute to rounded, 
1.5-3.2 mm long, 4.5-9.5 mm wide; staminodia 
white, to ca. 9.5 cm long, ca. 1.5 mm diam., the 
distal ends apiculate, relatively flattened and at- 
tenuated but otherwise identical to an anther; styles 
at anthesis 0.9-2 mm tall, in fruiting stages l- 


E 


mm long, 


2.5 mm tall; stigmas convex in side view and slight- 
ly uncinate, extending between the tepals, at an- 
thesis 1.2-3 mm long, 0.7-2 mm wide, in mature 
fruiting stages 3.5-4.2 mm long, 2-3.7 mm wide; 
stigmatic crest seen from above linear, papillate 
and sometimes sulcate; placentae 4, parietal. Seeds 
fusiform but moderately flattened, 1.4— 

long, 0.3-0.6 mm wide, dark amber-colored; seed 
coat with tannin and mucus; embryo lacking starch. 


Figure 1 


Distribution and habitat. 
wet forest in the Andean foothills of southwestern 
Colombia and northern Ecuador. In Colombia the 
species has been found only on the Pacific slope 


Known only from 


Distribution of Dianthoveus cremnophi- 


at mid elevations, but in Ecuador it is known from 
between 150 to 1,200 m elevation and occurs on 


both sides of the Andes. 


CoLoMBiA. NARIÑO: road from El Espino 
to Tumaco, ca. 16 km W of Ricaurte, 850 m, 18 Nov. 
1986 (sterile), B. Hammel & R. Bernal 15735 (MO); 
7 Dec. 1988 ( infr), B. Hammel 17135 (COL, MO). 
ECUADOR Quevedo, 
Contulit rin 13 km E of iden Pilar, 300 m, l 
Dec. 1986 (infr), B. Hammel & J. Trainer 15846 (MO); 
Rio Palenque Science Center, 2 km S of Patricia Pilar, 
common plant on creek bank, 150-220 m, 5-14 Mar 
1977 (young infr), C. Dodson 6645 P 19 July- 11 
Aug. 1977 (old infl and young infr), C. Dodson 
6730 (SEL); 30 Nov. 1986 (infr), B. po mel & C. 
Dodson 15845 (MO); 18 Nov. 1979 (reri) Schupp 
50 (SEL). NAPO: new road to Loreto, 18 km E of junction 
with Baeza-Tena Road, 1,200 m, 21 Dec. 1988 (infr), 
B. Hammel & G. Wilder 17272 (COL, MO, QCA, . 
Baeza to Tena, ca. 58 km S of Baeza, ca. 1,200 m, 17 
Dec. 1986 (sterile), B. Hammel 15971 (MO); Tena to 
Puyo, 3 km S of Tena, 500 m (sterile), B. Hammel 
16017 (MO). PICHINCHA: Reserva ENDESA, km 113 on 
Quito to Puerto Quito road, 450 m, 16 Dec. 1988 (infr), 
B. Hammel & G. Wilder 17234 (GB, MO, PSO, QCA); 
Santo Domingo de los Colorados to Quito, Rio Pilatón 
valley, 13 km E of Santo Domingo, Tinalandia resort, on 
steep slope along stream in forest, 800 m, 30 Nov. 1986 
(old infl and young infr), B. Hammel & J. Trainer 15820 
(CAS, K, MO, US); 20 June 1987 (infl), B. Hammel & 
G. Wilder 16066 (MO); (infl and infr), B. Hammel & G. 
Wilder 16067 (COL, F, MO, QCA); 18 km E of Santo 
Domingo, on vertical slope just behind CEPE gas station, 
750 m, 22 June 1987 (inf, B. Hammel & G. Wilder 
16080 (MO, US); 32 km E of Santo Domingo, N of 
highway and Rio Pilatón, 850 m, 19 June 1987 (infl), 
B. Hammel & G. Wilder 16059 (MO); 34.5 km E of 
Santo Domingo, on steep slopes S of highway, 1,100 m, 
21 June 1987 (infl and Mew. B. Hammel Wilder 
16074 (COL, GB, GH, MO, QCA); same locality as type, 
6 "AES 1986 (infr), B. Hammel l & J. Trainer 15890 
(MO, SEL). 


Paratypes. 


Dianthoveus cremnophilus is robust, short- 
stemmed, and terrestrial (Fig. 3) but, as indicated 
by its epithet, it is a “cliff-loving” plant (Fig. 4). 
At many of the sites along the Rio Pilatón we were 
able to collect plants only by pulling ourselves up 
nearly vertical slopes, using roots and stems as 
hand holds. Several large populations, including 
the type, were growing on dripping and mossy rock 
walls. All of the sites were along small streams or 
gullies or on wet, shaded slopes in isolated patches 
of relatively undisturbed forest. Even at Río Palen- 
que, where the terrain is essentially a rolling plain, 


exposed by recent inns of forest cover and showed 
signs of sun dam 

Although Dian omie is locally abundant and 
probably formerly occurred throughout the wet 
lowlands and foothills of the Andes in northern 


Volume 76, Number 1 Hammel & Wilder 
1989 Dianthoveus 


FIGURES 3, 4. Habit and habitat of Dianthoveus cremnophilus, at type locality.— 3. Habit; divisions on tape 
are 10 cm each. — 4. Habitat; plants were often abundant on wet, vertical slopes. 


118 


Annals of the 
Missouri Botanical Garden 


Ecuador, it seems not to occur outside of that area. 
Attempts to find it have failed on the Amazonian 
side of the Andes in southern Colombia near Mocoa 
in Putumayo Province, and in the Pacific lowlands 
of southern Colombia (near Barbacoas) in habitat 
similar to that of the adjacent Ecuadorian localities. 


Phenology. Nearly all of the collections of 
Dianthoveus have been made from June through 
December, which spans the primary dry season for 
humid areas of northwestern Ecuador (Dodson & 
Gentry, 1978). Our search of numerous large pop- 
ulations at five different localities during June re- 
vealed four phenological classes: (1) most plants 
bore no outward sign of fertile structures (N — ca. 
200) except for some with old peduncle stubs; (2) 
next most abundant were plants with remnants of 
old infructescences, primarily long-rotten pedun- 
cles (N = ca. 30); (3) we saw and collected only 
12 individuals with young to mature infructes- 
cences, and (4) five individuals with immature to 
old inflorescences. At the type locality, only fruiting 
plants were observed in December of 1986 and in 
June of 1987. The single mid wet season collection 
has a young infructescence. Individuals apparently 
bear a single inflorescence per rhizome at one time. 
Our studies suggest that Dianthoveus flowers and 
fruits sporadically throughout the year. From the 
relatively frequent plants with peduncle remnants 
during June, we infer a somewhat higher flowering 
frequency during January through March, i.e., in 
the middle of the rainy season, a common pattern 
in other Cyclanthaceae. 


Development and morphology of fertile struc- 
tures. The peduncles are most flexible at the base 
and are held erect at anthesis by the tightly over- 
lapping leaf sheaths. Later, as the infructescence 
becomes heavier, the elongating peduncle bends so 
that the mature infructescence is held either hor- 
izontally or nearly pendent. The peduncle is per- 
sistent long after the fruits have been shed, and 
plants are often found with the fibrous remains of 
peduncle firmly attached to the rhizome. Both con- 
spicuous and inconspicuous spathes persist into very 
late flowering stage with the largest, lowermost 
spathe persisting longest. Spathe scars are indis- 
tinct. 

The fruits of Dianthoveus remain green or be- 
come light brown at maturity and apparently offer 
little attraction to dispersers; they are neither fra- 
grant nor very juicy. We found seeds germinating 
in the debris of an old infructescence at the base 
of a plant, and often seedlings were found on wet 
rock walls below the mother plants. Such gravity 
or "flush dispersal” is probably usual for the genus; 


however, we found several fruits on one infructes- 
cence with the distinctive prints of bat teeth. In 
Evodianthus the berries become bright pale orange 
and very juicy at maturity and are undoubtedly 
dispersed by birds or bats. 

Staminate flowers are tightly crowded among 
the pistillate flowers throughout development and 
persist into the early stages of fruiting; remains of 
the pedicels can be found even among mature 
fruits. Tepals often vary in length within one flower 
and usually with a tendency for the longest tepals 
to be together, making the flower slightly asym- 
metrical. This asymmetry, however, appears to be 
random with respect to the associated pistillate 
flower; the longest tepals, for example, may or may 
not face the pistillate flower. The tepals are thin 
and become brown at anthesis. They are always 
without resin glandules. 

Tepals of the pistillate flowers are quite similar 
in shape and texture to those of the staminate 
flowers, an unusual condition for the family. Their 
apices are thin, nearly filamentous, and at first lie 
closely appressed to the stigmas and to the stamino- 
dia. In fruiting stages the tepals become much 
broader—only acute to rounded—and the tips often 
break off, leaving an irregular ridge around the 
stigmas, which no longer protrude and are often 
turned upward rather than uncinate. The sides of 
the fruit converge basally towards the long axis of 
the spadix into a narrow, vertical, lenticular shape. 


Relationships. A simple comparison of the 
new genus with others in the family suggests that 
it is closely related to Evodianthus. Most notably, 
both genera and no others have leaves that are 
scabrous when dry. Anatomically, this translates 
into “styloid sacs [which] are normally elongated 
in all directions within the mesophyll, routinely 
crossing cell layers" (Wilder, 1985). All other Cyc- 
lanthaceae with styloid sacs have them elongated 
only parallel to the longitudinal files of parenchyma 
cells and confined to one layer. Only Evodianthus 
and Dianthoveus have styloid sacs in roots. Pollen 
of both genera is the smallest in the family and 
inaperturate (Figs. 5, 6), in contrast to the aper- 
turate pollen of all other genera. Both genera have 
very short or no filaments and lack basal bulbs. 
Both have unicostate or cryptically subtricostate 
leaves, separate fruits, symmetrical staminate flow- 
ers, and terminal inflorescences, characters un- 
usual in the family but shared with other genera 
(Table 3). 

Features of staminate flowers and seeds have 
been especially important for delimiting supra- 
specific taxa within the Cyclanthaceae (Gleason, 


Volume 76, Number 1 Hammel & Wilder 
Dianthoveus 


FIGURES 5-8.— 5. Pollen. - Dianthoveus cremnophilus. Bar — 5 um; Hammel y Wilder 16067. —6. Pollen 
of Evodianthus funifer. Bar m. — 7. Seed of Dianthoveus BA kA Bar = 
16058. —8. Receptacle of staminate flower of Dianthoveus cremnophilus. À — base of [i m B = cupule-forming 
papillae; C = point of attachment of stamen; D = point of attachment of tepal. Bar = 0.2 mm; Hammel & Wilder 
6067. 


120 


Annals of the 
Missouri Botanical Garden 


TABLE 3. Hierarchy of unusual characters of Dianthoveus and the genera sharing them. 
Character Shared with 
Papillate receptacles none 
Long-acuminate tepals none 
Inconspicuous spathes Cyclanthus 
Eglandular staminate tepals Dicranopygium subg. Gleasonianthus 
Apiculate anther appendages Schultesiophytum 
Scabrous leaves Evodianthus 
Styloids in roots Evodianthus 
Evodianthus 


Inaperturate pem grains 
Sessile anther 

Spinules on wall of seed coat 
Moderately flattened seeds 
Separate fruits 

Terminal inflorescences 
Clustered spathes 


Evodianthus 

Carludovica, Thoracocarpus 

Carludovica, Schultesiophytum 

Evodianthus, Schultesiophytum, Sphaeradenia 

Evodianthus, Carludovica, Cyclanthus 

Evodianthus, Carludovica, Schultesiophytum, Dicranopygium 


1929; Harling, 1958; Wilder, 1978). In the new 
genus, characters of both these structures differ 
markedly from those of Evodianthus, from which 
the former differs by its lack of a biseriate perianth 
(unique to Evodianthus), its lack of a funnelform 
receptacle in the staminate flowers (found in Evo- 
dianthus and several other genera), and by its only 
moderately flattened and narrowly elliptic seeds 
(Fig. 7). Evodianthus, as well as Asplundia and 
Thoracocarpus, have broadly elliptic, strongly flat- 
tened seeds. The seed coat structure also differs 
markedly from that of Evodianthus (see below). 
The new genus is further distinguished by the eglan- 
dular, acuminate tepals of the staminate flowers 
(which are similar only to those of Dicranopygium 
subgenus Gleasonianthus), the apiculate laminar 
appendages of the anthers (shared only with Schul- 
tesiophytum), and especially the peculiar papillate 
receptacle of the staminate flowers (Fig. 8). 

Seed coat structure is variable among genera of 
Cyclanthaceae and was an important consideration 
in Harling's delimitation of genera in the family. 
We have interpreted the wall layers by focusing 
on the inner and outer cuticles, described by Har- 
ling and present in all genera. The seed coat of 
Dianthoveus (Fig. 9) is similar to those of Car- 
ludovica and Thoracocarpus; the innermost layer 

of the outer integument in all three genera has 
relatively large thin-walled cells with spinulose wall 
ingrowths on the inner tangential walls, whereas 
Evodianthus lacks these spinules (Harling, 1958: 
38-43) 


Evodianthus occurs rarely as a terrestrial plant 
and rarely with leaf blades nearly as large as those 
of the new genus. With fresh material, the less 
glossy (more satiny) aspect of the upper leaf sur- 
face, the larger size, and constantly terrestrial and 


short-stemmed habit of Dianthoveus distinguish it 
vegetatively from Evodianthus. Due to the simi- 
larity of these two genera, Evodianthus specimens 
(and all other Cyclanthaceae) at COL, F, MO, QCA, 
and SEL were examined to see if the new genus 
was included among them; it was not. 


PHYLOGENETIC ANALYSIS 


The Cyclanthaceae are widely considered to be 
monophyletic (Dahlgren et al., 1985; French et 
al., 1983), and for the most part each of the 11 
genera also has at least one clear apomorphy sup- 
porting its monophyly. Within Cyclanthaceae, Cy- 
clanthus possesses many unique and highly derived 
features (Dahlgren et al., 1985; Harling, 1958; 

ider & Harris, 1982) and apparently shares 
only ancestral characters with other genera. The 
similarity of seeds of Cyclanthus to those of Pan- 


FicunE 9. 
lus. — A. 
nulose wall ingrowths. — B. 


Seed coat of Dianthoveus cremnophi- 
Outer integument; lower arrow points to head 
Outer cuticle. — C. Inner in- 


tegument. — D. Inner cuticle; Hammel & Wilder 16058. 


Volume 76, Number 1 
1989 


Hammel & Wilder 121 


Dianthoveus 


1 
Asplundia Group 


CYCLANTHUS 


pp rere ee ee ee ee @ SCHULTESIOPHYTUM 


CARLUDOVICA 
ASPLUNDIA 
THORACOCARPUS 


EVODIANTHUS 


DICRANOPYGIUM 


LUDOVIA 


SPHAERADENIA 


Sphaeradenia Group [ 


1 


A ius MM 


FiGURE 10. 


Phylogenetic scheme of Cyclanthaceae. Dashed line is Harling's conven 


on for indicating highly 
8). 


uncertain position of Schultesiophytum —staminate flowers were unknown. Adapted from Harling (195 


danaceae, especially in the penus Freycinetia, fig- 
ured strongly in Harling’s “geneological tree” 
(1958) for the genera of Ar cladthacsae (Fig. a 
including division of the family into 
Furthermore, a preliminary analysis using Frey- 
cinetia (Pandanaceae) as the outgroup resulted in 
three trees, all of which supported the Cyclan- 
thoideae/Carludovicoideae subdivision of the fam- 
ily S unpubl.). 

hus, as a starting point we accepted a sister 
group sons for the two subfamilies of Cy- 
clanthaceae and postulated hypothetical ancestor 
for the Carludovicoideae based on the presumably 
ancestral characters of Cyclanthus, the only genus 
in the Cyclanthoideae. Cyclanthus contains only 


two species and is monomorphic for the characters 
analyzed; variability within the hypothetical ances- 
tor was therefore not a problem. Characters not 
obviously comparable between the two subfamilies, 
e.g., floral symmetry (Cyclanthus lacks distinct, 
meristic floral units at maturity) were coded either 
by looking outside the family to the Araceae, Are- 
caceae, and Pandanaceae, where Harling and oth- 
ers have indicated the most likely affinity with the 
Cyclanthaceae, or by designating them as missing 
characters for Cyclanthus 

Harling's (1958) lecum relationships among 
genera in his *Asplundia group," which would 
include the new genus, by his own admission were 
quite tentative (Fig. 10). The main purpose of our 
analysis was to make an explicit statement as to 
the placement of Dianthoveus within the family, 
using characters from Harling and new characters. 
Discussion of those aspects of the character anal- 
ysis not directly pertinent to the new genus and 


comparison of the phylogeny produced with these 
methods to that of Harling will be presented else- 
where (Hammel, unpubl.). 


Character coding. Several characters were 
coded as missing in Cyclanthus, the hypothetical 
ancestor for this analysis. For example, althoug 
the pistillate **units" seem to demonstrate a re- 
markable extreme of fusion among parts, they 
readily dehisce from each other at maturity. Fur- 
thermore, it is difficult to judge an immediate an- 
cestral state for this character from outgroup com- 
parison; fruits are commonly fused in Pandanaceae, 


that whe g 

(1983) sd "iudei c fruits" as a possible syn- 
apomorphy for their Areciflorae complex (Areca- 
ceae, Cyclanthaceae, Pandanaceae), the berries in 
most species of Cyclanthaceae are dehiscent by 
apical caps (Thoracocarpus, all but one species of 
Asplundia, and most species of the Sphaeradenia 
group; Hammel, unpubl.). Live, mature infructes- 
cences of Pandanaceae, especially Freycinetia, 
need to be examined for comparison. 

Characters of the staminate flowers were like- 
wise difficult to polarize with reference to Cyclan- 
thus, which at maturity has only amerous rows of 
stamens. Pandanaceae also have amerous and ape- 
talous staminate inflorescences. Flowers were cod- 
ed as primitively symmetrical by reference to Ar- 
aceae and Arecaceae. Presence/ absence of glands 
on tepals and shape of perianth were coded as 
missing data. 

The phyllotaxy in Cyclanthus is spirodistichous 
(Wilder, 1981), making the coding of this char- 


122 


Annals of the 
Missouri Botanical Garden 


r CYCLANTHUS 


IIS 


] C DH _ N S-c 


GHI 


BDE 


” DICRANOPYGIUM 


| HH 


LUDOVIA 


FIGURE 11. Cladogram of the NARRA Wide 
half-bars are synapomorphies of character 
Le above bars refer to characters (Table 


nown autapomorphies not included in analysis 


acter problematic; genera of the Carludovicoideae 
have either orthodistichous (in a few species spi- 
rodistichous) or spiral phyllotaxy. The Pandana- 
ceae have tristichous phyllotaxy. For this analysis 
spiro- and orthodistichy were both subsumed under 
the category "'distichous," and phyllotaxy was cod- 
ed as having only two states. The terminal inflo- 
rescences of Cyclanthus seem well supported as 
ancestral; terminal inflorescences are the rule in 
Araceae (Ray, 1987 and pers. comm.) and in Pan- 
danaceae (Tomlinson & Wilder, 1984) and are 
characteristic of the “primitive” coryphoid palms. 
In palms, however, terminal inflorescences are nev- 
er evicted by axillary shoots (Tomlinson & Wilder, 
1984). States for seed characters were coded di- 
rectly from Cyclanthus as ancestral; the similarity 
of seeds of Cyclanthus, Dicranopygium, and Lu- 
dovia with those of some members of the Pan- 
danaceae is one of the main features that Harling 
(1958) saw as indicating a close relationship be- 
tween Pandanaceae and Cyclanthaceae. 

The branched order of character states for **hab- 
it" (Table 1) is rationalized by the observation that 
variable taxa almost always include terrestrial 
members. It then seems unlikely that any nonter- 
restrial form was derived one from another. 


RESULTS AND DISCUSSION 


From our data matrix (Table 2), PAUP found 


sister-group relationship between Dianthoveus and 


ba 
states found elsewhere in the family or reversed within 
1), numbers below 
rphies discovered by analysis. Length = 48; consistency index = 0.604; originally 


ASPLUNDIA 


— THORACOCARPUS 


CARLUDOVICA 


O P R T EVODIANTHUS 


DIANTHOVEUS 


SCHULTESIOPHYTUM 


SPHAERADENIA 


FH. os 


rs are uniquely derived and consistent synapomorphies; 
the defined group. 


to character states. Length of terminal clade is 


Evodianthus is supported by all of these analyses. 

urthermore, when the data matrix is changed to 
Become different possibilities for states coded 
as "missing" or different interpretations of prob- 
lematic characters for the hypothetical ancestor, 
such as fruit fusion and phyllotaxy, this relationship 
remains unchanged even when the overall topology 
changes. 

The highly corroborated sister-group relation- 
ship between Evodianthus and Dianthoveus allows 
a rational discussion of homologies of the unusual 
staminate flowers in the new genus. On the basis 
of our hypothesis (Fig. 11), Dianthoveus appears 
to retain more relatively primitive character states 
than Evodianthus, such as the terrestrial (vs. 
climbing) habit. The moderately flattened seeds are 
also relatively primitive as compared with the 
strongly flattened seeds of Evodianthus. The lam- 
inar-apiculate anther appendages of Dianthoveus 
could be symplesiomorphous with those of Schul- 
tesiophytum, but that assumption would require 
an additional step; it is more parsimonious to as- 
sume that this kind of anther evolved independently 
in the two genera. The biseriate perianth of Evo- 
dianthus—an apomorphy for Evodianthus not in- 
cluded in the analysis— appears to be derived with- 
in the family. 

From this basis we can argue that the inner 
cycle of the biseriate perianth of Evodianthus might 
be an elaboration of the unusual papillate recep- 
tacle of Dianthoveus. By this hypothesis, the lobes 
of the inner perianth of Evodianthus are not ap- 
pendicular organs (leaf homologues), but rather 


Volume 76, Number 1 
1989 


Hammel & Wilder 123 


Dianthoveus 


emergences, equivalent developmentally to the pa- 
pillae, which compose cupules in Dianthoveus. 
In all cases, regardless of coding, eglandular 
staminate tepals appear to have arisen twice within 
the Carludovicoideae—in Dicranopygium subge- 
nus Gleasonianthus and in Dianthoveus. How- 
ever, according to Harling (1958; see also Wilder, 
1978), the eglandular tepals of subgenus Gleason- 
ianthus are not “real perianths,” but rather are 
homologous with stamen filaments; he found that 
lobes of this ““false perianth" sometimes bear re- 
duced anthers. In addition, the filaments of anthers 
in subgenus Gleasonianthus are somewhat flat- 
tened and acute, resembling the lobes of the false 
perianth. In Dianthoveus we see no equivalent 
basis for hypothesizing a staminodial nature for the 
eglandular tepals. We found none with reduced 
anthers, and they certainly bear no resemblance 
to filaments of stamens (Dianthoveus lacks fila- 
ments). Based on their position, they appear simply 
to be homologous with the glandular outer whorl 
of the biseriate perianth of Evodianthus and the 
"real perianth” of other Cyclanthaceae. Under this 
reasoning, the eglandular tepals of Dianthoveus 
are similar to those of Dicranopygium subgenus 
Gleasonianthus only by convergence. 


CONCLUSION 


Before our cladogram or any other is presented 
as a phylogeny for the family, several of the char- 
acters and character states need to be re-evaluated 
and the hypothesis tested with additional charac- 
ters. For example, the character of spathe position, 
crucial for placing Dicranopygium in the clade 
definied by clustered spathes, is somewhat subjec- 
The spathes of Carludovica, Schultesio- 
phytum, and Dianthoveus are not much closer 


tive. 


together than many species outside the clade. Which 
other, if any, multistate characters (e.g., seed shape) 
might logically be ordered? One also wonders if 
characters such as inflorescence position and phyl- 
lotaxy, which seem to encompass a large suite of 
characters, should be given extra weight or broken 
into several characters. All of this needs to be 
evaluated within the context of a hypothetical 
ancestor postulated on the basis of taxa outside the 
Cyclanthaceae. 

though a close relationship between Dian- 
thoveus and Evodianthus was suggested initially 
by the observation that only these two have sca- 


brous leaves, a phylogenetic analysis using parsi- 
mony demonstrates the hypothesis as supported b 
numerous characters. It also provides a logical basis 
for suggesting the El homologies of the 
unusual floral features of t ew genus, in par- 
ticular, the papillate je (possibly homolo- 
gous with the inner perianth of Evodianthus), the 
laminar apiculate anthers (evolved in parallel to 
those of Schultesiophytum), and the eglandular 
tepals (convergent with those of Dicranopygium 
subgenus Gleasonianthus). 


LITERATURE CITED 
DAHLGREN, R. & F. N. RASMUSSEN. 1983. Mon 


yledon evolution: characters and cala A 

mation. Evol. Biol. 16: 255-395. 

, T. H. CLirFORD & P. F. Yeo. 1985. The 

Families of Monocotyledons: Structure, Evolution and 
Taxonomy. Springer-Verlag, Berlin 

Dobson, C. H. & A. H. GENTRY. 1978. The flora of 
the Río Palenque Science Center. Selbyana 9: 87- 


ERIKSSON, be Chorigyne, a new genus of the Cyclan- 
bes e from Central America. Nordic J. Bot. (in 


PE H C., K. CLANcY & P. B. TomMLINSON. 1983. 
anaes patterns in stems of the Cyclanthaceae. 

r. J. Bot d E 1400. 
Gua H or noteworthy monocoty- 
ledons from ara HMM Bull. Torrey Bot. Club 


56: 
GRAYUM, M. H. & B. E. HauMEL. 1982. Three new 
species of Cyclanthaceae from E Caribbean low 
d. Rica. Syst. Bot. 7: 
Hann G. : Monograph of id Criatori: 
qt Berg. 18: 1- 
BE. A. R. & R. Riccins. 1987. The nature of 
cladistic data. Cladistics 3: 201-209. 
Rav, T. 1987. Diversity of shoot organization in the 
Araceae. Amer. J. Bot. 74: 1373-1387. 
Sworronp, D. L. 1985. PAUP (2.4) Users Manual. 
Illinois Natural History em Champaign, Illinois. 
ToMriNsoN, P. B. & G. J. WiLD m oa Systematic 
anatomy of be ficas prenden TEA otyledoneae)—an 
overview. Bot. G E rice de 535-549. 
WILDER, C. . Two new species and a ne 
i adm of Cyclanthaceae. J. Arnold rs. 59: 74- 


= 


1981. Structure and development of Cyclan- 
thus bipartitus Poit. (Cyclanthaceae) with reference 
to other Cyclanthaceae. I. Rhizome, inflorescence, 
root, and symmetry. Bot. Gaz. (Crawfordsville) 142: 
96-114. 


1985. Anatomy of noncostal portions of lam- 
ina in the Cyclanthaceae (Monocotyledoneae). I. Epi- 
dermis. Bot. pe ee 146: 82- 
z D. Ee Is. 1982. Laticifers in t 
bip Hd "Poit. (Cyclanthaceae). Bot. Gaz 
mies e 143: 84-93. 


ANATOMIA REPRODUCTIVA 
DE LACANDONIA 
SCHISMATICA 
(LACANDONIACEAE)' 


J. Márquez-Guzmán,? M. Engleman,? 
A. Martínez-Mena,? E. Martínez* 
y C. Ramos? 


RESUMEN 


andonia schismatica es una familia, género, y 


desarrollado. El fruto es indehiscente 


especie nuevos para la ciencia. El carácter más conspicuo lo 


o. Tal disposición no había 
r 


a el micrópilo. El endospermo es de tipo nuclear. El embrión en la semilla madura es poco 


ABSTRACT 


candonia schismatica comprises a new family, genus, and species. lts most conspicuous character is the central 


position of the androecium, surrounded by the apocarpous 


as not been recorded before 


er 
endosperm 


is of the nuclear type. The embryo of the mature seed is poorly developed. The fruit is indehiscent. 


La nueva familia Lacandoniaceae del orden 
Triuridales (Martínez y Ramos, 1989) es afín a 
Triuridaceae Gardner (Maas y Rübsamen, 1980) 
pero se diferencia por el gineceo que rodea al 
androceo y la dehiscencia introrsa de las anteras. 

La inversión de posición de los órganos repro- 
ductores fue lo suficientemente interesante para 
iniciar los estudios estructurales. A continuación 
se presentan los primeros resultados al respecto. 


MATERIALES Y METODOS 


Se estudió material de dos colectas (E. Martinez 


M. 19310 y 21822) de Lacandonia schismatica 


E. Martinez y C. H. Ramos. Se fijaron muestras 
de botones florales, flores, y frutos en diversas 
etapas de desarrollo en el sitio mismo de colecta 
(selva Lacandona, Chiapas, México), y las muestras 
se trasladaron al laboratorio para su procesamiento 
posterior. 

Se usaron tres fijadores diferentes: a) FAA (for- 
malina, ácido acético, etanol al 70%; 5:5:90); b) 
glutaraldehido al 5% en amortiguador de fosfatos 

pH 6.8 con sacarosa al 0.25 M; c) glutar- 
aldehido al 5% + paraformaldehido al 4% en amor- 
tiguador de S colidina 0.1 I 

Después de la fijación algunas muestras fueron 

deshidratadas en series graduales de alcohol etilico 


! Agradecemos al Biól. Braulio Centeno, Biól. Pablo Robles, y 


la M. C. Marcela Aguilar su valiosa ayuda técnica 


? Facultad de exo en Nacional Autonoma de M Departamento de Biologia, Ciudad Divum. 


C.P. 04510 


.F., 
3 Centro ls b. Colegio de Posgraduados, C.P. 56230, Chapingo, México 
* [nstituto de Puce oe Nacional Autonoma de México, Departamento de Botánica, Ciudad Universitaria, 


C.P. 04510, D.F., Méx 


Ficuras 1-4.—1. 
posición entes] La epidermis 


EP) se continúa con 
posición externa al androceo. Tépalos (TP). y 


> 
Corte Mp e de botón floral con el pedicelo a la izquierda. Las anteras (A) ocupan la 


ejidos del receptáculo (R). Los carpelos (C) ocupan una 


los 
— 2. Corte nal de antera joven. La pared formada por epidermis 


ANN. MISSOURI Bor. GARD. 76: 124-127. 1989. 


Volume 76, Number 1 Márquez-Guzmán et al. 125 
1989 Lacandonia schismatica 


PA 1 " x = | 
ES. ae 
EN a EP m 


4 
S 


b 
nt 


C 


tu 
4 
e< 


(EP), endotecio (ET), y tapete (T).—3. Corte longitudinal de antera madura. La pared esta formada por la epidermis 
(EP) y el endotecio fibroso (ET). El septo separa los loculos (L).—4. Corte transversal de antera madura. Epidermis 
(EP), endotecio (ET), y restos del tapete. La punta de un tépalo (TP) se ha metido entre las anteras. Escalas = 100 
um. 


126 Annals of the 
Missouri Botanical Garden 


P pot 


H Ta 
E ani 


i. 


r 

FicURAS 5-10.— 5. El grano de polen (G) está formado por tres células. Escala = 10 um.—6. Corte transversal 
de antera. La flecha señala la posible zona de dehiscencia donde el endotecio (ET) queda interrumpido. Escala — 100 
um . Corte longitudinal de un carpelo. El óvulo posee dos tegumentos (TG). El interno forma el micrópilo (M). 
Saco embrionario (S). Escala = 50 um.— 8. Corte longitudinal de un carpelo. El estilo (E) está en posición subapical. 
Escala — 100 um.— 9. Corte longitudinal del fruto. El endospermo (ED) con células de paredes gruesas. Escala — 
100 um.— 10. Corte longitudinal de un proembrión (PE) de dos células. La división del cigoto ha sido transversal. 
El micrópilo (M) se dirige hacia la derecha. Escala — 50 um. 


y otras en acetona, y se incluyeron en parafina, incluídas en resinas sintéticas un ultramicrótomo 
plástico de JB4 o metacrilatos. con navajas de vidrio. Los cortes de 1 a 2 um se 

Para seccionar las muestras incluidas en parafi- tiñeron con safranina /verde rápido o con azul de 
na se utilizó un micrótomo rotatorio, y para las  toluidina. 


Volume 76, Number 1 
1989 


Márquez-Guzmán et al. 127 


Lacandonia schismatica 


RESULTADOS 


Posición de los verticilos florales. El verticilo 
más externo està constituido por seis tépalos papilo- 
sos. La posición del androceo en el centro de la 
flor con el gineceo apocárpico alrededor se muestra 
claramente en cortes longitudinales de botones flo- 
rales (Fig. 1). La continuidad de la epidermis que 
se Mi" nd la base de las anteras hasta el 
udas sobre su posición central. 


F pta J 


Androceo. 
sarrollan y maduran antes que los carpelos y per- 
sisten, sin caerse, después de que los frutos se han 
dispersado. 

El androceo està formado por tres anteras (una 
flor entre cincuenta y cuatro observadas poseia 
cuatro anteras). Las anteras son biloculares, y la 
pared de la antera inmadura está formada por tres 
capas: la epidermis, el endotecio, y el tapetum (Fig. 
2). El tapetum desaparece en la antera madura 
(Figs. 3, 4). Los estambres no poseen vasculari- 
zación. El grano de polen maduro posee tres células 
(Fig. 5). La dehiscencia de las anteras es introrsa 
(Fig. 6 


Se observó que las anteras se de- 


Gineceo. El gineceo es apocárpico. La ori- 
entación de los carpelos es variable, de tal manera 
que la mayoria de ellos no tiene el estilo en el lado 
orientado hacia los estambres, en el centro del 
receptáculo. El estilo parte de la región superior 
del carpelo en el mismo lado que corresponde al 
hilo de la semilla (Fig. 8). 

Cada carpelo contiene un solo óvulo sésil, basal, 
anátropo (Fig. 7), y bitégmico; ambos tegumentos 
son biestratificados, y el interno forma el micrópilo. 
El desarrollo del endospermo es de tipo nuclear, y 
el endospermo maduro posee células de paredes 
gruesas (Fig. 9). El embrión más desarrollado que 
se ha observado posee dos células; el cigoto se 
dividió transversalmente (Fig. La cubierta 
seminal tiene una capa con inclusiones en forma 
de partículas. 


El fruto es indehiscente. La epidermis del fruto 
posee células globosas cuando está en antesis. 


DISCUSION 


Las estructuras reproductoras de Lacandonia 
schismatica son comparables a las mencionadas 
por Tomlinson (1982) y por Maas y Rübsamen 
1986) para la familia Triuridaceae. 

La dehiscencia introrsa de las anteras es una 
caracteristica que define esta especie. Sin embargo, 


— 


el hecho más conspicuo, y nunca antes mencionado 
para Triuridaceae u otras angiospermas, es la posi- 
ción central del androceo con respecto al gineceo 
(Cronquist, 1981). Esta disposición se observa más 
claramente en cortes medianos de botones florales 
inmaduros 

La continuidad celular, en especial la epidér- 
mica, que se observa con los tejidos del centro del 
receptáculo es la prueba más fehaciente de su 
posicion. Sin embargo muchos detalles de la for- 
mación del óvulo y la semilla se desconocen y aun 
no se sabe que tipo de embrión representan. La 
germinación no ha sido observada. Se pone de 
manifiesto la necesidad de realizar estudios em- 
briológicos, además de estudios anatómicos com- 
parativos de especies de la familia Triuridaceae. 
Tales resultados contribuirán a ubicar filogenéti- 
camente a esta familia y dar luz sobre el Pap d 
evolutivo de la inversión en la posición de los ór 
ganos reproductores. 


LITERATURA CITADA 


orae A. 1981. An Integrated System of Classi- 
on of Flowering Plants. Columbia Univ. Press, 
Ne w o: 
Maas, P. J. M. & T. E JBSAMEN. 
Fl. Neotrop. 40: 1-55. 
MARTÍNEZ, E. & C. H. Ramos. 1989. Lacandoniaceae 
(Triuridales una nueva Hen i México. Ann. 
Missouri Bot. Gard. 76: (this i 

TOMLINSON, P. B. Helobiae a EN In: C 
R. Metcalfe (editor), Anatomy of the Monocotyledons 
VII. Clarendon Press, Oxfor A 


1986. Triuridaceae. 


LACANDONIACEAE 
(TRIURIDALES): 

UNA NUEVA FAMILIA 
DE MEXICO! 


Esteban Martínez? 


y Clara Hilda Ramos? 


RESUMEN 


Se describen una familia, un género y una especie de Triuridales nuevos para la ciencia. La nueva familia es afin 


a Triuridaceae Gardner, de la cu 


al se diferencia por el gineceo rodeando al androceo; posición insólita me se describe 


por primera vez, y la dehiscencia introrsa de las anteras. Este es el primer registro del orden para Méx 


ABSTRACT 


A new family, genus, and species are described. The new family is placed in Triuridales. It differs from Triuridaceae 
the 


y the unexpected an 


unknown elsewhere position of the gynoecium, bli 


surrounds the androecium, and by 


band dehiscent anthers. This is the first record of the order for Mex 


Entre las plantas colectadas para el proyecto 
Flora Mesoamericana (CONACyT PCECBEU- 
001109), se encontraron plantas del orden Triuri- 
dales que consideramos son familia, género, y es- 
pecie nuevos para la ciencia y se describen a con- 
tinuación: 


Lacandoniaceae E. Martinez et C. H. Ramos, 
fam. nov. ordinis Triuridalium. 


Flores hermaphroditi gynoecio apocarpo androecium 
cingenti, tribus staminibus centralibus antheris bilocula- 
ribus dehiscentia introrsa, apertura singularis 


Flores hermafroditas; gineceo apocárpico ro- 
deando al androceo, tres estambres centrales y 
anteras biloculares con una sola linea de dehiscen- 


cia introrsa. 
Lacandonia E. Martínez et C. H. Ramos, gen. 
nov. 
Herbae saprophyticae, albidae, aphyllae. Caules sim- 


plices. Rhizoma glabrum horizontale radicibus pubescen- 
tibus. oras racemosa. Flores hermaphroditi. Te- 


pala (4-)6, apicibus caudatis, dense papillosa. Gynoecium 
apocarpum, carpellis 60- js Styli subapicales. Stamina 
(2-)3(-4), centralia, marce i 
antherae bilocularia e lobis 2 
longitudinaliter dehiscentibus. Fructus aggregatus e multis 
acheniis constatus. 


Hierbas saprofiticas, afilas, blanquecinas. Ri- 
zoma horizontal de crecimiento indefinido, con ra- 
ices fibrosas y esparcidamente pubescentes. Tallos 
aéreos simples, glabros. Inflorescencia racemosa, 
bracteada, de crecimiento simpodial; pedicelos as- 
cendentes. Flores perfectas; tépalos (4-)6, den- 
samente papilosos cuando frescos, cada uno con 
una cauda linear-laminar en el ápice. Gineceo y 
androceo sobre un receptáculo deprimido central- 
mente, con los carpelos rodeando al androceo. Ci- 
0-80 carpelos papilosos; estilo sub- 
apical. Estambres (2-)3(-4), centrales, persistentes, 
insertos en el borde de la depresión del receptáculo; 
anteras biloculares con 2(-3) lóbulos, introrsas, una 
sola linea de dehiscencia longitudinal, polen intec- 
tado, inaperturado, verrugado. Fruto poliaquenio. 


radecemos a F. Chiang C. la ayuda solicita que nos brindó durante el desarrollo F este trabajo, asi como la 


Agr 
nea critica del manuscrito y las descripciones latinas; a G. Davidse y a M. Sousa 


momento; a E. Martinez H. bul p de polen; a M. 


senor, T. We 


70-367, 04510 México, D.F. Mexi 


Veith po 
edina la elaboración del dibujo y 
la mecanografia del texto; al S. Cabrie ] Aguilar su inapreciable ayuda en el bo de campo; a . Gon 
i Villa ndt, y R. Weber sus valiosas sugerencias; a G. An a B. Hammel, 


dg en todo 


de ding 


apoyo 
r producir las os de SEM 
sus valiosas sugerencias; a M. 

zález Medrano, 


afi 
Biologia, Universidad Nacional Autónoma de México, Departamento de Botánica, Apartado Postal 


* Facultad de Ciencias, Universidad Nacional Autónoma de México, Departamento de Biologia, Ciudad Universitaria, 
Méx 


C.P. 04510 México, D.F. 


ANN. MISSOURI Bor. GARD. 76: 128-135. 1989. 


Volume 76, Number 1 Martínez & Ramos 129 
1989 


Lacandoniaceae 


0.2 mm 
4) 
Su 
c 
0.25mm 
bw eq 
AN 
P a 
> 
yw. 
HAS 
y 
DY 
c AÌ 
A A Y ES 
ANA b. 
e NM 
Smm | M 
0.2mm | oy 
vi 


FIGURA —a. Hábito. — b. Flor, pedicelo, y bráctea.—c. Flor vieja mostrando los estambres persistentes sobre el 


disco. —d. pss en vista "m udi ui (abierto con 2 loculos y un septo); ventral, mostrando la línea de 
dehiscencia. —e. Ovario. —f. Fru g. Semilla. 


130 


Annals of the 
Missouri Botanical Garden 


FicuRAS 2-5.— 2. Botón floral con los tépalos en proceso de antesis. — 3. Bot 
desarrollo. i kde recién abierta en la cual se distinguen tres estambres centrales rodeados por los ovarios inmaduros. 


o de desenrrollamiento de las caud 


Se ve el p 


as.— 95. 


ones florales en dde etapas de 


Flor abierta con las caudas de los tépalos erectas y los 


ovarios tug das que ocultan los estambres. Fotografías de Alejandro Martinez Mena. 


Semilla elipsoide con ornamentación reticulada, en- 
grosada distalmente y con una corta proyección 


basal. 


Lacandonia schismatica E. Martinez et C. H. 
Ramos, sp. nov. TIPO: México. Chiapas: Mpio. 
Ocosingo, Crucero Corozal, 170 km al SE de 


Palenque, camino a Boca Lacantum, sobre la 


carretera fronteriza del sur, 30 enero 1987, 
E. Martínez S. 19310 (holotipo, MEX U; isoti- 
pos, CR, ENCB, K, LE, MO). Figuras 1-12. 


Herbae tenues, usque ad 9 cm. Inflorescentia usque 
ad 4 cm longa, bracteis ie ca. 0.6-1.0 mm longis 
et 0.4-0.8 mm latis. Flores (1-)3-7(-13), pedicelli 3- 
4(- 7) mm longi; tepala 6, deltata, ca. 1 mm longa et lata; 


Volume 76, Number 1 Martínez & Ramos 
1989 Lacandoniaceae 


FicuRAS 6-8.— 6. Flor con los fruticulos bien desarrollados y las anteras centrales (material fijado). — 7. Flor vieja 
* ein. los estambres marcescentes.— 8. Rizoma con raíces saliendo de dos nudos. Fotografias de Alejandro 


Martinez Mena 


132 Annals of the 
Missouri Botanical Garden 


8888 10609.0U 1.INC 


FIGURAS 9, 10.— 9. Vista posterior de la flor en la que se observan los haces vasculares que divergen del pedicelo 
hacia lo Edi A bad se distinguen los frutículos (material fijado). Fotografía de Alejandro Martinez Mena. — 
Vista de la flor. Se observan la apariencia papilada y los tres estambres centrales rodeados por los ovarios. 


Fotografí a de Ricardo Peralta y Fabi. 


Volume 76, Number 1 Martínez & Ramos 133 
1989 Lacandoniaceae 


A -—. (ee ài 
d aM d "dt 
Ficuras 11, 12.— 11. Apice de un tépalo con la cauda erecta. Se distinguen las papilas. — 12. Fruticulo. Se ven 
las papilas y el estilo persistente. Fotografias de Ricardo Peralta y Fabi. 


cauda 1(-2) mm longa et 0.1 mm lata. Gynoecium car- Hierbas hasta de 9 cm de alto. Rizoma glabro, 

pellis obovoideis, dense papillosis. ja 3, filamentis 0.2-0.3 mm de diámetro, con raices filiformes, 1.0— 
| mm longis, antheris circa 0. ongis, 0.1 mm 3.5 i de l 0.1 de dibisiet Tall 

latis. Achenia obovata, 0.45-0.6 mm a 0.28-0.35 «2 om de largo, ca ¿ul PL O0 OECD MM 


mm lata, dense papilloso. rollizos, usualmente 3-5 cm de largo, 0.4-0.5 mm 


134 


Annals of th 
Missouri Botanical Garden 


de diámetro. Inflorescencia hasta de 4 cm de 
largo con brácteas trilobadas, 0.6-1.0 mm de 
largo, 0.4-0.8 mm de ancho en la base, subam- 


(1-)3-7(-13), en espiral, 3.8-4.1 
metro; tépalos 6, deltoides, fusionados desde la base 
hasta la parte media, ca. 1 mm de largo, 0.9-1.1 
mm de ancho, las caudas ca. 1(-2) mm de largo 
y 0.1 mm de ancho. Carpelos obovoides, 0.48- 

.6 mm de largo, 0.2-0.25 mm de ancho; estilo 
papilado 0.2-0.25 mm de largo; estigma de color 
oscuro, ca. 0.04 mm de largo y ancho. Estambres 
3, con los filamentos de 0.1 mm de largo; anteras 
sub-basifijas, ca. 0.3 mm de largo y 0.1 mm de 
ancho. Aquenios obovoides 0.45—0.6 mm de largo, 
0.28-0.35 mm de ancho, con el estilo persistente, 
papilosos, ligeramente esponjosos; semilla 0.4—0.5 
mm de largo, 0.2-0.25 mm de ancho, pardo os- 
cura. 

Distribución y hábitat. Sólo conocida de la 
localidad tipo, en selva alta subperennifolia inunda- 
ble de Calophyllum brasiliense Camb. con Ter- 
minalia amazonia (J. Gmelin) Exell y Cryosophila 
argentea Bartlett. Muchos de los elementos de esa 
zona son de nuevo registro para la región, como 
Chionanthus domingensis Lam., Guettarda ti- 
kalana Lundell, Schizaea poeppigiana Sturm, 
Actinostachys germanii Fée, y Guzmania lin- 
gulata (L.) Mez var. minor (Mez) L. B. Smith & 
Pittend, entre otros. Vive sobre substratos de ma- 
teria organica muy humedos, pero no inundados, 
en lugares muy sombreados con poca oscilación de 
temperatura. La localidad tipo se sitúa en el ex- 
terior del borde de la reserva de Montes Azules, 
área que se talará para crear potreros. 

Fenología. Florece todo el año si las condi- 
ciones de humedad son adecuadas. Muy abundante 
en noviembre y diciembre, después de la temporada 
de lluvias. 


Material adicional examinado. MEXICO. CHIAPAS: 
Mpio. Ocosingo, Crucero Coral 170 km al SE de Palen 
que, camino a Boca Lacantum, sobre la carretera foi 
teriza del sur, 15 Sept. 1985, E. Martínez S. 13996 
(MEXU); 6 Nov. 1985, Martínez 14982 (MEXU); 29 
mayo 1987, Martínez 21381 (MEXU); 27 Ago. 1987, 
pria 21822 (MEXU); 10 Dic. 1987, Martínez 
21921 (MEXU). 


Lacandoniaceae comparte con Triuridaceae los 
siguientes caracteres: gineceo apocárpico, óvulos 


anátropos, polen inaperturado, flores trímeras, té- 
palos con apéndices, hábito saprofítico, y micorri- 
zas endotróficas. Sin embargo, difiere por: a) La 
disposición espacial de los órganos reproductores, 
con los estambres ocupando la parte central de la 
flor. Tal posición no habia sido contemplada antes 
para angiospermas (A. P. de Candolle, 1813; 
Hutchinson, 1959; Takhtajan, 1969; Cronquist, 
1981). b) La dehiscencia introrsa de las anteras, 
extrorsa en Triuridaceae (Tomlinson, 1982; Maas 
& Rübsamen, 1986). 

El análisis cromatográfico comparativo entre 
Triuris sp. y Lacandonia schismatica indica cla- 
ras diferencias en el tipo de metabolitos secundarios 

elgado, inéd.). 

Cronquist (1981) colocó en el orden Triuridales 
a las familias Petrosaviaceae Hutchinson y Triuri- 
daceae Gardner. Petrosaviaceae comparte con La- 
candoniaceae la dehiscencia introrsa de las anteras, 
el hábito herbáceo, y la micotrofia, pero difiere 
por presentar estomas, tépalos en dos series, car- 


— 


pelos fusionados basalmente con muchos óvulos, 
nectarios septales, y granos de polen monosulcados. 

Dahlgren et al. (1985) consideraron en Triuri- 
dales únicamente a Triuridaceae. 

La relación espacial que presentan los órganos 
reproductores es difícil de explicar a la luz de los 
conocimientos actuales. En ausencia de informa- 
ción previa, se están llevando a cabo estudios anató- 
micos (Márquez-Guzmán et al., este tomo) y fito- 
químicos que ayuden a esclarecer su posición 
taxonómica. 

El nombre genérico está dedicado a la región 
Lacandona, asiento del pueblo Lacandón, en el 
estado de Chiapas, México. El epíteto especifico se 
refiere a los problemas que origina su morfología 
en la sistemática. 


LITERATURA CITADA 


CANDOLLE, A. P. 1 peca Théorie Elémentaire de la 
otanique ou Expo on des Puis ua de la Classi- 
fication Naturelle. Déterville, 
CRONQUIST, A. 1. An Int eae pre of Classi- 
cation E Flowering Plants. Columbia Univ. Press, 


DAHLGREN, R. M. T., H. T. CLIFFORD & P. F. YEo. 1985. 
The Families of the Monocotyledons. c Evo- 
lution, and Taxonomy. Springer -vorhig Be rlin 


HUTCHINSON, J . 1959. The Families of Flowering Plants, 
Volume 1: 23. Clarendon Press, Oxford. 

Maas, P. . & T. RÜBSAMEN. 1986. Triuridaceae. 
Fl. Neotrop. 40: 1-55. 


Volume 76, Number 1 
1989 


Martínez & Ramos 135 


Lacandoniaceae 


MARQUEZ-GuzMAN, J., M. ENGLEMAN, A. MARTÍ. 
NEZ-MENA, E. MARTÍNEZ & C. Ramos. 1989. Ana- 
tomia reproductiva de Lacandonia schismatica (La- 
candoniaceae). Ann. Missouri Bot. Gard. 76: 124- 
127. 


TAKHT eal A. 1969. Flowering Plants. Origin and 
Dispersal. Smithsonian Inst. Press, Washington, D.C. 

NR P. B. 82. Helobiae CM rna tide). In: € 
R. Metcalfe (editor), Anatomy of the Monocotyledons 
VII. Clarendon Press, Oxford. 


SYSTEMATICS OF THE 
AMAZON LILIES, EUCHARIS 
AND CALIPHRURIA 
(AMARYLLIDACEAE) 


Alan W. Meerow? 


ABSTRACT 


Eucharis and Caliphruria are closely related, neotropical genera of petiolate- a white-flowered eam mia 


found in the understory of primary tropical rain forest. 
and Caliphruria form a monophyletic group within * 


Together wit 
"infrafamily Pancratiidina” 
+) 


th the Per 


vian endemic Urceolina, Euchar 
on the basis of leaf and de 
di | H A 


| hy 


1001 pholog 
in Euchar 


'eolina a 
He Miss is included. 


s. Eucharis, marked by its crateriform or campanulate flowers, eae per 
usually dod into a staminal cup, and unicellular stigmatic papillae, is distributed fro 
es of the eastern Andes. Subgenus Heterocharis represents 


An alternative hypothesis of generic relationships is tested, and 


rianth tube, pigmented androecium 
m Guatemala to Bolivia, 


ves less par 


Recognizing the close phenetic relationship of subg. Heterocharis to the rest of Eucharis, the high levels of al si 
within the cladogram, and ambiguity caused by missing character state data, a less parsimonious phylogeny is accepted 
as the basis for classification. Keys and descriptions are provided for all species of Eucharis and Caliphruria. Eucharis 
caucana, a novel hexaploid (2n — 138) species, is described from Colombia. 


The closely related genera Eucharis Planchon 
& Linden and Caliphruria Herbert (Amaryllida- 
ceae), the Amazon lilies, comprise respectively 17 
and 4 species of bulbous, rain forest geophytes 
adapted to the low light conditions of the forest 
understory. Together with the Peruvian endemic 
Urceolina Reichenb., nom. cons., Eucharis and 
Caliphruria form a monophyletic group delimited 
by petiolate leaves with distinctive cuticular stria- 
tion; a turgid seed with a lustrous, usually black 
testa; and complete fidelity to the rain forest under- 


story niche. The species are distributed from Gua- 
temala to Bolivia. The major center of distribution 
for Eucharis is in the western Amazon basin (in- 
clusive of major tributary systems, e.g., the Napo, 
Pastaza, and Huallaga) and the adjoining lower 
slopes of the eastern Andean cordillera. With the 
exception of single Peruvian species, Caliphruria 
is restricted to the Cordillera Occidental and Cor- 
dillera Central of Colombia. The species of both 
genera are nowhere abundant, and are found grow- 
ing only in primary, rarely secondary, forest from 


! I thank the curators of AAU, B, BM, COL, CUVC, F, FTG, G, GB, GH, GOET, HUNT, K, LE, M, MG, MICH, 
MO, MPU, NA, NY, OXF, P, QCA, S, SEL, SP, U, UC, US, and VEN for loans of herbarium specimens, and the 


following individuals or institutions who gen 


Phil Silverstone- -Sopkin, James Watson, Mark W 
Hortorium, Royal Botanic Gardens at Kew and Edin 


erously provided living material: James 
Dodson, Robert Dressler, Mark Elliot, Thomas Fennell, Fred 
hitten, MA Williams, = bci Marcia Wilson, Bailey 
urgh, Longwo 


Bauml, Libby Besse, ME 
uchs, Harry Luther, Fred Meyer, Timot y Plowma 


ood Garden eley Botanical Garden, 


Honolulu Botanical Gardens, Huntington Botanical Garden. Fairchild Tropical Garden, Marie Selby Bo an a -— 


Missouri Botanical Garden, and Strybing Arboretum. Bijan 
Judd, Charles 
John Kress, and Christiane o Lai helpful criticisms to earlier versions 

cies. Portions of this 


of this work was completed and, along with Walter 


Meerow illustrated a number of the 

Improvement grant BSR- a EA a ie Club 

This paper represents part o h.D. diss 

Florida Agricultural Experime 
* University of Florida, Institut 


x Station Journal Se 
e of "Foo d 


ehgan provided the Pied environmen much 
Guy, Norris Williams, R. Mitche i Beauchamp. 
a Fisher- 
ork were su Doctoral Discos 


ported 


of een World Wildlife Fund Fellowship in Tropical Botany. 
ertation submitt 
eries No. 
and Agricultural Sciences, Research & Education Center, 


a to the graduate school of the University of Florida. 
7795. 


3205 SW 


ege Avenue, Fort Lauderdale, "Florida 33314; and Fairchild Tropical Garden, 11935 Old Cutler Road, Miami, 
U.S.A. 


Colle 
Florida 33156, 


ANN. Missouni Bor. GARD. 76: 136-220. 1989. 


Volume 76, Number 1 
1989 


Meerow 137 
Eucharis and Caliphruria 


ca. 50 to 1,800 (rarely to 2,000) m elevation on 
soils of high fertility. Large-scale deforestation has 
proven catastrophic to these plants, which are un- 
able to adapt to the higher light intensity of the 
clearings and soon perish. At least several species 
are probably extinct, and this number will only 
increase with time as rain forest destruction con- 
tinues unabated. 

The Amazon lilies are marked by their ever- 
green, petiolate leaves; white, often pendent, some- 
times fragrant flowers with a frequently conspic- 
uous staminal cup or false corona formed by the 
basal connation of the staminal filaments; obtusely 
trilobed stigma; and large, turgid, globose or ellip- 

soid seeds with a black, brown, or metallic blue 
testa. Eucharis amazonica Linden ex Planchon is 
widely known in horticulture, often erroneously as 
E. grandiflora Planchon & Linden (Meerow & 
Dehgan, 1984a). Neither genus has ever been crit- 
ically treated in the taxonomic literature. Baker 
(1888) provided a key and descriptions for all 
species known at the time in his Handbook of the 
Amaryllideae, and Macbride (1936) treated the 
known Peruvian species of Eucharis for the Flora 
of Peru. Though species of Eucharis have contin- 
ued to be described well into the present decade, 
the delimitation of these species from previously 
described taxa has consistently remained vague. 
No assessments of variation at either the population 
or species level have been attempted. 

As two of only three genera of neotropical Ama- 
ryllidaceae (the other being the related Urceolina) 
exhibiting complete fidelity to a primary rain forest 
niche, systematic understanding of Eucharis and 
Caliphruria offers important information relating 
to the evolution of the pancratioid Amaryllidaceae 
(“infrafamily Pancratioidinae" Traub (1957, 
1963)),* centered in the central Andean region of 
South America. 

No taxonomic scheme can encompass all of the 
information about the group of organisms under 
study (Holsinger, 1984, 1985; Ornduff, 1969; Ra- 
ven, 1 . In genera such as Eucharis and Cal- 
iphruria, rare in the wild and often accessible only 
with difficulty, and which often exhibit cryptic pat- 
terns of variation, this observation becomes much 
more acute. By accumulating data from as many 
diverse sources as possible, I have attempted to 
construct an evolutionary systematic classification 
of the Amazon lilies that is both practical yet re- 
flects their evolutionary relationships as accurately 
as these data allow. 


MATERIALS AND METHODS 
FIELD STUDIES 


Fieldwork was conducted in Peru and Ecuador 
in July-August 1982, and in Colombia and Ec- 
uador in July-August 1984. Herbarium specimens 
and living material were collected, and ecological 
observations of natural populations were made. 


ANATOMICAL STUDIES AND 
SCANNING ELECTRON MICROSCOPY (SEM) 


Material was prepared and examined as de- 


scribed in Meerow (1987c) 


CLADISTIC ANALYSES 


Cladistic analyses were run using PAUP version 
2.4.1 by David L. Swofford (Illinois Natural History 
Survey). All species (and two subspecies) of Fu- 
charis and Caliphruria (22 evolutionary units, 
EUs) excluding hybrid taxa were analyzed using 
forty-one characters (Tables 1—3). For analyses 
with PAUP, global branch swapping with the MUL- 
PARS option was used in all dep for the most 
parsimonious cladograms. Because of the unavail- 
ability of material, all desc n. species of Urceo- 
lina could not be included in the analysis. Urceo- 
lina was therefore entered in the analysis at the 
generic level, with character state data drawn from 
study of living material of U. microcrater Krànzlin 
and specimens of U. urceolata (Ruiz & Pavón) 
Green and U. ayacucensis Ravenna. Character 
states were determined from examination of living 
and dried material. Character state polarities were 
assessed via outgroup comparison (Maddison et al., 
1984; Watrous & Wheeler, 1981). Selection of 


outgroups is discussed later with the results of the 


analyses 
Of ihe 41 characters used in the analysis, 23 
were simple, two-state characters. The 18 remain- 


ing multistate characters (14 three-state, 2 four- 
state, | five-state, and 1 six-state) were treated in 
preliminary analyses as unordered (Fitch, 1971; 
Hartigan, 1973) using this option of PAUP, rather 
than attempting to create any a priori transfor- 
mation series. This resulted in a preponderance of 
weak monophyletic groups based on single apo- 
morphies and/or character state reversals, as well 
as unresolved polytomies among many terminal 
taxa. In the analyses presented, 11 of these char- 
acters were treated as ordered transformation se- 
ries (the remaining seven as unordered), with one 


* This unofficial rank is used in this paper as a label of convenience, since there is as yet insufficient evidence to 
ly. 


recognize this group of genera as a distinct subfami 


138 


Annals of the 
Missouri Botanical Garden 


(number 41, chromosome number) coded in ad- 
ditive binary. Justification for these treatments is 
discussed with the results of the analyses. Multistate 
characters were adjusted to equal weight with two- 
state characters using the "weights scale" option 
of PAUP. Additionally, an alternative hypothesis 
of generic relationship was tested by submitting a 
user-coded tree topology to PAUP for analysis. 


HERBARIUM STUDY AND LIVING MATERIAL 


Loans or gifts of herbarium specimens were re- 
ceived from the following herbaria: AAU, B, BM, 
COL, CUVC, F, FTG, G, GB, GH, GOET, HUNT, 
K, LE, M, MG, MICH, MO, MPU, NA, NY, OXF, 
P, QCA, S, SEL, SP, U, UC, US, VEN. 

Neither Eucharis nor Caliphruria is well rep- 
resented in herbarium collections, and critical mor- 
phological characters are often obscured by the 
drying process. Consequently, a living collection of 
over 100 accessions representing fifteen species 
and several hybrids was accumulated from botan- 
ical gardens, individuals, and field collections by 
myself and various colleagues. Vouchers for living 
material are listed with specimens cited for each 
species. 


TAXONOMIC HISTORY 


Herbert (1844) described the new genus Cali- 
phruria from collections made in New Granada 
(Colombia) near Guaduas by Hartweg, placing Cal- 
iphruria in the “section” Pancratiformes of ““sub- 


united by the single character of staminal conna- 
tion. The small, white, funnelform flowers were 
marked by the presence of a bristle or slender tooth 
at either side of the filament. Herbert (1844) made 
no mention of basal connation of the filaments. The 
orthography of the name, a single "I" in the Greek 
stem *'calli-," was emended by subsequent workers 
(e.g., Baker, 1877), but was returned to Herbert's 
original by Meerow & Dehgan (1984b). 

Planchon & Linden (1852) introduced another 
new pancratioid genus, Eucharis. The first species 
described, E. candida Planchon € Linden, col- 
lected in New Granada by M. Schlim, was char- 
acterized by its crateriform flowers with a con- 
spicuous staminal cup and a widely spreading 
perianth limb. If these authors were aware of Cal- 
iphruria, they did not note any relationship be- 
tween it and Eucharis. 

Bentham & Hooker (1883) placed Caliphruria 
and Eucharis in the tribe Cyathiferae. Caliphru- 


ria subedentata Baker was combined with Eucha- 


ris in their treatment, while C. hartwegiana Herb. 
was retained. Baker (1888) accepted this treat- 
ment, yet described another new species as C. 
tenera. He described Caliphruria as "nearly allied 
to Eucharis.” 

Baillon (1894) described the species C. castel- 
naeana and declared it precisely intermediate be- 
tween Caliphruria and Eucharis. He further sug- 
gested that Eucharis should be treated as a section 
of Caliphruria. Macbride (1931) later transferred 
C. castelnaeana to Eucharis. 

Nicholson (1884) transferred C. hartwegiana 
(the type species of Caliphruria) to Eucharis, 
ignoring the former's nomenclatural priority. Traub 
(1967) made the formal transfer of the remaining 
species of Caliphruria, C. tenera Baker, to Fu- 
charis and combined the monotypic genus Pla- 
giolirion Baker with Eucharis. He had previously 
listed Caliphruria, Plagiolirion, and Mathieua 
Klotzsch (another monotypic genus) as synonyms 
for Eucharis in his Genera of the Amaryllidaceae, 
citing Baillon (1894) as a “special reference" 
(Traub, 1963: 74). The nomenclatural priority of 
Caliphruria Herbert was overlooked. No proposal 
for the conservation of Eucharis Planchon & Lin- 
den over Caliphruria Herbert has ever been pro- 
posed previous to that of Meerow & Dehgan 


Traub (1971) later combined Eucharis with Ur- 
ceolina, a small Andean genus with petiolate leaves 
and markedly urceolate, brightly colored flowers. 
He designated five subgenera: Urceolina, Eucha- 
ris, Caliphruria, Mathieua, and Plagiolirion. 
Traub (1971) offered no explanation for the com- 
bination of Eucharis and Urceolina, but presum- 
ably his decision was prompted in part by reports 
in the literature of two intergeneric hybrids between 
Eucharis and Urceolina: x Urceocharis clibranii 
Masters, an artificial hybrid, and x U. edentata C. 
H. Wright, putatively discovered in Peru. Plagi- 
olirion and Mathieua are monotypic genera allied 
to Hymenocallis Salisb. and Stenomesson Herbert 
respectively (Meerow, 19872). 


VEGETATIVE MORPHOLOGY 
BULBS 


The globose or subglobose bulbs of Eucharis 
and Caliphruria, composed of concentric and 
modified leaf bases (scales), are characteristic of 
most members of the Amaryllidaceae. The bulbs 
are sympodial in growth habit (Arroyo, 1984). The 
outer scales of Kucharis and Caliphruria bulbs 
are modified into an either brown or tan paper 
tunic. The bulb scales of most species of Eucharis 


Volume 76, Number 1 
1989 


Meerow 139 
Eucharis and Caliphruria 


FIGURE 1. 
SEL). — 


C. subedentata (Meerow 1156, —E. E. mo 


—B. E. bouchei var. dressleri (Meerow 1108, a 


. amazonica (Schunke 
les. 


D 
14179, FLAS). p = proximal, m = ela. d = distal. pebeta indicate secondary vascular bundles 


and Caliphruria are apically articulated into a 
neck or pseudostem of variable length. Distally, 
the pseudostem grades into the petiole of individual 
leaves. The neck of Eucharis and Caliphruria 
bulbs always remains below the soil surface. Length 
of the neck may therefore be more a factor of bulb 
depth in the substrate rather than of any phylo- 
genetic pd Bulbs of most species of Eu- 
charis Caliphruria offset regularly and vig- 
4 Me forming sizable clumps in time i 
undisturbed. 


LEAVES 


Leaves of Eucharis and Caliphruria are uni- 
formly long-petiolate, with a well-developed elliptic, 
ovate, or lanceolate lamina. Petiolate leaves are 
characteristic of a number of genera of Amaryl- 
lidaceae, either completely or in part, and have 
undoubtedly evolved independently several times 
from the linear or lorate leaf morphology typical 
of the family. 

The petiole of the leaf of Eucharis and Cali- 


phruria is usually as long or longer than the lamina. 


140 


Annals of the 
Missouri Botanical Garden 


It tends toward subterete cross sections (Fig. 1), 
rounded abaxially, and flattened adaxially, becom- 
ing slightly channeled proximal to the sinus. The 
petiole is winged proximal to the sinus by atten- 
uation of the lamina. The midrib is pronounced 
abaxially along the entire length of the lamina, and 
slightly channeled adaxially, continuous with the 
petiole. 

Leaf shape only rarely provides taxonomically 
useful information. Length / width ratios are subject 
to considerable variation even among the leaves of 
a single bulb. Herbarium specimens will frequently 
include only a single leaf, with no indication of its 
developmental age. Taxonomic consistency of leaf 
shape is exceptional, but useful in the few cases 
where it occurs. For example, leaves of E. ulei 
Kranzlin are consistently narrowly elliptic. Eucha- 
ris amazonica (leaf length / width ratio greater than 
2) may be delimited from E. moorei (Baker) Mee- 
row (1/w less than 2). 

The leaves of Eucharis and Caliphruria are 
completely glabrous and nonglaucous, with a single 
exception. Eucharis bonplandii (Kunth) Traub, a 
rare tetraploid species from central Colombia, de- 
velops a glaucous bloom that gives the leaf a blue 
cast in strong light. 

The leaf apex of all species is shortly acuminate, 
the base attenuate. Coarse undulation of the margin 
will sometimes make the lamina appear cordate at 
the base. Leaf margins of Caliphruria are uni- 
formly nonundulate. 

Leaf venation of Eucharis and Caliphruria is 
parallelodromous (Hickey, 1973), with a great 
number of transverse, commissural veins intercon- 
necting the primary vasculature. All species of 
Caliphruria have smooth, nonplicate leaves. Eu- 
charis is variable for this character, but the ma- 
jority of species have plicate leaves. 

The adaxial epidermis of most species of both 
genera is lustrous, dark green; the abaxial surface 
appears lighter, or silvery green. Only E. astro- 
phiala (Ravenna) Ravenna has diverged markedly 
from the typical morphology and has a uniquely 
nonlustrous, bullate-pustulate leaf texture. 


CUTICLE 


Cuticular striation is prominent on the abaxial 
leaf surfaces of most Eucharis and Caliphruria 
species (Figs. 2-6, 8-12). Striae are thickest in 
C. subedentata (Fig. 11). Arroyo & Cutler (1984) 
recognized eight cuticular sculpturing classes in a 
survey of 25 genera of Amaryllidaceae. The most 
common cuticular morphology of Kucharis and 
Caliphruria fits their class VII: "thick striae, par- 
allel or not, interlocking, + transverse.' 


In a few species of Eucharis (E. amazonica, 
E. moorei (Fig. 10), E. bouchei (Fig. 9)), the 
striation is much less pronounced. Caliphruria kor- 
sakoffii (Traub) Meerow (the sole representative 
of Caliphruria outside Colombia) has the most 
aberrant cuticle morphology (Fig. 12), correspond- 
ing more or less to type V of Arroyo & Cutler 
(1984): *central, thick axial striation with less pro- 
nounced striae running from it, directly to anticlinal 
walls." The adaxial cuticles of Eucharis and Cal- 
iphruria are either smooth or rarely much more 
finely striate than the abaxial surface, the striations 
entirely axial in orientation. The adaxial cuticle of 
C. korsakoffii (Fig. 13) has several, thick, trans- 
verse striations across each cell, and the epidermis 
is unusually flat. 


STOMATA 


Leaves of Eucharis and Caliphruria are pre- 
dominantly hypostomatic. Stomata occur adaxially 
only along the midrib and vicinity (Fig. 14A), and 
occasionally in the proximity of primary veins. 
Stomata are usually absent from the abaxial midrib 
(Fig. 14B). Intercalary stomata were regularly ob- 
served only in E. cyaneosperma Meerow (Figs. 7, 
15D). The stomata of Eucharis and Caliphruria 
are anomocytic, as is typical for Amaryllidaceae 
(Arroyo & Cutler, 1984; Dahlgren & Clifford, 
1982), though E. astrophiala exhibits at least 
slight differentiation of cells neighboring the sto- 
mata (Figs. 2, 3) from other epidermal cells. These 
cells are more densely and regularly striate than 
other epidermal cells, as well as slightly more raised. 
The guard cells of Fucharis and Caliphruria are 
oriented with their longest axis parallel to that of 
the leaf. 


EPIDERMAL CELLS 


Abaxial epidermal cells range from rectangular 
to irregular in shape. Adaxial epidermal cells are 
in almost all cases rectangular. Kucharis astro- 
phiala (Fig. 15A) has the most irregularly shaped 
cells of both the abaxial and adaxial surfaces. In 
the vicinity of the midrib on both surfaces of the 
leaf (Fig. 14), epidermal cells become conspicu- 
ously elongated, and anticlinal walls are straight. 
Epidermal cells of the midrib are extremely long 
and narrow. 

The epidermal cells of all species of Eucharis 
subg. Eucharis have strongly undulate anticlinal 
walls (Fig. 15), as noted by Asatrian (1984) for 
the few species he surveyed. End walls of both the 
abaxial and adaxial cells of all species range from 
oblique to rounded. Abaxial cells are more strongly 


Volume 76, Number 1 Meerow 141 
1989 Eucharis and Caliphruria 


FiGURES 2-7. SEM photomicrograph of Euchoris ben ate luces. 2-6. Abaxial leaf surfaces. — 2, 3. E. astrophiala 
(Meerow 1111, FLAS). — . E. plicata subsp. plicata (Meerow 1025, FLAS).—6. F. idee. qued (Meerow 
1092. FLAS). —T. MET a surface of E. c y E (Meerow 1032, FLAS). All scales = 25 u 


142 Annals of the 
Missouri Botanical Garden 


Ficures 8-13. SEM mcg of Eucharis and Caliphruria leaf surfaces. 8-12. Abaxial surfaces.— 
8. E. bakeriana (Meerow 1108, FLAS).—9. E. = chei var. dressleri (Meerow 1107, FLAS).—10. E. moorei 
(Meerow 1141, FLAS).—11. C. subedentata (Meerow 1109, FLAS).— 12. C. we (Meerow 1096, FLAS). — 
13. Adaxial surface of C. korsakoffii (Meerow 1096, FLAS). All scales = 25 


Volume 76, Number 1 


Meerow 143 
Eucharis and Caliphruria 


undulate than those of the adaxial surface. Epidermal 
cells of Caliphruria (Fig. 15F) are more weakly 
undulate than those of Eucharis. The anticlinal 
walls of adaxial cells of C. korsakoffii are com- 
pletely straight. Eucharis subg. Heterocharis is 
polymorphic for anticlinal wall morphology. Ku- 
charis amazonica and E. sanderi have strongly 
undulate walls, while E. moorei (Fig. 15E) has 
essentially straight walls. 


LEAF ANATOMY 


In petiolar transverse section, a single arc of 
vascular bundles is usually observed (Fig. 1). Me- 
dian bundles are the largest. In petioles of E. moorei 
(Fig. 1E) and the closely related E. amazonica 
(Fig. 1F), both in subg. Heterocharis, small sec- 
ondary bundles were observed near the adaxial 
surface. These bundles are most conspicuous in £F. 
moorei; they are markedly smaller in E. amazon- 
ica. These secondary vascular traces disappear 
above the middle of the petiole. Asatrian (1984), 
who reported on petiole anatomy of three Eucharis 
and Caliphruria species, did not observe these 
bundles in E. amazonica (cited as E. grandiflora). 

The internal morphology of leaves of Eucharis 
and Caliphruria (Figs. 16-22) is largely invariant 
across both genera. No well-defined palisade layer 
is evident, the absence of which is characteristic 
of most genera of "infrafamily" Pancratioidinae 
(Arroyo & Cutler, 1984). Mucilage cells, common 
throughout the family (Arroyo & Cutler, 1984), 
are often present near the leaf surface, and ra- 
phides are occasionally observed in epidermal cells. 
The mesophyll consists of several layers of chlo- 
renchyma both ad- and abaxially, and a thicker 
region of spongy, slightly aerenchymous tissue. 
Small air cavities occur regularly only directly be- 
low stomata. Vascular bundles are surrounded by 
a sheath of one or two layer(s) of parenchymous 
cells. The only xylem elements present are tra- 
cheids with annular thickenings (Fig. 19). 


FLORAL MoRPHOLOGY 
INFLORESCENCE 


The inflorescence of Eucharis and Caliphruria 
is a naked scape typical of Amaryllidaceae. The 
scape is subterete in cross section and has a solid 
pith. Vascular bundles are distributed in several 
irregular, concentric rings within the pith. A layer 
of collenchyma cells occurs just below the epider- 
mis. 
The scape is terminated by two valvate-imbri- 
cate, ovate-lanceolate bracts that enclose several 


iS 


FIGURE 14. 


Leaf epidermal cell configurations in the 
vicinity of the midrib of representative Eucharis species. — 


A. E. formosa (Schunke 14174, FLAS), adaxial sur- 


face.—B. E. plicata subsp. plicata (Meerow et al. 1025, 
FLAS), abaxial surface. 


secondary bracts and the flower buds before an- 
thesis. These bracts vary from green (E. subg. 
Heterocharis) to greenish white (most species of 
subg. Eucharis) and are soon marcescent after 
opening and spreading laterally. Each flower is 
subtended by a linear-lanceolate bracteole. 

The inflorescence of the Amaryllidaceae is tra- 
ditionally described as **umbellate." Mann (1959) 
and Stout (1944) demonstrated that the superfi- 
cially simple umbel of Amaryllidaceae actually rep- 
resents a complex series of reduced, helicoid cymes. 
Anthesis occurs in a strict sequence within each 
cyme from the developmentally oldest flower to the 
youngest. The peripheral cymes flower first; the 
central cymes flower last. 

Flower number varies in Eucharis and Cali- 
phruria from 2 to 10, rarely as many as 12 (C. 
korsakoffii, rarely in E. castelnaeana). Number 
of flowers is often a taxonomically useful character, 
though any species characterized by 8-10 flowers 
is capable of producing a depauperate inflorescence 
with fewer florets. An increase in flower number 


144 


Annals of the 
Missouri Botanical Garden 


GAS UA Vero 
RAS EY 
ISA ee 
IR 


a RA 
PESOS 


pi Y 
DAT 


FIGURE 15A-C. Leaf epidermal configurations of representative Eucharis and Caliphruria species in the inter- 


costal area of the leaf. — A. E. 
E. castelnaeana (Schunke 14156, FLAS). 


generally does not occur. In some species of subg. 
Eucharis (E. astrophiala, E. bouchei, E. ulei), a 
flower number of five has become virtually fixed. 
Reduction in flower number is usually considered 
the derived state in Amaryllidaceae (Traub, 1962, 
1963). 


FLOWER SIZE 


The largest flowers in Eucharis are found in 
subg. Heterocharis, where they average 7-8 cm 
in length. Flowers of Caliphruria are the smallest, 
never exceeding 4 cm long. Subgenus Eucharis, 
the largest of the two subgenera of Eucharis, is 


astrophiala (Madison 3792, SEL).— B. E. bonplandii (Bauml 686, HUNT).—C. 


variable, with flowers 3-7 cm in length. Within a 
fairly broad range, flower size can be used to dis- 
tinguish phenetic species complexes within subg. 
Eucharis; however, most species of this subgenus 
are quite variable in size (Meerow, 1987 d, in press). 


FLORAL FRAGRANCE 


Subgenus Heterocharis is the only subgenus of 
Eucharis with uniformly fragrant flowers. The fra- 
grance of all species of subg. Heterocharis is in- 
tense and sweet. Flowers of Caliphruria do not 
emit any detectable fragrance. Most species of 
Eucharis subg. Eucharis also lack noticeable fra- 


Volume 76, Number 1 
1989 


Meerow 
Eucharis and Caliphruria 


145 


FicuRE 15D-F. Leaf epidermal configurations of representative Eucharis and Caliphruria species in the inter- 
FLAS 


costal area of the leaf.— . cyaneosperma (Meerow 


1032, )—E. E. moorei (Meerow 1141).—F. C. 


eaf. 1 
subedentata (Meerow 1123, FLAS). ab = abaxial, ad = adaxial. 


grance. In the few fragrant species of this subgenus 
(E. bakeriana N. E. Brown, E. castelnaeana (Bail- 
lon) Macbr., E. formosa Meerow, and E. plicata 
Meerow subsp. brevidentata Meerow), the odor is 
not intense. In £. formosa the odor is slightly fetid. 


PERIANTH 


The perianth of Eucharis and Caliphruria con- 
sists of six tepals in two whorls, basally connate 
into a tube of varying length and morphology. The 
tube of Eucharis subg. Eucharis (Fig. 23E) is 


cylindrical for almost its entire length, abruptly 
dilating near the perianth throat. The tube of subg. 
Eucharis is also strongly curved, either abruptly 
just above the ovary (E. bakeriana, E. cyaneo- 
sperma) or gradually throughout the proximal half 
of its length (all other species). The curving of the 
tube results in the pendent habit of most species 
of subg. Eucharis. The tube is white for its entire 
length. 

The tube of subg. Heterocharis (Fig. 23C, D) 
is tinted green proximally (for as much as half its 
length). The tube is curved, though not as markedly 


146 


Annals of the 
Missouri Botanical Garden 


FicuRES 16-22. Transverse sections of Eucharis m" ra leaves.— 16. E. bonplandii (Bauml 686 
U 


T).—17. E. d uie artes 3792, SEL). — 
bouchei 


19. E. formosa (Meerow 1103, FLAS). 19. Tracheid 


with annular thickening var. ae p 1107, FLAS). —21. C. subedentata cid 
1123, FLAS).— 22. C. EE (Meerow 1096, FLAS). All scales = 100 um except 25 um in Figure 1 


as that of subg. Eucharis, and the habit of the 
flowers is either declinate (E. moorei, E. sanderi 
Baker) or subpendulous (E. amazonica). The tube 
is cylindrical for 4 to % of its length; it abruptly 


dilates in the distal 4% to 4%. The tube morphology 
of xCalicharis butcheri (Traub) Meerow (Fig. 
23B), putatively an intergeneric hybrid between £. 
sanderi and C. subedentata, is intermediate be- 


Volume 76, Number 1 


Meerow 147 
Eucharis and Caliphruria 


1989 
Cc 
n lcm | 
£ 
FIGURE 23. Perianth tube morphology of Eucharis and Caliphruria species or hybrids. — A. C. subedentata 


cero 1098, FLAS). — B. 
F).—D. E. amazonica (Schunke 14179, FLAS). 


tween Caliphruria (Fig. 23A) and Eucharis subg. 
Heterocharis (Fig. 23C, D). 

The tube of Caliphruria (Fig. 23A) is straight 
and dilates gradually from base to throat. It is either 
subcylindrical (C. korsakoffii) or funnelform (all 
other species). The tube is tinted green proximally 
(in C. subedentata, for V$—75 of its length). 

The tepals of Eucharis and Caliphruria flowers 
are white. Those of the outer series are almost 
always longer and narrower than the inner tepals. 
The outer tepals are apiculate. The apiculum fre- 
quently has a small, papillate horn on the adaxial 
surface in Eucharis subg. Eucharis. The inner 
tepals vary from acute to obtuse, sometimes mi- 
nutely apiculate, at the apex. 

e tepals of most species of subg. Eucharis 
spread at an angle of 90? or more from the throat. 
Perianth morphology of subg. Eucharis is thus 
predominantly crateriform. At times the tepals may 
be reflexed strongly above the midpoint of their 
length, or rarely for their entire length. Tepal habit 
varies even among flowers of the same inflores- 
cence and shows no taxonomic consistency. If ex- 
posed to strong light, the abaxial midrib of the 
tepals of some species of subg. Eucharis may be 
lightly pigmented yellow. 

The perianth of Caliphruria is infundibular. The 
tepals remain imbricate for half their length and 
spread distally at angles of only 45-60”. The tepals 
of subg. Heterocharis are also for the most part 
imbricate proximally and spread at 45-60? from 
the throat. The perianth is more or less campan- 


X Calicharis butcheri (Meerow 1110, FLAS). 
— E. E. astrophiala (Madison 3792, SEL). 


—C. E. sanderi (Cuatrecasas 16380, 


ulate. One species, E. amazonica, has the crater- 
iform perianth characteristic of A Eucharis 
with a wide-spreading (ca. 90?) limb 


ANDROECIUM 


The stamens of Eucharis and Caliphruria are 
variously connate proximally. In all but three species 
of Eucharis, a conspicuous staminal cup is present 
(Figs. 24, 25). In Caliphruria, the cup is reduced 
to a short, membranous, connate portion of the 
filaments near the perianth throat (Fig. 26). Fu- 
charis sanderi (subg. Heterocharis) has a reduced 
staminal cup similar to that of Caliphruria. Stam- 
inal connation is one of the major characteristics 
of *'infrafamily" 
workers considered the staminal cup of pancratioid 
genera homologous to the corona of Narcissus L. 
(e.g., Pax, 1888). The corona of Narcissus is 
generally considered to be of perianthal origin 
(Eichler, 1875; Arber, 1937), while the staminal 
cup of pancratioid taxa has been described as com- 
posed entirely of androecial tissue (Arber, 1937; 
Singh, 1972). Einsiedel (1987) and Müller-Doblies 
(pers. comm.) consider this distinction artificial and 
claim that both structures arise from the same 
meristematic potential, citing putative instances of 
polymorphy for both types within the same genus. 

Stamens of Eucharis and Caliphruria may be 
dentate, edentate, or irregularly toothed. Both types 
of staminal morphology may occur in the same 
species, and variation may occur even among flow- 


Pancratioidinae. Some taxonomic 


148 


Annals of th 
Missouri Ecos Garden 


FIGURES 24-26. 
and Caliphruria species. — 24. 
3792, SEL).—25. E. amazonica (Schu 
FLAS).— 26. C. subedentata (Meerow 1109, FLA 


Androecial uma d of Eucharis 
astrophiala (Madison 


ers of a single clone (Meerow, 1986). The presence 
or absence of staminal dentation has frequently 
been overweighted in the alpha-taxonomic litera- 
ture relating to these genera (e.g., Ravenna, 
but can still be of taxonomic significance (e.g., E. 
astrophiala (Fig. 24), the only species of subg. 
Eucharis that always has an edentate staminal 
cup). 
A variable pattern of green or yellow pigmen- 
tation is present in the androecium of all species 
of subg. Eucharis and Heterocharis. Stamens of 


- 
No) 
eo 


Caliphruria are completely white. In subg. Het- 


erocharis, the green (rarely yellowish) pigmenta- 
tion is largely restricted to the interior of the cup, 
and extends into the dilated portion of the tube as 
well (Fig. 25). The coloration is concentrated along 
the filamental traces, but the tissue between the 
traces is also suffused with green. In subg. Kucha- 
ris, pigmentation is present on both the exterior 
and interior surfaces of the cup, does not extend 
into the dilated portion of the tube, and takes the 
form of either broad spots below each free filament 
or a uniform band of color at the basal V$- V4 of 
the cup. In this subgenus, the pattern is of limited 
taxonomic significance. Whether the pigmentation 
functions as nectar guides for pollinating animals 
is unknown. 

The stamens of most species of subg. Eucharis 
constrict distally into a broadly subulate portion 
(greater than 1 mm wide for most of its length) of 
varying length. Only in E. astrophiala (Fig. 24) 
and E. bouchei (in part) do the stamens constrict 
gradually from the rim of the staminal cup to the 
apex of the filament. The free filaments of Cali- 
phruria are narrowly subulate (less than 1 mm 
wide for most of their length, Fig. 26). The free 
filaments of E. sanderi (subg. Heterocharis) are 
narrowly subulate and slightly incurved. Those of 
E. moorei and E. amazonica are broadly subulate. 

Anthers of Eucharis are introrse, dehiscing lon- 
gitudinally, and either dorsifixed or sub-basifixed. 
They are most frequently oblong but are linear in 
subg. Heterocharis. At anthesis, the anthers of 
Caliphruria and Eucharis subg. Eucharis are erect 
but become versatile as they age. In subg. Het- 
erocharis, the anthers are versatile at anthesis. 


STIGMA AND STYLE 


Of the sixteen species of Kucharis and Cali- 
phruria observed in cultivation, all but Æ. castel- 
naeana are protandrous. Stigma receptivity does 
not occur until the second or third day following 
anther dehiscence. In some cases, the stigma does 
not fully expand until the perianth has begun to 
senesce. Successful greenhouse pollinations have 
been accomplished after the onset of periant 
nescence. In the single known autogamous species, 
E. castelnaeana, stigma receptivity and anther 
dehiscence coincide 

The styles of Eucharis and Caliphruria are 
usually exserted beyond the anthers, most fre- 


FIGURES 27-39. 


1111, FLAS).—29. E. plicata (Plowman 13941, 


=> 


SEM photomicrographs of aa and rad stigmas. — 27, 28. E. di i da 
FLAS). — : NE 
korsakoffii (Meerow 1096, FLAS). — 33, 34. E. ieri a a 1127, FLAS).— 


. subedentata (Meerow 115 
35. E. moorei ia 


Volume 76, Number 1 Meerow 149 
1989 Eucharis and Caliphruria 


1141, FLAS).— 36. E. sanderi (Cuatrecasas 16350, F).—37. E. amazonica (Schunke 14179, FLAS).— 38, 39. 
x Calicharis butcheri (Meerow 1110, FLAS). All scales = 50 um. 


150 


Annals of the 
Missouri Botanical Garden 


42 


FIGURES 40-43. 
(Schunke 14174, FLAS). —41. E. 
bouchei (Meerow 1125, FLAS). 


Fruits and seeds of Eucharis 
castelnaeana (Schur 


43. E. castelnaeana (Schunke 14156, FLAS 


quently 0.5-1 cm. In subg. Heterocharis, the styles 
are somewhat assurgent away from the stamens 
and are exserted well over 1 cm past the anthers. 
In two species of subg. Eucharis, E. castelnaeana 
and E. plicata, the style is included within the 
cup. In the former species, self-pollination occurs 
regularly, and stigma receptivity coincides with 


anthesis. 


rad Eucharis. 


Mature capsules.— 40. E. 1 
42, 43. Seeds. —42. E. bouchei var 
). 


rss 


> 14156, F 


Stigmas of Eucharis and Caliphruria (Figs. 27 
29, 30, 32, 33) are obtusely trilobed. Trilobed 
stigmas are relatively rare in the Pancratioidinae, 
and Urceolina—sister group to Eucharis and Cal. 
iphruria—has a capitate, entire stigma. Traub 
(1963) and Traub & Moldenke (1949) considered 
a trilobed or trifid stigma the ancestral state in the 


Amaryllidaceae. 


Volume 76, Number 1 
1989 


Meerow 151 
Eucharis and Caliphruria 


The stigmatic papillae of Eucharis are unicel- 
lular (Figs. 28, 34-37), while those of Caliphruria 
(Fig. 31) are multicellular, consisting of both a 
stalk cell and globose head cell. x Calicharis 
butcheri, putatively a natural hybrid of E. sanderi 
and C. subedentata, has the multicellular stigmatic 
papillae (Figs. 38, 39) characteristic of Caliphru- 
ria. Heslop-Harrison & Shivanna (1977) charac- 
terized the stigmas of Eucharis and Caliphruria 
as dry-type. 


OVARY AND OVULES 


The ovary of Eucharis and Caliphruria is in- 
ferior and contains septal nectaries. It is green, 
with the exception of two species, E. astrophiala 
and E. castelnaeana (subg. Eucharis), in which 
the ovary is white at anthesis. Ovaries of Eucharis 
and Caliphruria range from oblong-ellipsoid (subg. 
Heterocharis) to globose or subglobose (Caliphru- 
ria and Eucharis subg. Eucharis). The ovary of 
subg. Heterocharis appears rostellate after senes- 
cence of the perianth. 

The ovules of Eucharis and Caliphruria are 
globose, anatropous, and borne on axile placentae. 
Ovule number is quite variable throughout both 
genera. Within limits, however, this number is 
characteristic of species or species complexes. Sub- 
genus Heterocharis has the largest ovule number 
in Eucharis, generally 16-20 per locule, but oc- 
casionally as low as 7 in E. sanderi (which oth- 
erwise has 16-20 throughout most of its range) 
and 9-12 in E. amazonica. In both Caliphruria 
and Eucharis subg. Eucharis, ovules do not num- 
ber more than ten per locule. Eucharis astrophi- 
ala, E. bouchei, E. bonplandii, E. cyaneosperma, 
and E. ulei characteristically have two ovules per 
locule, but rarely as many as five. 


FRUIT 


The mature fruit of Eucharis and Caliphruria 
is a triloculicidal capsule typical of the nonbaccate- 
fruited Amaryllidaceae. In fruit, the pedicel elon- 
gates to two or more times its length at anthesis. 
In Caliphruria and Eucharis subg. Heterocharis 
(E. moorei), the capsule is thin-walled and green, 
sometimes turning yellow or brown at dehiscence. 
In subg. Eucharis, however, the capsule is leathery 
and bright orange (Fig. 40), contrasting vividly 
with the shiny black or blue seeds at dehiscence. 
It is probable, though unsubstantiated, that the 
combination of fruit and seed color functions mi- 
metically to attract avian dispersal agents (sensu 
van der Pijl, 1982). In the single known exception 
to this characteristic fruit morphology in subg. 


Eucharis, the fruit of E. castelnaeana (Fig. 41) 
remains green until shortly before dehiscence, at 
which time it yellows and finally turns brown. It is 
often tardily dehiscent and sometimes abscises be- 
fore opening, though the seeds within are ripe. 


SEED 

Regardless of the number of ovules per locule 
in any species of Eucharis and Caliphruria, all 
but a few abort as the fruit matures. Generally one 
to three seeds are present per locule in mature 
capsules, but as many as four have been observed 
(Fig. 40) 

The seeds of both Eucharis and Caliphruria 
are globose or ellipsoid and turgid, the consequence 
of copious, oily endosperm and a high moisture 
content. Left at room temperature, the seeds will 
shrink away from the testa somewhat as moisture 
is lost but remain capable of germination. Long- 
term viability has not been tested. 

The seed of subg. Eucharis (Fig. 42) is char- 
acteristically ellipsoid and has a shiny, smooth black 
(blue in E. cyaneosperma) testa. The single ex- 
ception so far known is again E. castelnaeana (Fig. 
43). The seed of this species is wedge-shaped by 
compression in the capsule, is less turgid than seeds 
of consubgeneric species, and has a dull, rugose 
testa. The seed of E. moorei (subg. Heterocharis) 
is globose to very slightly compressed and has a 
brown, slightly rugose testa. 

In Caliphruria, the seeds of only C. korsakoffi 
and C. subedentata are known. Seeds of C. kor- 
sakoffii are globose, turgid, and have a smooth, 
lustrous brown testa. Seed of C. subedentata is 
slightly compressed, with a lustrous black but ru- 
gose testa. 

The testa is alveolate in all species examined 
(Figs. 44-49). In E. bouchei var. dressleri Mee- 
row (Fig. 45), abundant wax extrusions are found 


Caliphruria seeds is composed of phytomelan 
(Huber, 1969), a simple, largely inert, carbona- 
ceous compound characteristically present in the 
seed coat of nonbaccate-fruited Amaryllidaceae 
(Dahlgren & Clifford, 1982; Huber, 1969). Wer- 
ker & Fahn (1975) reported the occurrence of 
phenolic quinones in the phytomelan layer of Pan- 
cratium seeds. In most species of Eucharis and 
Caliphruria, the phytomelan layer is all that re- 
mains of the integuments (Figs. 51, 57). In £. 
bouchei, however, there is an additional layer of 
integument tissue about five cells thick, interposed 
between the phytomelan and the endosperm (Fig. 
55). Whether this may be a consequence of the 


152 Annals of the 
Missouri Botanical Garden 


Tam 
FIGURES 44-49. SEM photomicrographs of reges xd Caliphruria ip surfaces.— 44. E. astrophiala 
(Meerow 1111, FLAS).— 45. E. bouchei var. dressleri (Meerow 1107, FLAS).— 46. E. formosa (Meerow 1103).— 
47. E. castelnaeana (Schunke 14156, FLAS). —48. C. korsakoffii (Meerow P FLAS).— 49. C. subedentata 
(Meerow 1152, FLAS). All scales = 50 pu 


Volume 76, Number 1 


Meerow 153 


Eucharis and Caliphruria 


Ae es 
5 


SAC 


SH T 


sAd >; J- ; 
Econ: 50-58. 


endosperm. — 52. 


- 


ES A MPs 
SETS 


Photomicrographs of mec d pa pieta seed Mu 50-54. C. korsakoff (Meerow 
—51. Tra 


verse section through testa and part o 


bryo.— 54. pi section through vascular initial of embryo. 55, 56. E. bouchei var. bouchei (Meerow 1125, 


FLAS). s Transverse section thr ough testa. 
tomelan layer. — 56. En Br A Note t 


(Schunke 14156. FLAS). — 


through embryo. All scales = dus um except 40 um in Figure 54. em — 


tetraploid condition of this species is unknown. 
Most of the seed body of Eucharis and Caliphruria 
is taken up by a copious quantity of endosperm 
characterized by abundant transfer cells (Fig. 56). 
At maturity, no remnants of the nucellus were 
observed. 

Most workers (e.g., Baker, 1888; Traub, 1963; 
Hutchinson, 1959; Dahlgren et al., 1985) have 


ansfer tissue with pitted walls and plasmodesmata. 57, 58. 
sioe. section through testa and part of endosperm. — 58. Transverse section 
testa. 


Note several layers of additional integument cells below outer phy- 


E. castelnaeana 


embryo 


en — endosperm, t — 


allied Eucharis and Caliphruria with Hymeno- 
callis, Eurycles, and Calostemma (e.g., tribe Eu- 
chareae, or Eucharideae) on the basis of “fleshy 
seeds." The last-mentioned three genera do indeed 
have fleshy, pora sometimes viviparous seeds, 
but they are not The “ps 
doseed” of Eurycles and Calos y 
an adventitious bulbil (Rendle, m The bulbi- 


154 


Annals of the 
Missouri Botanical Garden 


form propagules of Hymenocallis, while true seeds, 
have thick, fleshy chlorenchymous integuments with 
a well-developed vascular system and a starch- 
storing embryo (Rendle, 1901; Whitehead & 
Brown, 1940). These important seed differences 
have been overlooked by most phylogenists of the 
Amaryllidaceae. 

The turgid seed of Kucharis and Caliphruria, 
despite a high moisture content when first ripe, 
cannot be accurately described as fleshy. This be- 
comes evident if the seed is allowed to dehydrate 
slightly at room temperature, and is most apparent 
in the hard seeds of E. castelnaeana, which, at 
capsule dehiscence, are less turgid than seeds of 
other species of subg. Kucharis. Seeds of Eucharis 
and Caliphruria have a reduced integument, rep- 
resented in most cases only by the compressed 
phytomelan layer, and have never been observed 
to germinate viviparously. Phytomelan is absent 
from the testa of the pseudoseeds of Kurycles and 
Calostemma. It is present in only a single species 
of Hymenocallis, H. quitoensis Herbert (and prob- 
ably its close relative H. heliantha Ravenna), which 
has been segregated into the separate genus Lep- 
idochiton Sealy (1937) on this basis. 

Seeds of Pancratium are structurally most sim- 
ilar to those of Eucharis and Caliphruria. Al- 
though variable in shape and surface features 
(Werker & Fahn, 1975), all species of Pancratium 
have a hard, turgid, ovoid or compressed seed body 
with copious endosperm (Meerow, unpubl. data; 
Werker & Fahn, 1975). All examined species of 
Pancratium have a phytomelanous and alveolate 
testa. Seeds of Eucharis and Caliphruria have a 
higher moisture content than those of Pancratium, 
all species of which occur in xeric to seasonally 


dry habitats. 


POLLEN MORPHOLOGY 


Pollen morphology of Eucharis and Caliphruria 
is reviewed in detail elsewhere (Meerow & Deghan, 
1988) and will only be summarized here. Pollen 
grains of all species of Eucharis and Caliphruria 
(Figs. 59-62) are boat-shaped elliptic, monosul- 
cate, heteropolar, and bilaterally symmetrical. The 
sulcus runs the length of the distal face of the 
grain. Exine sculpturing is semitectate/reticulate 
in all species examined. 

Pollen of most Eucharis falls into Walker & 
Doyle's (1975) large size class (longest equatorial 
diameter 50-100 um). A single species, E. cau- 
cana, has very large pollen (longest equatorial di- 
ameter as much as 110 um). Pollen of Eucharis 
has average longest equatorial diameters greater 


than 60 um, with two exceptions: E. castelnaeana 
and E. plicata subsp. brevidentata, the two small- 
est-flowered species. Most species of Eucharis have 
pollen grains with longest equatorial diameters be- 
teen 65 and 75 um. Pollen of Caliphruria falls 
into the medium size class of Walker & Doyle 
(1975), with average longest equatorial diameters 
near 50 um. Polar diameter of pollen of most 
Eucharis species ranges from 45 to 60.6 um— 
diameters less than 40 um are rare. Polar diameter 
of pollen of Caliphruria is always less than 40 
um. Exine reticulation of Eucharis is character- 
istically coarse (Figs. 59-61), while that of Cali- 
phruria is fine (Fig. 62). 


CHROMOSOME CYTOLOGY 


A somatic chromosome number of 2n = 46 
largely characterizes both genera (Meerow, 1987b). 
Two tetraploid species (2n = 92), E. bonplandii 
from Colombia and E. bouchei from Central Amer- 
ica, are known. Eucharis caucana, described in 
this paper from Colombia, is a hexaploid with the 
largest chromosome number in the genus (2n — 
138). Eucharis amazonica, 2n — 68, is the only 
known departure from these 2x, 4x, or 6x karyo- 
types. À somatic chromosome number of 2n — 46 
(or derivations thereof) is characteristic of most 
genera of neotropical Pancratioidinae (Di Fulvio, 
1973; Flory, 1977; Meerow, 1987b, c; Williams, 
1981). 


EcoLoGY AND PHYTOGEOGRAPHY 
ECOLOGY 


All species of Eucharis and Caliphruria exhibit 
high fidelity to a primary forest habitat, and severe 
disturbance of the forest canopy is catastrophic to 
the plants. In recently cleared forest sites, the bulbs 
persist for a few seasons, but the leaves developed 
in sunlight show chlorosis and necrosis. Wilkins 
(pers. comm.) reported that leaves of E. amazonica 
are damaged at light levels above 5,000 foot can- 
dles, an observation confirmed by Rees (1985). 
Fidelity of these genera to mesic, low-light habitats 
suggests a strongly evolved adaptive complex. Only 
two other genera of pancratioid Amaryllid are 
completely adapted to forest understory: Eurycles, 
a small Australasian genus, and Urceolina, sister 


group to Eucharis and Caliphruria. 

Eucharis populations are often largest on flood- 
plains of small rivers and creeks where frequent 
short-term inundation is likely. Eucharis is also 
found in more upland sites, but usually in less 
abundance. No fully deciduous species of subg. 


Volume 76, Number 1 
1989 


Meerow 155 
Eucharis and Caliphruria 


FiGURES 59-62. 
FLAS). — 59. Whole 


ain, proximal polar view. — 60. 


pu pollen grains of Eucharis and Caliphruria. 59, 60. E. formosa (Meerow 1103, 
Detail of the exine surface 


.—61. E. sanderi (Killip 35401, 


T, soy lateral aeiia view.—62. C. subedentata (ex hort. s.n., K), proximal polar view. All scales = ca. 


Eucharis have been observed, though E. astro- 
phiala, endemic to the western slopes of north- 
central Ecuador, enters a season of dormancy when 
growth ceases. Yet several leaves may persist for 
the duration. 

The rarity of Eucharis and Caliphruria species 
throughout their range is a striking characteristic 
of their distribution. Single, widely dispersed clumps 
of bulbs are more the rule than the exception. 
Herbarium collections are usually unicates and often 
indicate the relative infrequency with which the 
plants were encountered in the forest. The largest 
population of Eucharis that I have observed con- 
sisted of about 30 individuals of E. moorei in an 
approximate half-hectare area. Throughout eastern 
Ecuador, populations of only two or three clonal 
clumps or single bulbs (and frequently as low as 
one) of E. candida, E. formosa, and E. moorei 
are common. In eastern Peru, a careful search 


through a five-hectare area turned up only two 
plants of E. cyaneosperma. Edaphic conditions are 
probably important in limiting colonization and es- 
tablishment of Eucharis species. Restriction to sites 
of high fertility is evident for all species of Eucharis 
and has been noted by floristic workers in the 
Amazon basin and Chocó region of Colombia (A. 
Gentry, pers. comm.). 

No species of Eucharis are found natively above 
1,800 m. The overwhelming majority of collections 
are from elevations below 1,000 m and, of these, 
more than half are below 500 m. Caliphruria is 
primarily collected at sites above 1,000 m (“selva- 
subandina" of Cuatrecasas, 1958). Most collec- 
tions of Caliphruria (C. subedentata) were from 
the Río Cauca River valley of western Colombia, 
an area now largely deforested. In July 1984, 1 
was not able to find members of this genus in any 
of its historical localities. 


156 


Annals of the 
Missouri Botanical Garden 


PHYTOGEOGRAPH Y 


More than half of the known species of Fucharis 
subg. Eucharis are found on the eastern slopes of 
the Andes. The majority of these species are con- 
centrated along the Amazon and its main tributar- 
ies, e.g., the Napo and Pastaza system in Ecuador 
and the Huallaga of Peru. Seven of the fourteen 
extant species of subg. Eucharis are endemic to 
the premontane Andean- Amazonas interface, but 
no species has been reported east of 68°W longi- 
tude. Thus the genus appears to be absent from 
the great, largely Brazilian expanse of lowland 
Amazonas. Five peripheral isolate species from the 
premontane Andean-Amazonian center of distri- 
bution show some degree of morphological novelty 
(see Taxonomic Treatment). Eucharis corynan- 
dra, very closely related to E. castelnaeana and 
E. plicata, and E. oxyandra, of uncertain phy- 
logenetic relationship, are both known only from 
the “ceja de la selva” forest formations of north- 
central eastern Peru. Eucharis astrophiala is en- 
demic to a relatively small area of western Ecuador. 
Eucharis lehmannii and E. caucana are restricted 
to western Columbia. Of the five species, only £. 
astrophiala is known from other than the type 
locality. 

Three of the four species of subg. Eucharis that 
occur above 2°N latitude are polyploids (chromo- 
some number is unknown for E. lehmannii). Eu- 
charis bonplandii (2n — 92), endemic to the Cor- 

illera Oriental and Cordillera Central of Colombia 
is rare and infrequently collected. The E. bouchei 
complex (2n — 92), a series of geographically isolat- 
ed and morphologically distinct population clusters 
(Meerow, 1987d), is endemic to Central America, 
chiefly Panama. Eucharis caucana is a hexaploid 
(2n = 138) 

Subgenus Heterocharis is widely dispersed from 
Colombia to Peru, but each of its three species are 

arrowly distributed. With the exception of a single 
E. moorei is not 


Eucharis found on both sides of the Andes. Eu- 
charis amazonica occurs natively within a narrow 
area of the Rio Huallaga valley in Peru. Eucharis 
sanderi is endemic to the Chocó region of Colom- 


ia. 
Caliphruria is almost completely western Co- 
lombian, most prominently in the Rio Cauca valley, 
with a secondary distribution in the Río Magdalena 
valley. Caliphruria korsakoffii is Peruvian. 


ETHNOBOTANY 


Labels of a number of herbarium specimens of 
Eucharis species indicate that lowland native peo- 


ple collect the bulbs for use as poultices. The bulbs 
are mashed, heated, and applied to sores and tu- 
mors. Lewis (1986) described the use of mucilage 
from Eucharis bulbs (the plant illustrated is Æ. 
formosa) by the Jivaro Indians of Peru for treating 
facial blemishes and acne. Indian women of the 
Pastaza valley in Ecuador reportedly quite actively 
collect the plants in flower for reasons they would 
not disclose (N. Whitten, pers. comm.). Most local 
inhabitants whom I met while collecting Eucharis 
in the Oriente were readily familiar with the plants 
when shown photographs. 

That various neotropical Amaryllidaceae have 
been cultivated in their native countries for many 
years cannot be discounted. Motifs recognizable as 
species of Zephryanthes Herbert and Stenomes- 
son Herbert appear on ceremonial Incan vases 
(Vargas, 1981). The seasonal mass-blooming of 
Hymenocallis amancaes (Ruiz & Pavón) Nichols. 
on the dry hills near Lima, Peru, has long been 
the nucleus of an annual festival named for that 
species (Herbert, 1839). I have suggested that 
frequent sympatric occurrence of E. candida and 
; y have been 
influenced by cultivation by transient agriculturists 
(Meerow, 1987e). Further circumstantial evidence 
for this hypothesis is the doen in nature E 
sterile hybrid taxa such as F. randiflora a 
X Calicharis butcheri. Almost all collections of i 
former are remnants of cultivation. Eucharis ox- 
yandra is known only from bulbs found uprooted 
by an abandoned dwelling in Peruvian rain forest 
(see Taxonomic Treatment). 


E. formosa in eastern Ecuador ma 


REPRODUCTIVE BIOLOGY 
PHENOLOGY 


Eucharis and Caliphruria species are largely 
seasonal flowering. Very well-collected species, such 
as E. candida or E. formosa, show flowering pat- 
terns skewed toward certain months (January to 

arch), but at least several flowering collections 
have been made throughout the year. Species in 
Amazonian Peru have been collected in flower most 
frequently from June to September. Each inflores- 
cence is moderately long-lived, two to three weeks, 
but usually no more than one to three flowers are 
open at any one time 

In the greenhouse, observations over a five-year 
period confirm an annual flowering for most Eu- 
charis and Caliphruria species. Eucharis ama- 
zonica, however, flowers two or three times during 
the year. Van Bragt & Sprenkels (1983) have 
regularly induced flowering in this species at any 
time of the year after treatment at 27?C for at 
least two weeks. Collection data for this species 


Volume 76, Number 1 
1989 


Meerow 157 
Eucharis and Caliphruria 


throughout its narrow range indicate that twice- 
annual phenology may occur in habitat as well. 
Eucharis castelnaeana mass-flowers in the green- 
house several times per year. Other well-collected 
species (e.g., E. candida and E. formosa) that 
show a peak period of flowering in certain months 
in nature are strictly annual-flowering in cultiva- 
tion. 


POLLINATION BIOLOGY 


Data on pollination ecology of Amaryllidaceae 
in general are scant and represent an area sorely 
in need of investigation. No information on polli- 
nation of Caliphruria is available. The large, white, 
heavily and sweetly fragrant flower of E. amazon- 
ica (subg. Heterocharis) was considered a model 
moth-pollinated flower by Percival (1965). She not- 
ed that the nectar level in the tube rises to a 
maximum height of 2376 tube length, thus effec- 
tively preventing access to all but long-tongued 
insects. The large, heavily fragrant flowers of subg. 
Heterocharis fit the syndrome of sphingid moth 
pollination (sensu Faegri & van der Pijl, 1979). 
Other related genera of *'infrafamily" Pancratioi- 
dinae with similar flower morphology are special- 
ized for hawkmoth pollination (Hymenocallis: 
Bauml, 1979; Grant, 1983; Pancratium: Morton, 
1965). 

Floral fragrance is rare in subg. Eucharis, oc- 
curring in only E. bakeriana, E. castelnaeana, E. 
formosa, and E. plicata subsp. brevidentata. 
However, floral fragrance is only weakly developed 

these species, and in E. formosa the odor is 
slightly fetid. Vogel (1963) reported euglossine bees 
visiting Eucharis bakeriana. Visitation by an un- 
identified euglossine has been reported for other, 
unidentified species of subg. Eucharis in Peru (J. 
Schunke, pers. comm.). Unfortunately, neither re- 
port indicated the sex of the visitor. The phenology 
of Eucharis (rarely more than one or two flowers 
open per inflorescence at any one time over à 
period of two or three weeks, and the often suc- 
cessive appearance of inflorescences from a clonal 
clump of bulbs) and the dispersed distribution of 
plants in the wild suggests pollination by a trap- 
lining organism (see Janzen, 1971) 


DISPERSAL 


The leathery, bright orange fruit of Fucharis 
subg. Eucharis is a major apomorphy that defines 
the subgenus. At dehiscence, the contrast between 
the capsule and the lustrous black or blue seeds 
creates a striking visual display in the forest under- 
story, but there are no recorded observations of 
visitation to the ripe capsules. 


BREEDING SYSTEM 


On the basis of greenhouse pollination attempts, 
virtually all species of Eucharis and Caliphruria 
demonstrate some degree of self-incompatibility. 
This observation suggests that most species are 
predominantly outcrossing. Capsules have been set 
readily with sibling pollen on all species in culti- 
vation with the exception of E. amazonica and 
functionally sterile hybrid taxa (E. x grandiflora, 
x Calicharis butcheri). Further evidence of out- 
crossing is the marked protandry of most Eucharis 
species (with the exception of E. castelnaeana, in 
which stigma receptivity coincides with anthesis) 
and the presence of putative interspecific hybrids 
in nature (E. candida x E. formosa (Meerow, 
1988), E. plicata subsp. plicata x E. ulei (Mee- 
row, in prep.)). Of all species in cultivation, only 
7. castelnaeana sets capsules with self-pollen. 
Autogamy in E. castelnaeana is associated with a 
number of other divergent character states for 
subg. Eucharis. This species has a mass-flowering 
phenology; the smallest flowers in the genus; a 
green, often tardily dehiscent capsule; a less turgid 
seed with a dull, rugose testa; and telocentric chro- 
mosomes. 

Reproductive biology of Eucharis and Cali- 
phruria is thus still largely conjectural. Annual 
flowering patterns, short-term pollen viability, and 
susceptibility to systemic diseases in cultivation 
impede greenhouse exploration of these questions, 
and the rarity of the plants is an obstacle to field 
studies. The disappearance of populations of these 
genera and their pollinating agents, concurrent with 
ain forest destruction, may be the final impediment 
to investigation of this aspect of their biology. 


~ 


PHYLOGENETIC RELATIONSHIPS 


Eucharis and Caliphruria have been variously 
treated as separate genera (Baker, 1888; Bentham 
& Hooker, 1883; Pax, 1888; Hutchinson, 1959), 
a single genus (Traub, 1963), and as subgenera of 
Urceolina (Traub, 1971). As a group, Eucharis, 
Caliphruria, and Urceolina appear to represent 
a distinct tribal assemblage within “‘infrafamily” 
Pancratioidinae Traub (1957, 1963), a group de- 
limited by various patterns of staminal connation, 
somatic chromosome number frequently of 2n — 
46, and an Andean center of diversity (Meerow, 
1985, 1986, 1987a, c). 


PHYLOGENETIC ANALYSIS 


Despite a lack of consensus on generic and spe- 
cific limits in Amaryllidaceae, cladistic analysis has 
only twice before been applied to such problems 


158 


Annals of the 
Missouri Botanical Garden 


FIGURE 63. — ta states of Urceolina. All photos are U. microcrater Kránzl. (Schunke 13633, FLAS) unless 


othe 
cell 


rwise stated. — lowers (Plowman & Kennedy 5721, GH). Photo courtesy T. Plowman. B, C. Leaf epidermal 
configurations. ^s Abaxial surface. — C. Adaxial surface. — D. SEM photomicrograph of abaxial leaf surface. — 


Volume 76, Number 1 
1989 


Meerow 159 


Eucharis and Caliphruria 


in the family, by Nordal & Duncan (1984) for 
Haemanthus L. and Scadoxus Raf., 
related, baccate-fruited African genera; and Mee- 
row (1987c) for Eucrosia Ker Gawler. The generic 
limits of Eucharis and Caliphruria in relation to 
Urceolina, as well as interspecific relationships in 
the former genera, seemed two areas that would 


two closely 


benefit from the application of cladistic analysis. 
Before assessing these relationships cladistically, a 
brief review of character states in Urceolina is 
necessary. 

Urceolina (ca. 6-8 species; Fig. 63) is so far 
known only from central and southern Peru on the 
eastern Andean slopes (Fig. 64) at elevations of 
700-2,000 m. The relationships of Urceolina have 
long been misunderstood, most workers (Baker, 
1888; Pax, 1888; Hutchinson, 1959) placing Ur- 
ceolina near Stenomesson Herbert. Traub (1957) 
was the first to recognize its affinities with Eucharis. 
The problem no doubt stemmed from the inclusion 
of a species of Stenomesson (S. miniatum (Her- 
bert) Ravenna) in Urceolina as U. miniata (Her- 
bert) Benth. & Hook. Despite the ventricose aspect 
of the corolla morphology of 5. miniatum, which 
is similar to that of Urceolina, the species clearly 
belongs to Stenomesson as evidenced by its narrow, 
subpetiolate leaves, the morphology of its staminal 
cup, and its numerous flat, black seeds (Ravenna, 


Urceolina is easily distinguished by its brightly 
colored, urceolate corolla formed by the coherence 
of the tepals throughout most of their length (Fig. 
63A). Vargas (1960) described the genus Pseu- 
dourceolina to accommodate a species of Urceo- 
lina in which the tepals are supposedly more laxly 
coherent. The morphology of the flower in Urceo- 
lina strongly suggests ornithophily (sensu Faegri 
& van der Pijl, 1979), i.e., vivid, "parrot" colors, 
ventricose corolla, pendulous habit, and absence 
of odor. As in Caliphruria, the staminal cup of 
Urceolina is reduced to a minute, membranous, 
basal connation of the filaments, which are oth- 
erwise linear throughout their length. The leaves 
of Urceolina, in thickness and lack of both surface 
plication and marginal undulation, resemble Cali- 
phruria more closely than Eucharis. At least one 
species is hysteranthous (U. microcrater Kranzlin), 
but all species enter a leafless dormant period in 
the wild. In size the pollen grains of Urceolina (Fig. 
63E) are similar to those of Caliphruria. The 


coarseness of the reticulum is intermediate between 
that of Eucharis and Caliphruria. In all species 
of Urceolina the small stigma is capitate and entire 
(Fig. 63G) versus the trilobed stigma found in Eu- 
charis and Caliphruria. Urceolina species have 
10-20 ovules. The ripe fruit of Urceolina is a thin- 
walled, yellow-green, turbinate capsule. The seeds 
of U. microcrater (Fig. 63H) are narrowly oblong, 
ca. 8 mm long, and 2.5 mm wide, and curved. 
The seeds have a unique anatomical feature: at 
the chalazal end, a dam of poorly differentiated 
tissue separates a "cap" composed of many small 
cells from the endosperm (Fig. 63J, L). The cells 
of the “cap” do not have pitted walls with plas- 
modesmata as is characteristic of the endosperm 
cells. There is no obvious surface feature on the 
seed corresponding to the area of this "cap," and 
its function is unknown. 

Outgroup selection. Preliminary cladistic 
analyses of the pancratioid Amaryllidaceae (Mee- 
row, 1985) indicate that within the Pancratioidi- 
nae, a large, white, fragrant, crateriform flower 
with a conspicuous staminal cup (““paneratioid,” 
cf. Pancratium L.), involved with sphingid moth 
pollination (Bauml, 1979; Grant, 1983; Morton, 
1965), may be symplesiomorphous. In other words, 
while the pancratioid flower was a major apomor- 
phy defining the Pancratioidinae as a distinct group 
within the Amaryllidaceae, it is the ancestral floral 
morphology from which all other pancratioid taxa 
have diverged. I have used the term “the pancra- 
tioid base" 
tioidinae with this type of flower morphology (Mee- 
5). These five genera are Eucharis, 
Hymenocallis, Pancratium, Pamianthe Stapf, and 
Paramongaia Velarde. All but Pancratium are 
entirely neotropical in distribution. The pancratioid 


to define the five genera of Pancra- 


row, 


flower correlates repeatedly with the largest pollen 
grain size within the Pancratioidinae (Meerow, 
1985; Meerow € Dehgan, 1985). Putatively basal 
complexes within each pancratioid lineage also have 
numerous ovules per locule (ca. 20 or more), a 
character state considered primitive in the Ama- 
ryllidaceae (Traub, 1963). 

Pollen morphology provides the best evidence 
for the monophyly of the Pancratioidinae. The 
large pollen grains of most species of the pancra- 
tioid base are uniformly coarsely reticulate, with 
some in each genus exhibiting a striking exine 


E. SEM photomicrograph of U. d pollen grain (Weberbauer 7822, US), proximal polar view. — F. Ovary.— 
. Seed. 


G. SEM ai of stig 
endosperm, t = testa = cap.— 
Internal chalazal “cap. " Scale bars B, C, G, K = 


I-L. Longitudinal transverse sections of seed. 
ai peer end of seed. — 
100 15.30 = 254 A 132 


em = embryo, en = 


=K. pi of embryo. — L. 
200 


alazal end of see 


160 Annals of the 
Missouri Botanical Garden 
— 
— 
A 
EUCHARIS subg. EUCHARIS 
ed E. subg. HETEROCHARIS 
| catipHruria 
p URCEOLINA 
0 400 800 1200 1600 
km SJ 
E oid 
90 | / 
110 90 78 50 30 
X \ i ERU 4 
FIGURE 64. Generalized distributions of Eucharis, Caliphruria, and Urceolina in Central and South America. 


dimorphism at the equatorial ends of the grain 
(Meerow, 1985; Meerow & Dehgan, 1985). None- 
theless, separate origins of the pancratioid flower 
in the Neo- and Paleotropics cannot be ruled out. 

That the neotropical tribes of the Pancratioidi- 
nae represent a monophyletic group is a much more 


robust hypothesis. All neotropical pancratioid gen- 
era have 2n = 46 as the most common somatic 
chromosome number (Di Fulvio, 1973; Flory, 1977; 
Meerow, 1985, 1987a, c; Williams, 1981). This 
number may be derived from 2n = 22, the most 

tic cl ber in the family 


Volume 76, Number 1 Meerow 161 
1989 Eucharis and Caliphruria 
ABLE l. List of EUs and label designations for cla- TABLE 2. Characters, character states, and transfor- 


distic analysis of Eucharis and Caliphruria. * = out- 
group. 


Desig- 
Taxon nation 
Eucharis amazonica Linden ex 
Planchon AMA 
E. astrophiala ede Ravenna AST 
E. bakeriana N. E. Brow BAK 
E. bonplandii (Kunth) Traub BON 
E. bouchei Woodson & Allen BOU 
E. candida Planchon & Linden CAN 
E. castelnaeana (Baillon) Macbride CAS 
E. caucana Meerow CAU 
E. corynandra (Ravenna) Ravenna COR 
E. cyaneosperma Meerow CYA 
E. formosa Meerow FOR 
E. lehmannii Regel LEH 
E. moorei (Baker) Meerow MOO 
E. oxyandra (Ravenna) Ravenna OXY 
E. plicata subsp. brevidentata Meerow PLI-B 
E. plicata Meerow subsp. plicata PLI-P 
E. sanderi Baker SAN 
E. ulei Kránzlin ULE 
Caliphruria hartwegiana Herbert HAR 
C. korsakoffii (Traub) Meerow KOR 
C. subedentata Baker SUB 
C. tenera Baker TEN 
*Pancratium L PAN 
Urceolina Reichb. URC 
Lepidochiton Sealy (Hymenocallis 
quitoensis & H. heliantha) LEP 


(Flory, 1977; Goldblatt, 1976; Meerow, 1984) via 
duplication or fragmentation of a chromosome, fol- 
lowed by doubling of the genome (Lakshmi, 1978; 
Sato, 1938). All paleotropical pancratioid genera 
have 22 or 20 chromosomes (Ponnamma, 1978; 
Zaman & Chakraborty, 1974). 

Pancratium (ca. 17 species) and two species of 
Hymenocallis (H. quitoensis Herbert and H. he- 
liantha Ravenna) were used as the primary out- 
groups in the analyses presented. Character state 
data on these taxa were accumulated from study 
of living material, herbarium specimens and liter- 
ature (Björnstad, 1973; Meerow & Dehgan, 1985; 
Morton, 1965; Ponnamma, 1978; Ravenna, 1980; 
Traub, 1962; Werker & Fahn, 1975). Pancra- 
tium is putatively the least derived genus of the 
pancratioid base (Meerow, 1985). It is the only 
paleotropical genus of the group, and the o 
member with 2n = 22 (Ponnamma, 1978), con- 
sidered by most workers as the ancestral somatic 
chromosome number in the family (Flory, 1977; 
Goldblatt, 1976; Meerow, 1984). 

The two species of Hymenocallis are undoubt- 
edly a monophyletic group. They are the only two 


mation series for cladistic analysis of Eucharis and Cal- 
iphruria. * = unordered multistate character. A linear 
transformation series is implied for all ordered multistéte 
characters except 41. 


: Au. linear or lorate, sessile; 1, petiolate. 
: 0, wp ae 1, deciduous and hysteran- 
thous; "d persistent. 3. Leaves: O, plicate; 1, smooth. 
4. Leaf margins: a nonundulate; 1, undulate. 5. Petiolar 
secondary bundles: 0, absent; 1, present. 6. Abaxial cu- 
ticular striations: 0, absent or nearly so; 1, dense, well 
developed. 7. Epidermal cell anticlinal walls: 0, more or 
less straight; 1, undulate. 8 wers: O, more than 7 
ong; 1, 5-7 cm long; 2, less than 5 cm long. 9. Floral 
fragrance: 0, heavy; 1, mild; 2, absent. 10. Flower num- 
ber: 0, more than 5; 1, ca. 5; 2, less than 5. 11*. Flower 
color: 0, white; 1, white or yellow; 2, yellow or orange. 
12*. Flower habit: O, erect or suberect; 
pendent by curving of tube; 2, declinate or pendent b 
xness of pedicel. 13. Pedicel length: 0, flower 
(sub)sessile; 1, less than 0.5 cm; 2, greater than 0.5 cm 
long. 14. Tube habit: 0, straight; 1, curved. 15. Tube 
length: 0, longer than tepals; 1, equal to or shorter than 
tepals. 16. Tube color: O, green; 1, concolorous with 
tepals. 17*. Tube morphology: 0, cylindrical proximally, 
dilating at 142-144 of its length; 1, funnelform, dilating 
gradually from the base; 2, cylindrical for most of its 
length, dilating abruptly near the throat. 18*. Perianth 
inis gy: 0, haere r. 1, campanulate; 2, funnel- 
for . 19. Staminal cup: 0, well developed; 
k ck 20. pec pigmentation: 0, present; 1, 
absent. 21. Staminal cup: 0, nonplicate; 1, plicate. 
22. Staminal cup: O, shallowly cleft between stamens ( « 
2 mm); 1, deeply cleft (> 2 mm). 2 ree filament: 0, 
linear (< 1 mm wide); 1, subulate (> 1 mm wide). 
24. Staminal dentation: O, present; 1, polymorphic; 2, 
absent. 25. Staminal teeth: 0, shorter than free filament; 
1, more or less equal to free filament; 2, longer than free 
filament. 26. Anthers: O, versatile at anthesis; l, erect 
at anthesis. 27. Pollen longest equatorial diameter: 0, 
greater than 80 um; 1, 76-80 um; 2, 66-75 um; 3, 
6 um; 4, 50-59 um. 28. Pollen exine reticulation: 
O, coarse; 1, moderately coarse; 2, fine. 29. Exine di- 
morphism: 0, present; 1, absent. 30. Style exserted: 0, 
greater than 1 cm beyond anthers; 1, 1 cm or less beyond 
anthers, but above the rim of the staminal cup; 2, to or 
below the rim of the staminal cup. 31. Stigma: 0, tri- 
lobed; 1, capitate, entire. 32. Stigmatic papillae: 0, uni- 
cellular; 1, multicellular. 33. Ovary color: 0, green; 1, 
white. 34. Ovary: 0, rostellate; 1, not rostellate. 
35. Ovule number (per locule): 0, 10-20; 1, 7-10; 2, 
3-6; 3, 2-3. 36. Ripe capsule: 0, green, relatively thin- 
walled; 1, leathery, thick-walled, orange. 37*. Seeds: 0, 
compressed; 1, globose or ellipsoid; 2, narrowly oblong, 
curved. 38*. Testa: 0, black; 1, blue; 
39. Testa: 0, dull; 1, lustrous. 40. 
1, rugose. 41. Somatic chromosome MA 
22. B (10000), 34; € (01000), 46; D (01010), 68; E 
(01100), 92; F (01101), 138. A > B A C > D, € 
> E, E-F 


1, declinate or 


; 3, urecola 


162 Annals of the 
Missouri Botanical Garden 
TABLE 3. Character state matrix for cladistic analysis of Eucharis and Caliphruria. Refer to Table 1 for key to 


EU abbreviations, to Table 2 for characters. 9 — missing or inapplicable character state data. * 


— outgroup. 


Character 


m 
c 
o 
ES 
en 
-1 


— 
=j 
— 
œo 
— 
ño) 
N 
© 
N 
— 


10 11 12 


— 
w 
— 
> 
— 
en 
— 
a 


E 

t 

x 
CORP mom Re Re ee 
COFP FP NNNNNNNNNNNNNNNNNNNNN IN 
cOoococooc-oooo-—-o—-—-—-ocoo--ooc|o 
cOocoocoooco-—-—-oo0-—-o---ooo-ecec- 
cOOcococooooooocooooooococoooooooc-- 
coc—-o0—-—-o0-—--—--—-o0o0-—--—-o0—-—--—-o--—---—-oo|o 
cOoc—-o—-oo0-—-—--—-oo0—--—-o0—-—--—----—--—-o0- 
DOOruNNN—NNN=rrn=nN-= ==" NOOO! 
CO Oro to to tolo ló—tlo oO —to Oto—tototo—oocoo|oc 


cCO—uwvcococococoooooocoooooocoocooco 
CON NNNNNNNNNNNNK NNR NNN e = - 
SOOO COOP HR HRP HB HR He Re eR eR RP RR RP Re Re He ee 
cOocc—-oco-ocoococooooooocoooooooco 
cOoc—-oocooc-—-—--—-o0o0—-—-—-—--—-—--—-—-—-oocc 
cct ——-—-—rt3t2t2t2t2t2t2t2clcrt2to2ot2t2ot2t2coc coc 
cOOcw—-—---oocoooooo-ooooocowwuwo 
Dorrerrrocoroocoorooooonor oo 
cOc—-—-—-—-—-oocoooooocooooooooco 
cOOcooo0ooo-—-oooo-o-ooo-ooco 


onoooocoorocororonoccrr O Nr 
C C to B2 b2 b2 R2 — — — — — e e ke Á Á- e o o oe om mM mM 


species of Hymenocallis with phytomelanous seed 
coats. All evidence suggests that possession of phy- 
tomelan is the ancestral condition in the Amaryl- 
lidaceae, and despite several independent losses, 
no reversal to the ancestral state has been proposed 
(Dahlgren & Clifford, 1982; Dahlgren & Ras- 
mussen, 1983; Dahlgren et al, 1985; Huber, 
1969). Hymenocallis quitoensis was segregate 
into a separate genus, Lepidochiton Sealy (1937) 
on the basis of phytomelanous seed. Both H. qui- 
toensis and H. heliantha are ephemeral compo- 
nents of the xeric flora of southwestern Ecuador 
and northwestern Peru, and differ only in flower 
color. Traub (1962) considered H. quitoensis a 
relict and the most primitive species of Hymeno- 
callis on the basis of ovule number (ca. 20 per 
locule; 2-10 occurs in all other species) and chro- 
mosome number (2n — 24, based on Snoad, 1952); 
however, collections from Peru and Ecuador that 
have examined have 2n — 34. These are the 
lowest reported chromosome numbers in the genus 
(Flory, 1976). These species will be called Lepi- 
dochiton in the following discussion. Additional 
analyses were run using other members of the 


pancratioid base as outgroups (Hymenocallis subg. 
Hymenocallis, H. subg. Ismene (Salisb.) Baker ex 
Traub, Pamianthe, and Paramongaia). The re- 
sulting cladograms were either equal in length or 
slightly less parsimonious than those presented here; 
they did not show changes in the resolution of 
terminal taxa from the cladograms presented. 


Ordered characters. Ten of the 18 multistate 
characters (Table 2) used in these analyses (8, 9, 
0, 13, 24, 25, 27, 28, 30, and 35) were ordered 
into linear transformation series. Study of char- 
acter state correlations among all genera of the 
pancratioid Amaryllidaceae (Meerow, 1985, 1987a, 
c) strongly suggests that flower size reduction (8), 
loss of floral fragrance (9), flower number (10), 
pedicel length (13), staminal tooth length (25), style 
length (30), and reduction in ovule number (35 
have evolved in this stepwise manner with no evi- 
dence of reversal. Numerous ovules per locule (20 
or more) characterizes all basal genera of the pan- 
cratioid Amaryllidaceae and is generally considered 
the ancestral condition for the family (Traub, 1962, 
1963; Traub & Moldenke, 1949). Though the 


Volume 76, Number 1 
1989 


Meerow 163 
Eucharis and Caliphruria 


TABLE 3. Continued. 


Character 


h 
N 
N 
w 
h 
an 
N 
en 
N 
[ey 
N 
~ 
N 
e 
N 
No) 
Ww 
[e] 
Ww 
— 


32 


w 
[e] 


37 


w 
œ 
w 
No) 
eA 
ED 


33 


w 
NN 
o 
on 


ceoocooocooococooocoodco 


200oO0NNOrooror=ronorroonmoo 
DO0ONVVrorno-ooovooroovvwvoo 
DDD --=P»>»rr> === - ooo 
CO O RRORGRGRIS UNIS SOIT OO BI t 09 — O20 MN 
OcOc-—ww-wcococo-oooooooooooocooc 
SOCORFP KF OPE RFP OOP RP RP Be HP Hh He Hee HOR He 
SOCP RP RP EP EF NNKF RFR eH eEDNKH OPH HCC O 


coc o0ooooo0-coccoco—-——-cococo--—-o-o0cc 
0000000 r--O00rrm- mim eB ee Hr HOOF 


ooococooocqcjeo 


© 


O o o- pes a D 


Oooococoooooooooooo-ocoo-ooc 
SCOP RF RP Re Pe RP eR OR eR Be eH ee ee HEH OCC OO 
SCOOSOWNNWWNHR RK KK WNNNNWWeE wooo 
oocoococoooco-—-—-oo-—-o—-o-----ooco0 
O-—twoo-o0-—--—--—-o0o-—--—-o0-o---o-56o0-2o 
Oto»oOoomoocooooco—-oooooocooomnvo 
SOF OSD CORP RFR OC OF KF OH ORR HHH GO 
Ooooco—-ooooooooooco-ococoooo-oco 


evidence from phenetic studies of Eucharis (Mee- 
row, 1987d, in press) and from study of other 
genera of the pancratioid Amaryllidaceae (e.g., 
Meerow, 1987c) indicates that androecial char- 
acters are among the most plastic morphological 
characters in this group, examination of the evo- 
lutionary patterns of staminal dentation (24) within 
various genera (Meerow, 1987c, in press) suggests 
that a reversal to the dentate state is unlikely once 
dentation is lost completely. Complete loss of stam- 
inal dentation often accompanies overall phenetic 
divergence among the pancratioid genera (Meerow, 
1985, 1987c; see E. astrophiala in this paper). 
Polymorphism for the character is not uncommon, 
however, and is conceived in the transformation 
series as the intermediate between the two fixed 
states of toothed or edentate stamens. Ordering of 
pollen size (27) and exine morphology (28) is based 
on data presented in Meerow (1985, 1987c) and 
Meerow & Dehgan (1985), as well as unpublished 
palynological data accumulated for many genera 
in the family. A single character, chromosome 
number (41), was coded in additive binary on the 
basis of much literature on amaryllid cytology (Flo- 


ry, 1976, 19777; Goldblatt, 1976; Lakshmi, 1978; 


Meerow, 1984, 1987b; Ponnamma, 1978; Sato, 
1938; Snoad, 1952), which strongly supports this 


concept of chromosome number evolution. 


Results and discussion. 
monious trees without any differences in topology 
resulted when either Pancratium or Lepidochiton 
was used as the outgroup. The twelve trees differed 
only in the branch lengths of a few internal nodes, 
and resolution of terminal taxa did not change 
among them. One of the twelve with Pancratium 
used as outgroup is illustrated and discussed in 
detail (Fig. 65). The cladogram has a length of 

2.4 steps (fractional lengths are a result of weight 
scaling), and a consistency index (Cl; Kluge & 
Farris, 1969) of 0.487. Lepidochiton is the first 
terminal EU to resolve in the cladogram after the 
outgroup, Pancratium. Character state changes 
from the outgroup to the ancestral node of Lepi- 
dochiton (flower number reduction, globose seeds) 
could not be polarized. Uniflory, perianth color 
polymorphism, brown seed testa, and 2n = 34 are 
the four apomorphies of Lepidochiton. The next 
internal node of the cladogram is the hypothetical 
ancestor to Eucharis, Caliphruria, and Urceolina, 


Twelve most parsi- 


164 


Annals of the 
Missouri Botanical Garden 


moea AMA  7* 23* 27+< 41 Eucharis subg. 
m 29* 40* . 
| l MOO  18* 38% Heterocharis 
| 
SAN 3* 10+ 18* 19* 24* 27* 
1 8* 33* 
2 L——424* 27« 
4 | 10+ 13*« 17 18* 19* 35« 41 
12 r- d 
m 13 | r—— CYA 38 
| 14 |10« t- - 4 
| 27 5* L - - ULE 
| 31< | 
| 41 9*| BON 24+< 27* 
| m 35*5——43« 4*« 41 
| | | BOU  6* 13*« 25* 30*« 
| | | 6 Y 3* | 
7* be---------- CAN . 
| | | 22 | en E. subg. Eucharis 
| L—__| 24 mH - FOR 
| | 39 | | 36 
| | | | mű BAK 4 27*« 
| | | | | 21 
| | | Ly 22« r CAS  3* 27 33* 36< 37* 39*« 40* 
| | | | 24+< ——]1 
| | | | | 8*| [——————— PLI-P 9* 25 27« 35« 
| | | 9* LLL] 27*0L——4 25* 
m~ 10* | m 10 |35 t----- PLI-B 
| | 37* | | | | 
| | | | | Mosen AA e deeem COR 
| | | | | 
| | | | | PASAS ce ae ee AS OXY 
| | | | 8 | | 4 
| | | | 13 | | 8* pmo mcn = AR 
| | | | 16 | | 9* m~ 24« 35* . . 
| | L 17 EA 18 28 TEN 2* 15* 25 Caliphruria 
| | | 26 | 19* |16< | 29* 37* 40* 
| | | 30 | 20 pool fo ee 
| | | 34 | 28 | 12 |18 24* 25* 32 35+ 39*« 
| | | 35 7< 15* 38* 
| | | | 2 
| | | URC  2* 8< 11 24+< 26« 30*< 31 35« 37 
| | | 
| | i LEH  24*« 25*-«-K E, subg. Eucharis 
| | 
| ETC LEP 10+ 11 38* 41 
| 
L PANO 
« - reversal 
*, + - parallelism 
- outgroup 
FIGURE 65. Cladogram of a and Caliphruria, based on data matrix in Table 3. Broken horizontal line 
8 P 


me zero-length branch. Numbers inside refer to apomorphies of adjacent EU or internal node. Refer to Table 


r EU designations. 


all of which thus form a monophyletic group. Nine 
apomorphies occur at this node. The most impor- 
tant are the evolution and fixation of the evergreen, 
petiolate leaf; the changes in flower habit, pedicel 
length, tube morphology, and pollen size; trilobed 


my concept of subg. Heterocharis, E. amazonica 
and E. moorei, linked by three apomorphies: sec- 
ondary petiolar bundles, loss of pollen exine di- 
morphism, and rugose testa (the state for the 
last-mentioned character is unknown for E. ama- 


zonica). The larger monophyletic group represents 
E. sanderi, Eucharis subg. Eucharis, Caliphru- 
ria, and Urceolina. Five apomorphies define the 
ancestral node: densely striate abaxial cuticle, un- 
dulate anticlinal walls (homoplasious with E. ama- 
zonica), deeply cleft staminal cup, polymorphism 
in staminal dentation, and lustrous testa. It is thus 
obvious that subg. Heterocharis is paraphyletic 
according to this cladogram. Eucharis moorei and 
E. amazonica also Pu cladistically as the most 
basal species of Euchar 

Eucharis sanderi is “cladistically basal to the 
remaining EUs with which it forms a monophyletic 


Volume 76, Number 1 
1989 


Meerow 165 
Eucharis and Caliphruria 


group. The six apomorphies of E. sanderi are 
plicate leaves, reduction in flower number, cam- 
panulate perianth (homoplasious with E. moorei), 
reduced staminal cup, loss of staminal dentation, 
and yet further reduced pollen grain size. 

The next monophyletic group is supported by 
eight apomorphies and comprises Eucharis subg. 
Eucharis, Caliphruria, and Urceolina. Apomor- 
phies at the ancestral node are reduced flower size, 
increased pedicel length, loss of green tube pig- 
mentation, change in tube dilation, erect anthers, 
decreased style exsertion, nonrostellate ovary, and 

ecreased ovule 
cladistically basal to the remaining EUs in the 
cladogram and is defined by two apomorphies: a 
reversal to dentate stamens and an increase in the 


number. Fucharis lehmannii is 


length of the teeth. The remainder of the cladogram 
forms two monophyletic groups. All but two species 
of Eucharis subg. Eucharis comprise one, and 
Caliphruria, Urceolina, and E. oxyandra consti- 
tute the second. Only two apomorphies define this 
clade: reduction in floral fragrance and an increase 
in flower number. 

The three apomorphies at the ancestral node of 
the Eucharis subg. Eucharis clade are subulate 
free filaments (homoplasious with E. amazonica), 
loss of exine dimorphism (homoplasious with the 
ancestor of E. amazonica and E. moorei), and an 
orange capsule. Within this clade, two monophy- 
letic groups are evident. The larger of the two is 

efined by a single apomorphy, plicate leaves. Eu- 
charis formosa is the first terminal EU to resolve 
but terminates a zero-length branch. Eucharis can- 
dida also terminates a zero-length branch, the an- 
cestral node of which is defined by two apomor- 
phies: loss of floral fragrance and reduction in ovule 
number. The next monophyletic group encom- 
passes all species of subg. Eucharis with about five 
flowers and about two ovules per locule. The hy- 
pothetical ancestor of Eucharis astrophiala and 
E. caucana forms an unresolved polytomy at the 
ancestral node with E. cyaneosperma, E. ulei, and 
the hypothetical ancestor of tetraploid species £. 
bonplandii and E. bouchei. 

Three apomorphies define the ancestral node of 
the second clade within subg. Eucharis: a plicate 
staminal cup, a reversal to shallowly cleft staminal 
cups, and a reversal from polymorphism in staminal 
dentation to fixed dentation. Eucharis bakeriana 
is the first terminal EU to resolve, on the basis of 
two apomorphies: loss of leaf margin undulation 
and increase in pollen size. The remaining species 
are the smallest-flowered species in the subgenus. 
The remaining apomorphies at the ancestral node 
are a decrease in pollen size and ovule number. 


The first terminal EU to resolve is E. corynandra, 
from a zero-length branch. It should be noted, 
however, that two autapomorphies of this species, 
a short staminal cup and club-shaped free fila- 
ments, were not included in the data matrix. The 
ancestor of the two subspecies of E. plicata form 
a monophyletic group with E. castelnaeana on the 
basis of style length. Subspecies brevidentata ter- 
minates a zero-length branch. Kucharis castel- 
naeana has the greatest patristic distance of the 
three EUs (seven apomorphies) 

The final large clade constitutes Caliphruria, 
Urceolina, and the cladistically basal E. oxyandra. 
Apomorphies are nonundulate leaf margins, small 
flowers, loss of floral fragrance, urceolate corolla 
(which does not apply to E. oxyandra, the corolla 
shape of which is either crateriform or campanulate 
but could not be coded as one or the other with 
certainty), reduction of the staminal cup, loss of 
androecial pigmentation, and moderately coarse 
exine. The character state of E. oxyandra is un- 
known for four of these seven. Within this mono- 
phyletic group, Urceolina and the hypothetical 
ancestor of Caliphruria resolve as sister groups 
on the basis of three apomorphies: declinate or 
pendent flowers via pedicel habit, straight tubes, 
and small pollen grains. Nine and eight apomor- 
phies define Urceolina and the hypothetical ances- 
tor of Caliphruria, respectively. Within Cali- 
phruria, the sole representative of the genus outside 
Colombia, C. korsakoffii is the first terminal EU 
to resolve, on the basis of three apomorphies: 
straight anticlinal epidermal walls, short tubes 
(homoplasious with C. tenera), and brown testa. 
Caliphruria subedentata terminates a zero-length 
branch from the hypothetical ancestor of the Co- 
lombian species of the genus. Apomorphies at the 
ancestral node are finely reticulate exines, loss of 
exine dimorphism, compressed seeds, and rugose 
testa (the states for the last two characters are 
known only for C. subedentata). Caliphruria hart- 
wegiana and C. tenera form a monophyletic group 
on the basis of toothed stamens and reduced ovule 
number, but C. hartwegiana terminates a zero- 
length branch. 

The clado 
70 parallel origins of character states and 29 re- 
versals (unordered characters not included). This 
may raise questions concerning the homology of 
characters that manifest low consistency. If the 
data matrix is analyzed with all characters removed 
that exhibit three or more independent origins for 
the same state (characters 3, 4, 8, 9, 10, 18, 19, 
24, 25, 27, 29, 35, 37, 38, 40), the three trees 


that result have a fourfold increase in zero-length 


gram presented in Figure 65 contains 


166 


Annals of the 
Missouri Botanical Garden 


branches and unresolved polytomies. Invariably, it 
is androecial characters (including pollen size) in 
which a great deal of homoplasy has occurred. 
Three of all the androecial characters, 24 (staminal 
dentation), 25 (length of teeth), and 27 (pollen 
size), are particularly homoplasious. When the data 
matrix was run with only these characters deleted, 
no change occurred in the resolution of terminal 
taxa from the cladogram in Figure 65. Finally, the 
data matrix was analyzed with all staminal cup 
characters (19, 20, 21, 22, 23, 24, 25), and pollen 
size (27) deleted. The resulting three cladograms 
(differing only by one internal branch length) placed 
Caliphruria and Urceolina within a monophyletic 
group including E. bakeriana, E. bonplandii, and 

. bouchei, on the basis of a single apomorphy, 
nonundulate leaf margins. 

Regardless of how the character-state data are 
manipulated, a number of monophyletic groups 
appear particularly robust. That Kucharis, Cali- 
phruria, and Urceolina represent a monophyletic 
group is uncontested. The monophyly of Eucharis 
moorei and E. amazonica is also strongly sup- 
ported, and these two species are the most cla- 
distically basal of the eucharoid lineage. These 
species have the greatest number of putatively 
primitive characters and seem to represent the 
more ancestral species in the genus. Nonetheless, 
according to the cladogram in Figure 65, retaining 
subg. Heterocharis in Eucharis renders the genus 
paraphyletic. Subgenus Heterocharis is a fairly 
heterogeneous group from the perspective of apo- 
morphic characters alone. Each of the three species 
may be characterized by autapomorphies, but only 
plesiomorphies join them. By including subg. Het- 
erocharis in Eucharis, Eucharis becomes para- 
phyletic according to this cladogram. However, it 
should be noted that it is unknown whether sec- 
ondary petiolar bundles (which require fresh ma- 
terial to observe), an important apomorphy which 
links E. amazonica and E. moorei, are present in 
E. sanderi. In the generalized sense, the distri- 
bution of subg. Heterocharis is quite broad, from 
Colombia to Peru (Fig. 64). However, each of the 
three species of subg. Heterocharis itself are only 
narrowly distributed. Fucharis sanderi is known 
only from the Chocó region of Colombia. Eucharis 
amazonica is found natively only in the middle 
Rio Huallaga valley of Peru. Eucharis moorei has 
been collected outside of Ecuador only once. More 
significantly, E. moorei is the only species of Eu- 
charis found on both sides of the Andes. I believe 
that all three species of subg. Heterocharis rep- 
resent the relictual remnants of the ancestral eu- 
charoid complex, each of which has remained iso- 


lated long enough to evolve its respective cohort 
of autapomorphies. 

The sister group relationship between Urceolina 
and Caliphruria also seems to be a robust cladistic 
Pee with E. oxyandra potentially ancestral 

ot 


S 
3^ 
Ie] 
> 
E 
m. 
an 
s À 
b: 
g 
3 
a 
i. | 
& 
a 
e 
- 
e 
z 
O 
5 
o 
E 
S 
| 
<= 


logical intermediacy (androecium and pollen mor- 
phology) between Eucharis and Urceolina. it is 
known only from bulbs found in local, transient 
cultivation in Peru, near the single recorded point 
of geographic sympatry between Eucharis (E. 
amazonica) and Urceolina (U. microcrater). The 
absence of much character state data, but partic- 
ularly those on fruit and seed morphology, further 
occludes resolution of its phylogeny, and any of 
several hypotheses concerning its origins are pos- 
sible (see Taxonomic Treatment). Kucharis leh- 
mannii, which usually resolves within a cladistic 
“no man’s land" between paraphyletic subg. Het- 
erocharis and all other clades, presents a similar 
problem, though Regel’s (1889) figure strongly 
suggests affinity to subg. Eucharis. In the initial 
analyses, in which all multistate characters were 
treated as unordered, this poorly known species 
resolved as sister group to E. bouchei. 1 believe 
that E. lehmannii is a sister species to E. caucana, 
to which it bears much phenetic resemblance, and 
that its unusual resolution in the cladogram is a 
result of the great deal of missing character state 
data. Fruit and seed morphology clearly place £. 
caucana in subg. Eucharis, even though it shows 
as much novelty as E. oxyandra or E. lehmannii 
in other characters. 

Excluding E. lehmannii and E. oxyandra, Eu- 
charis subg. Eucharis forms 
phyletic group. Species with five flowers and two 
ovules per locule (E. astrophiala, E. cyaneosper- 
ma, E. ulei, along with the tetraploids, E. bon- 
plandii and E. bouchei) form a very natural group 
with karyotypic similarity as well (Meerow, 1987b). 
The sister species relationship of Eucharis cau- 
cana to E. astrophiala within this group may be 
as equivocal as the basal resolution of E. lehman- 


1 1 
Cical 1) mono- 


nii, since it is based on two apomorphies (pollen 
size and lack of staminal dentation), the latter of 
which is highly homoplasious. I believe the reversal 
in ovule number necessary for E. caucana to share 
common ancestry with E. astrophiala is very un- 
likely. 

Eucharis candida and E. formosa, two often 
sympatric (and possibly sibling) species, are cla- 
distically close and basal to the five-flowered clade. 
The small-flowered species (E. castelnaeana, E. 
corynandra, and E. plicata) form another very 


Volume 76, Number 1 
1989 


Meerow 167 
Eucharis and Caliphruria 


r AMA 18 23* 27+< 
5 6*< 29* 40* Euc hats subg. 
VR MOO 7*< 38* Heterocharis 
[9< 10* 13*« 17* 18* 34< 
TUI SAN 3* 10* 19* 24+ 27* 
f AST  8* 24# 33* 
| 
35 24@< 27* 
4 | m 3< 4*< 41 
12 | | L.— — BOU  6*« 13*« 25+ 30< 
aues ite LL 27 
31*« p —34 10* | r——— CYA 38 
27 Las 
3* LE C Aun, 
Qe 
22 r CAU 10+ 13*< 17* 18* 19* 24# 
16* |24*  L————125* 44 45 
23* 35 L LEH 9< 23+< 248« 35 
26* 
29* i CAN 
30% L--] 
35 L FOR 9< 35 
12 6 36 
39* y BAK 4*« 27+< 
12 13 17 | 27 33* 36 
27 34 41 | Fra CAS E. subg. Eucharis 
L——— 21 974 39« 40* 
| |27* COR  30< 
| 30 
| | 135 r PLI-P 9* 25+ 27« 35 
L—————] 8* L—— —]25* 
| leo o o o > PLI-B 
| 
L- OXY 19* 23+< 24* 28* 29 
mm3 || | p--2---------- HAR 
15« 
mH l SUB 24+ 35 A : 
liis 15|29* 35 37* 40* Caliphruria 
18! TEN 2* 25+ 
|9* 18 ES 26* 30* 31*< 32 35 
19* pes KOR 7*« 24+ 38* 
(272 
i URC — 2* 11 16* 37 39* 
LEP 10 11 38* 41 
PANO 
« - reversal 
*, +, €, # = parallelism 
o - Outgroup 
FIGURE 66. User-generated cladogram of Eucharis and Caliphruria. Broken horizontal line indicates zero-length 


branch. Numbers inside refer to apomorphies of adjacent EU or internal node. Refer to Table 1 for EU designations. 


natural group. The position of E. bakeriana as 
basal to this clade, largely on the basis of a plicate 
staminal cup, may be equivocal. Karyotype anal- 
ysis (Meerow, 1987b) suggests putative relation- 
ship of E. bakeriana to E. formosa. 

An alternative hypothesis of phylogenetic rela- 
tionship is presented in Figure 66. The cladogram 
is a user-generated tree in which Caliphruria and 
Urceolina are placed as sister groups to a mono- 
phyletic Eucharis. The topology of subg. Eucharis 
in this cladogram represents my concepts of their 
evolutionary relationships based on phenetic re- 
semblance, phytogeography, and chromosome data. 


The major differences from the cladogram in Figure 
65 are the sister species relationship of E. caucana 
and E. lehmannii, the ancestral node of which 
forms a monophyletic group with the five-flowered, 
two-ovuled species complex; and the positioning of 
E. oxyandra as part of the small-flowered species 
group of subg. Eucharis. This user-generated 
cladogram had a length of 100.6 steps, with a CI 

of 0.443. There are 82 parallelisms represented 
among the character state changes, and re- 
versals. The user-generated tree (Fig. 66) is thus 
somewhat less parsimonious than the cladogram in 
Figure 65. Of particular interest in the user-gen- 


168 


Annals of the 
Missouri Botanical Garden 


erated cladogram is the resolution of the larger 
monophyletic groups constituting genera or sub- 
genera, and the accompanying character state 
changes within these clades. 

In Figure 66, a monophyletic subg. Hetero- 
charis is supported by seven apomorphies, four of 
which are reversals (unordered characters not 
counted). Some of these reversals are very unlikely 
(e.g., reversal to larger, heavily fragrant flowers; 
see previous discussion of ordered characters) and 
would thus represent retention of ancestral states. 

In the user-generated tree, a monophyletic subg. 
Eucharis is supported by a greater number o 
apomorphies (eight in Fig. 66) than in the parsi- 
mony cladogram (three). Caliphruria as a mono- 
phyletic group is best supported by Figure 66 (ten 
apomorphies) versus eight in Figure 

The results of these various cladistic treatments 
illuminate the problems inherent in phylogenetic 
reconstruction of large, character state 
data sets, and particularly with taxa whose evo- 
lution has involved large-scale homoplasy. The user- 
generated cladogram in Figure 66 represents my 
preferred hypothesis of phylogeny within the Eu- 
charideae; nonetheless I cannot deny the com- 
pelling evidence that Eucharis, by the inclusion of 
subg. Heterocharis, may be paraphyletic. 

From an orthodox cladistic perspective (Wiley, 
1981), Traub's (1971) concept of a single poly- 
morphic genus, Urceolina, is supported by the 
criterion of parsimony. From a phenetic (and prag- 
matic) standpoint, I find this unsatisfactory due to 
the close phenetic relationship between subg. Het- 
ris. This relationship is most 
conspicuous if one compares Eucharis bakeriana 
and E. formosa (the most basal species of subg. 
Eucharis) with E. amazonica and E. moorei (subg. 
Heterocharis). | prefer to recognize Urceolina and 
Caliphruria as distinct genera, despite the atten- 
dant risk of a paraphyletic Éucharis. I do not 
believe that Urceolina and Caliphruria necessarily 


erocharis and Eucha 


share an immediate common ancestor—the three 
apomorphies that define their sister group rela- 
tionship in Figure 65 (reversal to straight tubes, 
flower habit, and smaller pollen grains) are homo- 
plasious within the Eucharideae as well as in other 
tribes of the Pancratioidinae (Meerow, 1987c; 
Meerow & Dehgan, 1985). Until further data allow 
a more accurate understanding of their relation- 
ships, I also prefer to maintain E. lehmannii and 
E. oxyandra as species of Eucharis subg. Eucha- 
ris, allied with the small-flowered species with nu- 
merous ovules per locule. Without study of living 
material, and particularly knowledge of fruit char- 


acters, I am unwilling to declare E. oxyandra a 
sister taxon to Urceolina and Caliphruria. 
Ashlock (1971, 1972, 1979, 1984) has re- 
peatedly argued for the acceptance of paraphyletic 
groups in evolutionary systematic classifications. 
He defined two subclasses of monophyletic groups: 
holophyletic, which contain all descendents of the 
stem ancestor, and paraphyletic, those which do 
not. Ashlock (1979: 449) rejected ““the automatic 
conversion of the cladogram into a classification.” 
Meacham & Duncan (1987) have elaborated on 
the term *'convexity," first proposed by Estabrook 
(1978), which attempts to resolve the concepts of 
monophyly, holophyly, and paraphyly. Meacham 
& Duncan concluded that Ashlock's (1979) ad- 
vocation of overa luti y similarity as a com- 
ponent of classification schemes, rather than strict 
cladistic topology alone, is sound, and they dis- 
cussed the mitigating factors surrounding cladistic 
methodology (e.g., uncertainty of character state 
polarities, the speculative nature of phylogenetic 
reconstructions, among others) which compromise 


a direct translation of cladogram into classification. 


and Humphries & Chappill (1988) have responded 
vociferously to Cronquist's (1987) critique. Don- 
oghue & Cantino (1988: 108) assail paraphyletic 
higher taxa as “artificial classes created by tax- 
onomists who wish to emphasize phenetic 'gaps', 
and that formal recognition of such taxa conveys 
a misleading picture of common ancestry and char- 
acter evolution." 

The enormous amount of homoplasy evident 
within the highly canalized Amaryllidaceae creates 
great difficulty in phylogenetic reconstruction 
(Meerow, 1985, 1987c). In the analyses of Eu- 
charis and related genera, missing character state 
data further weaken the argument for adopting ad 
hoc the most parsimonious phylogeny as the basis 
for classification. At present, I prefer to accept a 
less parsimonious phylogeny as represented by the 
user-generated topology in Figure 66 until such 
time as cladograms can be generated with consis- 
tency indices greater than 0.5. Towards this end, 
work is continuing with careful application of se- 
lected character weighting (Neff, 1986; Wheeler, 
1986). 

In three neotropical lineages of the Pancratioi- 
dinae, parallel trends in the evolution of floral mor- 
phology have occurred (Meerow, 1985, 1987c). 
In each case, taxa with smaller, tubular or ven- 
tricose, brightly colored flowers with reduced stam- 
inal connation, and without noticeable fragrance 


Volume 76, Number 1 
1989 


Meerow 169 


Eucharis and Caliphruria 


have apparently diverged from taxa possessing a 
large, white, fragrant, crateriform flower with a 
staminal cup (Meerow, 1985, 1987c). Each lin- 
eage appears to be a monophyletic group on the 
basis of vegetative and ovarian morphology, as well 
as chromosome number (Meerow, 1985, 1987c; 
Traub, 1963). A similar pattern occurs in all three 
lineages: 1) floral morphology of “derived” taxa 
3 gap an Lo TR pollination syndrome, 
and 2) "derived" taxa are found, entirely or in 
part, at iid elevations than apparent ancestral 
taxa. Eucharis and Urceolina, as documented in 
this paper, present one such case. Pseudosteno- 
messon Velarde, submerged by Traub (1980) into 
Hymenocallis as section Artema, presents a par- 
allel situation within the Hymenocallis lineage 
(Meerow, 1985; Meerow & Dehgan, 1985). Tribe 
Stenomesseae contains two small genera, Pa- 
mianthe and Paramongaia, with ancestral floral 
morphology and contains a large genus, Steno- 
messon, with derived morphology (Meerow, 1987c). 
In the Hymenocallis and Eucharis lineages, the 
pancratioid floral morphology has radiated to a far 
greater extent than the putatively ornithophilous 
divergence (Pseudostenomesson an rceolina, 
respectively). In the Pamianthe- Paramongaia/ 
Stenomesson lineage, the derived genus, Steno- 
messon (35-40 species), has speciated to a great- 
er degree than taxa with the ancestral pancratioid 
flower (Pamianthe: 2 species, Paramongaia: 
monotypic). 

he high level of divergence in neotropical pan- 
cratioid tribes may be primarily a factor of two 
causes, 1) the uplift of the Andes in the Pliocene 
(van der Hammen, 1974, 1979), creating much 
opportunity for geographic isolation, and 2) greater 
genetic adaptability via tetraploidy to new ecolog- 
ical zones. The occurrence of rare hybridization in 
nature between the relict complex represented by 
Eucharis subg. Heterocharis and both Caliphru- 
ria (X Calicharis butcheri) and Urceolina (x Ur- 
ceocharis edentata) may be further indication that 
the evolution of these genera was geologically re- 
cent. 


TAXONOMIC TREATMENT 
SUBSPECIFIC TAXA 


The rank of subspecies is used once in this 
treatment (E. plicata) to designate strong mor- 
phological divergence coupled with geographical 
isolation that I accept within specific limits. The 
rank of variety is used in a case (E. bouchei) where 
both the morphological and geographic components 


of divergence are weaker but appear to represent 
the first stages of speciation within a tetraploid 
semispecies complex (Meerow, 19874). 


CRITICAL MEASUREMENTS 


Measurements of vegetative and floral parts in 
the following species descriptions are derived from 
examination of dried material (floral parts after 
rehydration in 3% solution of Aerosol OT brought 
to boil) supplemented with examination of fresh or 
FAA-preserved material when available. 1 found 
less than 5% difference between measurements of 
dried and fresh or spirit-preserved material of the 
same collection when available. 


A NOTE ON THE KEYS 


Species of Caliphruria and penas subg. Het- 
erocharis are separable by di ters very 
amenable to key construction. dia the keys to 
these two groups, any worker should be able to 
identify material referable to described taxa wheth- 
er in the field or the herbarium. The enormous 
degree of variation exhibited by species of subg. 
Eucharis, however, made key construction difh- 
cult. A number of the species overlap to at least 
some degree with one or more other species in 
virtually all quantitative morphological characters. 
As a consequence, sometimes I have had to rely 
on characters observable only with living material. 
With perseverance, it should be possible to key out 
all but the most depauperate herbarium specimens 
of subg. Eucharis. Collectors should note such 
characters as leaf plication and undulation, floral 
fragrance, tepal habit, and pigmentation and pli- 
cation of the staminal cup in their notes when 


preparing specimens of Eucharis. 


KEY TO EUCHARIS AND CALIPHRURIA 


la. Leaf margins usually undulate; flowers declinate 
he tube; peri- 


tube cylindrical, at least below middle, abruptly 
dilating at or above the midpoint of its length, 
curved, 25-50 mm long; staminal cup con- 
spicuous, basally pigmented green or yellow, 
exserted from the throat of the perianth or 
adnate to the dilated portion of the tube; stig- 
matic papillae unicellular ................................ Eucharis 
Leaf margins nonundulate; flowers declinate to 
subpendulous via the curving of the pedicel; 
perianth funnelform; tube funnelform, dilating 
gradually from base (rarely subcylindrical), 
straight or only slightly cernuous, 25 mm or 
less long; staminal cup inconspicuous and un- 
pigmented, reduced to a membranous, basal 


E 


170 


Annals of the 
Missouri Botanical Garden 


connation of the filaments; stigmatic Aem 
multicellular Caliphruria 


Eucharis Planchon et Linden, Linden's Ann. Cat. 
Hort. 8: 3. 1852; Fl. des Serres Jard. Eur. 
Ser. 1, 8: 107. 1853. TYPE: Eucharis candida 
Planchon et Linden. Urceolina subg. Eucha- 
ris (Planchon & Linden) Traub, Pl. Life 27: 
57-59. 1971. 


Evergreen, bulbous geophytes of rain forest 
understory. Bulb tunicate, usually offsetting vig- 
orously. Leaves petiolate, persistent, glabrous; pet- 
iole subterete, somewhat concave adaxially proxi- 
mal to the sinus, convex abaxially, light green, 
winged distally by attenuation of the lamina; lamina 
ovate, elliptic, ovate- or elliptic-lanceolate, usually 
thin, predominantly hypostomatic, usually lustrous, 
dark green adaxially, light or silvery green abax- 
ially, smooth or plicate between the parallel veins, 
cuticle of the abaxial epidermis variably striate, 
margins frequently undulate, apically acute or acu- 
minate, basally attenuate to the petiole, rarely ap- 
pearing subcordate. Inflorescence scapose, umbel- 
late (composed of 1 to several reduced helicoid 
cymes); scape solid, terete or slightly compressed, 
glaucous, terminating in 2 green or greenish white, 
ovate-lanceolate, valvate-imbricate, marcescent 
bracts enclosing the flowers and several secondary 
bracts before anthesis. Flowers 2-10(-12), pedi- 
cellate (rarely subsessile), each subtended by a lan- 
ceolate bracteole, pendent or declinate, sometimes 
fragrant, white, usually protandrous; perianth cra- 
teriform or campamiate;, tube cylindrical; dilating 
abruptl green 
proximally; limb of 6 tepals i in 2 series spreading 
widely from the throat or imbricate for half their 


length, the outer series usually longer, narrower 
and apiculate; the inner series acute, obtuse, or 
minutely apiculate. Stamens 6, variously connate 
below, pigmented yellow or green proximally; free 
filament linear, subulate, or otherwise petaloid; an- 
thers oblong to linear, sub-basifixed or dorsifixed, 
eventually becoming versatile, introrse, dehiscing 
longitudinally; pollen grain boat-shaped elliptic, 
monosulcate, the exine reticulate. Style filiform, 
included within, equal to, or exserted from the 
staminal cup; stigma obtusely 3-lobed, papillose. 
Ovary inferior, green or white, globose-ellipsoid, 
oblong, occasionally trigonous, 3-locular; septal 
nectaries present; ovules 2-20 per locule, placen- 
tation axile, globose, anatropous. Fruit a 3-lobed, 
loculicidal capsule, thin-walled or leathery, green 
or bright orange; seed globose or ellipsoid, some- 
times angled by pressure, turgid, with copious en- 


dosperm, few per locule; testa thin, phytomelanous, 
lustrous (rarely dull) black, dark brown or blue, 


usually smooth. Chromosome numbers: 2n = 46, 


KEY TO THE SUBGENERA OF EUCHARIS 


la. Flowers 3-7 cm long, usually pendulous, only 


drical below, dilated abruptly ju 
bo) (rarely in the upper %4), Nub strongly 

urved, white; limb segments usually spreading 
widely (c 90* de aa fia staminal cup 
inserted at throat of tu ximally pat 
or stained yellow W, piat or c yallow-ors ange IK 


E 


gigs of filament usually wide 
wide) or otherwise petaloid; anthers + 
erect at anthesis; ovules 2-9(-10) per locule; 
capsule leathery and bright orange (rarely green 
in which case tardily dehiscent) 
Eucharis subg. Eucharis 
lb. Flowers 7-8 cm long, declinate or subpendu- 
lous, strongly fragrant, funnelform-campanu- 
l 


anthesis; ovules (7, 9-)16-20 per locule ...... 
Eucharis subg. Heterocharis 


Eucharis subg. Eucharis 


Leaves glabrous, petiolate, persistent; lamina 
ovate, elliptic, or lanceolate, mostly thin, margins 
usually undulate, variably plicate between the par- 
allel veins, apically acute or acuminate, basally 
attenuate to the petiole or rarely subcordate, mostly 
dark green and lustrous adaxially, light or silvery 
green abaxially, the abaxial epidermis variously 
striate; petiole subterete, somewhat channeled 
adaxially proximal to the sinus. Inflorescence sca- 
pose, umbellate, terminating in 2 greenish white 
marcescent bracts. Flowers pedicellate, (3-)5-10, 
only rarely with noticeable fragrance, mostly pen- 
dulous, 3-7 cm long, crateriform (rarely slightly 
campanulate); perianth tube usually strongly 
curved, cylindrical below, dilated just below the 
throat or rarely at Y length, white; limb of 6 white, 
ovate to ovate-lanceolate tepals usually spreading 
widely from the throat, often recurved above the 
middle, the outer 3 usually longer, narrower, and 
apically apiculate, the midvein often faintly yellow 
in strong light. Stamens connate into a conspicuous 


Volume 76, Number 1 


Meerow 
Eucharis and Caliphruria 


or rarely reduced staminal cup, usually exserted 
from the rim of the throat; staminal cup apically 
white, marked green or yellow (rarely yellow-or- 
ange) basally, variously toothed, lobed or entire; 
distal portion of the filaments petaloid and variously 
shaped; anthers oblong or linear, subdorsifixed or 
sub-basifixed, + erect at anthesis, finally becoming 
versatile; pollen grain 45-60 um (polar axis), 55- 

6 um (longest equatorial axis), the exine coarsely 
reticulate. Style filiform, white; stigma obtusely 
trilobed, glandular pubescent. Ovary globose, ellip- 
tic or trigonous, green or white; ovules globose to 


KEY TO THE SPECIES OF EUCHARIS SUBG. EUCHARIS 


la. Perianth tube 25 mm or more lon 


ellipsoid, axile, superposed, 1-12 per locule, most 
often 2-4. Fruit a loculicidal capsule, orange and 
leathery whán ripe (rarely remaining green); seeds 
1-3(-4) per locule, ca. 1 cm long, turgid, ellipsoid 
(rarely somewhat compressed), with a lustrous black 
or blue testa. 


Distribution. Fourteen species, (Guatemala) 
Costa Rica to Bolivia, concentrated on the lower 
slopes of the northeastern Andes and western Ama- 
zonas, usually below 1,000 m 


2a. Flowers (7-)8-10 (or very indi 5); ovules 3- : hice pns (or very rarely 2). 
3a. 


Flowers not fragrant; perianth tube (25- 
long; staminal cup 8-1 


mm long to apex e het or lobes; ovules (2-)3-5(-7) per pow" ERAN 
l. 


m long; outer tepals (20-)25-30(-33) mm 


ca 
3b. Flowers mildly E perianth tube 35-45(-50) mm long; outer S. (28-)32-45(- i mm 
mm long to apex of teeth or lobes; ovules 
4a. Floral ran c slightly fetid; flowers pendent; staminal cup less than 15 mm lon 
between each stamen, nonplicate; staminal teeth (if present) 


long; staminal cup 10- 


of teeth or lobes, cleft 3-5 mm 


2-4)7-9 per locule. 
g to apex 


much less than half the length of the subulate portion of the filament; style exserted ca. 1 
2. 


cm beyond the anthers 


E. formosa 


4b. Floral iere sweet; flower perpendicular to vertical axis of scape; staminal cup more than 


ong to apex of teeth, cleft very shallowly (less than 2 mm) be 


tween each stamen, 


wapa staminal teeth half the length of the subulate portion of the filament; EA. exserted 


s than 0.5 cm beyond the anthers 
2b. Flowers 2- 5C 7); ovules 2-3(-5) per locule. 
a. Leaves nonplicate, s 


. bakeriana 


somewhat succulent, margins nonundulate, length-to-width ratio usually less 


than 3; petiole usually shorter than the lamina; plants of central and western Colombia or Central 


America 


6a. Leaves slightly cage adaxially; abaxial ve densely striate; perianth tube 25-33 mm 


ong; staminal cup irregularly to 


toothed, proximally pigm 


mented pale yellow; stamens ses 


constricted sali Sis a narrow subulate ai style exserted à than 0.5 cm 
ombi 


anthers; ovary not trigonous; plants o 


ond 
ia 4. E. A 


6b. DER not glaucous; abaxial cuticle largely devoid of striations; perianth tube (25- Ln 45 


mm long; staminal cup m 


most frequently edentate, proximally pigmented green; stame 


s often 


pee gradually from apex to ae style exserted 0.5-1 cm beyond anthers; z often 


ous; plants of Central 


. E. bouchei 


trig eric 
9b. Leaves Rae: thin, margins undis EIE length-to-width ratio usually greater than 3; petiole 


usually equal to or longer than the 
7a. Leaves bullate-pustulate in texture 


lamina; plants o 
, nonlustro 


estern E Colombia, or Amazonas. 


of w 
rous; stamin. 


mally; stamens deltoid, dilating gradually from apex to Vus ovary white; plants of western 
6 


E. astrophiala 


cuador ; 

7b. Leaves not bullate-pustulate in texture, lustrous; staminal cup pigmented yellow or 

stamens not deltoid, constricted distally into a narrow 
plants of western Colombia or Am 


ovary gre 


green; 
(less than 2 mm wide) subulate portion; 


en; 
8a. Perianth "funnelform-campanulate, tube dilated distally at y donor Y4 its length; 


staminal cup inconspicuous; filaments connate proximally for 3 mm 


westani Cola om ia 


E plant of 
. E. caucana 


iE Eos ex d 


1 1 2 a + + 


8b. tub 


ie connate proximally fo for 


DCIOW i l P 
or more; plants of Amazona 


9a. Perianth tube curved gradually throughout the proximal half; SEIT cup usually 
bidentate but sometimes obtusely lobed (rarely one to several stamens quadrate); 
k 


ovary globose-elliptic at anthesis; testa lustrous blac 
. Perianth tube usually curved abruptly above the ovary, then more or less straight 
for the rest of its length; staminal cup usually quadrately lobed, but 


Ke) 
c 


8. E. ulei 


t one or several 


stamens occasionally toothed; ovary deeply trigonous at anthesis; testa lustrous blue 
Y: EE 


cyaneosperma 


lb. Perianth tube equal to or less than 25 (or very rarely to 30) mm | 
10a. 


aminal cup usually less than 1 cm 


m long to apex of teeth or pim but connate portion of filaments 
always much shorter than free, subulate portion. 


172 


Annals of the 
Missouri Botanical Garden 


11a. 
ovules 10 per locule, plants of western 
llb. 


Teeth of the staminal cup acutely long-lanceolate, as long as the subulate portion of the filament; 
la 10. 


E. lehmannii 


Teeth of the staminal cup (when present) obtuse, much shorter than the subulate portion of 


the filaments; ovules > al than 10 per locule; plants of the Peruvian east An 


12a. Staminal cup 3.5 


m long (to apex of teeth), obtusely bidentate between each stamen; 
free filaments club- la (appearing elliptic-lanceolate in dried material), 1.8-2 
l 


mm 
E. corynandra 


wide, ovules 4-6 per locule 


12b. 


Staminal cup reduced to a basal connation of the filaments 0.8-1.5 mm long, edentate 


or obtusely bidentate; free filaments narrowly subulate, ca. 1 mm wide; ovules - 8 per 
l 


] E. oxyandra 


10b. 


D 


with a lustrous, smooth black testa 


ocule 
Staminal cup usually greater than 1 cm long to apex of teeth or lobes, but connate portion of filaments 


m wide; staminal cup campanulate, gradually dilating distally, plicate along 
the filamental traces; ovary green; capsule leathery, bright orange, dehiscent; seeds elli 


. Leaves 3-6 c 


3. 
cm wide; staminal cup funnelform- duet to cylindrical, often abruptly dilated 
distally at 4%2-% length, plicate between the filamenta 
thin-walled, often oed dehiscent; seeds wedge- ral with a dull, rugose black testa 
l 


l traces; ovary whitish; capsule green, 


4. E. castelnaeana 


1. Eucharis candida Planchon et Linden, Lin- 
den’s Ann. Cat. Hort. 8: 3. 1852; Fl. des 
Serres Jard. Eur. Ser. 1, 8: 107. 1853. TYPE: 
ex hort. Linden, supposedly imported from 
Colombia, no other data, Planchon s.n. (MPU). 
Urceolina candida (Planch. & Lind.) Traub, 
Pl. Life 27: 57-59. 1971. Figures 67-69, 
70A. 


Bulb subglobose, 3-5(-6) cm long, 3-4(-5) cm 
diam., neck 1-2.5(-4) cm long, 1-2.5 cm wide; 
tunics brown. Leaves 1-2, petiole (15-)18-30 
(-35) cm long, ca. 7 mm thick proximally, ca. 3- 
4 mm thick distally; lamina elliptic, (18-)30-35 
cm long, (7-)8.5-11.5(-12) cm wide; acuminate, 
deeply plicate, dark green but only slightly lustrous 
adaxially, light green abaxially, abaxial cuticle 
striate, the margins coarsely undulate. Scape (4-) 
5-6 dm tall, 8-10 mm diam. proximally, 4-5 
mm diam. distally; bracts 25-45(-50) mm long, 
ca. 5-6 mm wide, ovate-lanceolate. Flowers (7-) 
8-10, rarely as few as 5, without noticeable fra- 
grance; pedicels (9-)15-20(-35) mm long; tube 
(25-)30-35 mm long, ca. 2 mm diam. for most 
of its length, abruptly dilated to (7-)10(-11.5) mm 
at the throat; limb spreading to 4.5-6 cm wide; 
tepals sometimes recurved distally, outer tepals 
(20-)25-30(-33) mm long, (9-)10.5-14(-15) mm 
wide, ovate-lanceolate, apiculate, the apiculum only 
slightly or obscurely tufted adaxially; inner series 
(20-)22-28(-32) mm long, (10-)12-15(-20) mm 
wide, ovate, acute to minutely apiculate. Staminal 
cup funnelform-cylindrical to slightly campanulate, 
rarely widely ampliate distally, (7-)8-11 mm long 
(to apex of teeth or lobes), (10-)13-16(-18 
wide, most frequently edentate and lobed between 
each stamen, at times bidentate or irregularly 
toothed, widely spotted green to greenish yellow 


) mm 


below each stamen; teeth, when present, 1-2 mm 
long, < 1 mm wide, acute or obtuse; cup cleft for 
(2.5-)3-5(-6) mm between each stamen; stamens 
(3.5-)4.5-5.5(-6) mm wide proximally; distal sub- 
ulate portion (3.5-)4.5-6(-6.5) mm long, 1.5-2 
mm wide at the base; anthers (3.5-)4-5(-6) mm 
long, oblong; pollen grain 46.8—50 um polar diam., 
68 um longest equatorial diam. Style (38-) 
45-55 mm long, exserted 0.5-1 cm beyond the 
anthers; stigma 2—d3 mm wide. Ovary globose-el- 
lipsoid, green, 5-7(-8) mm long, 4-6(-7) mm 
diam.; ovules (2-)3-5(- 7) per locule. Capsule (1.5-) 
1.8-2.4 cm long, (2-)2.5-2.9 cm wide, bright 
orange, leathery; pedicel 2-4 cm long; seeds 1- 
2(-3) per locule, ellipsoid, ca. 1 cm long, ca. 0.5 
cm diam., with a lustrous, smooth black testa. 
Chromosome number: 2n — 46 


Distribution. | Understory of primary rain for- 
est chiefly in the Oriente of Ecuador, particularly 
the Rio Napo valley, occasional in north Peru and 
southeast Colombia (Figs. 71, 72) at (100- 
180)240-550(1,000-1,600) m. Flowering at any 
time of the year, most frequently in February, 
March, and August. 

Vernacular name. — Cebolleta. 

Additional specimens examined. COLOMBIA. 
AMAZONAS: Puerto Naririo, 24 Jul 1965, Lozano et al. 
594 (COL); Trapecio amazonica, Loretoyacu River, ca. 
100 m, Oct. 1946, Schultes £ Black 8478 (US). META: 
sabanas de San Juan de Arama, margen izquierda del Rio 
Gúejar, alrededores del aterrizaje Los Micos,” 500 m 
22 Jan. 1951, Idrobo & Schultes 1208 (COL, GH, NY, 
US). PUTUMAYO: vicinity Mocoa, Rio Rumiyaco, 1,100 
m, 8 Oct. 1959, Barclay & Juajibioy 9461 (MO). 
ECUADOR. NAPO: Tena, wet forest, 27 Sep. 1939, Asplund 
8853 (S); El Napo, 1931, Benoist 4717 (P); Rio Yasuni, 
12 km from mouth at Rio Napo, 0%57'S, 75?25'W, 200 
m, 25-26 Aug. 1982, Balslev & Alarcon 2966 (QCA); 


Volume 76, Number 1 Meerow 173 
1989 Eucharis and Caliphruria 
FIGURE 67. Eucharis formosa ds 1103, FLAS) and E. eno (Dodson et al. 14095, m — A. Flowers. 


i, E. posue ii, E. candida. — 
tepal; iv, inner tepal. — C. inse] cups. i, E. formosa; 


formosa; ii, E. candida. 


10 km downstream from Coca at Anangu, 260 m, 8-11 
Aug. 1982, Besse et al. 1598 (SEL); Tena-Puyo road, 
550 m, Aug. 1982, Besse et al. 1643 (SEL); Coca- Lago 
Agrio road, 45 km north of Coca, Rio Palanda Yacu, 7 
June 1983, Bohlin & Bohlin ue wei Isla Pompeya, 
19 Apr. 1983, Dinua 42 (QCA); km 5, Cotunda-Coca, 
1,130 m, 19 June 1963, | ido et al. 14095 (SEL); 
orilla i sper del Río San Miguel, Puerto Nuevo, 26 

jin V. 2687 (COL); Santa Rosa at Río 
Neo, ca. ale eb. 1972, Pei | 11090 (GB); 
Hacienda Cot tapin e nm m, 19-20 Feb. 
1968, Harling et = 7121 (FLAS, GB, lower Rio Aguari- 


. Tepals. i, ii. E. formosa. i ou 


B 


er tepal; ii, inner tepal. iii, iv. E. c a. iii, outer 
dida. 


id 
il, E. can —D. Ovaries, ceiling section. i, E 


co (above Puesto militar Puerto Loja, 7 Mar. 1968, 
Harling et al. 7400 (GB); ye ¿rf and rastrojos 
near the village along roa o Agrio, ca. 250 m, 2 
Feb. 1974, Harling & M ol 11682 (GB); Canon 
de los Monos, ca. 12 km north of Coca, 250 m, 4 Feb. 
1974, Harling & Andersson 11719, FLAS specimen 
(FLAS); Misahualli at Río Napo, 28 Mar. 1969, Holguer 
925 (FLAS, GB); environs of Limoncocha, 240 m, June 
1978, Madison et al. 5326 (F, SEL); between Río Napo 
and Rio San Miguel, 0°30'S, 76-77°W, 300 m, Aug.- 
Sep. 1978, Navarrete 78. PASTAZA: Mera, forest on shore 
of Río Pastaza, ca. 1,000 m, 30 Jan. 1956, Asplund 


Annals of the 
Missouri Botanical Garden 


M ces 68, 69. 
. Flower with wide-spreading staminal cup.— 


19120 (Sy Puyo-Arajuno Road, 1-5 km SW Diez ed 
Agosto, ca. 900 m, 4 Feb. 1980, Harling & Andersson 
16862 (GB); 68 km north of Puyo on road to Tena, 
along creek, ca. 500 m, 26 July 1982, flowered in cul- 
tivation, 15 Jan. 1985, Meerow 1144 (FLAS). 
MORONA-SANTIAGO: Huamboya, 1,500-1,600 m, 15 Feb. 
Acosta-Solis 7469 (F). PERU. AMAZONAS: 400 m 

atras de La Poza, Rio Santiago, 180 m, 23 Aug. 1979, 
Huashikat 164 (MO). LORETO: Maynas, Río Ampiyacu, 
Pebas and vicinity, approx. 3?10'N, 71?49'W, behin 
Pebas on trail north of town, 10 Apr. 1977, Plowman 
et al. 6724 (F); Maynas, Santa Maria de Nanay, Colonia 
San Phen de Indies Yaguas, 1.5 km del Fundo Balcon, 
Rio ud 110 m, 15 Nov. 1984, Schunke 14155- 
B (F, FLAS 

anis hybrids with E. formosa: ECUADOR. NAPO: 
km 23 Lago Agrio- Baeza road, 350 m, July 1982, Besse 
et al. 1558 (SEL); 35 4 south of Rio Aguarico, Lago 
Agrio-Coca road, July 1982, Besse et al. 1563 Sen 
Rio Coca, 10 km eo from ferry crossing, 250 m 
28 Nov. 1983, Besse et al. 1949 (SEL). 


x 


a 


Eucharis candida was originally described from 
cultivated material supposedly originating from Co- 
lombia, a country in which it has been encountered 
only rarely. The species previously has been de- 
limited by the absence of staminal dentation; how- 


related species E. formosa and E. ulei. Peruvian 
populations of E. candida are variable in the shape 
of the staminal cup (Figs. 68, 69), even among 
flowers of the same inflorescence. 

Eucharis candida is most common throughout 
the upper Napo Valley in eastern Ecuador and is 
very often sympatric with the larger-flowered and 


Variation in a single Peruvian population of Eucharis candida pu 14155-B, FLAS).— 
69. Flower with cylindrical staminal c 


more widely distributed E. formosa (Figs. 71, 72). 
In one instance, two specimens (Harling & An- 
dersson 11719), one each of the two species, were 
collected under the same number. To date, no 
species of Eucharis other than these two have been 
collected north of the Pastaza valley in eastern 
Ecuador. On the basis of herbarium study alone, 
Ecuadorean populations of these two taxa form a 
mosaic that seemed taxonomically insoluble until 
living material of both species from several pop- 
ulations was collected and flowered. Phenetic and 
karyotype analyses (Meerow, in press) support their 
recognition as distinct species and suggest that E. 
E. formosa have hybridized in at 
least one area of sympatry. 

Eucharis candida may be separated from E. 
formosa by its smaller leaves and flowers, complete 


candida an 


absence of fragrance (E. formosa produces a mild, 
"sour" odor), and generally fewer ovules per locule 
(though both species are variable in ovule number). 
On the basis of known collections, Eucharis for- 
mosa is slightly better represented in the Pastaza 
valley than E. candida. 


2. Eucharis formosa Meerow, Sida 21: 29- 
49. 1987. TYPE: Ecuador. Morona-Santiago: 
Road Limón-Macas, ca. km 20 from Limón, 
eru rain forest and rastrojos, 700-900 

, 26 Mar. 1974, Harling & Andersson 
12915 (holotype, GB; isotype, FLAS). Figures 
67, 70B, 73-78 


Volume 76, Number 1 Meerow 175 
1989 Eucharis and Caliphruria 
1cm 
FIGURE 70.  Staminal cup variation in four species of Eucharis. — À. E. candida. i, Besse et al. 1598 (SEL); ii, 


Harling et al. 7400 (GB); iii, Schultes & Black 8476 (US).— B. E. formosa. i, Penland 142 (US); ii, Holguer 


2655 (CB); iii, Harling et al. 7201 ( 
120 (US). iii. Variety dressleri (Meerow 1107 


E. ulei. i, Plowman & Kennedy 5811 (GH); ii holotype, 


Bulb subglobose, 4-7 cm long, 3-5 cm diam., 
neck 2-5 cm long, ca. 1 cm thick, tunics brown. 
Leaves 1-2(-3); petiole 25-38(-42) cm long, 8.5- 
1 mm thick proximally, 5-6 mm thick distally; 
lamina elliptic, (21-)30-45(-52) cm long, (8-)1 1- 
15(-16) cm wide, usually conspicuously plicate, 
dark green and only slightly lustrous adaxially, light 
green abaxially, abaxial cuticle striate, margins 
coarsely undulate. Scape (5-)6- 7(-8) dm tall, ca. 
l cm diam. proximally, 5-6 mm diam. distally; 
bracts ovate-lanceolate, (36-)43-60(-85) mm long, 
10-15 cm wide at the base. Flowers 8-10, very 
rarely fewer, pendent, emitting a mild “sour” odor; 
pedicels (8-)12-18(-30) mm long; tube 35-45 
(-50) mm long, ca. 2-2.5 mm wide for most of 
its length, abruptly dilated to (9-)10—-13(-14) mm 
at the throat; limb spreading to (55-)60-70(-80) 


cm; tepals sometimes recurved distally; outer tepals 


). — C. E. bouchei. 
FLAS). iv. Variety darienensis (Gentry & Mori 13945, MO).— 


i, ii. Variety bouchei. i, Lewis et al. 2617 (MO); ii, Allen 
D 


Ule 5737A (By; iii, Schunke 1887 (F). 


narrowly ovate, (30-)35-45(-47) mm long, (10-) 
15-18(-20) mm wide, apiculate, apiculum con- 
spicuously horned adaxially (Ecuadorean popula- 
tions); inner tepals ovate, (28-)31-40(-45) mm 
long, (15-)18-22(-25) mm wide, acute to mi- 
nutely apiculate. Staminal cup funnelform-cylin- 
drical, 10-13(-15) mm long (to apex of teeth or 
lobes), (15-)17-20(-22) mm wide; flushed green- 
ish yellow proximally, with the greatest concentra- 
tion of pigment below each free filament, rarely 
only widely punctate; bidentate, irregularly toothed, 
lobed or quadrate between the distal portion of the 
filament; cup cleft between each stamen for 3-5 
mm; teeth when present acute or obtuse, € 2 mm 
long; each stamen (5-)6-7(-7.5) mm wide tooth 
to tooth or lobe to lobe; distal portion of filament 
subulate, (4.5-)5-6.6(-7) mm long, (1.8-)2-2.5 


(-3) mm wide at point of dilation; anthers oblong, 


176 Annals of the 
Missouri Botanical Garden 
L— — 
-1— EN 
0 25 50 75 100 
m 
T— km S 
| ^ 
n i 
FIGURE 71. Distributions of Eucharis candida (squares) and E. formosa (circles) in Ecuador. X = E. candida 


x E. formosa. 


4.5-5.5(-6) mm long, gray-brown; pollen grain 
47.7-53.4 um polar diam., 65.5- 73.8 um longest 
equatorial diam. Style 5.5—6(—6.5) cm long, exsert- 
ed ca. 1 cm beyond the anthers; stigma ca. 2-3 
mm wide. Ovary globose-ellipsoid, 6-8.5(-10) mm 
long, (4.5-)5.5-7(-7.5) mm diam., green; ovules 
(2-)5-7(-8). Capsule 1.5-2 cm long, 2-3 cm wide, 
bright orange, leathery; pedicels 3-4 cm long; seeds 
(1-)2-4 per locule, ellipsoid, 8-10 mm long, 5- 
6 mm diam., with a lustrous, smooth black testa. 
Chromosome number: 2n — 46 


Distribution. Rich, moist soil in the under- 
story of premontane and lower montane rain forest, 
chiefly in the Napo and Pastaza drainage of Ec- 
uador (Fig. 71); less frequent in Amazonian Peru 
and Colombia, the lower “ceja de la selva” of north- 
central Peru, and upper Huallaga valley of Peru 
(Fig. 72); rare in central Colombia (a single, poorly 
documented, and very disjunct collection (Killip 
s.n., COL) from near Popayan may be of cultivated 
origin), 100-1,800 m. Flowering most commonly 
January-March. 


Volume 76, Number 1 Meerow 177 
1989 Eucharis and Caliphruria 


| | 


FiGURE 72. Distributions of Eucharis bakeriana ane E. candida (squares), and E. formosa (circles) in 
northwestern-central South America exclusive of Ecuad 


178 


Annals of the 
Missouri Botanical Garden 


FIGURES 73-78. Variation in Eucharis formosa. — (3, 
(FLAS). —78. Schunke 14174 (FLAS). 


Vernacular names and uses. Cebolla de la 
selva, sugkip, sacha cebolla. A poultice of the bulbs 
is used to treat tumors (Lawesson et al. 39632). 


Additional specimens examined. COLOMBIA. AMA 
ZONAS: confluencia de los rios Amazonas y Lore PEU: 
12 Apr. 1975, Cabrera 3336 (COL); Trapecio amazon- 
ico, Loretoyacu River, ca. 100 m, Sep. 1946 (fr), Schultes 
& Black 8342 (US); same locality as preceding, Oct. 
1946, Schultes & Black 8410 (GH, US). CAQUETA: Mo- 
relia, 150 m, 5 Oct. 1941 (fr), von Sneidern s.n. (S). 
CAUCA: Popayan, 25 Jan. 1935, Killip s.n. (COL). Ecua- 


74, 76. Meerow 1103 (FLAS). — 75, 77. Schunke 14171 


DOR. MORONA-SANTIAGO: 15 km N of Macas on road to 
Rio Upano, 2?7'S, 78*8'W, 1,250 m, 20 Feb. 1987, 
Bohlin et al. 1498 (GB); road Limón-Macas, ca. km 20 
from Limón, primary rain forest and rastrojos, 700-900 
m, 26 Mar. 1974, Harling & Andersson 12915 (FLAS, 
GB). NAPO: Nuevo Rocafuerte, Rio Napo, W of Rio Ya- 
suni, Laguna Jatuncocha, Alarcon 106 (QCA); Misa- 
hualli, Guacamayos, Puerto Francisco de Orellana, Alar- 
con 19509 (QCA); Napo, forest, 6 Oct. 1939, Asplund 
9122 (Sy; Tena, marshy forest, 21 Oct. 1939, Asplund 
9488 (S); Limoncocha, 300 m, 22 Jan. 1977, Dodson 
6636 (SEL); 45 minute walk by trail from Santa Cecilia 
up Río Aguarico, ca. 350 m, 28 Mar. 1972 (fr), Dwyer 


Volume 76, Number 1 
1989 


Meerow 179 


Eucharis and Caliphruria 


& MacBryde 9699 (MO, QCA); Santa Cecilia, rain forest 
off runway, 340 m, 30 Mar. 1972 (fr), Dwyer & Sim- 
mons 9743 (MO); Cañon de los Monos, ca. 12 km north 
of Coca, 250 m, Harling & Andersson 11719 (CB); 
path from Rio Bueno to Santa Rosa, Harling et al. 7201 
(GB); Río Jivino, Limoncocha, 13-15 Mar. 1968, Har- 
ling et al. 7673 (FLAS, GB); ge Vieja at Rio Napo, 
ca. 12 km SW of Coca, 12 Jan. 1973, Holguer 2655 
(FLAS, GB); Caron de los Monos, (nx Coca- Lago Agrio, 
ca. 12 km north of Coca, 24 Jan. 1973, por 2960 
(GB); Santa pe Lago Agrio-Baeza, ca. 16 km west 
7 Feb. 1973, Holguer "3532 (FLAS, 


— 


o 
1979 (fr), Holm- Adai et al. 20168 (AAU); Rio Agua- 
ee Jaramillo & Coello 2145 
( road to Las Yucas, 300 m, 
31 Jan. 1980, pasa & Coello 2191 (QCA); Río 
Cuyabeno, Puerto Bolivar, 6 July 1980, Jaramillo & 
Coello 2852 (QCA); Dureno, UR bank of Río Aguarico, 
10 July 1980, Jaramillo & Coello 2962 (QCA); Anangu, 
Rio Napo, 76°23'W, 0°32'S, 260-350 m, 27 June 1983, 
Lawesson et al. 39632 (AAU); 4.2-7.5 km west of Lago 
8.2 km east of Rio Conejo) near Lago Agrio- 
1972, MacBryde & 
Dwyer 1387 (MO); ex hort., orae of SEL Acc. 78- 
1099, collected vicinity Limoncocha, 240 m, 15 Dec. 
1982, Meerow 1103 (FLAS). PASTAZA: Mera, ca. 1,100 


m, 3 1956, Asplund 19571 (S); 3 km NW of Mera 
on road m Mera-Rio Anzu, Hacienda San Augustin, 
00-1,300 Dodson et al. 15717 (MO); Curaray 


(Jesús Pitishka), virgin rain forest near the posto militar, 


ca. 200 m, 18 Mar. 1980, Harli 


Holguer 1504 ( 
nelos, 400 m, doean 5 35, 
es mou Mon of Rio Queran 
ep. 1985, Neill & Palacios 6828 
(MO); 2 km NE of d Hacienda m. ape del Baron 
von — 1?27'S, 78*6'W, 1,3 de E B 
Mar. , Palacios et al. 75 (M o) “Na apo 

of Me 16 Fe b. 1953, Prescott 438 (NY). TUNGURAHUA: 
valley of Pastaza Ri ver, between Baños and Cashurco, 8 
hours east of Baños, 1,300-1,800 m, Hitchcock 21891 


142 (US). PERU. AMAZONAS: Quebrada Huampami, lugar 
tseasim, monte al lado nayumpin, 800 ft., 3 Apr. 1973, 
Ancuash 161 (MO); Quebrada de priedi Rio Ce- 
nepa, 720 ft., Kayap 597 (F, MO); Quebrada Cunup, 
monte cerca a la chacra, 800-850 ft., 24 July 1974 (fr), 
ep 1298 (MO); Rio Cenepa, vicinity of Huampami, 

5 km E of Chavez Valdivia, ca. 78*30'W, 4?30'S, 3 
a 1978, Kujikat 154 (MO); Rio Cenepa, vicinity of 
Huampami, ca. 5 km east of Chavez Valdivia, ca. 78°30'W, 
4°30'S, Quebrada Aintami, 17 Aug. 1978, Kujikat 415 


Napo, 72*48'W, 3?28'S, 120 m, 18 Aug. 1980, Gentry 
et al. 29867 (MO); same locality, 130 m, 18 Feb. 1981, 
Gentry et al. 31418 (MO); Maynas, Iquitos, Rio Ampi- 
acu, 4 vueltas de Monona Cocha, 4 Aug. 1976, Revilla 
990 (MO); Alto Amazonas, Yurimaguas, [S Om a “Shun- 
” al sur-este de Puerto Arturo, cerca a Yurimaguas, 
O m, 1 Dec. 1984, Schunke 14157 (FLAS). SAN 
MARTÍN: road between Moyobamba and Chachapoyas near 
km marker 415, 13.5 km W of Naranjos, 5?38'S, 


77°25'W, 700 m, 12 Apr. 1984, Croat 581724 (MO); 
Mariscal Caceres, Tocache Nuevo, Camino a Shunté, 12 
Mar. 1970, Schunke 3856 (F); Lamas, Alonso de Al- 
varado, San Juan de Pacaizapa, km 72, carretera Tarapo- 
to- Moyobamba, 1,000-1,050 m, 9 June 1977, Schunke 
9675 (F); Lamas, AME de Alvarado, Fundo Las Mal- 
vinas, carretera Moyobamba- Tarapoto, km 43, 850 m, 
6 Dec. 1984, P 14174 (FLAS); San Roque, in 
humid loam, 1,350-1,500 m, 5 Feb. 1930, Williams 
7748 (F 


Eucharis formosa is the most commonly en- 
countered species in eastern Ecuador (Fig. 71). It 
extends into Amazonian Peru and Colombia and 
occurs in the lower “ceja de la selva" forests of 
north-central Peru (Fig. 72). Like the closely re- 
lated E. candida, E. formosa has a wide elevati 
range, possibly in part from cultivation oe 
1987e). The flowers emit a mild and unpleasant 
“sour” odor. Eucharis formosa is larger in all parts 
than E. candida and generally has more ovules 
per locule. The conspicuous horn on the apiculum 
of the outer tepal is characteristic of Ecuadorean 
populations of E. formosa (Fig. 67Ai); this char- 
acter is not obvious in Peruvian collections. Forms 
with toothed or edentate staminal cups occur 
throughout the range of this species without any 
observable geographic pattern (Fig. 70B). Flowers 
of the same inflorescence can vary for this char- 


In floral morphology (Fig. 78), however, it is in- 
distinguishable from other Ecuadorean material. A 
second collection (Schunke 1417 1), from the same 
general vicinity of Peru as Schunke 14174, has 
only shallowly plicate leaves and reduced pigmen- 
tation of the staminal cup (Figs. 76, 77). At pres- 
ent, too little is known about E. formosa in Peru 
to justify recognition of subspecific taxa. 


3. Eucharis bakeriana N. E. own, Gard. 
Chro 


1890, s.n., in part (holotype, K). Urceolina 
bakeriana (N. E. Brown) Traub, PI. Life 27: 
57-59. 1971. Figure 79. 


Bulb to ca. 5 em diam., the tunics brown. Leaves 
2-4; petiole 15-17, 25-30 cm long, (5-)10-11 
mm wide; lamina elliptic, 24-29.5, 44-55 cm 
long, (10-)17-20 cm wide, somewhat succulent, 
smooth. Scape 6-8 dm tall, ca. 1 cm diam. proxi- 
mally, 5-7 mm diam. distally; bracts ovate-lan- 
ceolate, 30-38 mm long, ca. 10 mm wide at the 
base. Flowers (5—)10, with a mild, sweet fragrance, 
pedicels 10-30 mm long; tube 35-40 mm long, 


180 


Annals of the 
Missouri Botanical Garden 


FIGURE 79. Eucharis bakeriana (Meerow 1108, FLAS).—A. flower. — B. Tepals. i. Outer tepal. ii. Inner tepal. — 


C. Staminal cup. — D. Ovary, longitudinal section. 


2-3 mm wide for most of its length, abruptly dilated 
near the throat to 9-9.7 mm wide, curved abruptly 
just above the ovary and straight for the rest of 
its length, thus perpendicular to the vertical axis 
of the scape; limb spreading to 50-60 mm wide; 
outer tepals 28.5-32 mm long, ca. (10-)16.8 mm 
wide, ovate-lanceolate to ovate, apiculate; inner 
tepals 26-30 mm long, (14-)20-22 mm wide, 
ovate, acute to minutely apiculate. Staminal cup 
subcylindrical to campanulate, ca. 16 mm long (to 
apex of teeth), 13-15 mm wide, slightly plicate 
between the filamental trace, very shallowly cleft 
between each stamen (< 1 mm), proximally marked 
green, obtusely bidentate between each free fila- 
ment; teeth 2-3 mm long, half the length of the 
subulate portion of the free filament; each stamen 


5-6.4 mm wide tooth to tooth; subulate portion of 
the filament 3-4.5 mm long, 1.5-1.7 mm wide; 
anthers oblong, 5.4-6 mm long; pollen grain ca. 
50.7 um polar diam., ca. 76.9 um longest equa- 
torial diam. Style 45-54.5 mm long, exserted just 
slightly past the anthers; stigma 2.4-2.8 mm wide. 
Ovary ellipsoid, 6.5-7 mm long, 5.3-6.5 mm diam.; 
ovules 2-3, 8-9 per locule. Capsule ca. 1.5-2 cm 
long, 2.5-3 cm wide, bright orange, leathery; seeds 
ellipsoid, ca. 1 cm long, 0.5 cm diam., with a 
lustrous, smooth black testa. Chromosome number: 
2n — 46 


Distribution. Very rare in the understory of 
lower montane rain forest in the middle Rio Hualla- 
ga valley of Peru, 800 m (Fig. 72). Living material 


Volume 76, Number 1 
1989 


Meerow 181 
Eucharis and Caliphruria 


from which the type specimen was prepared was 
reportedly collected in Colombia, but I have seen 
no Colombian material. Flowering season not known. 


Additional specimens examined. PERU. SAN MARTÍN: 
17 km NE of Tarapoto on road to Yurimaguas, trail along 
stream to waterfall, wet premontane forest on rocky hills, 
6°30'S, 76°20’W, 800 m, 21 July 1982 (fr), Gentry et 
al. 37852 (MO); vicinity of Tarapoto, no other data, 
flowered in cultivation from material collected by L. Besse, 


Meerow 1108 (FLAS). 


The type of E. bakeriana was prepared from 
living material supposedly collected in Colombia. 
When I examined the type specimen, only the 
several large flowers present in the fragment packet 
resembled the figure that accompanied Brown's 
(1890) description of E. bakeriana. The mounted 
material was referable to the smaller-flowered F. 
candida. At the time, I thought that E. bakeriana 
might represent an aberrant form of E. candida. 
Several years later I received a bulb of a Eucharis 
collected near Tarapoto, Peru, by Libby Besse of 
SEL. The flowers that grew from the bulb bore 
an exact resemblance in habit and staminal cup 
morphology to E. bakeriana, though with consid- 
erably more ovules per locule. At present, E. baker- 
iana is known only from the type, the Besse ma- 
terial, and a fruiting specimen referred to this species 
on the basis of leaf size 

Eucharis bakeriana is distinct from E. formosa, 
its closest phenetic relative, by its nonpendent flow- 
ers that are perpendicular to the vertical axis of 
the scape, very shallowly cleft staminal cup (« 1 
mm, vs. 2 mm in E. formosa), short subulate 
portion of the stamen, and sweet floral fragrance 
(slightly fetid in E. formosa). In leaf size and karyo- 
type, E. bakeriana is very similar to Peruvian 
material of E. formosa (Schunke 14174) but dif- 
fers by its greater number of subtelocentric chro- 
mosomes and the submetacentric morphology of 
the second-largest pair (Meerow, 1987b). The leaves 
are only shallowly, if at all, plicate and are thicker. 


4. Eucharis ARM (Kunth) Traub, Pl. Life 
7: 40. 1951. Hymenocallis bonplandii 
Kunth, B Pl. 5: 666. 1850. TYPE: Co- 
lombia. Cundinamarca: Rio Magdalena, near 
Nares, Bonpland 1657 (holotype, P, photo 
of type, NY). Caliphruria bonplandii (Kunth) 
Baillon, Bull. Mens. Soc. Linn. Paris 143: 
1136. 1894. Urceolina bonplandii (Kunth) 
Traub, Pl. Life 27: 57-59. 1971. Figure 80. 


Bulb subglobose, 41-46 mm long, 29-32 mm 
wide, neck ca. 18 mm long and wide, tunics brown. 
Leaves 2, somewhat succulent; petiole 8-14(-18) 


FIGURE 80. 
FLAS) 


Eucharis bonplandii (Meerow 1098, 


cm long, 6-8 mm thick proximally, 3-4 mm dis- 
tally, always shorter than the lamina; lamina ellip- 
tic, (16-)18-24(-26) cm long, 8.5-11.5 cm wide, 
bluish green, especially in strong light, slightly glau- 
cous adaxially; lighter green abaxially, the abaxial 
cuticle densely striate; apex acute to shortly acu- 
minate; attenuate at the base. Scape 4.5-5.8 dm 
tall, 6-8 mm diam. proximally, 3-4 mm diam. 
distally; bracts (25-)33-40 mm long, ovate-lan- 
ceolate, greenish white. Flowers 5-7, nonfragrant, 
pendent; pedicels 18-25 mm long; tube 25-33 
mm long, 1.8-2.5 mm wide for most of its length, 
abruptly dilated at the throat to 7-9(-10) mm wide; 
limb spreading to 47-55 mm wide; outer tepals 
ovate-lanceolate, 25.7-30.5 mm long, 8-10 mm 
wide, apiculate; inner tepals 23-28 mm long, 11.5- 
14 mm wide, acute to minutely apiculate. Staminal 
cup subcylindrical, (11.5-)12.5-14.3 mm long (to 
apex of teeth), 11.5-13 mm wide, stained pale 
yellow proximally, irregularly bidentate between 
each free filament, one stamen occasionally only 
lobed or quadrate, cleft 2.6-4 mm between each 
stamen; teeth variously acute or obtuse, 1-2 mm 
long; each stamen 3.6-4.5 mm wide tooth to tooth; 
free portion narrowly subulate, (3.8-)4.5-5.8 mm 
long, ca. 1.8 mm wide; anthers 4-4.8 mm long, 
oblong, grayish-brown; pollen grain ca. 43.5 um 
polar diam., ca. 63 um longest equatorial diam. 
Style 50-60 mm long, exserted just beyond the 


182 Annals of the 
Missouri Botanical Garden 


-0— 


F10— 


,—— ^ 


| | | 


FIGURE 81. Distributions of Eucharis bonplandii a E. caucana (snowflake), E. cyaneosperma (squares) 
and EF. ulei (circles) in northwestern-central South Americ 


Volume 76, Number 1 
1989 


Meerow 183 
Eucharis and Caliphruria 


anthers; stigma 2-2.7 mm wide. Ovary subglobose, 
ca. 5-6 mm diam.; ovules 2-3 per locule. Capsule 
ca. 1 cm long, 2 cm wide, bright orange, leathery; 
seeds 1-2 per locule, ellipsoid, ca. 1 cm long, ca. 
0.5 mm diam., with a lustrous black testa. Chro- 
mosome number: 2n — 92. 


Distribution. Rare in central and western Co- 
lombia (Fig. 81), in the understory of lower mon- 
tane rain forest, 400-600(- 1, m). Flowering 


February-March, May-June, August. 


Additonal specimens examined. COLOMBIA. Depart- 
ment unknown: La Mejita (?), June Bi x Goudot s.n. 
(K, P). CALDAS: Cauca Valley, Tabeja, west of Armenia, 
| 100-1,300 m, 23 July 1922, Pennell et P 8604 (GH, 
Viotá, Quebrada Cachinibulo, 


as preceding, 600 m, 19 Feb. 1876, y o5» 1721 (K); 
ex hort. i iem: collected by J. Paxton near Bogotá, 

m, ved from Foster Gardens, Hawaii, 14 
May 1982, Meda 1098 (FLAS). TOLIMA: idem del Alto 
^ my vereda La Chamba (municipio del ee 


O m, 3 Mar. 1963 (fr), Uribe 4218 (COL 


Eucharis bonplandii is one of only two tetra- 
ploid (2n — 92) species so far known in the genus. 
The species is known only from Colombia, and has 
been rarely collected. Among Amazonian species, 
E. bonplandii has the greatest phenetic relation- 
ship to E. ulei, with five similarly sized flowers and 
two ovules per locule in common. It may be sep- 
arated from £. ulei by having succulent, glaucous 
leaves and short petioles. The staminal cup of F. 
bonplandii is pigmented pale yellow at its base; 
that of E. ulei is marked green. Eucharis bon- 
plandii and E. bouchei are the northernmost species 
of subg. Fucharis, and thus may form a mono- 
phyletic, tetraploid group (Meerow, 1987d). Both 
species have relatively short petioles. From PF. 
" m E. bonplandii differs primarily by its leaf 
glaucousness and longer pedicels. 


Y 


. Eucharis bouchei Woodson & Allen, Ann. 
Missouri Bot. Gard. 24: 181. 1937. Urceolina 
bouchei (Woodson & Allen) Traub, Pl. Life 
27: 57-59. 1971. TYPE: Panama. Coclé: El 
Valle de Antón, 500-700 m, 23-27 July 
1935, Seibert 466 (holotype, MO). Figure 82. 


Bulb subglobose, 30-45(-85) mm long, (25-) 
30-40(-50) mm diam.; neck short, to 25 mm 
long, 10-20 mm wide; tunics brown. Leaves 1-3 
(-4); petiole (9-)15-25(-28) cm long, 7-8 mm 
wide proximally, 5-6 mm wide distally; lamina 
widely anes delata, (17-)20-25(-40) cm long, 
(7-)8-10(-14) cm wide, shortly acuminate, slight- 
ly E. lustrous bright green adaxially, dull 


pale green abaxially, smooth, margins mostly non- 
undulate; abaxial cuticle largely devoid of striation. 
Scape ca. (4-)5.5 dm tall, ca. 1 cm diam. proxi- 
mally, ca. 5 mm diam. distally; bracts ovate- lan- 
ceolate, 25-36(-47) mm long, (5-)7-10 mm wide 
at the base. Flowers (3-)5(-6); usually pendent, 
sometimes only declinate, not fragrant; pedicels 5— 
10(-20) mm long, very rarely less than 5 mm; 
tube (25-)33-45 mm long, cylindrical and (1.5-) 
2-2.5(-3) mm for most of its length, abruptly 
dilated near the throat to (7-)8-10(-12) mm wide, 


al 


ly curved 


usually curved gradually, but 
abruptly at the base, in which case nearly straight 
for most of its length; tepals spreading widely from 
the throat (ca. 90%) or sometimes only at an angle 
of 45-607; outer tepals ovate-lanceolate, (18—)21 — 
28(-32) mm long, 8-11(-15) mm wide, apiculate; 
inner tepals ovate, (16-)20-26(-32) mm long, 
(10-)12-15(-17) mm wide, obtuse to acute. Stam- 
inal cup subcylindrical, 9-12(-15) mm long to 
apex of filament, (10-)12-15(-18) mm wide, 
deeply cleft between each stamen to 3-5 mm, 
usually edentate but variably lobed, acutely or ob- 
tusely bidentate, or irregularly toothed between 
each stamen, marked pale green to greenish yellow 
proximally; each stamen (3.5-)4-5 mm wide at 
the base, 4-6(-7) mm long, either trapezoidal in 
shape (in which case dilating gradually from apex 
to base), or abruptly dilated at 44-14 of its length 
(in which case the upper portion narrowly subulate 
and 2-3 or 3-4 mm long, 1.5-2 mm wide); anthers 
oblong, 3.5-4.5 mm long; pollen grain 45.7-49.65 
um polar diam., 66.8-68.43 um longest equatorial 
diam. Style (30-)45-60 mm long, exserted 0.5- 
1 cm beyond anthers; stigma 2-3 mm wide. Ovary 
globose or ellipsoid and deeply trigonous, rarely 
not trigonous, 5-8 mm long, 4—6(—6.5) mm diam., 
usually wider than long when deeply trigonous; 
ovules 2-3(-4, very rarely 5) per locule, super- 
posed in the lower half of the cell. Capsule 1.5-2 
cm long, 2-3 cm wide, bright orange, leathery; 
pedicels 15-27 mm long; seeds 1-2 per locule, 
ellipsoid, ca. 1 cm long, 0.5 cm diam., with a 
lustrous, smooth, black testa. Chromosome num- 
ber: 2n — 


KEY TO THE VARIETIES OF £. BOUCHEL 


la. Staminal cup edentate, or occasionally with one 
obscure tooth at the base of one stamen; sta- 
mens trapezoidal and gradually dilated from 
apex to base or, if obscurely constricted distally, 
subulate portion more than e; Coclé 
and Colón provinces of Panama, rare in Pina 
Province and in Costa ra xd Guatemala ..... 

chei var. bouchei 
uy 


184 Annals of th 
Missouri ion Garden 


FIGURE 82. Eucharis bouchei. — A. Flowers. i. Variety dressleri (holotype, Meerow 1107, FLAS). ii, iii. Variety 
bouchei. ii, Meerow 1125 (FLAS); iii, Meerow 1157 (FLAS).— B. Tepals, variety bouchei. i, ii. Meerow 1125. i, 
outer tepal; ii, inner tepal. iii, iv. Meerow 1157. iii, outer tepal; iv, inner tepal. —C. Staminal cups, variety bouchei. 
: "rae w 1125; ii, Meerow 1157. —D. Ovaries, variety bouchei, longitudinal section. i, Meerow 1125; ii, Meerow 


constricted distally at 14-14 length into a subu- da. Eucharis bouchei var. bouchei. Figures 

late portion 1.5-2 mm wide 2 70Ci-ii, 82Aii-iii, B. 

2a. Outer tepals 26-32 mm long; staminal cup 
irregularly toothed, the teeth acute; subu- Perianth tube (30.8-)34-45 mm long; outer 
late portion of filament 3.5-4.5 mm long; tepals (18-)24-28(-35) mm long, (8.8-)9.4-15.5 
vary not deeply cnc ovules 2-4 per mm wide; inner tepals 21-26(-31) mm long, 11- 
locule; Coclé Province of Panama near ran i , 
Ve ras 5b. E. bouchei var. dressleri 17.5 mm wide. Staminal cup (8, 9-)11.5-15 

2b. Outer tepals 20-26 mm long; staminal cup (716.7) mm long (to apex of filaments), (9-)12- 
obtusely bidentate or lobed; subulate por- 15.5(-16, 18) mm wide, edentate or rarely with 
tion of filament up to 3.5 mm long; ovary a single obscure tooth between one or several sta- 


deeply trigonous; ovules 2-5 per locule; 

Panamá and Darién provinces of Panama, 

rare in Guatemala 
Sc. 


mens; each stamen trapezoidal in shape, dilating 
gradually from apex to base, or, if obscurely con- 
E. bouchei var. darienensis stricted in the distal 2-3 mm, then the subulate 


185 


"Pureueq Ul 7242n0Q S11DYINZG jo uonnquis(] “Eg INJA 


Eucharis and Caliphruria 


Meerow 


T T T "T 
BL 08 18 08 
l | | | 
= L 4 
eerie 
00] 06 = 0 
> 
- 
^ 
|40/$$01p ien ¡9yINOQ “3 (S) ghee 
$ 4 
y x 
sysueuessep se, Jeuyonoq “30 1 4 
E ) € 
[9YINOGQ ‘18, JOYINOQ sj18uong e p": y 
J 
- pe 
/ 
l 
\ 


\ 
Le 
A — 05 e 
“ag = a2 V 6 
Qe 


(707 


1,- ) 
e o”; d 
ER PET et Ne tJ Y. 
Vu. vr MN 


Volume 76, Number 1 


1989 


186 Annals of the 
Missouri Botanical Garden 


o 200 400 
i HA A 
i km 
l 
i 
l 
CA t 
m 15 = Pi — 
O 1 r ARA 
9 277 ~ 1 
» ^ ^ LAS r 
ir 
Ss CENE A 
m 10 =e @ — 
a 
e 


| | | 


90 85 
d l | 


E84. Distribution of Eucharis P depo in Pu America exclusive of Panama. Circles = Eucharis bouchei 
var. bouchei; square = E. bouchei var. darie 


on steep slopes; (200-)500-1,000 m. Flowering 
(March) June-August, October- December. 


portion wider than 2 mm. Style exserted ca. 1 cm 
beyond the anthers. Ovary trigonous; ovules 2- 
SUE Ber qune: Additional specimens examined. Costa Rica. 
"-— . . PUNTARENAS: Canton de Osa, hills near Palmar Norte, Rio 
Distribution. Understory of primary, pre- Gra nde de Terraba, 2.000 i Allen 5347 (F, K, E 
montane and lower montane rain forest in Coclé eb. 1 
and Colón provinces of Panama (Fig. 83), partic- 
diri mais El Valle de dió dth s.n. ae ). GUA JATEMALA. SUC HITEPEQU EZ: Fin ca Mock: steep, 
— pisas zm ue venu E tee sies slope, 3,300 ft., 31 Oct. 1934, Skutch 1585 (F). 
Rio Guanche valley; rare in Panamá Province, PANAMA. COCLÉ: lower 'Río Antón, vic. El Valle de Antón, 
Costa Rica, and Guatemala (Fig. 84); frequently — 800-1,000 m, 30 Dec. 1936, Allen 120 (GH, MO); vic. 


Volume 76, Number 1 
1989 


Meerow 187 
Eucharis and Caliphruria 


of El Valle de Antón, 600-1,000 m, Allen 1228 (GH, 
MO); vic. El Valle de Antón, ca. 600 m, 10 Dec. 1939, 
Allen 2063 (MO); hills north of El Valle de Antón, 100 
m, 14 Aug. 1940, Allen 2182 (MO); region north of El 
Valle de Antón, ca. 1,000 m, 21 Aug. 1946, Allen 3641 
(G); La Mesa, above El Valle, 600-800 m, 18 Jan. 1968 
(fr), Duke & Dwyer 15180 (NY); 1-3 mi. W of Por- 
tobello, Gentry 1750 (MO); foot of Cerro Pilón, 11 Jan. 
1972 (fr), Gentry & Dwyer 3634 (F, MO); foot of Cerro 
Pilón, 11 Jan. 1972 (fr), Gentry & Dwyer 3636 (MO); 
El Valle de Antón, along Río Indio Trail, 500-700 m, 
(fr), Hunter & Allen 338 (G, MO, P, US); el Valle de 
Antón, 1,000-2,000 ft., edge of cloud forest and road- 


Amarillo, north of El Valle, 17 Oct. 1975, Witherspoon 
& Witherspoon 8736 (MO). COLON: Rio Guanche, 16 
Nov. 1975, D'Arcy 9679 aja hills just north of Rio 
Guanche, 1-200 m, 16 Nov r) Davidse & 
D'Arcy 1 0096 (MO); Cerro B ex hort., collected by 
R. Dressler, flowered in cultivation, July , Meerow 
1157 (FLAS); il south of Rio 
Cerro Pan de Azucar, m, Mori & 
(AAU); Rio dedi 6 Nov. 1974 (fr) Mori & Kallun ki 
3019 (AAU). P Á: mountains above Torti i gui ie 
Dec. 1977, (ies et al. 6582 (AAU, BM, MO); n 
Cerro Campana, on Wap duos from end of road ker 
passes Campana water tank, 23 Aug. 1967, Kirkbride 
& Hayden 305 (MO. "NY) 


5b. Eucharis bouchei var. dressleri Meerow, 
var. nov. TYPE: Panama. Coclé: El Valle de 
Antón, from bulbs collected by R. Dressler in 
Panama, flowered in cultivation, 17 Mar. 
1983, Merrow 1107 (holotype, FLAS). Figure 
82Ai. 


l z IN WM 2$ 23 


acute 


dentatis ad 1. a 2. 7 mm distale concratis, ovario. nos 
trigono, et 


Perianth tube 30-41 mm long; outer tepals 
26.2-32 mm long, 6-10 mm wide; inner tepals 
24-28 mm long, 9-13.5 mm wide. Staminal cup 
10-16 mm long (to apex of filaments), 9.5-11.5 
mm wide, irregularly toothed (some stamens lobed 
or quadrate), the teeth acute and 1.5-2.7 mm 
long; each stamen distally constricted abruptly at 
14 length, the subulate portion 3.5-4.5 mm long, 
ca. 1.8 mm wide. Style exserted ca. 0.5 cm beyond 
the anthers. Ovary not trigonous; ovules 2-4 per 
ocule. 


This variety is named in honor of Robert L. 
Dressler, well-known tropical biologist. 

Distribution. Rare rain forest understory herb 
in Coclé Province of Panama (Fig. 83), in the 
vicinity of El Valle de Antón, ca. 900 m. Flowering 
in June. 


Additional specimens examined. PANAMA. COCLÉ: El 
Valle de Antón, 900 m, 4 June 1939, Alston 8727 (BM). 


5c. Eucharis bouchei var. darienensis Mee- 
row, var. nov. TYPE: Panama. Darién: valley 
between Cerro Pirre and next most southerly 
peak, July 1977, Folsom 4402 (holotype, MO). 
Figure 70Civ. 


Varietas a Eucharis bouchei var. dressleri affinis sed 
differt staminibus obtuse dentatis vel lobis et parte sta- 
minea distale subulata breviore. 


Perianth tube 25.5-36(-43.4) mm long; outer 
tepals 20-26 mm long, 8-11 mm wide; inner 
tepals 10.5-24 mm long, (11-)12-15(-16.5) mm 
wide. Staminal cup 9-10(-13, 16) mm long (to 
apex of filaments), 12-14.5(-18) mm wide, ob- 
tusely bidentate or lobed, the teeth when present 
ca. 1-1.5 mm long; each stamen distally constrict- 
ed abruptly at 14-14 length; the subulate portion 
a. 1.7-3 mmlong, 1.5-2 mm wide. Style exserted 
less than 0.5 cm beyond the anthers. Ovary trigo- 
nous; ovules 2-5 per locule. 


eÓ 


The varietal epithet refers to Darién Province 
of Panama, where E. bouchei var. darienensis 1s 
most frequently collected. 

Distribution. _Understory of primary rain for- 
est in Darien Province of Panama (Fig. 83); rare 
in Panamá Province and in Guatemala (Fig. 84); 
480-1,450 m. Flowering January-February and 
Mec 


Additional specimens examined. GUATEMALA. 
SOLOLA: pian t. Pedro and Sta. Lucia, 20 Jan. 1857, 
sir 207 (COET). PANAMA. DARIÉN: La Boca de 

rre, p Oct. 1967 (fr), Bristan 1285 (MO); vicinity of 
eie ana gold mine, 480 m, disturbed forest, 29 
July iore. Es 37956 (AAU); Cerro Pirre, cloud forest 
and/or mossy forest, 2,500-4,500 ft., Aug. 1967, Duke 
& Elias 3661 (GH, MO, USy Cerro Tacarcuna, south 
slope, 1,250-1,450 m, 26 Jan. 1975, Gentry & Mori 
13945 (MO); 0-2 mi. E of Tres Bocas, along shortest 
headwaters of Rio Cuasi, p orest, 28 Mar. 
1968 (fr), pa & Duke 1175 (MO); vicinity Cana, 
1,750 ft., rare on forest floor, 23 June 1959, Stern et 
al. 499 (G Hi) gold mine at Cana, 480 1 m, 26 July 1976 
fr), Sullivan 626 (MO); trail northwest of Cana, 600 m, 
28 July 1976 (fr), Sullivan 7 18 (MO); gold mine at Cana, 
480 m, 29 July 1976 (fr), Sullivan 753 (MO); Cana- 
Cuasi trail, Chepigano, 2,000 ft., 12 Mar. 1940 (fr), 
Terry & Terry 1533 (F). pr stream flowing out of 
Serranía de Maje, 10 es 19 7 (fr), Folsom & Collins 
1725 (MO); Maje, 5 m m o Maje, steep forested 
ridge above Chocó Lad trail, 400 m 19 Nov. 1970, 
Kennedy 680 (MO, US); woods around La Eneida, 1,000 
m, 5 Aug. 1970 (fr), iai & Kennedy 1761 (E); nas o 
slopes and trail to Cerro Campana, 13 Sep. 1975, 
erspoon & Whereas 8372 (MO ) 


— 


188 


Annals of the 
Missouri Botanical Garden 


Eucharis bouchei is the northernmost species 
of Eucharis, and the only species found north of 
the Darién Gap. It is also the most variable species 
in the genus, in characteristics that elsewhere jus- 
tify specific delimitation. Patterns of variation in 
floral size and tube and limb habit form a complete 
mosaic throughout the range of E. bouchei that 
shows little or no geographic consistency (Meerow, 
19874). Staminal cup morphology (Fig. 70C) does, 
however, demonstrate a fair degree of geographical 
consistency, and it is chiefly on this basis that I 
have recognized var. bouchei and var. darienensis. 
Variety dressleri, rarely enountered mixed with 
var bouchei, presents a special case, discussed 
below. 

Variety bouchei, most common around El Valle 
de Antón in Coclé Province, is recognized by its 
largely edentate staminal cup in which the trape- 
zoidal free filament is not markedly constricted 
distally into a narrow subulate portion (Figs. 70Ci, 
ii, 82C). The staminal cup of "a darienenis, 
found both in Panamá and Darién provinces, is 
obtusely bidentate or lobed (Fig. 70Civ) The free 
filament constricts distally into a narrow (< 2 mm) 
subulate portion. These two varieties occur in close 
proximity in one location, near Cerro Campana in 
Panamá Province (Fig. 83). 

The rare var. dressleri (Fig. 82Ai), with its 
acutely toothed staminal cup (Fig. 70Ciii) and shal- 
lowly trigonous ovary, occurs close to populations 
of var. bouchei, which is an unstable tetraploid, 
producing at least some cells with diploid (2n — 
46) chromosome number (Meerow, 1987d). This 
variety also does not express either heterozygous 
or additive banding phenotypes for isozymes o 
aspartate amino transferase, which otherwise char- 
acterizes electrophoretic phenotypes of E. bouchei 
(Meerow, 19874). 

The unprecedented degree of variation in £. 
bouchei is likely the result of two main factors: (1) 
the species is tetraploid, and (2) it probably rep- 
resents a geologically recent colonization of Central 
America by this primarily northern Andean and 
Amazonian genus (Meerow, 19874). 

Northwesternmost populations representing var. 
bouchei have the most derived androecial mor- 
phology (Fig. 70Ci, ii) relative to southeasterly 
populations (var. darienensis, Fig. 70Civ). The 
latter have staminal cups similar to the generalized 
morphology characteristic of Andean and Ama- 
zonian species of subg. Eucharis. This suggests 
that general movement of E. bouchei in Central 
America has been away from the Colombian bor- 
der. The occasional presence of E. bouchei in Costa 
Rica is not surprising, but the two reported collec- 


tions from Guatemala (Wendland 207 and Skutch 
1585) represent a substantial disjunction, or two 
disjunctions, since the two collections represent 
different varieties. Given the history of cultivation 
of Amazonian Eucharis by Indian people for me- 
dicinal, ceremonial, and possibly ornamental use, 
the same may have held true in Central America. 


6. Eucharis astrophiala (Ravenna) Ravenna, 
Phytologia 57: 95-96. 1985. Urceolina as- 
trophiala Ravenna, Pl. Life 38: 49. 1982. 
TYPE: Ecuador. Cotopaxi: Quevedo- Latacun- 
ga road, km 46 from Quevedo, 79?11'W, 
0*55'S, 600 m, 4 Apr. 1973, Holm-Nielsen 
et al. 2851 (holotype not seen, AAU; isotype, 
S). Figure 85. 


Bulb globose, 4-5 cm long, 3-4 cm wide, usually 
without an appreciable neck, tunics tannish brown. 
Leaves 2-4 at anthesis, elliptic- or ovate-lanceo- 
late; petiole 10-20 cm long, 4-5.5 mm thick; 
lamina 15-25 cm long, 5-10 cm wide, thin, non- 
lustrous, deeply plicate and pustulate, adaxial sur- 
face light green, the white midrib conspicuous; 
abaxial surface whitish green; margin slightly un- 
dulate; apically acuminate; basally attenuate to the 
petiole. Scape 3-4(-5) dm tall, ca. 5 mm diam.; 
bracts 29-35(-40) mm long, lanceolate. F Hm 
5-8(-10); pedicels 8-14 mm long; tube 28-35 
mm long, ca. 2 mm wide for most of its length, 
dilating to (4-)5-6 mm at the throat, strongly 
curved; perianth limb spreading to 4-5 cm wide; 
outer tepals 25-30 mm long, (7-)10 mm wide, 
anceolate, apiculate; inner tepals 25-28 mm long, 
10-12(-14) mm wide, ovate-lanceolate to ovate, 
acute. Staminal cup funnelform-cylindrical, (10-) 
12-14 mm long, 8-12 mm wide, edentate, stained 
orange-yellow basally, cleft between each stamen 
for Y,-Y, of its length; each free filament (5-)6.4— 
6.9 mm long, ca. (2.5-)3.5-4.5 mm wide at the 
base, deltoid; anthers oblong, 5-5.5 mm long; av- 
erage pollen grain 58.6-60.6 um polar diam., 83.0- 

6.1 um longest equatorial diam. Style 50-55 mm 
sl exserted 5-10 mm beyond the staminal cup; 
a 3-lobed, ca. 2 mm wide. Ovary globose- 
aoe 3.9-4.5 mm long, 3.2-4 mm wide, white 
at anthesis; ovules 2-3(-4) per locule, medially 
superposed. Capsule ca. 1-1.5 cm long, 2-2.5 cm 
wide, bright orange, leathery; seeds 1-2 per locule, 
ellipsoid, ca. 1 cm long, 0.5 cm diam., with a 
lustrous, smooth black testa. Chromosome number: 


Distribution. Endemic to the western decliv- 
ity of the Andes in north-central Ecuador (Fig. 86), 
particularly in contiguous areas of Cotopaxi, Los 


Volume 76, Number 1 


Meerow 
Eucharis and Caliphruria 


189 


1989 
LLLI 22 
Sau 7 wm 
SR SS ER ÉS = 
AS ` 
RAS EON SS 
uL ETE E 
GS 
RS N 
IM 
FicURE 85. Eucharis astrophiala, adapted from a drawing by Boots N. bid: of Dodson et al. 
(SEL). — A. Habit. —B. Inflorescence. — C. Flower, longitudinal section. — D. Stigma.— E. Ovary, longitudinal dle sN 


Ríos, and Pichincha provinces, occupying the 

underst of lower montane rain forest 
(250- 400 800(-1,100) m elevation. Sporadic 
flowering may occur at any time but is concentrated 
in the wetter months of the year. 


Additional specimens examined. ECUADOR. BOLÍ- 
vaR: Limón, 800-1,100 m, 14 Oct. 1943, Acosta-Solís 
6374 (F). CHIMBORAZO: km 52-53 on Quevedo-Lata- 


Dodson & Gentry 13793 (MO, SEL); Puente de Chimbo, 
250 m, June 1876, Lehmann 7775 (K). COTOPAXI: km 


40 on road from Quevedo to Latacunga, 600 m, 6 Mar 
1975, Dodson 5864 (MO, SEL, US); 3 km E of El Palmar 


& Gentry 12815, 750 m, sterile, 15 Aug 
& Meerow 1140 (FLAS). Los níos: oeste hills 12 km 
1977 


km 113, 00%05'N, 79%02'W, 10 km al 
retera principal, A m, 27-29 Dec. 198 
Balseca 4695 (NY, QCA); same locality, 28 Feb. 1984, 


et al. 7122 (M 


O, SEL); 2 km SE of Santo Domingo de 
los Colorados along Rio Verde, 530 m, 5 Feb. 


1979, 


190 Annals of the 
Missouri Botanical Garden 

—_ —— À 

-2——— E 
0 25 50 75 100 
HA AH 

4 km — 

80 18 16 
FIGURE 86. Distribution of Eucharis astrophiala in Ecuador. 


Dodson & Duke 7714 (MO, SEL); road from Patricia 


985, Harling & An- 
dersson el di (GB); Pocero Fe d Endesa, Rio Silan- 
e, km 3 Quito-Puerto Quito, m N of main 


highway, pl N, 79°2’W, 650-700 m, 25 Feb. 1984, 


Jaramillo 6439 (GB); Endesa, km 113 Quito-Puerto 
Loy 750 m, 23-24 Apr. 1983, Rodriguez et al. 91, 
& 93 (QCA); Santo Domingo de los — Rancho 
tai n, ca. 10 km NW of the town road to Es- 
meraldas, 400 m, 31 Mar. 1967, Sparre 15216 (S). 
Eucharis astrophiala is easily separated from 
the other small-flowered species of subg. Eucharis 
by its uniquely bullate-pustulate and nonlustrous 
ovate-lanceolate leaves; edentate and deeply cleft 
staminal cup with deltoid free filaments (Fig. 85B, 
C); and the second-largest pollen grain in the genus 
(the largest is that of E. caucana). The largest 


Volume 76, Number 1 
1989 


Meerow 191 
Eucharis and Caliphruria 


1cm 


FIGURES 87, 88. — 


uration of E. caucana (2n — 138, type collection). Thirteen chromosomes are out of the figure frame 


chromosome pair of E. astrophiala is submeta- 
centric, unlike the metacentric pair of all other 
species of the genus I have examined (Meerow, 
1987b). It is the only species of the subgenus found 
exclusively on the western slopes of the Andes south 
of Colombia. It occurs sympatrically in some lo- 
calities (fide Dodson & Gentry 12815 and Meerow 
& Meerow 1140) with E. moorei, though the latter 
grows at slightly higher elevation in these areas. 
Eucharis astrophiala is the only species of subg. 
Eucharis that enters a definite rest period during 
the short dry season of the northwestern and cen- 
tral western Ecuadorean Andes. New growth com- 
pletely ceases, though one or two leaves may persist 
for the duration. 


7. Eucharis caucana Meerow, sp. nov. TYPE: 
Colombia. Valle de Cauca: between La Paila 

and Zarzal, Hacienda El Medio, 975 m, 14 
Apr. 1987, Silverstone-Sopkin et al. 3064 
(holotype, FTG; isotypes not seen, CUVC). 

igure 

Species nova affinis E. lehmannii Regel sed floribus 
breviter pedicellatis hour et paucioribus, stami- 


87. Eucharis caucana (type collection). —88. Root-tip mitotic metaphase chromosome config- 


nibus edentatis, et ovulis paucioribus in quoque loculo 
differt. 


Bulb subglobose, 52-60 mm long, ca. 34 mm 
diam., without an appreciable neck, tunics brown. 
Leaves 2-3; petiole 21-25 cm long, 7.2-10 mm 
wide; lamina ovate—elliptic, ca. 25.5 cm long, 10- 
11 cm wide, plicate, the margins undulate, acu- 
minate, attenuate at the base. Scape 4-5 dm tall, 
ca. 1 cm diam. proximally, 5 mm diam. distally; 
bracts 26-27 mm long, 8 mm wide, ovate-lanceo- 
late. Flowers 2-3, slightly declinate, funnelform- 
campanulate, not fragrant; ue ca. 6.5 mm 
long, ca. 2 mm diam.; tube . 32 mm long, 2 
mm wide proximally, dilating in its distal 44 to ca. 
9.2 mm at the throat, abruptly curved near its 
base, the dilated portion stained green internally; 
limb spreading to ca. 3.5 cm, the tepals diverging 
from the throat at angles of 75—90°; outer tepals 
ca. 20 mm long, 9.5-10.3 mm wide, ovate, apic- 
ulate; inner tepals ca. 18-19 mm long, 11.5 mm 
wide, ovate, acute to obtuse. Staminal cup ca. 3 
mm long (to apex of lobes), edentate, stained green 
proximally; stamens connate only in the proximal 
—] mm; each filament ca. 10 mm long, narrowly 
subulate and ca. 1 mm wide in the distal 4-5 mm, 


> 


192 


Annals of the 
Missouri Botanical Garden 


then abruptly dilated proximally to 3 mm wide; 
anthers oblong, ca. 4 mm long, gray-brown; pollen 
white, the exine coarsely reticulate, 55-75 um 
polar diam., 98-110 um longest equatorial diam. 
Style white, ca. 50 mm long, exserted ca. 1 cm 
beyond the anthers; stigma ca. 2 mm wide, greenish 
white. Ovary globose-trigonous, 7-7.5 mm diam., 
ovules 4-6 per locule. Capsule leathery, orange, 
ca. 1.5 cm long, 2.5 cm wide; seeds 2 per locule, 
ellipsoid, ca. 10 mm long, ca. 5 mm wide with a 
smooth, lustrous black testa. Chromosome number: 
= 138 


Named for the Rio Cauca valley of Colombia to 
which the species is endemic. 


vitiis Endemic to Valle de Cauca De- 

in the vicinity of La Paila in western 

een (Fig. 81). The type location is a 12.5- 

hectare remnant of primary forest planted with 

cacao in which canopy trees were retained for 

shade (Anacardium excelsum, Nectandra cau- 
cana). 


Additional specimen examined. COLOMBIA. VALLE 
DE CAUCA: La Paila, 30 May 1853 (fr), Holton s.n. (NY). 


The recent receipt of material collected by Phil 
Silverstone-Sopkin and coworkers is the first col- 
lection of the subgenus in the largely deforested 
Cauca valley since 1922. 

Eucharis caucana has the largest pollen grains 
in the genus and is the only known hexaploid species 
of Eucharis, with a somatic chromosome number 
of 2n — 138 (Fig. 88). As with most other South 
American peripheral isolates from the Amazonian 
center of distribution for subg. Eucharis (E. asto- 
phiala, E. corynandra, E. lehmannii, and E. ox- 
yandra), E. caucana exhibits some degree of mor- 
phological novelty; in this case, reduction of staminal 
connation, funnelform-campanulate perianth, and 
very large pollen grains. Nonetheless, E. caucana 
has fruit and seed morphology typical of subgenus 
Eucharis. By its reduced staminal cup E. caucana 
shows affinity with E. lehmannii from which it 
differs by having larger flowers, shorter pedicels, 
straighter tubes, edentate androecia, and fewer 
ovules per locule. By its high polyploid chromosome 
number, short pedicels, and flower size, E. caucana 
shows similarity to certain populations of E. bouch- 
ei var. bouchei. Some collections of the latter from 
Colón Province exhibit funnelform-campanulate 
perianth morphology and floral tube habit like E. 
caucana. The discovery of this hexaploid species 
in Colombia supports the hypothesis that a poly- 
ploid complex of Eucharis may have existed in 


Colombia in the recent geologic past (Meerow, 
987d). 


8. Eucharis ulei Kranzlin, Bot. Jahrb. 50, Beibl. 
111: 4-5. 1913. Urceolina ulei (Kranzl.) 
Traub, Pl. Life 27: 57-59. 1971. TYPE: Bra- 
zil. Amazonas: Juruá Miry, June 1901, Ule 
5737a (holotype, B), non Ule 5737 b (in fruit) 
(B) vel Ule 5737 (MC). Figure 89B, C 


Eucharis ipariensis (Ravenna) Ravenna, Phytologia 57: 
95. 1984. Urceolina ipariensis Ravenna, Pl. Life 

38: 50-51. 1982. TYPE: Peru. Huánuco: Pachitea, 
Honoria, Bosque Nacional de Iparia, Río Pachitea, 

20 km above confluence with Rio Ucayali, near Miel 

de Abeja, 1 km from Tuernavista, 26 Apr. 1967, 
Schunke 1887 (holotype not seen, NY; isotypes, F, 

L, G, US). 

a moana (Ravenna) e Phytologia 57: 95. 
nna, Pl. Life 38: 50. 
o Moa at Serra da Moa 


ii type, NY; isotypes, K, MO, not seen: herb. Ra- 


Eucharis boliviensis Ravenna, Phytologia 64: n 1988. 
olivia. El Beni: Covendo, 600 m, 19 Aug. 
1921. White 930 icit NY; NS. K). 


Bulb subglobose, (2.5-)3.5-4.5(-5) cm long, 2- 
3.5(-4.5) cm in diam.; neck 1-2 cm long, ca. 1.5 
cm wide; outermost tunics gray-brown, inner tunics 
tan. Leaves 2-3; petiole (10—)18-30(-35) cm long, 
5-6 mm thick; lamina (narrowly) elliptic (average 
length: width > 3), 18-25(-33) cm long, (5-)7- 
10(-12.5) cm wide, acute to shortly acuminate, 
attenuate at the base. Scape (35-)40—58 cm tall, 
8-10 mm diam. proximally, 3-4 mm diam. dis- 
tally; bracts lanceolate to ovate-lanceolate, (25-) 
30-37(-54) mm long, greenish white. Flowers (3-) 
(-7), pendent, without fragrance; pedicels (8-) 
11-15(-20) mm long, ca. 2 mm diam.; tube (25-) 
28-35(-37.5) mm long, curved gradually for most 
of its length, 1.5-2(-2.5) mm wide for most of its 
length, abruptly dilated just below throat to (6-) 

mm; limb spreading to 40—45(-55) mm wide; 
outer tepals ovate-lanceolate, 24—28(—32) mm long, 
(6.5-)8-10(-11) mm wide, apiculate; inner tepals 
ovate, 23-27(-30) mm long, (9-)10-13(-15) mm 
wide, acute. Staminal cup funnelform-cylindrical, 
10-12 mm long (to apex of teeth or lobes), 11- 
13(-16) mm wide, usually bidentate between each 
free filament, rarely edentate, irregularly toothed, 
or the teeth obscure (in which case the stamens 
quadrately lobed), cleft between each stamen for 
(1.5-)2-3(-4) mm, with a + rectangular, green 
zone in the proximal half of each stamen; teeth 
acute or obtuse, 0.5-0.7 mm long; each stamen 
3.5-4.5(-5.5) mm wide from tooth to tooth; free 
filament subulate, (3—)4.5—6 mm long, (1-)1.5-2 


Volume 76, Number 1 
1989 


Meerow 193 
Eucharis and Caliphruria 


FiGURE 
FLAS).— 


Eucharis cyaneosperma and E. u 


mm wide at its base; anthers oblong, 3-3.8 mm 
long; pollen grain ca. 49.35 um polar diam., ca. 

9.85 um longest equatorial diam. Style (40-)45— 
50(-60) mm long; stigma 1.5-2.5 mm wide. Ovary 
globose-ellipsoid, 6-8.5(-10) mm long, 4-5.5 mm 
wide; ovules 2(-4) per locule, superposed in the 
lower half of the cell. Capsule 1.2-1.5 cm long, 
2-2.5 cm wide, bright orange, leathery; pedicels 
2-4 cm long; seeds 1-2 per locule, ellipsoid, 8- 
10 mm long, ca. 5 mm wide with a smooth, lustrous 
black testa. 


Distribution. Understory of primary rain for- 
est in the Amazon Basin and eastern Andean foot- 
hills, most common in Peru but sporadically en- 
countered north to Colombia and south to Bolivia 
(Fig. 81), on fertile, usually noninundated, soils; 
100-300(-1,000) m. Flowering at any time of the 
year, most frequently June-September. 


Additonal specimens b coda BRAZIL. AMAZONAS: 
basin of Río Juruá, Foz da a, Yuma, rare, on varzea 
land, 1 June 1933, Krukoff 4613 (NY) basin of Rio 
Juruá, near mouth of Rio Embira, 7?30'S, 70°15’W, 3 
June 1933 (fr), Krukoff 4637 (G, GH, K, NY, S, US); 


Río Purus, Río Itaxi, Seringal Jurucua, 120 km south of 


OLOMBIA: AMAZONAS: 
a de Río Loretoyacu con el Río Ama- 
Diaz-M 15 (COL); 


Trapecio amazonico, Boiauassu River, 100 m, Oct. 1946, 


89. . Flower of E. cyaneosperma (holotype, Meerow 1032, 
— B. Flower of E. ulei (Schunke 1887, F). id pe ary of E. ulei, longitudinal section (Schunke 1887, F). 


Schultes & Black 8608a (US); Aeon Narino, mouth of 
72 (fr), Plowman 3216 


(CO ). PERU. AMAZONAS M ntiago, ca a 
uebrada Caterpiza, 200 11 Sep. 1979 (fr), Huashi 
cat 524 (MO); valle de Río Santiago, ca. 65 e 


de Pinglo, Quebrada Caterpiza, 2-3 km co 

munidad de Caterpiza, 200 m, 13 Oct. 1979 (fr), Huashi- 

ocali c. 1979, Huashikat 

1553 (MO); Monte del Isla, la ed ^ km L3 de La P. 

Rio Santiago, 180 m, 8 Aug. 1 

same locality, 15 Aug. 1979 
s 1979 (fr), Pena 9 ( 


Do 19289 TES a 
ayuga, edge of forest, 20 b 1972 (fr), Croat 20521 
(MO); near or Araguana, upper Rio Mazan, ca. due 
north of Santa Maria de Nanay, noninundated forest, 9 
Nr pro do, Gentry & Revilla 16586 (MO); Maynas, 
Quebrad omono, Explorama Tourist Camp, halfw way 
between "dia: iana and mouth of Rio Napo, mature nonin- 
undated forest on laterite, 4 Nov. 1979 (fr), Gentry et 
al. 27438 (MO), Yanomeno, Explorama Tourist 
f 8 


Gentry et al. 37204 (M 
ca. 5?2'S, 74°30'W, 160 m, umido restinga forest, 
O m, 3-11 


lower Rio Huallag 135 m, 
Killip & Smith 27656 (US); Río Marañon Valley, San 
Lorenzo, between Fury: of Río Pastaza and Río Hualla- 


ga, 150 m, 20 Aug.-9 Sep. 1929, Killip et al. 29227 
(US); Iquitos, Muena Caño, 105 m, 9 Feb. 1932, Mexia 
6504a (F, UC; M , Río Amazonas near Tamishi- 


yacu, 3 Sep. 1 1976, Revilla 1281 (MO); Pebas on Amazon 


194 


Annals of the 
Missouri Botanical Garden 


€ - July 1929 (fr), Williams 1751 (F); Pebas on 
y 1929 (fr), Williams 1787 (F); La Vic- 
inm on ia 929, Williams 2629 (F); 
same locality as preceding, 29 Aug. 1929, Williams 2938 
(F); alto Río Itaya (San Antonio), 145 m, Sep.-Oct. 1929 
e, E illiams 3398 (F); Puerto Arturo, Yurimaguas, low 
uallaga, 155-210 m, 16 Nov. 1929 (fr), Wil. 
liams. = 1486) PASCO: Oxapampa, Palcaz valley, Cabeza 
, 9-6 km Iscosacin, 17-20 Apr. 1983, 
Smith 3787 (MO); SAN TE Mariscal Cáceres, To- 
cache Nuevo, Quebrada de Cachiyacu de Huaguisha, 570 
m, 16 July 1982 (fr), Meerow et al. 1023 (FLAS); Mar- 
iscal rc Tocache a de Cachiyacu 


jo de Tocache Nuevo, 


uallaga), 21 Apr. 1971 
UCAYALI: Coronel Portillo. Yarina Cocha, = “El Pe 
cador," cerca al Caserio Nuevo Destino, e de Yarina 


Cocha, 150 m, 31 Oct. 1984, Schunke 14153 (FLAS). 


Eucharis ulei is among the more widespread 
Amazonian taxa of subg. Eucharis, extending north 
from its Peruvian center of distribution into Co- 
lombia and south to Bolivia (Fig. 81). The species 
is best recognized by its primarily narrow-elliptic 
leaves, chiefly five-flowered inflorescence, tube 
length of ca. 3-4 cm, limb spread of 4-5 cm, and 
reduced ovule number (generally two per locule, 
Fig. 89C). Both flower and ovule number have 
become nearly fixed throughout the range of the 
species. Eucharis castelnaeana is sympatric with 
E. ulei at times but tends to occur on seasonally 
inundated soils. Flower size and number, ovule 
number, and chromosome morphology (Meerow, 
1987b) suggest close relationship to E. cyaneo- 
sperma, which has different tube morphology, a 
deeply trigonous ovary, and blue-coated seeds. 

Ravenna (1982: 50) described Urceolina moana, 
citing the “absence of lobes, or teeth, in the cup." 
The single, poorly preserved, fragmentary flower 
of the holotype, however, had stamens at least 
shortly dentate to quadrate. As in E. candida and 
E. formosa, androecial toothing in E. ulei has little 
taxonomic significance. The androecial morphol- 
ogy of E. moana is well included within the range 
of variation for this character in £. ulei (Fig. 70D)— 
several collections of E. ulei have completely eden- 
tate staminal cups (e.g., Meerow et al. 1024, Plow- 
man & Kennedy 5811, Prance et al. 13915). 
Much the same argument applies to the likewise 
synonymous E. boliviensis (Ravenna, 1988), rep- 
resented by the only specimen of E. ulei collected 
in Bolivia. The only other collection assigned by 
Ravenna (1988) to E. boliviensis (Cardenas 1553) 


belongs to E. cyareosperma. Eucharis ipariensis, 
inexplicably described by Ravenna (1982) as allied 
to E. mastersii (= E. x grandiflora of subg. Het- 
erocharis), is also indistinguishable from numerous 
collections of E. ulei. 

Krànzlin (1913) and Macbride (1936) noted the 
dissimilarity of the two specimens comprising the 
holotype of E. ulei (Ule 57376), one in flower, the 
other in fruit. I have assigned the fruiting specimen 
(Ule 5737b), with blue seeds, to E. cyaneosperma 
Meerow. When a putative isotype of Ule 5737 
was received from MG, it proved to represent E. 
castelnaeana. Ule thus collected three species un- 
der a single number, an unsurprising occurrence 
in areas where several Eucharis species are sym- 
patric. 


9. Eucharis cyaneosperma Meerow, Sida 12: 
29-49. 1987. TYPE: Peru. San Martin: 20 
km N of Tocache Nuevo on road to Tarapoto, 
Rio Canuto, 520 m, 17 July 1982, Meerow 
et al. 1032 (holotype, FLAS). Figure 89A. 


Bulb subglobose, 3-5 cm long, 3-3.5 cm diam., 
tunics light brown. Leaves 2-4; petiole (10-)15- 
30(-35) cm long, 5-6.5 mm thick; lamina 
(ovate-)elliptic, 18-25(-30) cm long, (6.5-)7-8 
(-13) em wide, apically acute to shortly acuminate, 
attenuate at the base. Scape (3-)4-5(-6.5) dm 
tall, 5-7 mm diam. proximally, 3-4 mm diam. 
distally; bracts ovate-lanceolate, (20-)27-35 mm 
long. Flowers (3-)5(-7), pendent, without fra- 
grance; pedicels (10-)15-25(-28) mm long, ca. 
1.7-2 mm diam.; tube 30-40 mm long, 1.5-2 
mm diam. for most of its length, dilating abruptly 
to 7-9 mm proximal to the throat, curved abruptly 
ca. 5 mm above the ovary and then + straight; 
outer tepals 23-28(-32) mm long, 8-10(-13) mm 
wide, ovate-lanceolate, apiculate; inner tepals 21- 
24(-30) mm long, 10-14(-15) mm wide, ovate, 
acute to minutely apiculate. Staminal cup cylin- 
drical, (8—)10—12 mm long (to tooth or lobe), 10- 
13(-15) mm wide, pale yellow or green proximally, 
quadrate or irregularly toothed between each free 
filament, the teeth when present < 1.5 mm long, 
cleft between each stamen 2-3 mm deep; each 
stamen 3.5-4(-5) mm wide at the base; the nar- 
row, subulate free filament (3-)3.5-4.5(-5.5) mm 
long, ca. 1.5 mm wide at the base; anthers ca. 3 
mm long, oblong; pollen grain ca. 47.95 um polar 
diam., ca. 67.55 um longest equatorial diam. Style 
4.5-6 cm long, exserted 0.5-1 cm beyond the 
anthers; stigma 2-2.5 mm wide. Ovary subglobose 
and deeply trigonous at anthesis, 5-7 mm long, 
7-10 mm wide, usually wider than long; ovules 


Volume 76, Number 1 
1989 


Meerow 195 


Eucharis and Caliphruria 


2(-3, 5) per locule, superposed in the lower half 
of the cell. Capsule 10-12 mm long, 15-20 mm 
wide, bright orange, leathery; pedicel 25-36 mm 
long; seed ellipsoid, 7-9 mm long, ca. 5 mm wide, 
with a lustrous, cobalt blue testa. Chromosome 
number: 2n — 

Distribution. Sporadic in the understory of 
premontane to lower montane rain forest of the 
Amazon basin and eastern Andean foothills, from 
Peru to Bolivia (Fig. 81), 120-800(-1,200) m 
elevation. Flowering at any time of the year, most 
commonly in August. 


Additional specimens p as BoLiviA. EL BENÍ: 
vicinity of Rurrenabague, 3 25 Nov. 1921, Car- 
denas 1179 (AA, NY, US); "een ua 500 
Oct. 1921, Cardenas 15534 (NY); San Antonio, 15 Nov 
1958, flowered in cultivation 30 Apr 
301 (MO); same collection as preceding, flowered in cul- 
tivation 4 Apr. 1961, Nelson 58-301 (MO). BRAZIL. ACRE: 
basin of Rio Purus, near mout i 


0 1943, Sandeman 
3724 (K, OXP.1 LORETO: eta, CERDA del Puerto Belen, 
i *58'S, 120 m, 16 Nov. 1978 (fr), 


120 = 29 May 1979 (fr), 
Cocha Iricahua, Rio 


nas, Yanamono, Explorama Tourist Camp, Rio Amazo 
halfway between Indiana and mouth of Rio Napo, 72°50'W, 
3°28'S, 120 m, 28 Aug. 1983, Gentry et al. 43758 
(MO); Flor de Yarina (Rio Samiria), 22 Aug. 1983 (fr), 
7 (MO); Ucayali, Canchahuayo (Isla 
low. je S, 200 m ov. 1985 (fr), 
997 (MO); lower Río Nanay, 24 May 


Aug. 1929 (fr), Williams 2619 (F); 
azon, 28 Aug. 1929 (fr), Williams 2878 (F, US). JUNÍN: 
Puerto Yessup, ca. 400 m, 10-12 July 1929 (fr), os 
& Smith 26394 (F, NY, US); Rio Negro to Satip, 

m, 17 Aug. 1960, Woytkowski 5830 (MO). MADRE DE 
pios: Tahuamanu, Iberia, 200 m, 15 Nov. 1973 (fr), 
Alfaro 1684 (MO); Iberia, Miraflores, vicinit Rio Ta- 
huamanu, 1 Sep. 1945, Silent 214 5 (US). 


Nuevo, Quebrada de Huaguisha (margen derecha de 
Huallaga, 400-450 m, 3 July 1974 (fr), Schunke 7146 
(F). UCAYALI: middle Ucayali, Cashiboplaya, 10°S, 1923, 
Tessman 3179 (G, NY, S). 


Eucharis cyaneosperma is the only species of 
Eucharis with blue-coated seeds. The species ap- 
pears very similar to E. ulei but differs by having 
usually shorter leaves, different tube morphology 
(abruptly curved at the base in E. cyaneosperma, 
sinuously curved in F. ulei), irregularly dentate to 
quadrate staminal cup (Fig. 89A), deeply trigonous 
ovary, and different seed color. In the southern 


part of its range, E. cyaneosperma occupies more 
upland sites than are usually characteristic of F. 
ulei. 


10. d Mer in Regel, Gartenfl. 38: 
13-314, 1300, fig. 1. 1888. Urceolina 
de (Regel) Traub, Pl. Life 27: 57-59. 
1971. TYPE: Colombia. Cauca: ex hort. Regel, 
from bulbs collected by Lehmann near Po- 
payan, Apr. 1888, Regel s.n. (holotype (frag- 
mentary), LE; photo of holotype, FLAS, K). 
Figure 90D-F. 


Bulb subglobose, not seen. Leaves 2; petiole 
45.5 cm long, 3.5 mm thick; lamina ovate-elliptic, 
24-25 cm long, 16 cm wide, short acuminate, 
basally subcordate and attenuate to the petiole. 
Scape not seen; bracts lanceolate; bracteoles linear- 
lanceolate. Flowers 4; pedicels to 30 mm long; limb 
patent, spreading to ca. 4 cm; tube ca. 25 mm 
long, ca. 1.2 mm wide below, dilating abruptly to 
8 mm at the throat; outer tepals 20-22 mm long, 
8-10 mm wide, ovate-lanceolate, acute-apiculate; 
inner tepals 18-20 mm long, 10-12 
obtuse. Staminal cup 7-8 mm long (to apex of 
tooth), deeply cleft between each stamen to < 2 
mm from the throat; each stamen bidentate, the 
teeth long-lanceolate, as long as E free aus d 
free filament linear, 7-8 mm mm 
wide; anthers sub-basifixed, ES, Style Anio 
exserted beyond the anthers. Ovary globose-ellip- 
soid, ca. 6 mm long, ca. 4 mm wide; ovules ca. 
10 per locule. Fruit and seed unknown. 


mm wide, 


Distribution. 
story of moist, lower montane forest of the Cor- 
dillera Oriental in Cauca Department of Colombia 
(Fig. 91), 1,200 m. 


Extremely rare in the under- 


Additional specimen examined. COLOMBIA. CAUCA 
Aganche, Río Orejas, 1,200 m, Lehmann 5883 (K). 


Eucharis lehmannii is known only from the 
fragmentary type (a single flower) and a single 
specimen at Kew with only one flower present. An 
Ecuadorean specimen mislabeled as the type of E. 
lehmannii, received from Kew (Lehmann 7775), 
turned out to be E. astrophiala (Ravenna) Ra- 
venna. Like E. caucana, to which it may be very 
closely related, E. lehmannii is a peripheral isolate 
of subg. Eucharis, with novel androecial character 
states. Despite the unusual staminal morphology, 
the plant figured in Regel's (1889) description re- 
sembles Eucharis subg. Eucharis by overall mor- 
phology and habit of the flower. In 1984, I was 
not able to collect this species successfully at the 


196 


Annals of the 


Missouri Botanical Garden 


FIGURE 90. 
al. 5983, UC). —A. Flower.—B. And 


lehmannii (after tab. 1300 in Regel, 1889 


Ovary, longitudinal section. 


locality of Lehmann 5883 in Colombia, an area 
now largely deforested. 


11. Eucharis corynandra (Ravenna) Ravenna, 
Phytologia 57: 95-96. 1985. Urceolina cor- 
ynandra Ravenna, Pl. Life 34: 80-81. 1978. 
TYPE: Peru. Cajamarca: Chinganza, between 
Aramango and Montenegro, 2 July 1973, 
Ravenna 2090 (holotype, herb, Ravenna, not 
seen; isotypes, K, not seen: NY, MO). Figure 
90G-I. 


Bulb globose, 47 mm long, 37 mm diam.; tunics 
light brown. Leaves elliptic-lanceolate; petiole 25- 


Eucharis corynandra, E. lehmannii, and E. oxyandra. A-C. E. oxyandra (isotype: Hutchison et 
roecium. Note the polymorphism. —C. Ovary, Zu section. 

and a photo of the type at LE. —D. Flower. — E. Stamin 

Ovary, longitudinal section. G-I. E. corynandra (isotype, Ravenna 2090, K). 


al cup. -F 
—G. Flower. —H. Staminal cup.— I. 


27 cm long, 3.5-5 mm thick; lamina 20-27 cm 
long, 5 cm wide, apically acute. Scape 50 cm tall, 
slender, ca. 2 mm diam. distally; bracts lanceolate, 
ca. 32 mm long. Flowers 8-10; pedicels 15-23 
mm long, thin; perianth tube curved, 17-18 mm 
long, ca. 1 mm wide for most of its length, abruptly 
dilated to 3-4 mm at the throat; limb spreading 
to 3-4 cm wide; outer tepals 27-29 mm long, 6- 
7 mm wide, ovate-lanceolate, acute-apiculate at 
the apex; inner tepals ca. 28 mm long, 8-10 mm 
wide; ovate, acute. Staminal cup short, funnelform, 
thick and waxy in texture, 3.5 mm long, 7.5 mm 
wide, with two obtuse teeth between each free 
filament, somewhat plicate; free filaments club- 


Volume 76, Number 1 Meerow 197 
1989 Eucharis and Caliphruria 


m 


O 200 400 600 
LI Li 1 


do | la 
| | | 
FIGURE 91. Distributions of Eucharis castelnaeana (circles), E. corynandra (squares), E. lehmannii (snowflake), 


E. oxyandra (stars), E. plicata ril d rd (closed triangles), and E. plicata subsp. brevidentata (open triangles) 
in northwestern-central South Americ 


198 


Annals of the 
Missouri Botanical Garden 


shaped, appearing narrowly elliptic-lanceolate when 
dry, ca. 6.2 mm long, 1.8 mm wide, abruptly 
dilated to ca. 1 mm at the base, inserted adaxially 
between 2 of the teeth; anthers oblong-linear, ca. 
3 mm long, versatile, black. Style 30-44.5 mm 
long, stigma trilobed. Ovary globose-ellipsoid, 6 
mm long, 3 mm wide; ovules 4-6 per locule, su- 
perposed. Fruit and seed unknown. 


Distribution. Known only from the type lo- 
cality (Fig. 91), in tropical forest on slopes, near 
a small ravine with Dieffenbachia sp., Heliconia 
sp., Clusia sp., and Xyphidium coeruleum Aubl. 


Elevation not reported. 


In size and number of the flowers, as well as 
ovule number, Eucharis corynandra shows affinity 
to E. castelnaeana (Baillon) Macbr. and to E. 
plicata Meerow. The short staminal cup and dis- 
tinctive club-shaped free filaments (Fig. 90H) are 
the principal unique characters of this species. I 
cannot, however, confirm Ravenna's (1978) de- 
scription of the anthers as densely pubescent. The 
collection locality suggests that it is an upland 
isolate from the Amazonian center of the subgenus, 
the plant having colonized the lower limits of the 
“ceja de la selva" forest formations in Cajamarca 
Department of Peru. 


12. Eucharis oxyandra (Ravenna) Ravenna 
[subg. — ae sedis], Phytologia 

57: 95-96. . Urceolina oxyandra Ra- 
venna, du x 251-253. 1983. TYPE: 
Peru. Huánuco: Huánuco, Rio Chinchao, be- 

low Carpish on rd. to Tingo Maria, 1,800 m, 

19 July 1964, Hutchison et al. 5983 (spec- 
imens prepared from bulbs flowered in culti- 
vation at UC, 26 Apr. 1967) (holotype, USM, 

not seen; isotypes, MO, UC). Figure 90A-C. 


Bulb not seen. Leaves (one seen) elliptic; petiole 
17 cm long, 4-6 mm thick; lamina 25 cm long, 
12 cm wide. Scape 27-28 cm tall; bracts 34-35 
mm long, ovate-lanceolate. Flowers 6-7, (crater- 
iform?), cernuous; pedicels ca. 19 mm long; tube 
curved, 25 mm long, 1.5 mm diam. below, dilated 
abruptly at the throat to 3.5 mm; tepals spreading 
to 27 mm wide; outer series 17-19 mm long, 6.7 
mm wide, ovate-lanceolate, acute-apiculate; inner 
series 17-18 mm long, 8-9 mm wide, ovate, ob- 
tuse. Staminal cup very short, 0.8-1.5 mm long, 

-3.5 mm wide, edentate or with 2 obtuse teeth 
between the free filaments; free filaments ca. 7 
mm long, ca. 1 mm wide at the base, narrowly 
subulate; anthers oblong, 3.3 mm long, sub-basi- 
fixed, eventually versatile; pollen grain ca. 42.36 


um polar diam., ca. 68.36 um longest equatorial 
diam. Style 32.8 mm long; stigma 2-2.5 mm wide. 
Ovary globose-ellipsoid, 6 mm long, 4 mm wide; 
ovules 6-8 per locule. Fruit and seed unknown. 


Distribution. Eucharis oxyandra is not known 
in the wild state. The collector's field notes (UC) 
indicate that three bulbs were found at the type 
locality (Fig. 91) “exposed on surface of ground 
with evidence of past cultivation, in deep shade 
near abandoned hut above road. The third bulb 
aue to be Urceolina urceolata (R. & P.) M. L. 
Gree 


This unusual species has the smallest staminal 
cup of any Peruvian species of Eucharis subg. 
Eucharis (Fig. 90B). The long, almost linear, free 
filaments are another unusual feature of the an- 
droecium. The polymorphism of the staminal cup 
(both edentate and obtusely dentate forms repre- 
sented in the isotypes, Fig. 90B) is puzzling and 
adds further credence to my suggestion that this 
character has been over-weighted as an indicator 
of species delimitation in the alpha-taxonomic lit- 
erature relating to the genus. Alternatively, it is 
possible that the two bulbs found by the collector 
and representing this species were of two different 
origins. 

In publishing Urceolina oxyandra, Ravenna 
(1983) was apparently unaware of the unusual 
situation in which the species was found, having 
examined only a duplicate at USM, which seems 
to have lacked the detailed field notes of the col- 
lector. He argued that the unusual morphology of 
the androecium (reduced staminal cup and long, 
narrowly subulate filaments), similar to that of Ur- 
ceolina, supported the inclusion of Eucharis within 
Urceolina (Traub, 1971), a position he recently 
reversed (Ravenna, 1985). This type of androecial 
morphology is characteristic of two putative inter- 
generic hybrids between Eucharis and Urceolina, 
X Urceocharis edentata Wright (1910) and x U. 
clibranii Masters (1892). 

Given that the plant was discovered as an artifact 
of cultivation in company with a bulb of Urceolina, 
and in a geographic area of loose sympatry for the 
two genera, I thought it possible that E. oxyandra 
might represent a hybrid between Eucharis and 
Urceolina. X Urceocharis edentata was suppos- 
edly collected in the wild in Peru (Wright, 1910). 
Pollen of E. oxyandra, however, stains 100% with 
Alexander's (1969) stain. Both x U. edentata and 
X U. clibranii produced very little pollen (Wright, 
1910), an observation confirmed when I examined 
specimens of both hybrids (ex hort. BM). Pollen 


grain size of the species is more like that of Eu- 


Volume 76, Number 1 
1989 


Meerow 199 
Eucharis and Caliphruria 


charis subg. Eucharis, but exine sculpturing re- 
sembles that of Urceolina (Meerow & Dehgan, 
1988). Fertile hybrids between closely related gen- 
era are not unknown in Amaryllidaceae (Traub, 
1963). 

Alternatively, Eucharis oxyandra might be a 
relict taxon close to the ancestor of Urceolina, a 
genus whose divergence from Eucharis may have 
been relatively recent (see previous section). Phy- 
logenetic analysis supports this latter hypothesis. 
At present, it seems best to treat E. oxyandra as 
a species of Eucharis, even though its known shared 
characteristics with Eucharis are plesiomorphic, 
since its phylogenetic relationships are obscured 
by the large amount of unknown character state 
data. As in E. astrophiala, E. caucana, E. cor- 
ynandra, and E. lehmannii, this species exhibits 
a pattern of morphological novelty characteristic 
of peripheral isolation in Eucharis. 


13. Eucharis plicata Meerow, Brittonia 36: 18- 
25. 1984. TYPE: Peru. San Martin: Mariscal 
Cáceres, Tocache Nuevo, Quebrada de Hua- 
guisha, right bank Rio Huallaga opposite To- 
cache Nuevo, 8?09'S, 76°27'W, 500-600 m, 
14 Dec. 1981, Plowman et al. 11394 (ho- 
lotype, FLAS; isotypes not seen, F, K, NY, 
USM). 


Bulb subglobose, 5-6 cm long, 4 cm wide, tunics 
brown. Leaves 2-4 at anthesis, petiole 25-35 cm 
long; lamina widely elliptic to ovate, (19-)20-30 
(-35) cm long, 7-12(-14) cm wide, short acu- 
minate, very shortly attenuate to the petiole, thin, 
lustrous dark green adaxially, silvery-green abax- 
ially, abaxial cuticle densely striate. Scape 40-60 
cm tall; bracts 29-30 mm long, ovate-lanceolate; 
bracteoles successively shorter and narrower. Flow- 
ers (7-)9-10, sometimes lightly fragrant; pedicel 
suberect, 10—15(—25) mm long; perianth tube 25- 
28 mm long, 2.5-3 mm wide throughout most of 
its length, dilating abruptly to 6-10 mm at the 
throat; limb spreading to 30-40 mm; tepals ovate, 
recurved at the apex, subequal; outer series ca. 
19-24 mm long, 9-14 mm wide, apiculate; inner 
series 16-23 mm long, 11-15.5 mm wide, apically 
acute. Staminal cup campanulate, ca. 8.5-1 
long (to apex of teeth), 10-12(-15) mm wide, 
apically white, basally pale greenish yellow, pli- 
cately folded on either side of the filamental trace, 
bidentate; each stamen 4.5-5.5 mm wide, the an- 
ther-bearing part subulate, 2-3.5 mm long, api- 
cally obtuse; anthers oblong-linear, sub-basifixed, 
erect at first, then becoming versatile, grayish 
brown, 3.5-4 mm long; pollen grains 41.3-43.5 


um polar diam., 59.9-68.9 um longest equatorial 
diam. Style 36-40 mm long; stigma ca. 2.5 mm 
wide. Ovary globose-ellipsoid, green, ca. 5-6 mm 
long, 3.4-4 mm wide; ovules 4-8 per locule. Cap- 
sule ca. 1 cm long, 2 cm wide, bright orange, 
leathery; seeds 1-2 per locule, ca. 1 cm long, 5 
mm wide; testa shiny black. Chromosome number: 


2n = 46 


KEY TO THE SUBSPECIES OF EUCHARIS PLICATA 


la. Flowers s uam e Pen perianth tube 
dilating the throat; staminal 
cup xd plicate, ca. 12 mm long to apex of 
biis teeth ot Ee adm late portion of fil- 
ment, co e, imbricate; style 
nd to half ik lent of the cup; ovules 
7-8 per locule . . E. plicata subsp. plicata 


lb. Wes E agant ‘perianth tube dilating 
o 7.5-10 at T throat; staminal cup only 
ml pice a. 8-10 mm long to apex of 


eet eeth sh 


just below the ben ovules 4-5(-7) per locule 
2b. E. plicata subsp. brevidentata 


12a. E. plicata subsp. plicata. Figure 92. 


Flowers without noticeable fragrance; pedicel 
suberect, 10-1 m long; perianth tube 22-24 
mm long, dilating abruptly to 6-7.5 mm wide at 
the throat; outer tepals 18-23 mm long, 9-12 
mm wide; inner tepals 16-20 mm long, 11-12 
mm wide. Staminal cup ca. 12 mm long and wide, 
deeply plicate on either side of the filamental trace; 
the anther-bearing portion of each filament inserted 
between 2 irregular, obtuse, coarsely serrulate teeth, 
each 4-5 mm long, adjacent pairs somewhat im- 
bricate and appearing as one; pollen grain ca. 43.45 
um polar diam., ca. 68.9 um longest equatorial 
diam. Style reaching to V4 the length of the staminal 
cup. Ovules 7-8 per locule. 


Distribution. Known only from the type lo- 
cality, where it is locally abundant (Fig. 91). This 
population appears to have hybridized with E. ulei 
and contains a range of intermediate morphs in 
addition to true E. plicata. The intermediates show 
reduced pollen stainability, the presence of non- 
homologous chromosomes, and extreme allozyme 
heterogeneity. It is unclear whether these represent 
a highly variable F,, the F,, or the results of in- 
trogression with E. plicata subsp. plicata. An anal- 
ysis of this population is in preparation. 

Additional specimens examined. PERU. SAN MARTÍN: 
same locality as the type, 16 July 1982, Meerow et al. 
LAS); same locality as the type, 4 Aug. 1974, 
Schunke 8046 (F). Putative hybrids with E. ulei: PERU. 


hu 
© 


200 Annals of the 
Missouri Botanical Garden 


/ 


B 
2. Eucharis ee subsp. plicata (Meerow et al. 1025, FLAS).—A. Habit.—B. Inner tepal. =C, 
em pesa — D. Stigma. — E. Staminal cup. — F. Ovary, longitudinal section. — G. Ovary, transverse section. 


SAN MARTÍN: same locality as the type, 16 July 1982, ical Garden from collections by Fred Fuchs, 
Meerow et al. 1030, 1031 (FLAS). Jr., 3 Oct. 1984, Meerow 1143 (holotype, 
FLAS). 


13b. E. plicata subsp. brevidentata Meerow, 


ubsp. nov. TYPE: Bolivia. El Beni [?]: no Subspecies nova differt a subspecies typica dentibus 
collection information, ex hort. Fairchild Trop-  staminalibus brevioribus et non serrulatis vel imbricatis. 


Volume 76, Number 1 
1989 


Meerow 
Eucharis and Caliphruria 


Flowers mildly fragrant; perianth tube 25-29 
mm long, dilating abruptly to 7.5-10 mm wide at 
the throat; outer tepals 20-24 mm long, 9-12 
mm wide; inner tepals 18.7-23 mm long, 12-15 
mm wide. Staminal cup 8-11 mm long (to apex 
of teeth), 10.5-14 mm wide, only shallowly plicate; 
teeth ca. 1.5-2 mm long, reaching to about half 
the length of the subulate portion of the filament, 
obtuse, entire, nonimbricate; pollen grain ca. 41.3 
um polar diam., ca. 59.9 um longest equatorial 
diam. Style reaching to just below the apex of the 
teeth. Ovules 4—6(-8) per locule. 


The epithet of this subspecies refers to the short 
teeth of the staminal cup. 


Distribution. Rare in the understory of pre- 
montane and lower montane rain forest of north- 
central Peru and Bolivia (Fig. 91), 200-420 m. 
Flowering August- October. 


Additional specimens examined. PERU. AMAZONAS: 
Rio Mond vicinity of Huampami, ca. 5 km E of Chavez 
Valdivia, 78°30'W, 4?30'S, 200-250 m, 11 jud 
1978 (fr), pote 1024 (MO); Rio Cenepa, 10-12 
NW of Huampami, ca. 420 m, 2 2 Oct. 1972, Berlin 148 
(NY); Río Cenepa, vicinity of Huampami, ca. 5 km E of 
Chavez Valdivia, Quebrada Huampami, ca. 78°30'W, 
4°30'S, 200- 250 m, 15 Aug. 1978, Kujikot 365 (MO). 


Eucharis plicata is phenetically and cladisti- 
cally closest to E. castelnaeana. The phenetic re- 
lationship between the two species is most evident 
in E. plicata subsp. brevidentata, which in char- 
acteristics of the staminal cup and ovule number 
is intermediate between subsp. plicata and E. cas- 
telnaeana. From the latter, E. plicata may be 
distinguished by its wider leaves and tepals, larger 
flowers, campanulate staminal cup which is plicate 
along the filamental traces (Fig. 92E), more nu- 
merous ovules, and fruit and seed morphology typ- 
ical of subg. Eucharis. 


14. Eucharis castelnaeana (Baillon) Macbride, 
Publ. Field Mus. Nat. Hist. Bot. Ser. 11 
1931. Caliphruria castelnaeana Baillon, Bull. 
Mens. Soc. Linn. Paris 144: 1133-1130. 
1894. Urceolina castelnaeana (Baillon) 
Traub, Pl. Life 27: 57-59. 1971. TYPE: Peru. 
Ucayali: Pampa del Sacramento, June 1847, 
Castelnau s.n. (holotype, not located; isotype, 
P). Figure 93. 


Eucharis narcissiflora Huber, Bol. Mus. Goeldi Para 4: 
543. 1906. Urceolina narcissiflora (Huber) Traub, 
7 a 


Sarayacu, Catalina, 25 Ju 
(holotype, MG; isotype (photo and fragment), F). 


Bulb small, subglobose, 2.2-3(-4.5) cm long, 


1.4-2.7 cm wide; neck 1-2 cm long, 1-1.5 cm 
wide; tunics tannish brown. Leaves 2-4; petiole 
(10-)13-17(-25) cm long, 3-6 cm wide; lamina 
narrowly ovate-elliptic, (10-)15-20 cm long, 5- 
7(-10) cm wide, apex shortly acuminate, lustrous 
dark green adaxially, lighter green abaxially, mar- 
gins undulate. Scape (2.5-)3-4(-5) dm tall, 5-6 
mm diam. proximally, ca. 3 mm diam. distally; 
bracts (2.5-)3-4 cm long, ovate-lanceolate, green- 
ish white. Flowers (7—)8—10, pendent, small, with 
a faint lemon fragrance; pedicels 10-18 mm long; 
tube 15-25(-30) mm long, 1-1.5(-2) mm wide 
for most of its length, abruptly dilated near the 
throat to 5-6(-8) mm wide; tepals patent, spread- 
ing to (2.5-)3-4 cm, often distally recurved, some- 
times strongly reflexed for their entire length; outer 
tepals (ovate-)lanceolate, 15-20 mm long, 5-7 mm 
wide, apiculate; inner tepals ovate, 11.5-18.7 mm 
long, 8-11 mm wide, acute. Staminal cup fun- 
nelform-cylindrical to cylindrical, usually subcylin- 
drical proximally and abruptly dilated at 14-94 of 
its length, (5.5-)7-8(-9.5) mm long (to apex of 
tooth), (7-)9-11(-12) mm wide, zoned greenish 
yellow in the proximal 4—%4, slightly incurved at 
the rim, waxy and slightly succulent in texture, 
bidentate between each free filament, conspicu- 
ously plicate between the teeth, very shallowly cleft 
between each tooth (< 1 mm) and as deep or more 
deeply cleft beteen the teeth and the free filament 
(> 1 mm); teeth 0.5-1 mm long, reaching to about 
half the length of the free filaments, obtuse, entire; 
free filament subulate, (1.5—)2—3(—4) mm long, ca. 
1.5 mm wide at the base, obtuse or acute; anthers 
3-4.5 mm long, oblong, grayish brown, sub-basi- 
fixed; pollen grain ca. 39.45 um polar diam., ca. 
55.8 um longest equatorial diam. Style 25-35 mm 
long, reaching the apex of the cup or slightly in- 
serted; stigma ca. 1.5 mm wide. Ovary globose, 
3.5-5 mm diam., white to greenish white; ovules 
(2-)4-5(-7) per locule. Capsule subglobose, shal- 
lowly trilobed, ca. 1 cm long, 1.5 cm wide, glaucous 
green, thin-walled, sometimes abscising indehis- 
cently; seeds 1-3 per locule, compressed-ellipsoid, 
ca. 1 cm long, 5 mm wide; testa dull black, rugose. 
Chromosome number: 2n = 46. 


Distribution. Understory of lowland and pre- 
montane, often seasonally inundated, primary rain 
forest in the Amazon basin, most commonly in 
Peru, rare in Colombia and Brazil (Fig. 91), 100- 
200 m. Flowering June-September (-December). 


Vernacular names. Amangay, sacha cebolla. 


Additional specimens examined. BRAZIL. AMAZONAS: 
Rio Jurua, Rio Miry, July 1901, Ule 5737 in part (MG). 
COLOMBIA. AMAZONAS: vicinity Leticia, ex hort. Fairchild 


202 


Annals of the 
Missouri Botanical Garden 


Eucharis castelnaeana. 
(Schunke 1 
D. Ovary (Schunke 14156), longitudinal section 


FicunE 93. 
habit of limb. — B. Tepals 


Minas? Vos from collections made by R. Buttons, 1 
pr. Watson 1868 iue AS). PERU. LORETO: Isla 

Santa ‘Maria, near "is s, Huallaga i 150- 

200 m, 16 Sep. 1 B, Ferreyra 4984 ( 

Quebrada Sucusari, Lacha apa camp 

side of Río Napo below M 

lateritic soil, 140 m, 6 Nov. 1 


n 
een Indiana and mouth of Río Napo, 
72°48'W, 3?28'S, seasonally inundated tahuampa, 120 
m, 28 July 1980, Gentry et al. 29203 (MO); Puerto 
Arturo, lower Rio Huallaga below Yurimaguas, ca. 135 
m, 24-25 Aug. 1929, Killip & Smith 27801 (NY, US); 
same locality, Aug. 1929, Killip & Smith 27844 (F, 
NY ; 


illip & Smith 28249 (US); Santa Rosa, lower Rio 


—C. Stamin 


—A. Flowers. i, Schunke 14154-A (FLAS); ii, Due 14156. Note varying 
56).— 


4156). i, outer tepal; ii, inner tepal. 


cup (Schunke 1415 


ere below Yurimaguas, ca. 135 m, Killip & Smith 

28886 (F, NY, US); Maynas, Santa María de Nanay, 
rine San Fransisco de M gn Yaguas, 1.5 km del Fundo 

Balcón Momon, 106-1 15 Nov. 1984, Schunke 
14154a (FLAS); Alto reee Yurimaguas, camino a 
"Schunguyce" al sur este de Puerto Arturo, near Yuri- 
maguas, 150-200 m, 1 Dec. 1984, Schunke 14156 
(BM, COL, F, FLAS, G, GB, GH, K, MO, NY, P, S, UC, 
US); Paba on the Amazon, 30 July 1929, Williams 
1898 (F, S, US); m Rio Huallaga, Sapotoyacu, Santa 
Rosa, 155-210 m, 11 Nov. 1929 (fr), Williams 4906 
(F); Puerto rene Yurimagu uas, lower Rio Huallaga, 155- 
210 m, 15 Nov. 1929, Williams 5051 (F). 


Eucharis castelnaeana has been collected most 
frequently in the vicinity of Yurimaguas in Peru 


Volume 76, Number 1 Meerow 203 
1989 Eucharis and Caliphruria 
(Fig. 91) and is sometimes locally abundant in rain Subgenus Euch ffini imis differt florib 


forest understory. It has the smallest flowers of 
any species in subg. Eucharis and is the major 
exception to the correlation of reduced flower size 
with loss of fragrance that otherwise characterizes 
Eucharis. Several living collections produce a mild 
lemon scent when ambient temperatures are high 
(as does E. plicata var. brevidentata). It is the 
only species of subg. Eucharis in which the ripe 
capsule is not leathery and orange. The glaucous, 
green, thin-walled capsule is tardily dehiscent and 
can abscise without opening, though the seeds with- 
in are fully ripe. 

Eucharis castelnaeana is autogamous. Under 
greenhouse cultivation it is the only species of 
Eucharis that successfully sets fruit with self-pol- 
len. Unmanipulated inflorescences regularly set 50- 
75% of their capsules. 

Eucharis castelnaeana is variable in leaf and 
flower size (Fig. 93A) and, to a lesser extent, in 
androecial morphology. In his description of E. 
narcissiflora, Huber (1906) made reference to £. 
castelnaeana, described from the same location as 
Huber's type of E. narcissiflora, but differentiated 
his new species solely on the basis of size. The two 
taxa represent the extremes of size diversity of a 
single species and differ by no other characters. 
Populations represented by Schunke 14154a (Fig. 
93Ai) have flowers almost twice as large as those 
of Schunke 14156 (Fig. 93Aii) but are otherwise 
indistinguishable. Shape of the staminal cup in the 
latter collection can range from nearly cylindrical 
to funnelform-c ylindrical. 

Eucharis plicata is phenetically the closest 
species to E. castelnaeana. Baillon (1894) consid- 
ered E. castelnaeana intermediate between Cali- 
phruria and Eucharis in his argument for com- 
bining these two genera. Baillon probably weighed 
flower size heavily in his judgment, one of only two 
characters by which E. castelnaeana resembles 
Caliphruria. In Baillon's time, most other small- 
flowered Eucharis species were not yet described. 
Additionally, the green, thin-walled capsule of £. 
castelnaeana is like the fruit of Caliphruria. Bail- 
lon (1894) gave no indication whether he was fa- 
miliar with the fruit of either Eucharis or Cali- 

hruria. Nonetheless, perianth and pollen exine 
morphology place E. castelnaeana squarely in Fu- 
charis despite its aberrant fruit and seed mor- 
phology. 


Eucharis subg. Heterocharis Meerow, subg. nov. 
TYPE SPECIES: Eucharis X grandiflora Plan- 
chon & Linden, Fl. Serres Jard. Eur. Ser. 1, 
9: 255. 1853. 


pleru mque non ata tubo dee bene infra fauce 
edes limbo ple rumque minus expanso, ovario grandius 
ovulis numerosis en quoque loculo. 


Large bulbous perennial herbs. Leaves petiolate, 
persistent; lamina ovate, thin, plicate or smooth 
between the veins, undulate margined, apically 
acuminate, subcordate basally and shortly atten- 
uate to the petiole, bright or dark green adaxially, 
light green abaxially, the abaxial epidermal cells 
variably striate; petiole subterete. Inflorescence 
scapose, umbellate, terminated by 2 green ovate- 
lanceolate bracts. Flowers subsessile or shortly 
pedicellate, 2-7, white, 7-8 cm long, strongly fra- 
grant, declinate (rarely subpendulous), funnelform- 
campanulate (rarely crateriform); perianth tube cy- 
lindrical to subcylindrical below, abruptly dilated 
at 14-14 of its length, curved, tinged green below; 
limb of 6 ovate tepals in 2 series, imbricate for 
half their length, subequal, spreading somewhat 
above, often slightly undulate. Stamens basally con- 
nate into short or long staminal cup partially adnate 
to the upper portion of the tube, striped green or 
yellow within along the filamental traces, variously 
toothed or edentate; distal portion of the filaments 
linear or subulate, sometimes incurved; anthers 
linear, versatile at anthesis; pollen grain 40-60 
um (polar axis), 60—76 um (longest equatorial axis), 
the exine coarsely reticulate. Style filiform, well- 
exserted beyond the staminal cup and slightly as- 
surgent; stigma deeply trilobed, often green. Ovary 
large, ellipsoid, trigonous, appearing rostellate when 
dry: ovules usually per locule, ellipsoid, 
biseriate. Capsule green, seeds blackish brown with 
a slightly rugose testa (E. moorei). 


Three species and two natural hybrids in western 
Colombia, Ecuador, and rarely Peru. 


KEY TO THE SPECIES AND HYBRIDS 
OF SUBGENUS HETEROCH ARIS 


la. Leaves deeply plicate; staminal cup reduced to 
basal connation of the filaments less than 7 
mm lon 
2a. Staminal cup acutely bidentate a 
each filament, teeth ca. 1.5 mm long ....... 
x Visdonp Mel 
a 


2b. Staminal cup edentate or r rarely wit 
s 


matic papillae unicellular .. 2. 
. Perianth tube straight or only slightly 
cernuous, subcylindrical proximally but 
distally dilating gradually towards the 


o2 
c 


204 


Annals of the 
Missouri Botanical Garden 


throat; free filament midi stig- 
matic papillae rhe 
m€— — x Ca nu butcheri 
lb. Leaves shallowly plicate or o —— ate; 
staminal cup well developed, greater than 1 c 
lon, 


ng. 
4a. Leaf length-to- width ratio usually equal to 


dentate, strongly recurved 

ovules 16-20 per locule; plants of Ecua- 
r, very rare in Peru E. 

; Leaf length-to-width ratio usually greater 

than 2; perianth crateriform, tepals spread- 

ing ca. 90° from t the throat; pie cup 

patina i l t t ongly 


D 
c 


ecurved at the margin; us 9-12 per 
ub: ded of the Huallaga valley in Peru 
5. E. amazonica 


1. Eucharis Xgrandiflora Planchon € Lin- 
den, Fl. des Serres Jard. Eur. Ser. 1, 9: 255. 
1853. Urceolina ro le (Planchon & 
Linden) Traub, Pl. : 57-59, 1971 
LECTOTYPE (Meerow & PARA 1984a): Fl. 
des Serres Jard. Eur. Ser. 1, 9: pl. 957. Fig- 


ures 94-96. 


Eucharis mastersii Baker, Curtis's Bot. Mag. 111: t. 
6831. 1885 Mrd d (Baker) Traub, 
da ei 27: eg. 59. TYPE: ex hort. Sander 

pe, K; photo, 

Eucharis | ia Baker, Gard. Chron. 13: 538-539. 1983. 
Urceolina lowii (Baker) Traub, Pl. Life 27: 57-59. 

71. TYPE: ex hort. Low (holotype, K; photo, F). 


Bulb 3-5 cm diam., neck 2-4 cm long, tunics 
light brown. Leaves 1-3; petiole 19-36 mm long, 
5-7 mm thick; lamina ovate or elliptic, 20-33 cm 
long, (10-)13-16 cm wide, apically acuminate, 
subcordate basally and shortly attenuate to the 
petiole, deeply plicate, adaxial surface lustrous dark 
or bright green, abaxial surface light green and 
densely striate, margins undulate. Scape 4-5 d 
tall, 5-6 mm diam., subterete; bracts ovate-lan- 

8 


form-campanulate, declinate, sweetly fragrant; 
pedicel short, 5-7(-10) mm long, 2.5-3.4 mm 
diam.; tube curved, 40-55 mm long, 1.5-2 mm 
wide below, dilating at 14-14 its length to 20-25 
mm at the throat, green in the proximal half, white 
distally; tepals ovate, imbricate for half their length, 
white, margins usually undulate, spreading slightly 
apically; outer series (30-)35-40 long, 
(18-)20-26 mm wide, acute-apiculate; inner series 
25-35(-40) mm long, 23-31 mm wide, obtuse. 
Staminal cup short, 5-7 mm long (to apex of teeth), 
23-25 mm wide, stained green or yellow where 


adnate to the dilated portion of the tube, with 2 
acute or obtuse teeth between each stamen, teeth 
ca. 1.5 mm long; stamens (5-)6.7-7.5(-10) mm 
wide at the base from tooth to tooth; free filament 
7-8.5(-10) mm long, linear or narrowly subulate, 
1-1.5 mm wide at the base; anthers oblong-linear, 
5.5-6.7(-7.5) mm long, slightly curved; pollen 
with only 10% stainability. Style filiform, (67 -)74—- 
85 mm long, green, assurgent, well exserted be- 
yond the stamens, stigma deeply 3-lobed, 2.5- 
3.5(-5) mm wide, white or greenish. Ovary oblong, 
rostellate, 12-19 mm long, 6-8 mm wide; ovules 
16-20 per locule, globose, superposed. Fruit and 
seed unknown, doubtfully ever formed. Chromo- 
some number: 


Distribution. Rare in the understory of pri- 
mary and secondary rain forest of southern Chocó 
and northwestern Valle de Cauca departments of 
Colombia (Fig. 99), 80-600(- 1,000) m elevation, 
but widely cultivated throughout western Colombia 
below 1,750 m. Rare and possibly introduced in 
Ecuador. The plant is functionally sterile. Most 
collections are ion cultivation, and even seem- 
ingly native populations may be remnants of cul- 
tivation. Flowering at any time of the year. 


Additional specimens examined. COLOMBIA. CHOCÓ: 
entre Istmina y 
100 m, 5 Aug. 1944, Garcia-Barriga 11525 (COL, US). 
VALLE DE CAUCA: Rio Cabrera, uve Río Sumapaz, Cun- 
dinamarca, 600-1,000 m, Feb. 1883, Lehmann 2644 


, US); Cauca, 1,000 m, Lehmann s.n. (K); 
cultivated 50 km SE of Buenaventura on old road to 

acific coast, Río Anchicaya, 500 m, 22 July 1984, 
Meerow & Teets 1127 (FLAS). ECUADOR. CARCHI: Chical, 
path from Juan Maldonado to Tobar Danuso, flowered in 
cultivation, 23 Dec 


C 4 
Mexia 6644 (GH, US): vicinity Duran, cultivated, Rose 
& Rose 23627 (US). E T.: voucher of Madison et 
al. s.n., Feb. 1983, ca 1104 (FLAS). 


Eucharis X grandiflora has long been confused 
with E. amazonica Linden ex Planchon (Meerow 
& Dehgan, 1984a). Meerow € Dehgan (1984a) 
re-established E. grandiflora (without hybrid des- 
ignation) as distinct from E. amazonica in Eucha- 
ris subg. Heterocharis. At the time, I considered 
E. lowii and E. mastersii to be distinct but closely 
related to E. x grandiflora. In 1984, I collected 
material in Colombia referable to E. lowii Baker 
and received living material of E. mastersii col- 
lected in Ecuador. According to Baker (1893), who 
described these two species from cultivated ma- 
terial, E. lowii had a staminal cup half as long as 


Volume 76, Number 1 
1989 


Meerow 205 


Eucharis and Caliphruria 


LÀ URES 94- 98. 
rsit” form 


in Eonia, — 96. Flower. 97, 98. x Calicharis Dute hak (Meerow 1098, FLAS). —97. Inflorescences. — 98. 


(Madison et al. s.n., SEL). 


that of E. “grandiflora,” 


amazonica or E. moorei (as E. grandiflora var. 


probably referring to E. 


moorei Baker). When I compared the staminal cup 
of E. lowii with that of the lectotype of E. x gran- 
diflora, the synonymy of E. lowii with E. X gran- 
diflora became evident. The two seemed identical 
in all respects. Baker considered E. lowii to be a 
hybrid of E. mastersii and E. “grandiflora” and 
E. mastersii Baker (1885) to be a hybrid between 
E. sanderi and E. “grandiflora.” Baker distin- 
guished E. lowii from E. mastersii merely by the 
former's slightly longer-exserted staminal cup, which 
does not hold up. I now believe that E. mastersii 
and E. lowii are elements of a slightly variable 
hybrid taxon, E. x grandiflora, a putative hybrid 


Natural hybrids of Eucharis subg. Heterocharis. 94-96. E. x grandiflora. —94. Flower of E. 
6. E. “lowii” 


form (Meerow & Teets 1127, FLAS).— 95. n 
Flow 


of E. sanderi and E. 


lected in western Colombia, E. moorei does occur 


moorei. Though never col- 
in contiguous northwestern Ecuador. It was likely 
a collection of this hybrid that Planchon & Linden 
(1853) described as E. grandiflora. 
X grandiflora appears intermediate in all respects 
to E. moorei and E. sanderi. Virtually all collec- 
tions of E. x grandiflora are from cultivated pop- 
ulations in Colombia and Ecuador according to 
specimen labels. Pollen stainability of either Co- 
lombian or Ecuadorean collections of E. x gran- 
diflora is never better than 10%, and I have not 
succeeded in obtaining seeds in cultivation with 


Eucharis 


either sibling pollen or pollen of other species. The 
hybrid is slightly variable in leaf morphology, num- 


206 Annals of the 
Missouri Botanical Garden 


b » ‘ 


FicuRE 99. Distributions of species and hybrids of Eucharis subg. Heterocharis in northwestern-central South 
America. Eucharis amazonica (closed circles), E. sanderi (open circles), E. moorei (closed squares), E. x grandiflora 
(open squares) and x Calicharis butcheri (closed iral s). 


Volume 76, Number 1 Meerow 207 
1989 Eucharis and Caliphruria 


E 
FIGURE 100. Eucharis sanderi (Cuatrecasas 16380, F). — A. Habit. — B. Flower. — C. Tepals. i, outer tepal; ii, 
inner tepal.— D. Detail of androecium. — E. Ovary, longitudinal section. 4 


ber of flowers, and color of androecial pigmentation ker) Traub, Pl. Life 27: 57-59. 1971. TYPE: 
throughout its range, the extremes of which were ex hort. (K). Figure 100. 

recognized respectively as E. mastersii (Fig. 94) 

and E. lowii (Figs. 95, 96) by Baker (1885, 1893). Bulb 42.5-49 mm long, 32-47 mm wide; neck 


The labels of all of Lehmann's collections indicate thick, 24-30 mm wide; tunics light brown. Leaves 
that they were prepared from material in Colom- petiolate, persistent; petiole 31-50 cm long, 5.5- 
bian gardens, and I encountered cultivated material 6(-8) mm thick; lamina ovate or elliptic, (23-)30- 
referable to this hybrid in the Cauca valley and 37 cm long, (10-)14-17 cm wide, thin, deeply 
Pacific slopes of the Cordillera Occidental. plicate between the veins, undulate margined, api- 
cally acuminate, subcordate basally and shortly 

2. Eucharis sanderi Baker, Curtis's Bot. Mag. attenuate to the petiole, bright green adaxially, light 
109: t. 6676. 1883. Urceolina sanderi (Ba- green abaxially, the abaxial surface intensely striate. 


208 


Annals of the 
Missouri Botanical Garden 


Scape subterete, 45-54 cm long, 5-6 mm diam.; 
bracts lanceolate, 45-65(-77) mm long. Flowers 
2(-3), subsessile (pedicels 5 mm or less long), 
strongly fragrant, declinate, funnelform-campan- 
ulate; perianth tube (45-)50-60(-70) mm long, 
cylindrical to subcylindrical and 2-3 mm wide be- 
low, abruptly dilated at !4 of its length to 20-27 
mm wide, curved, tinged green proximally; tepals 
white, ovate, subequal, imbricate for half their 
length, spreading somewhat distally; outer series 
26-32(-37) mm long, 16-20 mm wide, apiculate; 
inner series 24-30(-33) mm long, 20-25 mm 
wide. Stamens basally connate into a short staminal 
cup partially adnate to the upper portion of the 
tube, ca. 
along the filamental traces, only shortly exserted 
beyond the rim of the throat, edentate or rarely 
with one to few obscure teeth; free filaments 
(6-)7-9(-9.7) mm long, 1.8-2.2 mm wide at the 
base, linear, incurved; anthers (5.6-)6-8(-9) mm 
long linear, versatile, often curved; pollen grain 
39.8 um polar diam., 61.2 um longest equatorial 
diam., the exine mostly coarsely reticulate. Style 
filiform, (66-)75-80(-90) mm long, well exserted 
beyond the staminal cup and slightly assurgent, 
white, sometimes flushed green; stigma deeply an 
obtusely trilobed, (2.8-)3-4 mm wide. Ovary large, 
ellipsoid, trigonous, appearing rostellate when dry, 
(10-)15-20 mm long, (5-)7-9(-11) mm wide; 
ovules (7-)10-20 per locule, ellipsoid, biseriate. 
Fruit and seed imperfectly known; capsule green; 
seeds several per locule. 


mm wide, striped green within 


Distribution. Endemic to western Colombia 
locally on sites with rich soil in the understory of 
wet, lowland rain forest, primarily in Chocó de- 
partment (Fig. 99), frequently along watercourses; 
occasional in similar habitats in Valle de Cauca 
Dept., (5-)30-300(-1,000) m. Flowering may oc- 
cur at any time of the year. 


Additional specimens examined. COLOMBIA. CHOCÓ: 
headwaters of Río Tutunendo, E of Quibdo, 20-21 May 
1931, Archer 2197 (US); An de San José del 
Palmar, Rio Torito, 630-830 m, 5 Mar. 1980, Forero 
et al. 6728 (MO); Rio San Juan between Dipurdu & San 
Miguel, ca. 100 m, 14 Aug. 1976, Gentry & Fallon 
17687 (MO); Taparalito, Quebrada Taparal, N of Pales- 
tina, 4?15'N, 77°12'W, 30 m, 30 Mar. 1986, Gentry 
et al. 53784 (FTG, MO); Concavada region, upper Rio 

an Juan, Yeracüi valley, 200-275 m, 24-25 Apr. 1939, 
Killip 35276 (US); Andagoya, 70-100 m, Apr. 1939, 
Killip 35401 (US). VALLE DE CAUCA: Río Calima, La 
Trojita, 5-50 m, 19 Feb.-10 Mar. 1944, Cuatrecasas 
16380 (F); Río Calima, Quebrada de La Brea, 30-50 m, 
18-22 May , Cuatrecasas 21195 (F); Río Telembi, 
12 Aug. 1880, Lehmann 65 (G); Río Dagua, 0-300 m 
11 Aug. 1884, Lehmann s.n. (G). CAUCA: El Tambo, La 
Costa, 1,000 m, 3 July 1935, von Sneidern 404 (S); El 


Tambo, 800 m, 7 July 1936, von Sneidern 1129 (S). 
CALDAS: Riseralda [?], Tatáma, Santa Cecilia [La Celia?], 
800 m, 1 Dec. 1945, von Sneidern 5208 (S). 


This species is characterized by its large, sweetly 
fragrant funnelform-campanulate flowers, large 
ovaries and capsules, and reduced staminal cups. 
“Natural” hybrids exist between E. sanderi and 

oth E. “moorei” (E. x grandiflora) and “Cali- 
phruria subedentata" (X Calicharis butcheri). 
Despite the many putatively primitive character- 
istics of E. sanderi (e.g., large flowers, strong fra- 
grance) in common with E. moorei, ovule number 
of E. sanderi is highly variable, and its leaves are 
deeply plicate. The reduced morphology of the 
androecium is the major advanced character of E. 
sanderi. 


3. xCalicharis butcheri (Traub) Meerow, 


1967. Urceolina 
butcheri (Traub) Traub, Pl. Life 27: 57-59. 
1971. TYPE: ex hort. Traub, purported to have 
been collected in Panama by J. N. Giridlian, 
Traub 1051 (holotype, MO). Figures 97, 98. 


Eucharis sanderi ai iei multiflora Baker, Curtis's 
. Mag. 111: t. 6831. 1885. Eucharis sanderi 
Baker subsp. pudiera (Bake) Traub, Pl. Life 23: 
65. Urceolina sanderi subsp. multiflora (Baker) 
Traub, Pl. Life 27: 57-59. 1971. TYPE: Colombia. 

No other data, Lehmann s.n. (holotype, K). 


Bulb 6-7 cm long, 3.8-5 cm wide, neck short 
and thick, tunics brown. Leaves 2-3; petiole 20- 
40 cm long, 5-6 mm wide; lamina elliptic, 21- 
26(-35) cm long, (10.5-)12-15 cm wide, shortly 
acuminate, subcordate-attenuate at the base, con- 
spicuously plicate, adaxial surface bright to dark 
green, abaxial surface lighter green, the cuticle 
intensely striate. Scape 5-6 dm long, 4-5.5 mm 

iam.; primary bracts lanceolate, 41—57 mm long. 
Flowers 4-6(-7), funnelform-campanulate, fra- 
grant; pedicels 4.5-5(-8) mm long; tube 35-42 
mm long, funnelform proximally, dilating gradually 
from 1.5 mm wide at the base to 3.5 mm wide at 
median length, then abruptly dilated in the distal 
half to 13-16 mm wide at the throat, green for 
most of its length, the interior of the dilated portion 
stained green, most prominently along the fila- 
mental traces; tepals white, imbricate for half their 
length, spreading eventually to 45-55(-60) mm 
wide, margins nonundulate; outer series 20-28 
-35) mm long, 12-17.5 mm wide, acute-apicu- 
late; inner series 19-27(-32) mm long, 18-25 
mm wide, obtuse. Stamens shortly connate below, 
edentate or rarely with one obscure tooth between 
some of the filaments, white; free filament linear, 


— 


Volume 76, Number 1 
1989 


Meerow 209 
Eucharis and Caliphruria 


straight or slightly curved apically, 6.7-8.7 mm 
long, 0.6-1 mm wide at the base where abruptly 
dilated to 2.5-3 mm; anthers (4.5-)6 mm long, 
linear-oblong, grayish brown, mostly devoid of pol- 
len. Style 6-7 cm long, overtopping the stamens, 
slightly assurgent, white distally, green proximally; 
stigma obtusely trilobed, (2-)2.8-3 mm wide, pa- 
pillae multicellular. Ovary globose-ellipsoid, 7.7— 
12(-15) mm long, (5-)7-9 mm wide; ovules 7- 
10(-12) per locule. Fruit and seed unknown, 
doubtfully ever formed. Chromosome number: 2n 


Distribution. Rare in western Colombia (Fig. 
99), along the Rio Dagua in Valle de Cauca De- 


partment, and the lower Cauca valley. 


Additional yy examined. COLOMBIA. VALLE 
DE CAUCA: Rio ua, 17 Mar. 1883, Lehmann 2736 
(BM, Ky) Rio Dus 200-500 m, Lehmann 7774 (F, 
K in part). CAUCA: El Tambo, La Costa, 1,000 m, 24 
1936, Von Sneidern 1269 (S) PROVENANCE 
UNKNOWN: no data, Jan. 1938, Cuatrecasas 1620 (F). 
EX HORT.: Kew acc. 430.53.43001, June 1984, Meerow 
111 E (FLAS). 

This putative hybrid between E. sanderi and 
Caliphruria subedentata was first described by 
Baker (1885) as Eucharis sanderi var. multiflora. 
It has been collected in the wild at the interface 
of p^ NUR dde of both parents and at inter- 


. The plant produces little pollen, 
none of it aid with Alexander's (1969) stain. 
Floral morphology is intermediate between both 
parents, although, like C. subedentata, the stig- 
matic papillae are multicellular. There is some 
karyological evidence of hybrid intermediacy as 


well (Meerow, 1987b). 


Heulia vati 


4. Eucharis moorei (Baker) Meerow, Sida 13: 
29-49. 1987. Eucharis grandiflora var. 
moorei Baker, Gard. Chron. 4: 628. 1888. 
TYPE: ex hort., 1888, Glasnevin s.n. (K). Fig- 
ures 101, 


Bulb 6-7 cm long, 2.5-4 cm diam., tunics brown. 
Leaves 2-3; petiole 2-4 dm long, 5-7 mm thick, 
with an anomalous arc of accessory vascular bun- 
dles near the adaxial surface; lamina broadly ovate, 
length/width ratio < 2, 17-30 cm long, 10-14 
cm wide, apex shortly acuminate, base appearing 
cordate at the base, margins coarsely undulate, 
lustrous dark green adaxially, lighter green abax- 
ially, abaxial cuticle mostly without striations. Scape 
5-7 dm tall, terete, 7-10 mm diam. proximally, 
ca. 9 mm diam. distally; bracts ovate-lanceolate, 
(25-)35-45 mm long, green. Flowers usually 4, 
rarely up to 7, + campanulate, declinate, sweetly 


fragrant; pedicels usually short, 3-10(-18) mm 
long; tube 40-52 mm long, cylindrical and 1.7- 
2 mm wide proximally, abruptly dilating at about 
its midpoint to (15.5-)18.5-25 mm at the throat, 
white except for a slight green tinge at the base; 
limb spreading less than 90? from the throat to ca. 
70 mm wide or less; tepals ovate, the margins 
undulate; outer series 3.3-4 cm long; 17-22 mm 
wide, apiculate; inner series 2.9-3.8 cm long, 22- 
27 mm wide, obtuse. Staminal cup broadly cylin- 
drical, (8-)10-15(-16.4) mm long (to apex of 
teeth), 20-25 mm wide, strongly recurved at the 
margins, white on the exterior, yellowish green on 
the interior, shallowly cleft between each stamen, 
bidentate between each free filament, teeth acute, 
2.5-3 mm long; each stamen ca. 7-8 mm wide 
tooth to tooth; free filament subulate, (5-)6-8.5 
mm long, 2 mm or less wide at the base; anthers 
5.5-6.5 mm long, oblong-linear, grayish-brown; 
pollen grain ca. 48.6 um polar diam., ca. 71.2 um 
longest equatorial diam. Style 6-7 cm long; stigma 
2.5-3.5 mm wide, white. Ovary ellipsoid-trigonous, 
(7-)10-13 mm long, ca. 5 mm ui yag 16- 
20, biseriate. Capsule globose m long, 
1.3 cm wide, slightly ble, a. E glau- 

cous; seeds 1-3 per locule, compressed globose, 

ca. Ó mm diam., turgid, testa blackish brown and 
slightly rugose. Chromosome number: 2n = 46. 


Distribution. An understory plant of primary 
and secondary lower montane rain forest of the 
Ecuadorean Andes (Fig. 99), in Morona-Santiago 
and Zamora-Chinchipe provinces on the east slopes, 
and Los Rios, Cotopaxi, and contiguous Pichincha 
provinces on the western declivity, (220-)600— 
1,200(-1,600) m; rare in the “ceja de la selva" 
of Cajamarca Department, Peru. Flowering is con- 
centrated January-March and again (June-) July- 
September 


Additional specimens examined. ECUADOR 
COTOPAXI: km 52-53 on rd. from Quevedo to Latacunga, 
Rio Pilalo, 800-950 m, 11 Aug. 1984, Meerow & Mee- 
row 1137 (FLAS); same locality, 13 Aug. 1984, Meerow 
& Meerow 1141 (FLAS). GUAYAS: Guayaquil, cultivated, 
Nov. 1926, Mille 40 (QCA). Los Rios: km 56 Quev 


N 
- 


an. 1981, Gentry et al. 30913 (MO, SEL); 
Indanza-Limón (General Plaza), 1,300-1,600 m, 23 Mar. 
1974, Harling & Andersson 12779 (GB); 3 km N of 
Tucumbatza = road Gualaquiza—Indanza, 1,200 m, for- 
est remnants, 19 Apr. 1985, Harli. ng & Andersson 24329 
(GB); Río Yunganza, rd. Limón-Mendez (78°19'W, 
2?49'S), 1,100 m, 23 Sep. 1979, Holm-Nielsen et he 
20393 & 20407 (AAU); Rio Gualaquiza and Rio B 

boiza, E Andes of Sigsig, 800-1,200 m, Lehmann 5882 


210 


Annals of the 
Missouri Botanical Garden 


FIGURE 101. 
B. Flower. — C. Ovary, longitudinal section 


(K); 10-20 km from Gualaquiza on rd. to Sigsig-Cuenca, 
1,300 m, 5 Aug. 1984, Meerow & Meerow 1135 (FLAS). 
PICHINCHA: Nanegal, W slope Andes, i 000 ft., Jameson 
9 (G, P). ZAMORA-CHINCHIPE: Yurupa , 3 June 
1947, Harling 1407 (CB); W side Rio Valladolid: 2,100- 
2,400 m, 15 Oct. Wes Steyermark 54717 (F). PERU. 
CAJAMARCA: Jaen, Rio Tabaconas men 900-1,000 m, 
May 1912, Waberbausr 6251 (GH, U 


when first 
received, was assigned to E. amazonica. In 1984, 


Herbarium material of E. moorei, 


2MM 


Eucharis moorei, after a drawing by Wendy B. Zomleffer of Dodson 5527 (SEL). — A. Habit. — 


I collected the species on both sides of the Ecu- 
adorean Andes. Morphological, anatomical, and 
karyological differences between these collections 
and the Peruvian E. amazonica became evident. 
Eucharis moorei is fully fertile, diploid (Zn — 46), 
and of fairly wide distribution throughout Ecuador. 
The anomalous arc of secondary bundles that ap- 
pear in petiolar transverse sections of E. moorei, 
and reduced, in Æ. amazonica, are not found in 


Volume 76, Number 1 
1989 


Meerow 
Eucharis and Caliphruria 


any other species of the genus investigated. Eu- 
charis moorei is putatively the most primitive 
species of the genus, and shares numerous plesio- 
morphic character states with other genera of “in 
frafamily" Pancratioidinae (e.g., large fragrant 
flower, short pedicels, and numerous ovules 

locule). Eucharis moorei is the only species in the 
genus that occurs on both sides of the cordillera. 


Y 


. Eucharis amazonica Linden ex Planchon, 
Fl. des Serres Jard. Eur. Ser. 2, 2: 1216- 
1217. 1857. LECTOTYPE (Meerow & Dehgan, 
1984a): Fl. des Serres Jard. Eur. Ser. 2, 2: 
t. 1216-1217. Figure 103. 


Bulb 3.5-6 cm diam., neck 2.8-4.5 cm long, 
bro Leaves usually 2-4; petiole 

T 325- 30(- -50) cm long, 5.5-9 mm thick; lam- 
ina long-elliptic, (20-)30-40(-50) cm long, (10-) 
12-18 cm wide, length/width ratio usually greater 
than 2, acuminate, 
attenuate to the petiole, lustrous, dark green adax- 


subcordate at the base and 


ially and shallowly or inconspicuously plicate, abax- 
ial surface lighter green, cuticular striations ob- 
scure or largely absent, margins coarsely undulate. 
Scape 4.5-7(-8) dm tall, ca. 
proximally, ca. 5 mm diam. distally, terete; pri- 
mary bracts (30-)35-58(-61) mm long, broadly 
ovate-lanceolate, green. Flowers (4-)5-6(-8), sub- 
pendulous, sweetly fragrant; pedicel (9-)10-15 
(-25) mm long; tube white throughout, curved, 
(41-)46-58 mm long, cylindrical below and 2-2.5 
mm wide, dilating abruptly at Y length to (15-) 
18-21(-24) mm at the throat; limb white, spread- 
ing widely to 60-90 mm wide; tepals ovate, outer 
series 35-45(-50) mm long, 21-30 mm wide, 
apiculate; inner series 30-40(-45) mm long, 25- 


1-1.5 cm diam. 


35 mm wide, acute. Staminal cup widely cylindri- 
cal, 11.2-13.8 mm long (to apex of teeth), (22-) 
28-30(-34) mm wide, margins slightly recurved, 
the interior stained green, particularly along the 
filamental traces, shallowly cleft between each sta- 
-9.5) mm wide at the base, 


men; stamens (-— 
each with 2 obtuse teeth, one on each side of the 
subulate free filament, rarely subquadrate; teeth 
2-3 mm long; free filament 6.5-8(-10) mm long, 
2.8-3.4 mm wide at the base; anthers oblong- 
linear, 6-7(-8) mm long, grayish; pollen ca. 51.6 
um polar diam., ca. 78.3 um longest equatorial 
diam. Style white, 66-78(-85) mm long, exserted 
1-1.5 cm from the staminal cup, slightly assurgent; 
stigma deeply 3-lobed, 3-3.5 mm wide when re- 
ceptive. Ovary oblong-ellipsoid, (8-)10-14 mm 
long, (4-)5-7.5(-8) mm wide, somewhat rostellate; 
ovules 9-12 per locule, superposed. Fruit and seed 


and E. 


Eu aoe moorei 
Meerow 1141, 
FLAS). — 103. E. amazonica Sake 14179, FLAS). 


FIGURES 102, ms 
VARI 2; 


zonica oorei (Mi 


imperfectly known, capsule reported to be green 
and the seed ellipsoid with a black testa. Chro- 
mosome number: 2n = 68. 


Distribution. An understory herb of lower and 
mid-montane rain forest of northeastern Peru, most 
prominently in the Huallaga valley in the vicinity 
of Moyobamba and Tarapoto (Fig. 99), 500-1,500 
m. Widely cultivated throughout the warm tropics 
and as a house and greenhouse plant in the tem- 
perate zone. Occasionally adventive in the West 
Indies and probably elsewhere. Flowering at least 
twice per year, (May-)July-August(-September) 


and December- March. 


Vernacular names. Amangais blanco, amangay, 
flor de cebolla, azuzena, Amazon lily, Eucharist 
lily. 

Additional specimens examined. PERU. HUÁNUCO: 
Huánuco, Tingo Maria, 20 Aug. 1940, Asplund 13214 
(S); Huisca, Tingo Maria, hwy. La Oroya-Tingo Maria, 


212 


Annals of the 
Missouri Botanical Garden 


Mar. 1977, Boeke 1196 (NY); Leoncio Prado, Rupa 
Rupa, Las Palmas, km 18.5 a la carretera Tingo Maria- 
Huánuco, 756-800 m, 22 July 1984, Schunke 14055 
(F, FLAS, MO); Leoncio Prado, Rupa Rupa, Castillo 
Grande, al oeste de Tingo María, 24 July 1984, Schunke 
14057 (COL, F, FLAS, GH, K, MO, NY, UC, US); 
Huánuco Puerte Durand north of Huánuco, Río Chinchao 
valley, 7 Nov. 1938, Stork & Horton 9880 (F, US); 
vicinity of Afilador, 670 m, Woytkowski 101 (F). SAN 
MARTÍN: Lamas, Lamas, on trail from Tabalosos to n 
along Rio Cumbaquiri, V6 hour W of Rio Mayo, ca. 1,500 
ft., 13-15 Sep. 1937, Belshaw 3416 (F); near Uchiza, 
Huallaga valley, 500-550 m, 6 Aug. 1948, Ferreyra 
5154 (MO); Sie ieee eee Moyobamba, 1,200-1,600 
m, Mar. 1934, Klug 35 
Pachiza, Río Huayabamba, 1 AE 1959, Mathia 
Taylor 3974 (F); Roque, 9 Aug. Tres ji om 92 T 
between Balsapuerto and Moyobamba, 3,000 ft., Sande- 
; Lamas, Lamas, Fundo San Rafael. sector 
ec. 1984, 


); Lamas, near Tarapoto, 
929, Williams 6348 (F, US); San Roque, 
1,350-1,500 m, Feb. 1930, Williams 7802 (F). 


Meerow & Dehgan (1984a) re-established Eu- 
charis amazonicas distinct from E. x grandiflora, 
a taxon with which it has been confused since its 
description by Planchon (1857). The species is 
indigenous only to Peru, in the upper and middle 
Huallaga valley, but is grown worldwide in tropical 
regions and may be sporadically adventive. The 
species has a somatic chromosome number of 2n 
— 68 (Meerow, 1987b; Nagalla, 1979) and is at 
least partially sterile. No specimen of E. amazon- 
ica, in contrast to most other species, has ever 
been collected in fruit; pollen stainability is only 
50-65%; and viable seed has never been produced 
in cultivation. It is therefore conceivable that all 
populations of E. amazonica constitute a single 
clone. Its distribution throughout its narrow v range 
cultur- 


possibly extinct mutual ancestor. The morpholog- 
ical differences between these taxa are cryptic and 
virtually impossible to discern in most dried ma- 
terial without dissection of the ovary. Their close 
relationship is further supported by the presence 
of secondary vascular bundles in the petiole of both 
taxa, although much smaller in E. amazonica. 


DOUBTFUL OR EXCLUDED NAMES IN EUCHARIS 


Eucharis galanthoides hort., Linden, usually 
cited as E. galanthoides (Klotzsch) Linden (e.g., 
by aS 1936; Traub, 1963), Ann. Cat. Hort. 
17: 4. 1862. This combination is based on the 
erroneous assumption that Linden's Kucharis ga- 
lanthoides was based on Mathieua galanthoides 


59 (F, G, GH, 2 NY, S M 


Klotzsch (Meerow, 1987a). The actual basionym 
was Pancratium galanthoides, a name under which 
Linden received material of a Eucharis collected 
by Gustav Wallis in Amazonas. Mathieua galan- 
thoides was collected by Warscewicz in xeric, 
northwestern Peru (Warscewicz s.n., o type 
has been located for Linden's plant, and the name 
E. galanthoides merely appeared as a listing in 
his catalog. It is likely referable to E. castelnaeana 


(Baillon) Macbride. 


Eucharis himeroessa Sandwith ex Standley, 
Herbertia 3: 4. 1936. This name was never validly 
published with diagnosis or description. It was ap- 
plied to a population of E. bouchei var. bouchei 
Woodson & Allen from San José Province in Costa 

ica. 

Eucharis paradoxa hort., T. Moore, Gard. 
Chron. 1: 242. 1876. A listed name, never de- 
scribed or illustrated, supposedly applied to the 
plant later described as Caliphruria subedentata 


Baker. 


Pancratium galanthoides hort., Linden, Ann. 
Cat. Hort. 17: 4. 1862. See discussion under Ma- 
thieua galanthoides. 


Caliphruria Herbert, Edwards's Bot. Reg. 30 
(misc. no. 83): 87. 1844. Urceolina le 
Caliphruria (Herbert) Traub, Pl. Life 27: 57- 
59. 1971. TYPE SPECIES: Caliphruria hart- 
wegiana Herbert. 


Small bulbous perennial herbs. Leaves glabrous, 
petiolate, persistent or rarely hysteranthous; petiole 
subterete; lamina ovate, ovate-elliptic or elliptic, 
slightly succulent, apically acute or acuminate, ba- 
sally attenuate to the petiole, margins nonundulate, 
dark green and smooth-surfaced adaxially, lighter 
green abaxially, the cuticle of the abaxial epidermis 
thickly striate or ridged, hypostomatic or with adax- 
ial stomata only near and upon the midrib. Inflo- 
rescence scapose, umbellate; scape slender, ter- 
minating in 2 marcescent bracts enclosing the 
flowers before anthesis. Flowers pedicellate, (3-)5- 
10(-12) on thin pedicels, each subtended by a 
linear-lanceolate bracteole, declinate or subpen- 
dulous by the laxness of the pedicel, 2-4 cm long, 
without noticeable fragrance, white, funnelform or 
funnelform-tubular, protandrous; perianth tube 
funnelform, dilating gradually from the base, tinged 
green below, or subcylindrical and white through- 
out, straight; limb of 6 lanceolate, ovate or elliptic 
tepals in 2 subequal series, the segments imbricate 
for half their length, diverging in their distal half. 
Stamens connate only at the base, forming a short, 


Volume 76, Number 1 
1989 


membranaceous staminal cup, variously toothed or 
edentate between each filament, distal portion of 
the filament narrowly subulate or linear, subequal; 
anthers oblong, erect at anthesis, eventually ver- 
satile; pollen grains medium-sized (longest equa- 
m), the exine finely reticulate. 
Style filiform; stigma trilobed, papillae multicellu- 
lar. Ovary globose, green, with septal nectaries; 
ovules ellipsoid, (1-)2-7 per locule, medially su- 
perposed, placentation axile. Fruit a thin-walled, 
leathery, yellow-green, loculicidal capsule; seeds 


torial axis ca. 50 p 


ew per locule; globose or compressed-ellipsoid, ca. 
3-5 mm long, testa black or brown. Chromosome 
number: 2n = 46 


Four species, three endemic to western Colom- 
bia, one to Peru 


KEY TO THE SPECIES OF CALIPHRURIA 


— 


a. Perianth more than 2 cm long; tube 9 mm or 
more long 
2a. Perianth tube 9-13.5 mm long; stamens 
5.8-6.8 mm long, bidentate between each 
stamen, length of en at a Y the riis 
of the free k. artwegiana 


never more than Y, he length of the fre 
filament 7. C. sube aid 
Perianth rs to or less than 2 cm long; tube 
less than 9 mm lon 
Ja. Leaves es thous; perianth tube fun- 
nelform; stamens less than 5 mm long, 
more irr fasciculate, bidentate, teeth 
much siens the E filament in aO 
free filamen m long, ovules 
per locule 3. C tenera 
3b. Leaves t perianth tube subcylin- 
drical; stamen m long, divergent, 
edentate, free dient 7-9 mm long, ovules 
4 


4-5 per locule C. korsakoffü 


E 


Ne 


[d 


. Caliphruria hartwegiana Herbert, Ed- 
wards's Bot. Reg. 30, misc. no. 83: 87. 1844. 
Eucharis hartwegiana (Herbert) Nicholson, 
Illus. Dict. Gard. 1884. Urceolina hartweg- 
iana (Herbert) Traub, Pl. Life 27: 57-59. 
1971. The species as described by Herbert 
from Hartweg collections made near Guaduas, 
Colombia, lacks a holotype, and I have not 
located any specimen which could be desig- 
nated as lectotype. The following neotype is 
herein proposed. NEOTYPE: Colombia. Huila: 
Rio Magdalena near Paicol, 600-1,200 m, 
Lehmann 6376 (neotype, K). Figure 104B. 


Bulb globose with a brown tunic, ca. 32.8 mm 
diam., often apically articulated into a slender neck 


Meerow 213 
Eucharis and Caliphruria 
B 
poem 


> 


m ape of EAE A 
289, COL). — B. C. hart- 

Sie Lean 5d K). C, 5 C * à 

itti Holotype, ex hor 


BC. korsakoffi (Meerow 1096, 


FIGURE 104. 
Beaufoy s.n. 
FLAS). 


to 19.6 mm long. Leaves 2-4; petiole 13 cm long, 
2.5-4.4 mm thick; lamina ovate to elliptic, 10— 
14 cm long, 3.8-6.3 cm wide, acuminate, shortly 
attenuate to the petiole at the base. Scape 28-31 
cm tall; bracts lanceolate, 13.5-16.8 mm long and 

. 3 mm wide at the base; inner bracteoles suc- 
cessfully smaller and narrower. Flowers 5-7; ped- 
icels 12.8-15.8 mm long, thin; perianth 19-26 
mm long, white, except for the proximal Y3-1% of 
the tube, this green; tube funnelform, dilating grad- 
ually from the base, 9-13.5 mm long, ca. 2-3 
mm wide at the base, 5-6 mm wide at the throat; 
limb spreading to 7.5-11 mm; tepals ovate-elliptic, 
the outer series ca. 14.8 mm long, 3.4 mm wide, 
acute-apiculate, the inner series ca. 12 mm long, 

mm wide, obtuse. Stamens 5.8-6.8 mm long, 
narrowly subulate for most of their length, dilated 
to ca. 2.5 mm basally where connate, bidentate 
between each free filament; teeth ca. 2 the length 
of the free filament; free filament ca. 1.5 mm wide 
at point of insertion to cup; anthers 5.2-5.4 mm 
long, linear, versatile. Style 25 mm long, exserted 
ca. 2.4 mm beyond the limb; stigma 1.8 mm wide; 
ovary globose, 2.8-3.1 mm diam.; ovules 2-3 per 
locule, axile, medially superposed. Mature fruit and 
seed unknown. 


214 Annals of the 
Missouri Botanical Garden 


FIGURE 105. Distribution of species of Caliphruria in northwestern-central South America. Caliphruria hart- 
wegiana (triangles), C. korsakoffii (star), C. subedentata (circles), and C. tenera (square). 


Volume 76, Number 1 
1989 


Meerow 215 
Eucharis and Caliphruria 


FIGURE 106. 
al; ii, inner tepal.— 
* nieder section. 


Distribution. Rare in western-central Colom- 
bia, in the understory of montane forests of the 


Río Magdalena Valley (Fig. 105), 600-2,350 m. 


Additional specimens examined. COLOMBIA. 
CUNDINAMARCA: Caparrapi, 1,275 m 
García-Barriga 7713 (COL). Ex HORT.: 
s.n. (GOET). HUILA: forest a Fines erg, 
2,350 m, 6 July 1984, D'Arcy et a 4 (MO); 
Municipio de La Plata, Vereda Agua Bonita, cupa Meren- 


A 
Caliphruria subedentata (Meerow 1123, FLAS).— epa 
— D. Longitudinal section through part of flower to show detail of decia =E 


—A. Habit.—B. Flower.—C. Te 


als. i, outer 
Ovary, 


berg, 1,200-1,300 m, 15 July 1975, Díaz et al. 534 
COL). 


This seldom-collected species is distinguished 
from C. subedentata by its smaller flowers and 
short stamens with long teeth interposed between 
the filaments (Fig. 104B) 


2. Caliphruria subedentata Baker, Curtis's 
Bot. Mag. 103: t. 6289. 1877. Eucharis sub- 


216 


Annals of the 
Missouri Botanical Garden 


edentata (Baker) Bentham & Hooker, Genera 
Plantarum 2: 731. 1883. Urceolina subeden- 
tata (Baker) Traub, Pl. Life 27: 57-59. 1971. 
TYPE: ex hort. Beaufoy s.n. (holotype, K; pho- 
to of type, COL). Figure 106. 


Eucharis fosteri Traub, PI. Life 7: 36. 1951. vro 
fosteri (Traub) Traub, Pl. Life 27: 57-59. 
TYPE: Colombia. Cauca: above Cali on road to e 
ventura, 2,000 ft., collected by M. B. Foster, 5 Dec. 
1946, Traub 17 (holotype, MO). 


Bulb globose-subglobose with a brown tunic, 
22-45(-63) mm long, 20-32 (-44.5) mm wide, 
often apically articulated into a slender neck 11.8- 
23.5 mm long and 10-15.7 mm wide, offsetting 
vigorously. Leaves 2-4; petiole 10-28(-37) cm 
ong, 4-7 mm thick; lamina ovate, ovate-elliptic, 
or elliptic, 14.5-19.5(-21) cm long, (4.5-)7-10 
cm wide, acuminate, shortly attenuate to the pet- 
iole. Scape 25-40 cm tall, slender, 2.5-5.5 mm 
diam.; bracts 13-25.5(-30.5) mm long, 3-5.2 mm 
wide at base; inner bracteoles 5-14 mm long. Flow- 
ers (3-)5-7(-8); pedicels (13.6-)15.2-28(-44.3) 
mm long, thin, ca. 1 mm diam.; perianth (24-) 
31.6-39(-44.8) mm long, white except for the 
distal 14-14 of the tube, this green; tube funnelform, 
dilating gradually from the base, 15-25 mm long, 
ca. 2 mm wide at the base, 5-8 mm wide at the 
throat; limb spreading to 12.5-21 mm wide, tepals 
ovate-elliptic; outer series (15-)17.5-22.5 mm 
long, 5-8 mm wide, acute-apiculate; inner series 
(13.5-)15.8-20 mm long, 7-10 mm wide, obtuse 
and minutely apiculate. Staminal cup less than 1 
mm long, edentate, rarely with 1-2 small teeth 
between all or some of the filaments, white; fila- 
ments narrowly subulate-linear for most of their 
length, (8-)9-11.5(-14.6) mm long, dilating to 
1.5-2.5 mm at their point of insertion; tooth, when 
present, 1 mm or less long; anthers (5-)6.2-7 
(-7.5) mm long, linear; pollen grain ca. 39.3 um 
polar diam., ca. 50.9 um longest equatorial diam. 
Style (28.9-)30-40(-45) mm long, exserted 4-6 
mm beyond perianth, white; stigma 3-lobed, 1- 
1.5(-2.5) mm wide. Ovary globose-ellipsoid, 3-5 
mm long, 2.5-5 mm wide; ovules (2-)3-5(-7) per 
locule, axile, superposed. Capsule ca. 1 cm diam.; 
seeds 1-2 per locule, compressed-ellipsoidal, 5-7 
mm long, ca. 3-5 mm diam.; testa black, rugose. 


Distribution. Understory of lower and middle 
montane forests of western Colombia in the cor- 
dilleras Occidental and Central, chiefly in the Rio 
Cauca valley (Fig. 105), (760-)1,100-1,800 
(-2,000) m. Flowering at any time of the year. 


Additional specimens examined. COLOMBIA. CAUCA: 
Santander de Quilichao; selva densa en “Río Chiquito,” 


2,000 m, 6 Oct. 1954, Fernandez 2795 (COL); Rio 
Cauca, 1,400 m, Sep. 1881, Lehmann 939 (BM, G); 
Caloto, 1,200-1,500 m, June 1883, Lehmann 2885 (G); 
Quebrada Guatica, 1,800 m, Lehmann 3269 (K); Ca- 
jamarca, Lehmann s.n. (K); Caloto, 1,200-1,500 m, 
June 1880, Lehmann s.n. (K); along Rio La Paila above 
Corrinto, Central Cordillera, 1,400 m, 19 Jan. 1906, 
Pittier 1009 (US); vicinity of Medellín, 23 Mar. 1927, 
Toro 111 (NY). VALLE DE CAUCA: Río Dagua Valley, La 
Margarita, ca. 760 m, Apr. 1939, Killip 34872 (US); 
"La Manuelita," Palmira, 1,090-1,100 m, May 1922, 
Pennell & Smith us (US). Ex HORT.: 3 Apr. 1906, 
Dammer s.n. (B, GH photo); Bailey Hortorium BH 71- 
813, May 1982, ue 1097 (FLAS); Bailey Hortorium 
BH 76-467, June 1984, Meerow 1109 (FLAS); Hun- 
tington Ere Garden HUNT 44356, June 1984, 
Meerow 1123 (FLAS), Bailey Hortorium BH 60-777, 
July 1985, Meerow 1153 (FLAS); ex hort. Bull, s.n. (K); 
ex hort. Linden, s.n. (K). 


Caliphruria subedentata is the most common 
and variable species of Caliphruria, at one time 
found throughout the moist lower and middle mon- 
tane forests of the slopes of the cordilleras Central 
and Occidental west of 76°W longitude (Fig. 105), 
and chiefly those surrounding the Cauca Valley. 
Much of this forest type has been destroyed in 
western Colombia so the species is no longer abun- 
dant. In the course of fieldwork in western Colombia 
in 1984, I was not able to find C. subedentata in 
its historical localities nor in the remnants of lower 
and middle montane forest that 1 explored. The 
species may yet persist in more inaccessible for- 
ested sites. Living material received from Mr. 
Thomas Fennell was said to have been collected in 
Ecuador, but this information is undocumented. At 
present, I have seen no Ecuadorean specimens of 
Caliphruria, nor encountered populations in the 

eld 

Within the range of C. subedentata, variation 
in leaf size, flower size, dentation of the androecium 
(Fig. 104C, D), and number of ovules is evident, 
but these patterns are for the most part continuous 
and/or mosaic in distribution, rendering recogni- 
tion of subspecific taxa inappropriate. Though Ba- 
ker (1877) characterized the species by the pres- 
ence of one or two small teeth between the free 
filaments, the flowers present on the type specimen 
of C. subedentata are completely edentate. Clono- 
typic material of the taxon described by Traub 
(1951) as E. fosteri is distinguishable from other 
living material of C. subedentata only by its slightly 
smaller leaves and flowers. 


3. Caliphruria tenera Baker, Handbook of the 
Amaryllideae, p. 112. 1888. Eucharis tenera 
(Baker) Traub, Pl. Life 23: 65. 1967. Ur- 
ceolina tenera (Baker) Traub, Pl. Life 27: 
57-59. 1971. TYPE: Colombia. Río Magda- 


Volume 76, Number 1 
1989 


Meerow 217 
Eucharis and Caliphruria 


FicunE 107. 


lena, Aug. 1844, Goudot s.n. (holotype: K; 
isotype: P). 


Bulb globose or ellipsoid, ca. 25 mm diam. or 
30 mm long x 1 
articulated into a short neck 10 mm long X 5 mm 
wide; tunic grayish brown or tan. Leaves hyster- 
anthous, not seen. Scape terete, slender, 16-27 
cm tall, 1-2 mm diam.; bracts linear-lanceolate, 
17-28.3 mm long; inner bracteoles linear, the 
longest to 14.5 mm. Flowers 5-10; pedicels 14- 
23.2 mm long, thin; perianth 17-19 mm long, 
funnelform, white; tube 8.5-9.5 mm long, fun- 
nelform, dilating gradually from 1.3 mm wide at 
the base to 3-4 mm at the throat; limb spreading 
to 15.5 mm; tepals ovate to elliptic, the outer series 
8.5-11.2 mm long, 2-2.5 mm wide, acute-apic- 
ulate; the inner series 8- 10 mm long, 2.8-3.8 mm 
wide, obtuse. Stamens 3-5 mm long, each stamen 
bidentate, 1.5-2.2 mm wide, the free filament in- 
serted between the teeth; teeth much exceeding 
the free filament, 3-4 mm long; free filament 0.8- 
1.5 mm long, linear; anthers 4-5.5 mm long, 
linear; pollen grain ca. 35.2 um polar diam., 53.7 
um longest equatorial diam. Style 18-21 mm long; 
stigma 3-lobed, 1 mm wide. Ovary globose, 1.8- 
2.7 mm diam.; ovules 1-2 per locule, axile, me- 
dially superposed. Fruit and seed unknown. 


Distribution. Colombia, valley of the Rio Mag- 
dalena (Fig. 105), 400 m 


7.5 mm wide, sometimes apically 


Additonal specimen examined. COLOMBIA. CUNDI- 
NAMARCA ?: “Prov. Bogota," Copo la Parada, 400 m 
July 1853, Triana 1289 (COL, US, photo GH). 


Caliphruria tenera is the smallest-flowered Co- 


Caliphruria korsakoffii (Meerow 1096, FLAS). 


lombian representative of Caliphruria. It is readily 
distinguished by its hysteranthous leaves and the 
long teeth of the androecium which greatly exceed 
the short free filament (Fig. 104A). Unfortunately, 
C. tenera has not been re-collected since 1853. 
The hysteranthous leaves of this species suggest 
adaptation to drier habitats than usually charac- 
teristic of the genus. Pockets of xeric vegetation 
do occur in the vicinity of the type locality (Cua- 
trecasas, 195 


4. Caliphruria korsakoffii (Traub) Meerow, 
comb. nov. Eucharis korsakoffii Traub, Pl. 
Life 23: 85-87. 1967. Urceolina korsakoffii 
(Traub) Traub, Pl. Life 27: 57-59. 1971. 
TYPE: Peru. San Martin: 40 km from Moyo- 
bamba, Hierra waterfalls, 1,500 m, ex hort. 
J. N. Giridlian from bulbs collected by Lee 
Moore, 16 July 1966, Traub 1060 (holotype, 
MO). Figure 107. 


Bulb + 
without an appreciable neck, tunic brown, offset- 
ting weakly. Leaves 2-4; petiole 4.5—10 cm long, 
3-4 mm thick; lamina ovate-lanceolate to narrowly 
elliptic, 13-17 mm long, 3.3-4.8 mm wide, acute- 
acuminate, basally attenuate to the petiole, cuticle 
of the abaxial epidermis thickly ridged. Scape 15- 
25 cm tall, 3.5-3.7 mm diam.; bracts lanceolate, 
19-25 mm long, 5.5 mm wide at the base; inner 
bracteoles 8.5-22 mm long. Flowers 10-12, 15- 
20 mm long, funnelform, white, declinate to slightly 
ascendent; pedicels 2-4 cm long; tube subcylin- 
drical, straight, 4-6 mm long, 2 mm wide at base, 
3.2 mm wide at throat; limb spreading to ca. 2 


globose, 31 mm long, 29.3 mm wide, 


218 


Annals of the 
Missouri Botanical Garden 


cm; outer tepals lanceolate, 10-18.5 mm long, 4— 
mm wide, acute-apiculate; inner tepals ovate, 
7.5-16 mm long, (5-)7-8 mm wide, obtuse and 
minutely apiculate. Stamens (7-)9.5-10 mm long, 
.7 mm wide at the base, edentate, diverging dis- 
tally, narrowly subulate, white; anthers 3-4 mm 
long, oblong, dorsifixed in lower third, versatile; 
pollen grain ca. 32.3 um polar diam., 
longest equatorial diam. Style 16-21 mm long, 
white, slightly longer than limb segments; stigma 
3-lobed, (1.2-)2.5-2.7 mm wide. Ovary globose- 
ellipsoid, 3.5-5 mm long, 2-3 mm wide; ovules 
4—5 per locule, medially superposed. Capsule ca. 
l cm diam., yellow-green, thin-walled; seeds 1 per 
locule, globose, ca. 3 mm diam., testa brown. Chro- 
mosome number: 2n — 46. 


Distribution. North-central Peru (Fig. 105) 
at 1,500 m. Known only from the type locality in 
the understory of dense lower montane rain forest 
in humic topsoil on steep slopes, often growing 
wedged in the crevices of rocks. 


Additional specimens examined. 
of type iR A n, s.n. (K); clone of ss bcn e 
n. ( MO} ret of type collection, May 
1982. Meerow 1096 (FL AS). 


Caliphruria korsakoffii is the only species of 
Caliphruria presently known outside of Colombia. 
In the size of the flower this species has affinity 
with C. tenera but can be distinguished by having 
evergreen leaves, subcylindrical tubes, and eden- 
tate, divergent, and longer stamens (Fig. 104E), 
coarser pollen exine reticulation, more numerous 
ovules, and higher altitudinal limits. The seeds of 
C. korsakoffii have a brown testa; those of C. 
subedentata are black. The ridged cuticle of the 
abaxial leaf surface of C. korsakoffii (Fig. 12) is 
quite different from the narrow cuticular striations 

C. subedentata (Fig. 11). The morphological 
novelties exhibited by C. korsakoffii suggest con- 
siderable isolation from the Colombian taxa of Cal- 
iphruria. 


LITERATURE CITED 
ALEXANDER, M. P. 1969. Differential staining of aborted 
and non-aborted pollen. Stain Technol. 44: 117- 


122 
ARBER, A. 1937. Studies in flower structure III. On the 
corona and androecium in certain Amaryllidaceae. 


Bot. (London) 1: 293-304 


A 
ARROYO, S. C. 1984. Contribución al conocimiento de 
los bulbos de pe Kurtziana 17: 55-70. 
& D. F. Pen R. 1984 volutionary and 


taxonomic aspec ee = internal mor 
Amaryllidaceae dh uth Amer 


Africa. Kew Bull. 39: 467- 498 


phology in 
erica and Southern 


` DONOGHUE, M. L& 


1984. Anatomical structure in some 


amily 


ASATRIAN, M. J. 
tropical and subtropical representatives of the 
Amaryllidaceae. Bot. Zurn. 69: 1342-1346 

— P. D. Ii, Monophyly and associated terms. 

3-69. 


1972. Monophyiy again. Syst. Zool. 21: 430- 


l A evolutionary systematist's view of 
classification. Syst. Zool. 28: 441-450. 

Monophyly: its meaning and impor- 

ance. Pp. 39-46 in T. Duncan & T. F. Stuessy 

(editors), Cladistics: Perspectives on the came: 


tion of Evolutionary History. Columbia Univ. Press, 
New York. 
BaiLLoN, M. H. 1894. Sur les limites du genre Cali- 


phruria. Bull. Mens. Soc. Linn. Paris 144: 1133- 
1136 


BAKER, J. G. 1877. Caliphruria subedentata. Curtis's 
Bot. Mag. 103: t. 6289. 
85. Eucharis mastersii and Eucharis san- 
deri var. multiflora. Curtis's Bot. Mag. 111:t. 6831. 
1888. Handbook of the Amaryllideae. George 
Bell & Sons, London 
. 1893. Euc iss lowii. Gard. Chron. 13: 538- 


539. 
BAUML, J. A. 1979. A Study of the Genus Hymenocallis 
(Amaryllidaceae) in Mexico. M.S. Thesis. Cornell 


ide 
M G. & J. D 1883. Genera Planta- 
m, Volume 2. J. Cramer, Weinheim. 
LM I. N. 3. evision of the genus Pan 
m L. d in South Africa south 
of the piei dietas. oe t. don 281-291 
BnowN, N. E. 1890. s ri j Gard. Chron. 
7: 416. 


epriana 


CRONQUIST, A. 1987. A botanical critique of cladistics. 
Bot. Rev. (Lancaster) 53: 1-52. 
oe J. Aspectos de la vegetacion na- 
ural de Colombia. Revista Acad. Colomb. Ci. Exact. 
10. 221-264 
DAHLGREN, R. M. T. & H. T. CLirFORD. 1982. The 
onocotyledons: A Comparative Study. Academic 
Press, London 
& F.N. RASMUSSEN. 983. Mon cotyledon 
evolution, rS de and phylogenetic piii on. 
Evol. Biol. 16: 255-395. 
H. T. CLirrorD & P. F. Yeo. 1985. The 
Families of the Monocotyledons. Springer-Verlag, 
Berlin 
Di FULVIO, T. E. 1973. Contribución al conocimiento 
cariológico de Amaryllidaceae. Estudio cromosómico 
en Hieronymiella y otras generos afines. Kurtziana 


7: 117-131 


P. D. Cantino. 1988. Paraphyly, 
ancestors, and the goals of taxonomy: a botanical 
defense of cladism. Bot. Rev. (Lancaster) 54: 107- 
128 


EICHLER, A. W. 1875. Bluthendiagramme I. Wilhelm 
Engelmann, Leipzig. 

EINSIEDEL, B. 1987. Morphological studies in the vari- 
ation of the corona in Amaryllidaceae. Proc. 14th 
Int. Bot. Cong., Berlin: 331. [Abstract. 

ESTABROOK, G. F. 19 Some concepts for the esti- 
mation of evohationary relationships in systematic 

: 146-150. 


AN DER PijL. 1979. The Principles 
of Pollination ees 2nd edition. Pergamon Press, 
Oxford. 


Volume 76, Number 1 
1989 


Meerow 219 
Eucharis and Caliphruria 


Frrca, W. M. 1971. 
evolution: minimum 
pology. Syst. Zool. 20: 
', W.S. 1976. s nii chromosome 2 
and types of various species and taxa of Hymen 
callis bou 19: 204-227. 
77. Overview of da evolution in 
the nda Nucleus 20: 70-88. 
GOLDBLATT, P. Chromosome E sib ogy of Hessea, 
Strumaria, and Carpolyza (Amaryllidaceae). Ann. 
Missouri Bot. Gard. 63: 314-320 
GRaNT, V. 1983. The systematic and geographical dis- 
tribution of hawkmoth flowers in the temperate North 
American flora. Bot. Gaz. (Crawfordsville) 144: 439- 
449. 


Toward defining the course of 
o or a specific tree to- 
6. 


HARTIGAN, J. A. 1973. Minimum mutation fits to a 
given tree. Biometrics 29: 53-65. 
HERBERT, W. 1 Ismene macleana. 

Mag. 65: t. 3675. 
. 184 E Caliphruria ls Edwards’s 
Bot. Reg. 30 (Misc. no. 83): 8 
HEsLoP- o Y. & K. R. E. VANNA. 1977. The 
receptive surface of the angiosperm stigma. Ann. 
Bot. 41: 1233-1258. 
1973. Classification of the architecture 
cotyledonous leaves. Amer. J. Bot. 60: 17-33. 
: 84. The nature of biological species. 
s. Sci. 51: 293-307. 

* 1985. A phenetic study of Clarkia ungui- 
culata Wer. VEMM) and its relatives. Syst. 
10: 15 
"e H. Die Sammenmerkmale und ver- 

wandtschafts verhaltnisse e Liliiflorae. Mitt. Bot. 
Staatssamml. München 8: 219-298. 
» ed 2 1906. SM narcissiflora. Bol. Mus. 
e Hist. Nat. 4: 543. 
HUMPHRIES, po J. & J. A. b 1988. "pela. 
ci : a response to Cronquist. Bot. Rev. (Lan 


Curtis's Bot. 


9 The Families of Flowering 

ants, Monasatiledou 2nd edition. Clarendon Press, 
Oxford. 

JANZEN, D. H. Euglossine bees as long-distance 
pollinators of tropical plants. Science 171: 203-205. 

JerrreY, E. C. 1917. The Anatomy of Woody Plants. 
Univ. of Chicago Press, Non 

KLUGE, S. G. & J. S. Farris. 1969. bre ai PAR 
letics and the evolution E anurans. Syst. Zool. 18: 

T32, 

d Nen F. 1913. Eucharis ulei. Bot. Jahrb. Syst. 

50: 4-5 


LAKSHMI, N. ` 1978. 


pip oe studies in two allopoly- 
ploid species of the genus Hymenocallis. Cytologia 
43: 555-563 
LEwis, W. H. The Jivaro Indians: notes on an 
expedition. Discovery 17: 
LINDEN, J. J. 1862. Eucharis dalla Ann. Cat. 
H 


ort. 17: 4. 
MACBRIDE, F. 1931. 
Mus 


Eucharis castelnaeana. Publ. Field 
. Nat. Hist., No IE 


73- 675 in Flora of 
. Ser. 13. 


labores, W. P., M. J. DONOGHUE & D. R. MADDISON. 
1984. yr ad analysis and parsimony. Syst. Zool. 
33: B3- 
Mann, L. K. 


l The Allium inflorescence: some 
species of the section Molium. Ame 
39. 


r. J. Bot. 46: 


T P ji E Urceocharis x clibranii. Gard. 
Maus [3 T ie T. Duncan. 1987. The necessity 
of gra groups in biological classifications. Syst. 
Bot. 12: 790. 
MEEROW, A. v . Karyotype quen in the 
Amaryllidaceae. Herbertia 40: 139-1 
1985. e evolutionary 3 xm of pan- 
baud floral morphology in bx Amaryllidaceae. 
Amer. J. Bot. 962. [Abstrac 
1986. A j^ etin of Becher and Cali- 
phruria (Amaryllidaceae). Ph.D. Dissertation. Uni- 
versity of Florida, Gainesville, Florida. 
19 The identities and systematic affinities 
f Mathieua Klotzsch and Plagiolirion Baker 
(Amaryllidaceae). Taxon 36: 566-572. 
1987b. Chromosome cytology of ero a 
Caliphruria, and Urceolina. Amer. J. Bot. 74: 1559- 
dian 


——— —. 1987c. A p. of Eucrosia (Amaryl- 
nee Syst. Bot. 13: 492. 

nbus of tetraploid Eucharis 

(Amaryllidaceae). Ann. Missouri Bot. Gard. 73: 278- 

296. 


——— New species of Phaedranassa and 

Eucharis (Amaryllidaceae). Sida 12: 29-49. 
iosystematics of two sympatric species of Eu- 
horis (Amaryllidaceae). Pl. Syst. Evol. (in press). 
DEHGAN. Re-establishment and 
lectotypification of Eucharis amazonica Linden ex 
Planchon (Amaryllidaceae). Taxon 33: 416-422. 

& 19 Proposal for the conser- 
vation of Eucharis Planchon, 1852 (Amaryllidaceae) 
against Caliphruria Herbert, 1844. Taxon 33: 516- 
517. 


& —————. 1985. The auriculate pollen grain 
of Hymenocallis quitoensis Herb. (Amaryllidaceae) 
and its systematic implications. Amer. J. Bot. 72: 


540-547. 


988. Pollen morphology of the 
Eucharideae (Amaryllidaceae). Amer. J. Bot. 75: 
870. 


The experimental taxonomy of 
the West African species of Pancratium L. (Amaryl- 
lidaceae). Kew Bull. 19: 337-347 

NaGALLA, L. 1979. Cytological studies in Euchar ja 
Ex Planch. & Linden. Microbios. Lett. 
89-9 


NEFF, N. hi 1986. A rational basis for a priori character 
weighting. Syst. Zool. 25: 110-123. 

NICHOLSON, G. 1884. Illustrated Dictionary of Garden- 

ing. L. Upcott Gill, London. 

NorDaL, I. & T. Duncan. 1984. A cladistic analysis 
of Haemanthus and Scadoxus. Nordic J. Bot. 4: 
145-153. 

OnNDUFF, R. The origin and relationships of 
Lasthenia bunte (Compositae). Amer. J. Bot. 56: 
1042-1047. 

Pax, F. 1888. Amaryllidaceae. /n: A. Engler & K. 
Prantl, Die Natürlichen Enden 2nd edi- 
tion. 5: 97-124. 

PercivaL, M. S. 1965. Floral Biology. Pergamon Press, 
Oxford. 

PLANCHON, J. E. 1857. Eucharis vur Vua Fl. des 
Serres Jard. Eur. Ser. 2, 2: 1216- i 
& J. J. LINDEN. 1852. Fucharis and E. can- 

dida. ip J. J. Linden, Ann. Cat. Hort. 8: 3. 


220 


Annals of the 
Missouri Botanical Garden 


——— € ——. 1853. Eucharis grandiflora. Fl. 
des Serres Jard. Eur. Ser. 1, 9: 255. 

PoNNAMMA, M. G. 1978. Studies on bulbous ornamen- 
tals I. Karyomorphology of diploid and triploid taxa 
E E triflorum Roxb. Cytologia 43: 717 


RAVEN, P. H. 1976. Systematics and plant population 
biology. Syst. Bot. 1: 284-316. 
RAVENNA, P. F. 1978. Contributions to South American 
MM VII. Pl. Life 34: 69-91. 
980. Hymenocallis heliantha. Bot. Not. 


133: E 98. 

82. Contributions to South American Ama- 
ryllidaceae IX. IV. Rev tare studies in the genus 
Urceolina. Pl. Life 38: 48-54. 

1983. Revisional yarn in the genus poo 
lina rr ger i II. Wrightia 7: 251-25 
985. Taxonomical notes in fs and 
TuS Phytologia 57: 95-96. 
198 new species of Eucharis (Amar ylli- 
daceae) from Bolivia. Phytologia 64: 218. 
Rees, A. R. 19 Eucharis grandiflora. Pp. 3 
3 in A. H. mr (editor), CRC Handbook ul 
Flowering. CRC Press, Boca Raton, Florida. 
REGEL, E. Pa Eucharis lehmannii. Gartenflora 37: 
3 


RENDLE, A. B 1901. The bulbiform seeds of certain 
Amaryllidaceae. J. Roy. Hort. Soc. 26: 89-96. 
SATO, l aryotype pela and phylogeny. 
IV. seca in Amaryllidaceae with special ref- 

erence to S romosomes. Cytologia 9: 203-242. 
SEALY, à R. 1937. Lepidochiton quitoensis. Curtis's 
ag. 163: t. 9491. 
1972. Floral morphology of the Amarylli- 
ae. I. Subfamily Amarylloideae. Canad. J. Bot. 


ade Y 


155 5. 
1952. Chromosome counts of species and 
varieties of garden plants. Annual Rep. John Innes 


1965. Culeular studies as an aid to plant 
taxonomy. Bull. Brit. Mus. (Nat. Hist.), Bot. 4: 3- 


STOUT, A. B. 1944. The synflorescence of amaryllis 
hybrids. Herbertia 11: 268-273. 
Traub, H. E Eucharis fosteri. Pl. Life 7: 36. 
Classification of the Amaryllidaceae: 
v ber and genera. Pl. Life 13: 76-81. 
962. Key to the subgenera, alliances and 
species of Hymenocallis. Pl. Life 1 -12. 
1963. Genera of the licor, Amer- 
ican Plant Life Society. La Jolla, California. 
66. rio in Hymenocallis mex- 
icana. Pl. Life 22: 
——. 1967. Asmara notes. Pl. Life 23: 65. 


1971. Amaryllid notes. Pl. Life 27: 57-59. 

. 80. Sections and alliances, genus Hyme- 

nocallis Salis. Pl. Life 36: 46-49. 
& N. MOLDENKE. 1949. Amaryllidaceae: 

Tribe Amarylleae. American Plant Life Society, La 
Jolla, California. 

Van Bract, J. & P. A. SPRENKELS. 1983. ES jo 
rures of Eucharis flowers. Acta. Hort. 147: 


173-178. 

VAN DER HAMMEN, T. 1974. The Pleistocene changes 
of vegetation and climate in tropical South America. 
J. Biogeogr. 1: 3- 

i History of the flora, vegetation and 
climate i in the Colombian Cordillera Oriental during 
the last five million years. arsen 

Holm-Nielsen (editors), Mropical Botany. 
Academic Press, London. 
VAN DER Pur, L. 1982. Principles of Dispersal i in Higher 
Plants, 3rd nm Springer-Verlag, Berlin. 
VARGAS, F. C Pseudourceolina. Bol. Fac. Sci. 
Univ. Cusco l: 7-8. 
81. Plant motifs on Inca ceremonial vases 
. Linn. Soc. 82: 313-325. 
63. Das sexuelle Anlockungsprinzip der 
Catasetinen- und Stanhopeen Blüten und die wahre 
Funktion ihres sogenannten Futtergewebes. Oesterr. 


Bot. Z. 2 : 308- 
.& 


WALKER, J. W J. A. DOYLE. 1975. The bases of 
angiosperm phylogeny: palynology. Ann. Missouri 
Bot. Gard. 62: 664-723. 

WATROUS, L. E. & Q. D. , WHEELER. 1981. The out- 
g P i analysis. Syst. 
Zool. 30: 1-11. 

WERKER, E. & A. ect 1975. Seed anatomy of Pan- 
cratium species from three different habitats. Bot. 


Gaz. (Crawfordsvile) 136: 396-403. 

WHEELER, Q. D . Character weighting and cla- 
distic analysis. Syst. Zool. dd iac 109. 

WhHITEHEAD, M. R. & C. A. Bro The seed 
of the spider lily, Hymenocallis occidentalis: Amer. 
J. Bot. 27: 199-203. 

Wiley, E. O. 1981. Phylogenetics: The Theory and 
Practice of xi iae Systematics. John Wiley & 
Sons, New 

WiLLIAMS, M. p. . Chromosome count for Par- 
amongaia weberbaueri Velarde. Pl. Life 37: 83- 

9. 

Wricut, C. H. 1910. Urceocharis Xedentata. Bot. 
Mag. 136: t. 8359. 

ZAMAN, M. A. & B. N. CHAKRABORTY. 1974. Cyto- 
genetics of Amaryllidaceae: I. Karyomorphology and 
meiotic behavior of inversion o Eurycles 
sylvestris Salisb. Bangladesh J. Bot. 3: 51-58. 


THE SYSTEMATICS AND Ching-I Peng? 


(ONAGRACEAE)! 


ABSTRACT 


Ludwigia sect. Microcarpium, which consists of 14 species distributed mainly in the southeastern United States, 
is the second-largest section in the genus, the largest being the phylogenetically central sect. Myrtocarpus of South 


merica. 

Plants of Ludwigia sect. Microcarpium are morphologically diverse; they form a polyploid complex of four diploids, 
eight intr two hexaploids, and one octoploi oid. They are facultatively autogamous herbaceous plants capable of 
reproducing both sexually and asexually (forming stolons late in the flowering season). Based on extensive crossing 
experiments as well as field and Reden studies, the relationships among species of this section iut been clarified. 
Internal genetic barriers iem species are nonexistent, and natural hybridization is fairly common. 

Among the diploid species, L. linifolia and L. linearis have very narrow leaves, petaliferous Add and elongate 
capsules. The two species, oweve r, differ chromosomally by one réim sall translocation. F, hybrids between them 
are vigorous, have ca. 50% sta huile pollen, and produce acces quantities of viable seeds. Ludwigia stricta, the 
Cuban endemic, is apparently most closely allied to L. linifolia, based on morphological evidence, but has not yet 
been studied biosystematically. udwigia microcarpa, in contrast, has spatulate-obovate leaves, small, apetalous 

owers, and minute obconical capsules. Artificial hybrids between L. microcarpa and either L. linearis or L. linifolia 
form neither stainable pollen nor seeds; for the most part, the chromosomes of War To do not pair in meiosis. 


crossed in any direction in the experimental ues and form fertile F, onn d Natural e in plants of 
this group are also fairly common and in some cases blur the boundaries of the species. Somewhat surprisingly, none 
of the extant diploid species are considered to be direct ancestors of the tetraploids, based on cytogenetic studies. 
The two hexaploid species, L. alata and L. simpsonii, are not closely related to each other. The affinity of L. 
lata is definitely with the tetraploid species group. Hybrids between L. alata and any other tetraploid species 
consistently exhibit a modal configuration of 16 II and 8 I in meiosis. Based on de morphology and the cytology of 
hybrids involving it, L. alata may have originated through hybridization between a tetraploid species, possibly L. 
lanceolata, and the diploid L. microcarpa, or populations ancestral to each of these Epa 
udwigia simpsonii, on the other hand, is closely related to L. curtissii, which is the only octoploid in sect. 
Microcarpium. The two species are unique in sect. Microcarpium in having loculicidal capsules and, together with 
the diploid L. microcarpa, in having spatulate-obovate cauline leaves. Cytogenetic and morphological studies suggest 
that the octoploid L. curtissii may have resulted from hybridization of L. s simpsonit and a petaliferous diploid species, 
possibly L. linifolia, L. linearis, or populations like them. Ludwigia microcarpa is involved in the formation of the 
hexaploid L. uS and ide ore that of the octoploid L. curtissii. The other progenitors of L. simpsonii are 
not clear, but some species in sect. Dantia or populations ancestral to them with apetalous FR T caite leaves, 
and a prostrate pem are likely andes 


! This Pu was supported by the U. S. National Science Foundation under Grant No. BSR82-14879 to Dr. 


C. Hoch for helpful discussions and assistance with the manuscript and to David E. Boufford for reviewing the 
manuscript. I thank P. P. Lowry II for photographing type materials at P; Karen Prescher for cataloging herbarium 
specimens and preparing distribution maps; Emily Colletti and Jim Henrich for care of greenhouse plants; Wanling 
Peng, Chien-Chu Chen, and Yu-Mei Chiou for their excellent “a TERDA and Gloria Hoch for expertly typing the 
manuscript. I am also thankful to David E. Boufford, Emily Wood, Jychian Chen, and my wife, Wanli ing Peng, who 


LL, LSU, M, MEXU, MISS, MISSA, MO, MSC, NA, NCSC, NCU, ND, NDG, NO, NY, P, PH, RSA, S, SMU, 
TAES, TENN, TEX, TI, U, UNA, UPS, US, USCH, USF, VA, VDB, W, WAG, WCUH, WU, Z, and East Carolina 
University Herbarium. Finally, I express my appreciation to the sw s "Mash Academia Sinica, Taipei, Taiwan, 
Republic of China, for granting me leaves of absence to work on this 

? Institute of Botany, Academia Sinica, Nankang, Taipei, Taiwan 11529. Republic of China. 


ANN. MISSOURI Bor. GARD. 76: 221-302. 1989. 


222 


Annals of the 


Missouri Botanical Garden 


summarize, recombination from iris differentiated desi followed by polyploidization and the 


To s 
maintenance of well- -adapted genetic strains by 


combination o ogam 


vegetative EDU appears to 


have played a central role in the evolution of p species of Ludwigia EA Missam 


Ludwigia is a distinct genus of Onagraceae and 
is the only member of Jussiaeeae. Fossil records of 
pollen suggest that it is archaic and has existed at 
least 50 million years since the middle Eocene in 
both the Northern and Southern hemispheres (Gon- 
zales Guzmán, 1967; Rouse, 1962; Bratseva, 
1969). Except for its chromosome structure and 
number, which appear to be relatively specialized 
(Kurabayashi et al., 1962), Ludwigia is one 
of the least specialized genera as to wood anatomy 
(Carlquist, 1975, 1982), floral anatomy (Eyde, 
1977, 1978, 1981), and leaf anatomy (Keating, 
1982). Based primarily on its unusual nectary po- 
sition, ovular vasculature, and independently de- 
rived epigyny, Eyde (1977, 1978, 1981) dem- 
onstrated that Ludwigia represents a distinct 
branch from the common ancestor of the family, 
with all other living genera on the other branch; 
in Hennigian terms, it is the sister group of the 
remainder of the family. 

The genus Ludwigia comprises 82 species that 
are diverse enough, especially in fruit, seed, and 
flower characters, to be classified in 23 sections, 
14 of which are monotypic (Raven, 1963; Ra- 
mamoorthy & Zardini, 1987). Plants of Ludwigia 
are adapted to wet habitats in the tropics and 
subtropics. They probably originated in South 
America, the most important center of diversity 
for Ludwigia, and the area where the most prim- 
itive species are found (Ramamoorthy & Zardini, 
1987). The genus, however, is also well represented 
in temperate North America and has three endemic 
species in temperate Asia. 

In North America, there are 23 native Lud- 
wigia species, characterized by having convex or 
elevated nectaries, tetramerous, haplostemonous 
flowers, pluriseriate ovaries, free seeds with narrow 
raphes, and herbaceous habits. Most of these plants 
are confined to the Atlantic and Gulf coastal plains 
of the United States, although some species are 
more widespread, and some extend into the islands 
of the Caribbean; one is endemic in Cuba. Based 
primarily on characters of capsules, mode of per- 
ennation, and phyllotaxy, the North American plants 
are classified into sect. Ludwigia (4 species), sect. 
Dantia (DC.) Munz (5 species), and sect. Micro- 
carpium Munz (14 species). Table 1 summarizes 
the diagnostic characters of these three sections. 

Based on his study of the structure and mor- 
phology of the capsules and nectaries of Ludwigia, 


Eyde (1981) considered sections Dantia and Mi- 
crocarpium to comprise a closely knit group de- 
rived from the phylogenetically central, diploste- 
monous Myrtocarpus complex, whereas sect. 
Ludwigia is different enough to have arisen in- 
dependently from the Myrtocarpus complex. 

Section Microcarpium was selected for the pres- 
ent revision because it is the second-largest section 
of Ludwigia and is very diverse, because it has 
not been revised in light of recent new data and 
evolutionary concepts, and because several species 
of this section are variable and taxonomically dif- 
ficult, with boundaries between species often blurred 
by intermediates. 

he purposes of this revision are to clarify species 

limits, to determine the degree and significance of 
natural hybridization, and to analyze natural affin- 
ities among species of sect. Microcarpium. Toward 
this end, more than 7,000 specimens from 44 
herbaria have been consulted for morphological 
studies and for mapping. Furthermore, I have made 
three extensive field studies in the coastal plains 
of the southeastern United States. Searches for 
possible natural hybrids were made in the field 
whenever taxa occurred sympatrically. Seeds and/ 
or clonal transplants of all species were propagated 
at the Missouri Botanical Garden for chromosome 
determination and morphological study. 


TAXONOMIC HISTORY 


After the establishment of the genus Ludwigia 
in 1753 by Carl Linnaeus, the first group of North 
American species of sect. Microcarpium was dis- 
covered by the British botanist Thomas Walter 
(1788). Four of his species are currently recog- 
nized: L. glandulosa, L. linearis, L. pilosa (in- 
cluding L. rudis), and L. suffruticosa. 

Jean Baptiste A. P. M. de Lamarck (1789, 
1792) published “Jussiaea brachycarpa” and 
“Ludwigia hirsuta," based on materials collected 
from South Carolina. These names are synonyms 
of L. glandulosa and L. pilosa respectively, as 
Lamarck himself suspected. 

The next botanist to study North American Lud- 
wigia was Andre Michaux (1803), who described 
nine species of Ludwigia in his Flora Boreali- 
Americana. Of these, L. microcarpa is a member 
of sect. Microcarpium 


In 1814, Jean L. M. Poiret correctly added the 


Volume 76, Number 1 
1989 


Peng 
Ludwigia sect. Microcarpium 


223 


TABLE 1. Comparison of the native North American sections of Ludwigia. 
Sect. Ludwigia Sect. Dantia Sect. Microcarpium 
Character (4 species) (5 species) (14 species & 1 subspecies) 

Habit Perennial. Plants Perennial. Plants creep- Perennial. Plants erect, usual- 
erect, lacking sto- ing, rooting at nodes; ly stoloniferous (rhizomatous 
lons; roots tuberous roots fibrous in one species); roots fi- 

rous 
Phyllotaxy Leaves alternate Leaves opposite Leaves alternat 
Capsules Capsules pedicellate, Capsules sessile or pedi- Capsules 2d or short- d 


Seed surface 


Petals 


Nectaries 


opening by a ter 
minal pore formed 
after abscission of 
style base 

Cells in parallel col- 
umns, transversely 
elongate to the 
seed length 

Petals 8-15 mm long 


4 shallow ciliate 
depressions on the 
raised summit of 
the ovary, alternat- 
ing with the sepals 


cellate, irregularly de- 
hiscent 


Cells in parallel columns, 


transversely elongate to 
the seed length 


Petals lacking or 2-10 


mm long 


4 raised and rounded 


lobes, surrounding the 

style base and alternat- 
ing with the sepals and 
stamens 


cellate, ring-, loculicidal-, 
peeled-dehiscent (see mut x 


Cells + isodiametric or in par- 
allel columns and either lon- 
gitudinally or tranversely 
elongate to the seed length 

Petals ae (in 3 species 3- 

ong); p" petals 
dias pres 
some primarily A etsi 
ecies 

4 raised and rounded lobes, 
surrounding the style base 
and alternating with the se- 
pals and stamens 


and stamens 


Chromosome numbers n — 8 (4 species) 


Pollen grains Shed as tetrads 


Genetic compatibility Self-compatible 


n = 8 (1 species) 
n — 16 (2 species) 
n — 24 (2 species) 


Shed singly or as tetrads 


Self-compatible 


— 8 (4 species) 

n — 16 (7 species & 1 sub- 
species) 

n — 24 (2 species) 

n — 32 (1 species) 

Shed singly or as tetrads 


Self-compatible 


new species L. linifolia, nte in the southern 
United States; it resembles but is t from 
L. linearis. 

It was not until 1821 that the species of sect. 
Microcarpium were described by American bot- 
anists. Of the four species in sect. Microcarpium 
that Steven Elliott (1816-1821) described, Lud- 
wigia alata, L. sphaerocarpa, and L. lanceolata 
are currently recognized, whereas L. cylindrica 
represents a synonym of L. glandulosa Walter. 
Although Elliott correctly differentiated L. alata 
from L. lanceolata, the striking superficial resem- 
blance of the two species, especially in dried her- 
barium specimens, has constantly been a source 
of confusion among botanists (Long & Lakela, 1976; 
i dfrey & Wooten, 1981; 
Wunderlin, 1982; Clewell, 1985). The great dis- 
tinctiveness of these two species in such characters 
as floral morphology, pollen and seed surface fea- 
tures, and chromosome numbers were only appre- 
ciated recently (Peng, 1988, and this study). 


In 1836, Ludwigia polycarpa, a distinctive 
member of sect. Microcarpium distributed mainly 
in the northcentral United States, was discovered 


in Kentucky by Charles Wilkins Short and Robert 


eter. 

Shortly thereafter, John Torrey and Asa Gray 
(1838-1840), in A Flora of North America, pro- 
posed two varieties in sect. Microcarpium, i.e., 
Ludwigia cylindrica 8 brachycarpa and L. cap- 
itata B pubens. The former is currently recognized 
as the small-fruited, western subspecies of L. glan- 
dulosa (Peng, 1986), whereas the latter represents 
a natural hybrid between L. suffruticosa and L. 
pilosa (Peng, 1988). 

In 1845, George Engelmann and Asa Gray de- 
scribed a densely pubescent variety of Ludwigia 
linearis from Texas. Owing to the complex vari- 
ation pattern and the many intermediate popula- 
tions found in the range of this species, however, 
the variety is not recognized in the present study. 

Ludwigia stricta, the only Cuban endemic in 


224 


Annals of the 
Missouri Botanical Garden 


sect. Microcarpium, became known when August 
Grisebach (1866) described /snardia stricta in Ca- 
talogus Plantarum Cubensium. The correct com- 
bination in Ludwigia was made by Francisco Adol- 
fo Sauvalle (1873) in the Flora Cubana. 

Alvin Wentworth Chapman added Ludwigia 
curtissii (1883) and L. simpsonii (1892) to the 
genus in his study of the flora of the southern 
United States. 

John Kunkel Small named Ludwigia simulata 
in 1903, not knowing that it represented the hybrid 
L. pilosa X L. lanceolata. In 1933, he grouped, 
for the first time, plants of Ludwigia sect. Micro- 
carpium under “Microcarpeae,” and described 
from the Everglades of Florida L. spathulifolia, 
which has now been merged with L. curtissii. 

Burghard Helwig in 1928 published Ludwigia 
cubensis from Cuba, which, however, is now con- 
sidered as conspecific with L. simpsonii from Flor- 
ida. 

Merritt Lyndon Fernald and Ludlow Griscom 
(1935) described three additional varieties for the 
highly polymorphic Ludwigia sphaerocarpa, which 
itself may have originated as a result of hybridiza- 
tion and which now intergrades with L. pilosa 
freely (Peng, 1988). These hybrids are not rec- 
ognized in this paper for reasons discussed else- 
where. 

Paul Munz's monograph (1944, 1965) con- 
tained the most comprehensive treatment of North 
American Ludwigia in his time. In the 1944 paper, 
he formally delimited sect. Microcarpium. His con- 
cept has since been widely accepted. James Allan 
Duke (1955) was the first author to indicate and 
to provide examples of natural hybridization among 
North American species of Ludwigia. 

In the course of my biosystematic study of Lud- 
wigia sect. Microcarpium, a number of atypical 
specimens initially identified as L. pilosa Walter 
were shown to be a new species, which I subse- 
quently named L. ravenii (Peng, 1984). This rep- 
resents the last named species of sect. Microcar- 
pium, and makes the total number of species of 
this section 14. 


MORPHOLOGY 
HABIT 


All species in Ludwigia sect. Microcarpium are 
erect perennial herbs ranging from about 15 to 
100 cm tall. Leafy, usually unbranched stolons are 
produced from the base of erect stems. The stolons 
creep along the surface of moist ground or float 
on water late in the flowering/fruiting season. Some 
species, however, may also send out stolons in the 


summer while they are actively flowering or even 
throughout their growing season. The stolons can 
be long (up to 2.5 m) and stout as in L. pilosa or 
short (5-15[-20] cm) and slender as in L. micro- 
carpa and L. ravenii. The stolons give rise to erect 
shoots from their tips when the growing season 
resumes. In L. polycarpa the stolons are normally 
up to 15 cm long, and with leaves densely con- 
gested at the apex of stolons, overlapping one 
another and thereby protecting the apical meri- 
stem. This is apparently an adaptation to the severe 
winter that plants of L. polycarpa have to endure 
in the northern States. In L. suffruticosa stolons 
are uncommon. Instead, perennation is mainly by 
thick, somewhat woody, branched rhizomes. New 
shoots arise from the nodes of the rhizomes in 
growing season. It may be that in drier sandy 
situations, the plants persist by rhizomes, whereas 
in wet situations, the plants send out stolons. 

In Ludwigia curtissii and L. simpsonii, stolons 
are rarely formed; the plants simply produce new 
shoots, which are ascending or decumbent at first 
and soon become erect from the base of the pre- 
vious year's stem. They may not need to overwinter 
with low-profile stolons, as they are distributed 
mainly in Florida and the Caribbean Islands where 
the growing season is long and the winter is not 
severe. 


STEMS 


Stems are usually well branched in most species 
of Ludwigia sect. Microcarpium. Plants of L. 
suffruticosa are, however, often simple. When they . 
branch, the branches are few and usually arise 
from upper leaf axils beneath the compact inflo- 
rescences and characteristically overtop the main 
flowering/fruiting stems. Unbranched flowering 
stems are seen also in some populations of L. cur- 
tissii, L. linifolia, L. microcarpa, L. simpsonii, 
and L. stricta. The stems of some plants of L. 
microcarpa are distinctly zigzag. The stems of most 
species are slightly ridged; those of L. alata and 
L. microcarpa are often conspicuously ridged or 
winged. Ludwigia pilosa is the only species with 
nearly terete stems. The lower part of the stems 
of all species, when entirely submerged, is swollen 
and spongy. 


PUBESCENCE 


Many species in Ludwigia sect. Microcarpium 
are glabrous to subglabrous; the margins of leaf, 
sepal, or bracteoles, and surfaces of fruit may 
sometimes be minutely strigillose or minutely pa- 
pillose. Only L. ravenii and L. pilosa are consis- 


Volume 76, Number 1 
1989 


Peng 225 


Ludwigia sect. Microcarpium 


tently hirtellous throughout. Ludwigia sphaero 
carpa, a species that intergrades substantially with 
L. pilosa, is variable in many aspects, especially 
pubescence. Populations of L. sphaerocarpa range 
from being completely glabrous to densely strigil- 
lose throughout. Similarly variable is L. linearis, 
but no interspecific hybridization has apparently 
played a role here. The pattern of the distribution 
of hairs in L. suffruticosa is especially interesting: 
some plants are completely glabrous; some plants 
are hirtellous either on the terminal inflorescence 
or on the basal stems (including stolons/rhizomes), 
or both, but are otherwise glabrous. 

The following are the main types of pubescence: 

l. Minutely puberulent. Hairs exceedingly 
short, fine, straight, erect to ascending, sparse or 
dense, 0.02-0.1 mm long, found occasionally in 
plants of Ludwigia linearis, L. sphaerocarpa, on 
fruits of L. microcarpa, and less commonly on 
fruits of L. simpsonii, L. curtissit, and L. suffru- 
ticosa. 

2. Strigillose. 
cellular, 0.1—0.6 mm long, appressed to ascending; 
found in some populations of Ludwigia sphaero- 
carpa and L. linearis. 

3. Minutely strigillose. Similar to strigillose 
pubescence, but the hairs more slender and shorter, 
ranging from 0.02 to 0.1 mm long. Frequently 
seen on sepals, bracteoles, and leaves of various 
species; also on the wings of capsules in Ludwigia 
lanceolata and capsules of both subspecies of L. 
glandulosa, and occasionally of L. polycarpa. 

Blunt-tipped, minute 
projections from the surface, ca. 0.02-0.03 mm 
in length. This hair type is found occasionally on 
the margins of leaves and sepals, on fruit walls, 
and on nectary discs. It may be difficult to distin- 
guish from minutely strigillose hair when the latter 
is exceedingly short. 

. Hirtellous. stiff, straight, 
erect to spreading, with an attenuate apex and a 
slightly broadened base, 0.25-0.95 mm long, 
translucent or spotted with light brown pigments. 
Hairs of this type are typical of Ludwigia pilosa, 
L. ravenii, and occasionally of L. sphaerocarpa, 
as well as part of the plant body of L. suffruticosa. 


Hairs coarse, translucent, uni- 


EN 
~. 
3 
& 
~ 
Sle 
S 
ms 
R 
Ta 
= 
e 
N 
D 


Hairs coarse, 


LEAVES 


The leaves are simple and alternate. Ludwigia 
simpsonii is exceptional in frequently having some 
opposite or subopposite leaves at the lower nodes 
or in the seedling stage, a feature that might be 
related to the derivation of one or more of its 
genomes from plants of the opposite-leaved sect. 


Dantia (see below). The leaves on the rhizomes of 

. suffruticosa are small and scalelike. Those on 
stolons are mostly elliptic- to rhombic-obovate or 
suborbicular (narrowly elliptic to narrowly oblan- 
ceolate in L. linearis and L. linifolia). These are 
usually broader and shorter than the cauline leaves, 
which are usually narrowly lanceolate to lanceolate, 
occasionally elliptic or oblanceolate, and often 
gradually reduced and narrower up the stem. Lud- 
wigia linearis and L. linifolia have sublinear leaves. 
In L. curtissii, L. microcarpa, and L. simpsonii, 
the leaves range from (narrowly) spatulate-oblan- 
ceolate to spatulate-obovate or spatulate. 

The leaves of all species of sect. Microcarpium 
are sessile or nearly so and either narrowly cuneate 
or attenuate basally. Ludwigia suffruticosa is ex- 
ceptional in having a rounded to obtuse leaf base. 

As in Ludwigia sect. Myrtocarpus sensu lato 
(Ramamoorthy & Zardini, 1987), the leaf margin 
is usually subentire, with a few glandular reddish 
or pink teeth that have an unusual hydathodal 
structure (Castells et al., 1979; Keating, 1982); 
such teeth are especially evident in L. microcarpa. 

The secondary veins of leaves are usually ob- 
scure adaxially and distinct abaxially in sect. Mi- 
crocarpium. In the closely allied Ludwigia lini- 
folia and L. stricta and in L. simpsonii and L. 
curtissii, the secondary veins are, however, faint 
on both sides of the leaves. Ludwigia linearis, 
having narrow leaves like the related L. linifolia, 
has, by contrast, on its abaxial leaf surface very 
evident secondary veins, which anastomose and 
form a pair of marginal veins along the margins. 


INFLORESCENCE 


The flowers are sessile or shortly pedicellate and 
borne singly in the (usually upper) leaf axils of the 
main stem and the branches. In Ludwigia suffru- 
ticosa, the flowers are congested in very compact 
terminal spikes or racemes. The leaves subtending 
the flowers are much reduced and bractlike. As 
stems of this species remain single or have only a 
few branches, there are only one to several such 
inflorescences per plant. In most other species, the 
stems are well branched, and the flowers are ax- 
illary and usually very abundant. In L. alata and 
L. pilosa, the flowers are sometimes quite con- 
gested on the upper parts of branches. In L. poly- 
carpa and L. glandulosa, one to a few very short 
branches heavily laden with flowers/ fruits are often 
seen from the very base of primary branches. This 
frequently gives an appearance of several flowering 

ranches arising simultaneously from the nodes of 
the main stem. 


226 Annals of the 
Missouri Botanical Garden 
PEDICELS PETALS 


The flowers and fruits are nearly sessile in most 
species. Pedicels (measured from the base of ma- 
ture fruits), when present, are usually up to 0.5 
(-1) mm long. Pedicels of Ludwigia suffruticosa 
and L. sphaerocarpa range 0.5-1.5(-2.3) mm 
long. Populations of L. linearis from west of the 
Mississippi River often have distinct pedicels up to 


3.5(-5) mm long. 


BRACTEOLES 


Bracteoles are present in pairs near the base of 
the ovary in all species. They are succulent and 
are flanked by a pair of reddish or dark purplish, 
glandlike stipels on their base. A small swollen 
emergence located immediately below the bracteole 
is apparently an extension of the bracteole base 
and is prominent in some species, especially Lud- 
wigia curtissii, L. lanceolata, L. microcarpa, and 
L. simpsonii. This character is more evident in 
live plants than in herbarium specimens. 

Despite often being variable in length, even with- 
in the same plant (in Ludwigia linifolia, the brac- 
teoles range (1.5-)2.5-9(-13) mm long), a few 
species, including L. glandulosa, L. microcarpa, 
and L. sphaerocarpa, consistently have shorter 
bracteoles (less than 1.5 mm long) than those of 
the other species (Table 2). The bracteoles are 
located either at the base or higher up on the sides 
of the capsules. In L. sphaerocarpa and occa- 
sionally in L. glandulosa, L. linearis, and L. suf- 
fruticosa, however, the bracteoles are found on 
the short pedicels. 


FLOWERS 


Of the 15 taxa of sect. Microcarpium, only 
three diploid species are petaliferous, although ves- 
tigial petals may occasionally appear in a few flow- 
ers of polyploid species (Table 2). Flowers are con- 
sistently tetramerous, pentamerous flowers being 
exceptional. 


SEPALS 


Sepals are entire, but may be minutely strigillose 
along the margins in some species. Unlike plants 
of sect. Myrtocarpus sensu lato (Ramamoorthy & 
Zardini, 1987), the sepals persist even after the 
capsules mature and dehisce. Sepals are greenish 
in most species, but they are creamy white on the 
adaxial surface in L. alata, L. pilosa, and L. 
suffruticosa and are yellowish on the adaxial sur- 
face in L. sphaerocarpa. 


Petals are yellow, small, narrowly obovate to 
subrotund, and range from 2.5 to 6 mm long and 
from 1.2 to 5 mm wide. They fall off a few hours 
after anthesis, which occurs early in the morning, 
and are usually gone by noon. All petaliferous 
species are able to effect mechanical self-pollina- 
tion. My field experience, however, did not allow 
me to judge whether these plants attract signifi- 
cantly more insect pollinators than other apetalous 
species with showy sepals. 


STAMENS 


Most species of Ludwigia are consistently either 
haplostemonous (stamens in one whorl) or diploste- 
monous (stamens in two whorls) (Ramamoorthy & 
Zardini, 1987). All species of sect. Microcarpium 
are haplostemonous, which is a derived condition. 
The stamens are antesepalous, situated between 
the sepal base and the nectary discs. 


ANTHERS 


The anthers vary from 0.1 mm long in Lud- 
wigia microcarpa to 2 mm long in L. linearis. 
Ludwigia linearis is unique in sect. Microcarpium 
in having polysporangiate anthers partitioned hor- 
izontally by septa that include both tapetum and 
parenchyma (Tobe & Raven, 1986). Such spe- 
cialization is shared only by L. latifolia, an un- 
related South American species of sect. Tectiflora, 
and five other genera of Onagreae. It is worth 
noting, however, that these septa in the anthers 
disintegrate and result in a continuous sporogenous 
tissue before the anthers dehisce. Many conspic- 
uous vestiges of the disintegrated septa are visible 
on the inner surface of the anther wall at this stage. 


POLLEN 


The pollen of Ludwigia sect. Microcarpium is 
quite uniform, being characterized by isopolar grains 
frequently with prominent colpal extensions and 
with a psilate exine (Praglowski et al., 1983). Five 
species of sect. Microcarpium — L. alata, L. cur- 
tissit, L. microcarpa, L. simpsonii, and L. suf- 
fruticosa—shed their pollen singly, whereas the 
other nine species shed their pollen in tetrads (Table 
2). This character is of some taxonomic value. For 
example, it can be used along with seed-surface 
pattern to distinguish L. alata, which sheds its 
pollen singly, from L. lanceolata, which sheds its 
pollen as tetrads. These two species are similar in 
general aspect and in their obpyramidal, winged 


227 


Volume 76, Number 1 


1989 


Peng 
Lud 


icrocarpium 


t. M 


wigia Sec 


"1e[n2au1t SI S[[99 reuurm][oo ASAYI jo UONBJUALIO AY] seurnaurog 'q1Zue| peas əy} 0} Á[os1oAsu?J] Jo [o[[Jered ojeZuo[o s[[oo jo ounixtur e SMOYS ooejuns PAG , 
'suone[ndod esojrguis a10ur ut 1uo[eAaud ore yi3ua] 

pees eq 0} Ajasroasuesy ojeGuo[o s[[oo seazaym 'suone[ndod snoiqe[8 aso Áq penquxo Ápueumuopaid oie yi3ua] pass ay} o1 [o[[ered eieguo[e s[[oo aoejms PAG , 
'peA1esqo səwyəwos s[ejed [erduso4 (p-)e- [ YUM S19MO[J - 


Á|as1eAsue1] o1e2uo[o pue (gq) y13ua] paas eui oi [oj[e1xed ayeSuoja suurn[oo ut “([) oujeurerpost SSA] JO AJOUI s[[99 IILS YIM spoos ajeorput | pu? ‘d ‘J sapos 19197 | 


L C'b-G'I 9jeutqun y Ge'v-c'c(-z2) snoJqe[^) E :— ZE 2u-'nsuamo 7] 
L S'z-1 9jeutq1nj Á[peo1g (Cz-)z-SI snoJqe[o E = pz = u ‘nuosduns 7] 
sīəU109 papunoi 
L (cTI-)1-€0 qua [eprure1Ádqo €'[-I snoiqey‘y S - g = u 'ndu»20121u "T 
L EPS) proAoqo-Zuopqo) £'c-tv(-£) sno[[o111H L = 91 = u ‘nuaa "T 
ptoaoqo 
I G L-S6 -Buojqo o1 asoqo[3qng G-G'£(-€) sneemH L > 91 = u *psopd "T 
L pu? d (c'z-)e 1-90 asoqo¡3qng C'P-Z asoy[isis AJasuap 0} snoJqe[o) L - 91 = u *ndapoo1anyds "T 
L SP-S pesuim-p ‘jepruesAdqg P-E snoJqe[?) S - pz = uomo *] 
I a t pasum-p *peprurez&dqo) s-v(-S'£) ibi tal L = Gq =u ugaeuuer "T 
snoj[oiuq 
Á[as1eds are yərym 
s19ul09 papunoi ‘aseq ura]s pue 2DUIISIL 
I (c9-)€s-v(-€) qua [eprurex&dqo b-S'€ -opur ayi 1deoxe snosqeys) S = 91 = u ‘wsonmaffns ^T 
d S'L-S'E pro^oqo-Suo[q() L-S'b(-p) snoJqe[^) L = 91 = u *ndavo&jod ^T 
yn] pue ulgjs gp =u 
L 80-70 oupugAoqng t-z uo aso[tgujs mq *snoiqe[o L — *pd1D944y9D1q “dsqns 
jjj pue was uo eso[t2uns QT = u *»sogpnpuvjg 
d c I-€0 9upurAoqng 8-S(-p) ÁA[9seds inq snouqej? L — ‘dsqns psonpun]3 "T 
Ld S'Z-S'0 [|eprurez&dqo-o1eSuo[q OI-9 esojidugs Ájəsuəp 01 snoiqeysy L + g = u 'sumauy `T 
I Se-1 oupun&oqng $'9-G6 Sue pep) L + 8 = u "DIONS "T 
I (£1-J01-£&(€ D oupun&oqng cI-9 SAPE L * 8 =u “oyofuy T 
UOT} (wu) edeug əmsdeg (ww) y3uəT 92uoosoqng [E1240 (1) (—) I9qUInN| 9UIOSOUIOlQ") 
-&juau() pue que] a[0919e1g apnsde”) spei  juasqy YIM uoxe] 
adeys 1129 -191 s? Jo (+) 
ooepmg PA 10 (Ç)  1uas 
Ajus — -euq 
pus I 
sulted‘) 
uəjjod 


"umdaip2o4oi “Was DIFINpN'T jo satoadsqns pue səarvəds jo 


S19]9B.18y9 ousouZeu] ‘s AMV, 


228 Annals of the 
Missouri Botanical Garden 


Ficures 1-15. Mature fruits of Ludwigia sect. Microcarpium. —1. L. linifolia. —2. L. stricta. —3. L. lin- 


earis. —4. L. glandulosa subsp. glandulosa. —5. L. glandulosa subsp. brachycarpa. —6. L. polycarpa.— 1. L. 
suffruticosa. —8. L. lanceolata. —9. L. alata. —10. L. sphaerocarpa. —11. L. pilosa. — 12. L. ravenii. — 
microcarpa. —14. L. simpsonii. — 15. L. curtissii. Drawn from living plants cultivated at the Missouri Botanical 
Garden. 


fruits, and are superficially difficult to separate. modes of pollen dispersal or pollen in very loose 

e mode in which pollen is shed is useful to detect — tetrads. Finally, the percentage of stainable pollen 
natural hybrids when their putative parents differ is useful to detect sterile hybrids or hybrids with 
in this respect. Such hybrids generally exhibit both reduced fecundity. 


Volume 76, Number 1 
1989 


Peng 229 
Ludwigia sect. Microcarpium 


16 17 


FIGURES 16-18. 


DISCS 


One of the important characters that allows Lud- 
wigia to be recognized as the sister group to all 
other genera of Onagraceae is that its nectary disc 
is situated on the summit of the ovary. By contrast, 
all other onagraceous plants have nectaries at the 
base of the floral tube; the nectaries, therefore, are 
androecial (Eyde, 1981). 

The discs of Ludwigia sect. Microcarpium are 
shallowly dome-shaped, and they occur on the apex 
of the ovary. They are more or less distinctly four- 
lobed, the lobes alternating with the stamens, and 

re antepetalous (in petalous species). They are 
usually yellow or greenish at anthesis and turn dark 
reddish purple in fruit. In most species the discs 
are glabrous. Ludwigia pilosa is, however, usually 
hirtellous around the style base and between the 
disc lobes. Plants of L. sphaerocarpa are some- 
times hirtellous between the disc lobes, too. In 
densely strigillose populations of L. linearis, the 
nectary discs tend to be strigillose throughout or 
between the lobes also. In L. linifolia and L. stric- 
ta, the disc lobes are minutely papillose throughout, 
as are their fruit and seed surfaces. 


CAPSULES 


Characters of the capsules have been very useful 
to systematic and phylogenetic studies in Ludwigia 
(Small, 1933; Munz, 1942, 1944; Raven, 1963; 
Eyde, 1978; Ramamoorthy & Zardini, 1987). 
When Small (1933) first grouped plants of Lud- 


Capsular dehiscence in Ludwigia sect. Microcarpium. — 


16. L. lanceolata. —17. L. linifolia. — 


18. L. curtissii. Scale line equals 4 mm. Drawn from living plants cultivated at the Missouri Botanical Garden. 


wigia sect. Microcarpium (as ““Microcarpeae”), 
the mode of capsular dehiscence was one of the 
main criteria. Although his observations on cap- 
sular dehiscence were incomplete and not all species 
in this group have capsules dehiscing by the sep- 
aration of flaps from the Es of the nectary disc 
(see below), his concept taken up by Mun 
(1944, 1965) who ferially puse nd d. 
lished sect. Microcarpium. 

The shape, size, and pubescence of capsules are 
very diverse within sect. Microcarpium (Table 2, 
Figs. 1-15). Capsule shape ranges from obpyrami- 
dal to subcylindric, oblong-obovate, turbinate, or 
subglobose, and size ranges from 1 mm long in 
Ludwigia microcarpa to 12 mm long in L. linearis 
and L. linifolia. The surface vestiture ranges from 
glabrous to minutely papillose, strigillose, or hir- 
tellous. These characters have been extremely use- 
ful for identifying species and natural hybrids. 

Furthermore, in Ludwigia sect. Microcarpium, 
three patterns of capsular dehiscence are recog- 
nized (Peng & Tobe, 1987), each of which has a 


distinct anatomical basis: 


Ring dehiscence. The capsule dehisces along 
an apical “ring” as a result of the disintegration 
of thin-walled cells located between the nectary 
disc and the sepal base (Fig. 16). This pattern is 
exhibited by Ludwigia alata, L. lanceolata, L. 
linearis, L. microcarpa, and L. suffruticosa. lt 
was previously thought of as the only way capsules 
of Ludwigia sect. Microcarpium dehisced (Small, 
1933; Munz, 1944; Raven, 1963). In these plants 


230 


Annals of the 


Missour 


f th 
i Botanical Garden 


Volume 76, Number 1 
1989 


Peng 231 
Ludwigia sect. Microcarpium 


the zonation pattern in the capsule wall is relatively 
simple, consisting of the outer, enlarged paren- 
chymatous exocarp and the inner spongy endocarp. 
Peeled dehiscence. The capsule dehisces by 
the irregular rupture of the outer capsule wall (Fig. 
17). This is exhibited by Ludwigia glandulosa, L. 
linifolia, L. pilosa, L. polycarpa, L. ravenii, L. 
sphaerocarpa, and L. stricta. These plants have 
a spongy mesocarp and an endocarp with conspic- 
uous inner hypodermis. When the seeds are dis- 
persed, they often carry along many hypodermal 
cells. Whether these cells are useful for the seeds 
to float awaits confirmatio 
Loculicidal delis The capsule dehisces 
by amd lone four longitudinally lenticular 
it h of the loculi (Fig. 18). Eventually 
the cansada is split into four basally united parts. 
This type of dehiscence is seen only in the closely 
related polyploids Ludwigia curtissii and L. simp- 
sonii. In these species, the zonation pattern of the 
fruit wall is basically similar to that of the “ring- 
dehiscent" species, except that the exocarp con 
sists only of the outer epidermis and a single-celled 
layer of outer hypodermis compactly aligned be- 
neath. These hypodermal cells are remarkably en- 
larged and radially elongate except along the me- 
dian longitudinal line facing each locule, where they 
are abruptly reduced or replaced by small, paren- 
chyma cells. It is exactly along these four lines 
that the capsules dehisce at maturity. 
To summarize, the fruit wall of all species of 
Ludwigia sect. Microcarpium is differentiated into 
a spongy zone (endocarp in ring-dehiscent and loc- 
ulicidally dehiscent species, or mesocarp in peeled- 
dehiscent species) and a continuous zone. As Eyde 
(1978) suggested, such differentiation undoubtedly 
aids seed dispersal by reducing the effective thick- 
ness and by the differential growth rate that could 
result in the separation of the outer layers from 


the inner. The specialized hypodermis in L. cur- 
tissil and L. simpsonii appears to represent a 
condition that has evolved a step beyond this point. 


SEEDS 


In all species of sect. Microcarpium, seeds have 
a narrow raphe and are free and pluriseriate in 
each locule of the capsule, with a narrow raphe. 
The seeds are small, from .7 mm long, 
and are cylindric, ellipsoid, reniform, or ovoid. 
They are mostly yellowish, light brown, or brown 
in color, although three species, Ludwigia linifo- 
lia, L. microcarpa, and L. stricta, nearly always 
have dark reddish seeds. 

Eyde (1978) was the first to point out that 
species of sect. Microcarpium are remarkably di- 
verse in surface cell orientation. Like capsule mor- 
phology, or rather together with capsule morphol- 
ogy, seed Eu [putem 1E ae excellent 
di gnosti c identifying the species and 
detecting natural hybrids. The seed surface cells 
are either subisodiametric or are elongate parallel 
or elongate transversely to the seed length, with 
minor variation on the two ends and areas near 
the raphe (Figs. 19-34). Although in different pop- 
ulations of both L. glandulosa and L. linearis the 
seed surface cells are oriented in two different ways, 
the pattern is not random. In L. glandulosa, the 
cells are elongate parallel to the seed length in 
subsp. glandulosa (Fig. 23) and transversely elon- 
gate in the western, short-fruited subsp. brachy- 
carpa (Fig. 24). Plants of L. linearis are extremely 
variable in pubescence, ranging from completely 
glabrous to densely strigillose. In general, seed sur- 
face pattern seems to be correlated with pubes- 
cence: subglabrous plants usually exhibit cells elon- 
gate parallel to the seed length (Fig. 21), whereas 
densely strigillose individuals generally have seed 
surface cells elongate transversely to the seed length 
ig. 

In Ludwigia sphaerocarpa the seed surface 
cells (Fig. 29) are not as regularly oriented as they 
are in the other species. They are arranged in 
columns both transversely elongate and parallel to 
the seed length, with the former alignment often 
dominant in the central part of the seed. Some 


= 


— 


FIGURES 19-34. 
(MO). — 20. L. stricta, Cuba, 


Photographs of seeds of Ludwigia sect. Microcarpium. — 19. L. linifolia, Florida, Dille 408 
Prov. Piñar del Rio, Britton et al. 6613 (NY).— 


—21. L. linearis, North Carolina, 


Chatham Co., Campbell 1077 (NCU).— 22. L. linearis, Louisiana, St. Tammany Parish, Dille 420 (MO).— 23 
glandulosa subsp. glandulosa, Florida, Santa Rosa Co., Dille 412 (MO).— 24. L. glandulosa subsp. brachycarpa, 


Louisiana, Cameron Parish, Peng 4367 (MO). —25. L. polycarpa, Missouri, Lincoln Co., 
L. suffruticosa, Florida, Hillsborough Co., Peng 4327 (MO).— 27. 
MO NA Morar 11 (FSU).— 29. L. "ar dor n South Carolina, Jasper Co., 
Dille 348 (MO). — 30. L. pilosa, Georgia, Emanuel Co., Peng 4025 (MO).— 
32. L. microcarpa, Florida, Collier ay uud 4281( 


. curtissii, Florida, Collier Co., 


.— 28. L. alata, Florida, Wakulla 
Co., Boufford & Wood 18898 (MO). — 
Florida, Collier Co., Peng 4266 (MO).— 34. 
mm, except in Figure 25; there scale bar — 0.4 mm 


Dille 443 (MO).— 
L. lanceolata, Florida, Highlands Co., Peng 4183 


. ravenii, North Carolina, Sampson 
MO). — 33. L. simpsonii, 
Peng 4357 (MO). M bar — 0.5 


232 


Annals of the 
Missouri Botanical Garden 


seeds of this species have variously oriented cells, 
a pattern that supports the suggestion of a hybrid 
origin for L.. sphaerocarpa (see below). 


REPRODUCTIVE BIOLOGY 


All 14 species in Ludwigia sect. Microcarpium 
are genetically self-compatible perennials that re- 
produce sexually and vegetatively. About two-thirds 
of them are modally autogamous—a statistic com- 
parable to that for the whole genus (Raven, 1979). 
Shortly after the flowers open in the morning, the 
stigma becomes sticky and receptive and the sta- 
mens dehisce, exposing yellowish pollen entangled 
by the viscin threads. Initially the stamens are held 
away from the stigma and spread apart. The fila- 
ments, however, gradually arch toward the center 
and finally the anthers become firmly appressed to 
the sides of the sticky stigma in one to several 
hours (usually before noon), thus effecting self. 
pollination. This occurs in petaliferous as well a 
apetalous species, with the exceptions discussed in 
the following paragrap 

In Ludwigia alata, L. pilosa, L. suffruticosa, 
and many individuals of L. sphaerocarpa, the an- 
thers also bend toward the stigma at late anthesis, 
but the distance between the androecium and the 
gynoecium is such that self-pollination is impossible 
without the intervention of a pollen vector. In the 
first three, apetalous, species the sepals are con- 
spicuous and creamy within; in L. pilosa they are 
often tinged with pink along the main veins and 
the edges. Presumably these petaloid sepals serve 
to attract insect pollinators. In addition, these three 
species have yellowish discs that produce abundant 
nectar. Ludwigia sphaerocarpa, by contrast, has 
sepals that are yellowish within and very bright 
yellow discs that are more conspicuous than those 
of any other species, with the possible exception 
of L. linifolia (see below). Outcrossing in all four 
above-mentioned species is therefore not reinforced 
by protogyny/protandry or by male sterility, but 
merely by the physical separation of the stigma 
and the anthers. The self-incompatibility found in 
the more primitive members of Ludwigia sect. 
Myr tocarpus sensu lato is not present in sect. 
Microcarpium 

ectary discs are present in all species in sect. 
Microcarpium, but they vary in size and in the 
quantity of nectar produced. Ludwigia microcar- 
pa. an apetalous species with tiny flowers (the 
ovary is less than 1 mm long), has a nearly flat, 
greenish disc that produces very little nectar, and 
is presumably highly autogamous. Despite this, the 
fact that several natural hybrids—e.g., L. micro- 


carpa X L. curtissii and L. microcarpa X L. 
simpsonii, and an intersectional hybrid, L. micro- 
carpa X L. palustris (n — 8; sect. Dantia)— 
were collected in the field where the parental species 
grow sympatrically (see below) suggests that insect 
pollinators do occasionally visit plants with minute, 
inconspicuous flowers. 

Living plants from two of the three petaliferous 
species were available for observation; Ludwigia 
stricta, endemic in Cuba, has not been studied as 
a living plant. Flowers of L. linifolia have very 
conspicuous nectary discs that produce copious 
nectar. This, along with the yellow petals, serves 
to make the flowers highly attractive to insects. 
Ludwigia linearis has a comparatively shallow 
ovary disc that secretes less nectar. This species 
is noteworthy in sharing with the unrelated South 
American L. latifolia (Benth.) Hara (sect. Tecti- 
flora) the distinction of being the only members of 
the genus Ludwigia in which the sporogenous tis- 
sue of the anthers is divided into packets horizon- 
tally by both tapetum and parenchyma (Eyde, 1977; 
Tobe & Raven, 1986). This character is also shared 
by five other genera of Onagraceae, as mentioned 
previously. 

Paper wasps of the genus Polistes have been 
observed to visit flowers of species of Ludwigia 
sect. Microcarpium grown in the netted experi- 
mental greenhouse at the Missouri Botanical Gar- 
den. In the field, more insects (usually oed 
honeybees, and wasps) were attracted to L. pilo 
and, less so, to L. sphaerocarpa. Moths and ants 
have also been observed visiting L. pilosa. Rela- 
tively few field observations of insect pollinations 
on other species have been made in my trips to 
the southeastern United States. Circumstantial evi- 
dence from the presence of abundant natural hy- 
brid populations suggests that insect pollination must 
be more common than I have observed. 

In addition to sexual reproduction, nearly all 
species of Ludwigia sect. Microcarpium produce 
sprawling stolons at the base of the erect, flowering 
stems late in their flowering season. This vegetative 
reproduction enables the plants to overwinter, when 
the erect stems die back; in spring, they can then 
produce a large colony before other species can 
invade. Some of the sterile populations of natural 
hybrids also persist by this means. In £. suffruti- 
cosa, however, stolons are not as common as un- 
derground rhizomes. In L. curtissii and L. simp- 
sonii, stolons are not apparent. These plants simply 
form new shoots from the lower nodes of the died- 
back stem of the previous year. These shoots are 
decumbent or ascending initially, but soon become 
erect. 


Volume 76, Number 1 
1989 


Peng 233 
Ludwigia sect. Microcarpium 


CYTOLOGY 


Chromosomes of Ludwigia are the smallest in 
the Onagraceae, and they may differ conspicuously 
in size within a single genome. The proximal ends 
of the chromosomes are heavily pycnotic and ap- 
pear in interphase nuclei as very distinct and def- 
inite chromocenters (Kurabayashi et al., 1962). 
The basic chromosome number of the genus is x 
= 8, with no aneuploidy but extensive polyploidy 
(Raven & Tai, 1979). Through the efforts of Greg- 
ory & Klein (1960), Kurabayashi et al. (1962), 
Raven & Tai (1979), and Peng (1984, 1988), the 
chromosome numbers of all species of Ludwigia 
sect. Microcarpium have been established. There 
are four diploid species with n = 8 (L. linearis, 
L. linifolia, L. microcarpa, and L. stricta), eight 
tetraploid taxa with n — 16 (L. glandulosa subsp. 
glandulosa, L. glandulosa subsp. brachycarpa, 
L. lanceolata, L. pilosa, L. polycarpa, L. ravenii, 
L. sphaerocarpa, and L. suffruticosa), two hexa- 
ploid members with n — 24 (L. alata and L. simp- 
sonii), and an octoploid with n — 32 (L. curtissii). 
For a review and discussions see Peng (1988). 


EcoLoGY AND GEOGRAPHICAL DISTRIBUTION 


With the exceptions of Ludwigia stricta, which 
is endemic to Cuba, and L. polycarpa, which is 
distributed mainly in the northcentral United States, 
Ludwigia sect. Microcarpium is confined primar- 
ily to the Coastal Plain of the southeastern United 
States. Several species also extend further south 
or east: L. alata to Jamaica; L. curtissii to the 
Bahamas; and L. linifolia disjunct to Tabasco, 
Mexico. Ludwigia simpsonii occurs in Cuba and 
Jamaica, and L. microcarpa in the Bahamas, Cuba, 
and Jamaica. Ludwigia stricta is exceptional in 
being endemic to Cuba. A detailed description of 
the ranges with maps for each species is given in 
the taxonomic treatment. 

Like Ludwigia species occurring in other parts 
of the world, plants of Ludwigia sect. Microcar- 
pium grow in wet habitats. They are commonly 
found along alluvial ground or in the shallow water 
of many areas, including ponds, lakes, 
streams, lagoons, sloughs, backwaters, swales, wet 


rivers, 


meadows or prairies, open swamp forests, drain- 
ages, and irrigation ditches. All species grow in 
sandy or occasionally peaty soils. A few species 
grow in brackish marsh or tidal flats (L. alata, L. 
linifolia, L. microcarpa, and L. simpsonii), in 
sinks in limestone prairies (L. alata, L. curtissii, 
L. microcarpa, L. simpsonii, L. sphaerocarpa, 
and L. suffruticosa), and in clay soils (both sub- 
species of L. glandulosa). None of the species in 


Ludwigia sect. Microcarpium exhibit the aggres- 
siveness characteristic of weedy plants, although 
the following taxa have rarely been observed grow- 
ing in disturbed habitats: L. linearis, L. micro- 
carpa, L. simpsonii, and both subspecies of L. 
glandulosa. 


EXPERIMENTAL HYBRIDIZATION 


Results from the extensive artificial hybridiza- 
tion among members of Ludwigia sect. Micro- 
carpium (Peng, 1988) are summarized as follows. 


l. There is generally little difficulty in obtaining 
hybrids between members of sect. Microcar- 
pium; most F, plants grow to maturity, and 
many set abundant seeds. Hybrids between 
plants of the same ploidy level are generally 
intermediate in morphology; hybrids between 
plants of different ploidy levels tend to be more 
similar to the parent with the higher chromo- 
some number. Character such as capsule shape 
and size, bracteole length, overall pubescence, 
presence or absence of petals, seed surface pat- 
tern, and pollen stainability are very useful in 
distinguishing species and identifying hybrids. 

. All diploid species can be crossed with one 
another and form floriferous F, plants. Hybrids 


N 


between the petaliferous diploids with narrow 
leaves (Ludwigia linearis and L. linifolia) are 
vigorous, have ca. 50% stainable pollen, and 
produce a moderate amount of viable seeds. A 
modal meiotic configuration of a ring or chain 
of four plus six bivalents are frequently ob- 
served in reciprocal F, hybrids. Hybrids be- 
tween either of the petaliferous diploids and the 
apetalous, diploid L. microcarpa are, in con- 
trast, highly sterile as to pollen and seed set. 
Nearly all chromosomes remain unpaired in 
meiosis in these hybrids. 

. All eight tetraploid members studied are inter- 


o2 


fertile: chromosome pairing in meiosis is nearly 
complete, and pollen stainability and seed set 
are high in hybrids between any two tetraploid 
species, no matter how different they are from 
each other morphologically. 
4. Although the two hexaploid (n — 24) species, 
Ludwigia alata and L. simpsonii, can 
crossed and produce agb 3 18> meiosis is 
irregular with ak 
The affinity of L. alata is apparentlk with species 
of the tetraploid group, based on morphological 
characters. Hybrids between L. alata and any 
tetraploid species are easily obtained. They pro- 
uce moderate seed set and have 43-71% 
stainable pollen. Meiosis in these F, individuals 


234 


Annals of the 
Missouri Botanical Garden 


typically exhibited a maximum of 16 bivalents 
and eight unpaired chromosomes. By contrast, 
the hexaploid Ludwigia simpsonii (n — 24) is 
clearly allied to the octoploid L. curtissii (n — 
32). Hybrids between them are highly fertile, 
having 80-90% stainable pollen and producing 
good seed set. These plants consistently exhib- 
ited a meiotic configuration of 24 bivalents and 
eight unpaired chromosomes. 

Seeds obtained from crossing species of any two 
of the following groups, (1) tetraploid species 
group (including the allied Ludwigia alata), (2) 
diploid species, and (3) L. curtissii and L. simp- 
sonii, often failed to germinate. Those that ger- 
minate may either be inviable or produce flow- 
ers that set negligible or no seed and tend to 


D 


have very low levels of stainable pollen. For a 
complete discussion of the experimental hy- 


brids, consult Peng (1988). 


NATURAL HYBRIDIZATION 


Experimental hybridization (Peng, 1988) re- 
vealed that vigorous and floriferous hybrids can 
readily be obtained between most members of Lud- 
wigia sect. Microcarpium. The general lack of 
postzygotic barriers and the facts that all species 
in this section are perennial and that most of them 
are sympatric, have similar preference for habitats, 
flower in the morning during the same season, are 
facultatively outcrossing, and have similar kinds of 
indiscriminate insect pollinators, strongly suggest 
that natural hybrids should be fairly common in 
the field. 

Field observations and herbarium study show 
that hybrid populations of Ludwigia sect. Micro- 
carpium are indeed abundant. Except for this and 
the companion paper (Peng, 1988), however, very 
little has been reported about the natural hybrid 
populations of sect. Microcarpium. The variation 
patterns generated by the intergradation among 
some members of sect. Microcarpium have pre- 
viously been detected by Torrey & Gray (1840) 
and Small (1903); such plants were given names 
at varietal and specific rank respectively, but no 
indication of their hybrid origins was mentioned. 
Torrey & Gray (1840) described L. capitata B 
pubens from a population of L. suffruticosa X L. 
pilosa from Georgia; Small (1903) applied the name 
L. simulata to a population of L. pilosa x L. 
lanceolata from North Carolina. 

Duke (1955) was the first to point out that some 
Ludwigia species show varying degrees and pat- 
terns of intergradation in North Carolina. In his 
informative study, he compared the morphological 


characters of a mixed population consisting of plants 
of L. pilosa, L. glandulosa, and the intermediates, 
and considered the latter as possible hybrids. Upon 
examination of his voucher collections, it is con- 
firmed beyond doubt that these plants represent 
uniform F, hybrids of L. pilosa x L. glandulosa. 
Duke (1955) further suggested that “‘self-pollina- 
tion is the rule in this genus, and that cross-polli- 
nation is a very rare exception to the rule," and 
speculated that phyllophagous insect caterpillars 
may have brought about cross-pollination while for- 
aging leaves. This, however, is not true, as at least 
four species in sect. Microcarpium have somewhat 
showy sepals and produce copious nectar and re- 
quire insect vectors to pollinate their flowers to set 
seeds. Such insects as bumblebees, wasps, honey- 
bees, moths, and even ants have frequently been 
seen obtaining nectar on the flowers of Ludwigia 
sect. Microcarpium. 

In a biosystematic study of Ludwigia sect. Dan- 
tia, a close ally of sect. Microcarpium with op- 
posite leaves and prostrate habit, Schmidt (1967) 
reported two intersectional hybrids, L. repens X 
L. simpsonii (as L. repens X L. curtissii) and L. 
palustris X L. glandulosa from North Carolina 
and Florida, respectively. His observations of the 
hybrid between L. palustris (n = 8; sect. Dantia) 
and L. glandulosa (n = 16; sect. Microcarpium) 
revealed very little association between chromo- 
somes, with 0-3 weakly joined bivalents in meiosis. 
The other natural hybrid, involving L. repens (n 
— 24; sect. Dantia) and L. simpsonii (n — 24; 
sect. Mesa. yielded 48 unpaired chro- 
mosomes at mesiosis, suggesting the dissimilarity 
in the genomes of plants in the two North American 
sections. 

Observation of individuals exhibiting a combi- 
nation of characters intermediate between distinct 
taxa initially suggests natural hybridization. Mem- 
bers of the following species pairs, Ludwigia alata 
(n — 24) and L. lanceolata (n = 16), L. pilosa 
(n = 16) and L. ravenii (n = 16), and L. simpsonii 
(n = 24) and L. curtissii (n = 32), however, are 
usually so similar in appearance, especially in dried 
herbarium specimens, that their hybrids cannot be 
recognized readily. 

esults from experimental hybridization suggest 
that capsule shape and size are the most important 
diagnostic characters for detecting hybrids in sect. 
Microcarpium. Other useful features include over- 
all pubescence, seed surface cell shape and ori- 
entation, the presence or absence of petals, whether 
pollen grains are shed singly vs. in tetrads, and 
pollen stainability. Seed surface pattern is useful 
only for the determination of hybrids in the tet- 


Volume 76, Number 1 
1989 


Peng 235 
Ludwigia sect. Microcarpium 


raploid group (including Ludwigia alata), as these 
nine taxa are interfertile, yield abundant seeds, and 
are diverse with respect to this character. Absence 
of developing fruits and low levels of pollen stain- 
ability, however, are characteristic of hybrids re- 
sulting from all other heteroploid crosses, with the 
exception of L. curtissii (n = 32) x L. simpsonii 

— 24). As a last resort, chromosome number 
and meiotic chromosome behavior are useful in- 
dicators of natural hybrids when the parents in- 
volved have different chromosome numbers or dif- 
ferentiated genomes. 

The natural hybrid populations mentioned below 
are discussed dnd poetas the treatment 
of the species in t tion. Full citation 
of these specimens of hissed is given in the com- 
panion paper (Peng, 1988). 

ield experience indicates that natural hybrid 


populations are fairly common wherever two or 
more species grow side by side. A total of 15 
intrasectional hybrids and seven intersectional 
(among plants of sects. Microcarpium and Dantia) 
hybrids are recognized (Peng, 1988). Hybrids re- 
sulting from crosses among members of the tet- 
raploid species group (including Ludwigia alata) 
are by far the most common. The following hybrid 
combinations are representative: L. alata X L. 
pilosa (four populations from Florida, one from 
Georgia, and one from Mississippi), L. alata x L. 
suffruticosa (seven populations from Florida), L. 
glandulosa subsp. glandulosa X L. glandulosa 
subsp. brachycarpa (one from Oklahoma and one 
from Texas), L. glandulosa subsp. glandulosa x 
L. pilosa (two from North Carolina, one from Geor- 
gia, one from Alabama, and two from Mississippi), 
L. glandulosa subsp. glandulosa X L. sphaero- 
carpa (one from South Carolina, one from Mis- 
souri), L. lanceolata X L. pilosa (three populations 
from Florida), L. lanceolata x L. suffruticosa (two 
from Florida), L. pilosa x L. sphaerocarpa (many 
populations from North Carolina, South Carolina, 
Georgia, Florida, Alabama, and Texas), L. pilosa 
X L. suffruticosa (two from North Carolina, three 
from Georgia, and two from Florida), and L. poly- 
carpa X L. sphaerocarpa (two populations from 
Indiana). Backcrossed generations or segregates of 
advanced generations are found of L. lanceolata 
X L. suffruticosa, L. pilosa X L. suffruticosa, 
and most abundantly of L. pilosa X L. sphaero- 
carpa. 

As in the case of artificial hybridization, the 
tetraploid hybrids are highly fertile as to seed and 
pollen; hybrids between the hexaploid Ludwigia 
alata and the tetraploid species have slightly lower 
levels of stainable pollen and seed set. It is note- 


worthy that in nearly all of the above hybrids at 
least one member of p pur of putative parents 
is unable to und If-pollination 

has somewhat qmm sepals. In other words, L. 
alata, L. pilosa, L. sphaerocarpa, and L. suffru- 
ticosa hybridize more frequently than other tet- 


raploid species that are modally autogamous. 
Sporadic occurrence of sterile triploid hybrids 
16) x L. lin- 
earis (n — 8) (one population from North Carolina 
and the other from Georgia) and L. sphaerocarpa 
(n = 16) x L. linearis (one population from Al- 
abama). One probably diploid hybrid population of 
L. linearis X L. linifolia was collected from Flor- 
ida in 1981. Attempts to re-collect and to study 
live plants of this hybrid have failed, because the 
habitat was destroyed (Popenoe, pers. comm.). 
Hybrids between Ludwigia curtissii (n = 32) 
and L. simpsonii (n — 24) very probably occur in 
the field but would be difficult to detect. Ludwigia 
curtissii hybridizes with L. linifolia (n = 8) and 


include Ludwigia glandulosa (n — 


with L. microcarpa (n — 8) to form sterile hybrids. 
At least four populations of L. simpsonii X L. 
microcarpa are known from Florida. It is of in- 
terest to note that among the putative parental 
species here concerned, only L. linifolia has some- 
what showy petals and produces copious nectar, 
whereas all other species are probably modally 
autogamous. 

Hybrids involving putative parents from sect. 
Microcarpium and sect. Dantia are also not un- 
common. Ludwigia repens (n — 24; sect. Dantia) 
hybridizes with L. simpsonii (n — 24) and L. cur- 
tissii (n = 32) in Florida. At least eight such 
populations are known, all producing flowers, but 
the ovaries abort after mechanical self-pollination. 
Ludwigia palustris (n = 8; sect. Dantia), a wide- 
spread and apparently autogamous species, hy- 
bridizes with L. glandulosa subsp. glandulosa (n 
— 16) and forms sterile hybrid populations in Vir- 
ginia (one population). It also hybridizes with the 
diploid, small-flowered and presumably autogamous 
L. microcarpa of sect. Microcarpium and forms 
sterile hybrids in North Carolina (one population), 
Georgia (one population), and Florida (three pop- 
ulations). In Kentucky, L. palustris hybridizes with 
L. polycarpa (n — 16) and likewise forms sterile 
hybrids. 

The only intersectional tetraploid hybrid is known 
from a single population of Ludwigia arcuata (n 
— 16; sect. Dantia) X L. pilosa (n — 16) collected 
from Alabama. Flowers of both putative parents 
are showy and attract many insect pollinators. Es- 
tablished hybrids, however, are very rare in nature. 
Although some seeds were observed in mature cap- 


236 Annals of the 
Missouri Botanical Garden 
TABLE 3. Comparison of diploid petaliferous species in Ludwigia sect. Microcarpium. 
Characters L. linearis L. linifolia L. stricta 


Plant height (cm) 


Cauline leaves 


(22-)50-100(-145) 
Very narrowly elliptic 
to sublinear 


12-55(-62 


Oblanceolate-linear to 


10-60(- 


5) 
Oblanceolate or elliptic to 


narrowly so 


Obscure Obscure 
2 3-)4-7 (1.8-)2-3 
1-1.7 0.9-1.5 

a 2 
0.65-1.1 0.5-0.7 
Lackin Lacking 
z te )1.5-2.5(-3) 0.75-1.1 
-2.5 0.55-0.9 


Abaxial marginal veins Distinct 
Sepal 
Length (mm) 2.3-5(-5.6) 
Width (mm) 1-3(-3.5) 
Length : width 1.5-2.5 
Anther length (mm) -2 
Septa in anther sac resent 
Filament length (mm) 1.1-2(-2.2) 
Style length (mm) (0.4-)0.7-1.5 
Stigma 
Shape Cylindric 
Length (0.6-)1-1.9 
Nectary disc Shallow 
Capsule 
Shape Elongate-obpyramidal 
Surface With a shallow 
longitudinal groove 
on each side 
Length (mm) 5-10(-12 
Thickness (mm) D 
Dehiscence Ring-dehiscent from top 


Bracteole length (mm) 0.4-4(- 7.5) 
Seed 
olor Yellowish 
Surface cell shape Columnar 


Depressed globose 
0.3-0.6 


Very prominent 


Subcylindrical 
Nearly smooth 


5-10(-12) 
1.3-2(-2.2) 
Peeling off from the sides 


(1.5-)2.5-9(-13) 


Reddish brown 


Subisodiametric 


Depressed globose 
0.25-0.5 
Prominent 


Subcylindrical 

Nearly smooth 

5-6(-7) 

1.6-1.8 

Peeling off from the sides 


0.7-2.5(-3) 


Reddish brown 


Subisodiametric 


sules of the herbarium specimen (Mobile Co.: Au- 
dubon Bird Sanctuary, Dauphin Island, Deramus 
0643 [DS, UNA], they are probably inviable. 
Such hybrids synthesized in the experimental 
greenhouse showed a remarkably high level of 
chromosome pairing (ca. 12-15 bivalents and 2- 
8 unpaired univalents; 1 or 2 trivalents occasionally 
seen) but had only 38% stainable pollen. Not a 
single seed was set despite many attempts at ar- 
tificial self-pollination (like L. pilosa, this hybrid 
cannot undergo mechanical self-pollination). 

o summarize, geographical isolation and self- 
pollination are the primary factors limiting natural 
sect. Mi- 
crocarpium. For example, L. polycarpa occurs in 
the central Midwest (Fig. 45), where it is isolated 
geographically from all other species of sect. Mi- 
crocarpium except L. sphaerocarpa. Where the 


hybridization among species of Ludwigia 


two species come into contact, hybridization oc- 
curs. Ludwigia microcarpa, an extreme selfer, 
has been hybridized successfully with most species 


of sect. Microcarpium in the greenhouse, and the 
resulting hybrids were vigorous. Ludwigia micro- 
carpa grows sympatrically with many other species, 
but natural hybrids have been found only with £. 
simpsonii, L. curtissii, and some members of sect. 
Dantia. 

In general, natural hybrids are usually frequent 
wherever two species occur together. This is par- 
ticularly true for plants in the tetraploid group, 
hybrids of which were often found intermixed with 

utative parents. The sorts of disturbed habitats 
suggested by Anderson (1948) do not appear to 
be necessary for the establishment of natural hy- 
rids in Ludwigia sect. Microcarpium 

As outlined above, natural hybridization is not 
limited to species within Ludwigia sect. Micro- 
carpium. At least seven hybrid combinations, some 
of which occur commonly in nature, have been 
found between members of sect. Microcarpium 
and sect. Dantia. Most of these have also been 
synthesized in the experimental greenhouse. None 


Volume 76, Number 1 
1989 


Peng 237 
Ludwigia sect. Microcarpium 


of the intersectional hybrids set viable seeds, al- 
though they are usually very vigorous and appear 
to compete well with their parents. Once estab- 
lished, these sterile hybrids may be able to persist 
in a given location due to their perennial habit. 
Furthermore, new colonies may be established veg- 
etatively if entire plants or fragments are trans- 
ported to suitable habitats, most likely by water. 


EVOLUTIONARY RELATIONSHIPS 


Ludwigia sect. Microcarpium consists of four 
diploids, eight tetraploids, two hexaploids, and one 
octoploid. 

Ludwigia microcarpa is distinct among the di 
loid species by the more or less spatulate cauline 
leaves and small, inconspicuous, apetalous flowers; 
and it is modally self-pollinated. All three other 
diploid species, L. linearis, L. linifolius, and 
stricta, have elongate, narrow leaves and relatively 
showy, petaliferous flowers and are facultatively 
outcrossing. The divergence of L. microcarpa from 
the rest of the diploids is further demonstrated by 
the cytogenetic study of artificial hybrids in this 
group. Artificial F, hybrids between L. microcarpa 
and either L. linearis or L. linifolia were com 
pletely sterile; in meiosis they revealed very few 
chromosome associations, with 0-3 weakly joined 
bivalents. 

Ludwigia linearis is unusual in sharing with the 
unrelated South American species L. latifolia 
(Benth.) Hara (sect. Tectiflora) the character of 
polysporangiate anthers in which the locules are 
divided by transverse septa composed of tapetum 
and parenchyma. Otherwise, L. linearis is appar- 
ently somewhat related to L. linifolia. The two 
species differ chromosomally by at least one recip- 
rocal translocation. F, hybrids between them have 
ca. 50% stainable pollen, and they produce mod- 
erate quantities of viable seeds. 

Ludwigia stricta, which is endemic to Cuba, is 
more closely related to L. linifolia than to any 
other species in sect. Microcarpium. It differs from 
L. linifolia only by the broader leaves, smaller 
floral parts, and shorter fruits (Table 3). The cross- 
ing relationships of L. stricta with other diploid 
species are currently being investigated. 

The eight tetraploid taxa (n — 16) are uniformly 
apetalous and are otherwise remarkably diverse in 
overall pubescence, shape and color of sepals, shape 
and dehiscence of capsules, bracteole length, seed 
surface cell pattern, and whether pollen is shed 
singly or as tetrads. Correlations in these features, 
however, are evidently nonexistent. It is therefore 
of great interest to find that these eight morpho- 
logically well-delimited taxa can be crossed in any 


combination and produce vigorous and fertile off- 
spring. The hybrids exhibit nearly complete chro- 
mosome pairing and high levels of stainable pollen. 
Natural hybridization among plants of this group 
is abundant, especially among those members with 
somewhat showy sepals. Most natural hybrids are 
F, plants that occur in more or less undisturbed 
habitats where the putative parents also grow. In- 
trogressed populations or segregates of advanced 
generation of Ludwigia pilosa X L. sphaerocarpa 
hybrids, however, are also abundant and wide- 
spread. Cytogenetic studies of experimental hybrids 
between the tetraploid species and the diploid species 
suggest that no extant diploids are believed to have 
been involved piani in the formation of the tet- 
e species grou 
udwigia alata, a piba (n = 24) species, 
fet the tetraploids morphologically. Super- 
ficially it is most similar to L. lanceolata in general 
aspect and especially in the characters of winged 
capsules. Nearly all of the reciprocal hybrids be- 
tween the hexaploid L. alata and any of the tet- 
raploid species exhibited a modal meiotic config- 
uration of 16 bivalents and 8 univalents, which 
suggests that they share two genomes in common. 
Judged from the morphology of the hexaploid and 
from the pairing of chromosomes in experimental 
hybrids, L. alata probably originated following hy- 
bridization between one of the tetraploids, perhaps 
L. lanceolata, and the diploid L. microcarpa, or 
populations ancestral to them. 
The affinity of the other hexaploid, Ludwigia 
simpsonii, however, is definitely with L. curtissit, 
the only octoploid in sect. Microc | wid These 


two species are unique in having 
and in having, along with the diploid L. microcar- 

a, more or less spatulate cauline leaves. Ludwigia 
curtissii and L. simpsonii are polymorphic and 
are sometimes difficult to distinguish morphologi- 
cally. Cytogenetic data suggest that the hexaploid 
chromosome complement of L. simpsonii includes 
three different genomes, one of which is identical 
with that of L. microcarpa. Artificial hybrids be- 
tween L. simpsonii and L. curtissii exhibited a 
meiotic configuration of 24 bivalents and eight 
univalents, which suggests that these two closely 
related species possess three genomes in common. 
Based on morphological characters and crossing 
relationships, the octoploid L. curtissii was prob- 
ably derived following hybridization between a pe- 
taliferous, narrow-leaved diploid lineage similar to 
L. linifolia and the hexaploid L. simpsonil. 

o summarize, following the differentiation of 
diploid species of Ludwigia sect. Microcarpium, 
some diploids have evidently become extinct. Nat- 
ural hybridization among the diploid lineages fol- 


238 


Annals of the 
Missouri Botanical Garden 


lowed by polyploidy has played a major role in the 
evolution of this group. Postzygotic genetic barriers 
do not separate most of the species of this section. 
Rather, the major limiting factor to natural hy- 
bridization appears to be the modally autogamous 
breeding system of most species. Geographical iso- 
lation is important only with respect to L. poly- 
carpa, which is distributed well to the north of 
nearly all the other taxa. 

Natural hybridization is prevalent within Lud- 
wigia sect. Microcarpium, and hybrids often grow 
intermixed with putative parental populations. All 
interploid hybrids are completely sterile except for 
alata (n — 24) and the tetra- 
ploid species (n = 16), and crosses between L. 
curtissii (n = 32) and L. simpsonii (n = 24). Even 
sterile hybrids, however, can persist and form large 
colonies, at least locally, by means of strong veg- 
etative reproduction via stolons. 

Natural hybrids are especially common among 
members of the tetraploid group (including Lud- 
wigia alata) and are nearly always vigorous and 
fertile. Especially complex is the pattern of vari- 
ation in the tetraploid L. sphaerocarpa, which is 
apparently comprised largely of widespread stabi- 
lized hybrid populations that exhibit a combination 
of characters distinguishing them from other taxa. 
As in the evolution of Epilobium in New Zealand 
(Raven & Raven, 1976), the recombination of 
genetic information from somewhat differentiated 


crosses between L. 


populations followed by the maintenance of well 
adapted genetic strains by a combination of autog- 
amy and vegetative reproduction appears to have 
played a central role in the evolution of the poly- 
ploid members of Ludwigia sect. Microcarpium. 


SYSTEMATIC TREATMENT 


Ludwigia sect. Microcarpium Munz, Bull. Tor- 
rey Bot. Club 71: 154. 1944; N. Am. Fl. ser. 
II, pt. 5: 42. 1965. Raven, Reinwardtia 6: 


KEY TO THE SPECIES OF LUDWIGIA SECT. MICROCARPIUM 


la. Pu uie elongate, at least twice as long as broad. 
Petals absent; leaves narrowly elliptic 


336. 1963. “Microcarpeae” Small, Man. S.E. 
Fl. 941. 1933. Type species: Ludwigia pi- 
losa Walter. 


Erect perennial herbs, perennating mostly by 
leafy, basal stolons (mainly by rhizomes in Lud- 
wigia suffruticosa). Stems usually profusely 
branched except in L. suffruticosa, becoming swol- 
len and spongy below when submerged. Leaves 
alternate, sessile or nearly so, subentire, usually 
with hydathodal glands along margins. Flowers ses- 
sile or shortly pedicellate, axillary (in very compact, 
short racemes or spikes in L. suffruticosa). Brac- 
teoles in subopposite pairs attached from the base 
of the pedicel to halfway up the ovary. Flowers 
4-merous, mostly apetalous (consistently with 4 
petals in 3 diploid species). Sepals mostly greenish 
(creamy white or yellowish in 4 apetalous species). 
Stamens haplostemonous; anther sacs undivided 
except in L. linearis, in which the polysporangiate 
anthers are partitioned horizontally by tapetums 
and parenchyma. Pollen shed singly or as tetrads. 
Nectary disc 4-lobed, usually shallowly raised above 
ovary apex (flat in L. microcarpa). Capsules with 
ca. 10-700 seeds, obpyramidal to subcylindric, 
oblong-obovoid, turbinate, or subglobose, dehiscent 
by the separation of the disc and the ovary wall, 
or forming 4 longitudinally lenticular slits opposite 
each locule, or the irregular rupture of the outer 
wall. Seeds yellowish, light tan to reddish brown, 
free and pluriseriate in each locule, with a narrow 
raphe; surface cells suborbicular, elongate parallel 
to or perpendicular to the seed length. Self-com- 
patible, modally autogamous in most species. Ga- 
metic chromosome number, n = 8, 16, 24, 32. 


Distribution. Atlantic and Gulf Coastal Plain 
of the United States (Ludwigia polycarpa is ex- 
ceptional by ranging into the northcentral United 
States), Bahamas, Cuba, Jamaica, and Tabasco, 

exico. 


4. L. glandulosa 


= Petals 4; leaves sublinear to linear. 
L 


a. Lateral and marginal veins on abaxial leaf surface distinct; ia elongate-obpyramidal, ring- 
on top; seeds yellowish; anthers 1-2 m 3. L. 
3b. on and marginal veins on abaxial leaf surface aa. da subcylindric, peeled-dehiscent 
from the sid vien cia 


dehiscent 


es; seeds re 


m long, the s 


o 


inearis 


0.5-1.1 mm long, the sacs not septate 
E mm long; 


L. linifolia 


rie Sepals (3.3-)4-7 mm qe es as long as wide; filaments (1.3-)1.5 
style 1.25-2.5 mm pem paeem. " pr )2.5-13 mm long; U.S. & Tabasco, CHE 
4b. Sepals 1.8-3 mm long, 2 times as Jong as wide; wes 0.75-1.1 mm long; ek 0.55- 


).9 mm long; bracteoles 0.7-2. is 
lb. Capsules various, be than twice as long as 
5a. Flowers in compact 


overtopping the main stem; rhizomes often prese 


mm long; Cu 


racemes or spikes 3 5 cm longi stems single or with a few deis ues 


. L. stricta 


suffruticosa 


Volume 76, Number 1 
1989 


Peng 239 
Ludwigia sect. Microcarpium 


5b. dde) axillary, in cae interrupted, leafy racemes or spikes 2-20 cm long; stems usually profusely 


nched; rhizomes a 
de Plants densely TO throughou 


7a. Sepal apex elongate-acuminate or PA reflexed; plants densely hirtellous; seed surface 
1 


cells suborbicular 


L. pilosa 


7b. Sepal hs acuminate, ascending; plants densely strigillose or hirtellous; seed surface cells 


colum 


8a. Pli hirtellous; capsules — ras sepals greenish adaxially; bracteoles 9- Ag - 


ng, at or near capsu 


.3 mm | 
8b. Plants Boe | inso Elsie sepals yellowish adaxially; bracteoles 0. E A 5m 
dicel 9. L 


long, usual short pedic 


. Plants subglabrous to Mid 


an 
[o 


L. ravenii 


. sphaerocarpa 


9a. Capsules obpyramidal un narrowly winged co 
10a. Stems nearly smooth or slightly ridged; aniey D about 4 as long as the capsule; 
KI ce wall flat between the wings; seed surface cells suborbicular; pollen grains shed 
ES 


L. lanceolata 


s tetrads 
10b. Sel Mis distinctly ridged or winged; sepals creamy whitish, nearly as long as capsule; 


capsule etween the wings bulging out ear is seed surface cells elongate 
parallel y The seed length; pollen grains shed singly 8. L. alata 
9b. Capsules various, not w inged. 


lla. Nectary disc Pm flat on ovary apex; fruits 1-1.5 mm na md 10-20 seeds, 
12. L 


often recog nizable as bumps from outside; seeds reddish bro 


O seeds, not reco 


. microcarpa 
m long, containing 40- 


y apex; fruits 1 


12a. Cauline leaves lanceolate, oblong-elliptic to T so; capsules dehiscing by 
nt. 


.5- 
nate, reflexed; capsules oblong-obovoid with a constricted 


er walls; stolons pres 
mm long; sepals me n, the apex cH acumi- 
DM S ss 


polycarpa 


a L. 
13b. Bracteoles 0.5-1.5 mm long; ME yellowish adaxially, the apex acuminate, 
n Y. L 


12b. Ca 


TE 


bianchi to very narrowly so; 


ioi yd 4 longitudinally dus slits on the wall opposite the loculi; puis 
not com 


vi Capsules 2.5-4.7 mm long; sepals 1.5-3 mm long; vestigial m occ 


onally present; lower or seedling leaves always alternate 
mm long; sepals 1.2-1.8 m 
lower or seedling leaves tending to be opposite or subopposi 


14b. eccl 1.5-2.5 m 


L. curb 
mm long; dan Lam rare; 


As in Figures 1-15 and 19-34, the sequence 
of the taxa in the taxonomic treatment reflects the 
possible phylogeny of Ludwigia sect. Microcar- 
pium. Diploid petaliferous species that are presum- 
ably more primitively outcrossing are arranged first, 
followed by tetraploid members and the related 
hexaploid L. alata. Ludwigia microcarpa, an 
apetalous diploid, which definitely played a role in 
the formation of the distinct L. curtissii complex, 
is placed last along with the complex. 


= 


. Ludwigia linifolia Poiret in Lam., Encycl. 
Suppl. 5: 513. 1817. /snardia pr (Poi. 
ret) Kuntze, Rev. Gen. i 891. 
LECTOTYPE: "Amerique," Poiret ES 
(P; lectotype here designated). Figure 35. 


Plants glabrous. Stems erect or ascending, usu- 
ally well branched, 12-55(-62) cm tall, aeren- 
chyma rarely seen. Stolons slender, 4-15(-30) cm 
long, 0.7-1(-1.5) mm thick, seen in flowering sea- 
son, the leaves narrowly obovate to narrowly ob- 


13. L. simpsonii 


lanceolate, sometimes spatulate, 4.5-20 mm long, 

m wide, petioles 0.5-5 mm long. Cauline 
leaves linear or linear-oblanceolate, 15-40 mm 
long, (0.65-)0.9-4(-6) mm wide, the apex taper- 
ing into a sharp point or acute, margin entire with 
obscure hydathodal glands, the base very narrowly 
cuneate, sessile. Stipules narrowly ovate to nar- 
rowly lanceolate, 0.2-0.3 mm long, 0.1-0.2 mm 
wide. Flowers many, in leaf axils, their subtending 
leaves not reduced. Sepals green, narrowly trian- 
gular, ascending, (3.3-)4- 7 mm long, 1.1-1.7 mm 
wide, glabrous or minutely papillose, the apex nar- 
rowly acute, the margin entire. Petals yellow, nar- 
rowly obovate-elliptic, 4-6 mm long, 2-4 mm wide, 
the apex obtuse or rounded, the base obtuse. An- 
thers 0.65-1.1 mm long; filaments yellowish, 
(1.3-)1.5-2.5(-3) mm long. Pollen shed as tet- 
rads. Nectary disc bright yellow, elevated 0.3-0.7 
mm on top of the ovary, 0.8-1.5 mm across, 
prominently 4-lobed, minutely papillose. Style yel- 
low, 1.25-2.5 mm long, glabrous; stigma yellowish, 
0.3-0.6 mm long, 0.6-0.8 mm thick, the apex 


240 


Annals of the 
Missouri Botanical Garden 


FIGURE 35. 
Lower stem with basal stolons; stems continue at arrows. — b. F 
e. Cross section of capsule. 


shallowly 4-lobed. Capsules subcylindrical, slightly 
narrowed toward base, 5-10(-12) cm long, 1.3- 

2(-2.2) mm thick, densely minutely papillose, oc- 
casionally also remotely minutely scaberulous, the 
hairs 0.05-0.15 mm long, sessile. Bracteoles at- 


e 
Ludwigia linifolia. Florida, Santa Rosa Co. 2 Bes did 65936 (LL). —a. Habit, erect stem.—a’. 


— c. Mature capsule. —d. Dehiscent capsule. — 


tached 0-1.5 mm above capsule base, very nar- 
rowly oblanceolate to linear, (1.5-)2.5-9(-13) mm 
long, 0.15-0.8 mm wide, the margin entire, as- 
cending or spreading. Seeds reddish, oblong-elliptic 
with curved ends, 0.55-0.6 mm long, 0.2-0.3 


Volume 76, Number 1 Peng 241 
1989 Ludwigia sect. Microcarpium 
FIGURE 36. Distribution of Ludwigia linifolia in the United States. 


mm thick, the surface cells nearly isodiametric. 
Self-compatible. Gametic chromosome number, n 
=8 


Distribution. Plants of L. linifolia are often 
found in drainage ditches, along margins of creeks 
or swamps, on bottom meadow, on open edges of 
cypress swamps, in moist pineland, and along edges 
of brackish lakes. Ludwigia linifolia ranges along 
the Atlantic coast from extreme southeastern North 
Carolina and eastern South Carolina through south- 
ern Georgia and peninsular Florida. To the west, 
its range extends through the Florida panhandle, 
to extreme southern Alabama and Mississippi. A 
disjunct population has been collected in north- 
central North Carolina (Fig. 36). It is also known 
from a disjunct population at Huimanguillo, Ta- 
basco, Mexico, on the Yucatán PORE. Flow- 
ering from late June through October; fruiting from 
July through November. 


"Mui arc specimens examined. U.S.A. 
BAMA: Baldwin Co., 3 mi. E of Southport, Webster * 
Wilbur 3559 (FSU, GH, SMU, US). Covington Co., ca. 
13.5 mi. S Opp. on US 331, Kral 32508 (DS, GA, MO, 
SMU, VDB). Mobile Co., W jct. ALA 188 and county 
59, Kral 37358 (FLAS, FSU, VDB). FLORIDA: Bay Co., 
ca. 3.4 mi. inland from Long Beach, Godfrey & Hen- 
de 'rson 62969 (FLAS, FSU, TEX). Bradford Co., near 
US 301, Radford 8253 (NCU). Brevard Co., Okeechobee 
region, Fredholm 5937 (GH, US). Calhoun Co., 

mi. NW of Blountstown, Thorne & Davidson aot 
(FSU). Charlotte Co., Caloosa Experimental Range, U.S. 
Forest & Range Stas Adams 198 (FSU). Collier Co., 
S of Bonita Springs, Atwater M-78 (FLAS). DeSoto Co., 

Lacy, West 171 (FLAS). Dixie Co., 4 mi. N of dal 
Godfrey 56177 (FSU, GH, NY, USF). Duval Co., near 
Jacksonville, Curtiss 922 (FI, F LAS, GA, GH, M, MO, 

, RB, S, . Escambia Co., ca. 3 mi. N 
Kral 17580 (VDB). Flagler Lo. i. 

odfrey & Reinert 61115 (FSU, SMU). Franklin Co., 
38.8 mi. W of jct. of US Hwys. 98 & 319, Peng et al 
4343 (MO). Gulf Co., 3 mi. S of Port St. Joe, Kral & 
Kral 7193 (FLAS, CH, USF, VDB). Hernando Co., Sec- 
tions 11 & 12, T21S, R17E, Cooley et al. 7049 (USF). 


242 


Annals of the 
Missouri Botanical Garden 


Hillsborough Co., 7 mi. NE of USF —— Lakela 
24117 (FLAS, GH, USF). Jackson Co., . W of 
Chattahoochee, Thorne 5956 (GH). lake Dd Sus of 
Eustis, Nash 1240 (F, GH, MO, MSU, ND, NY, 
Lee Co., Myers, Hitchcock 119 (E, F, GH, MO, NY, 


un 
T 


of Orangedale, along FL m Godfrey 70630b (FSU. MO, 
NCU). St. Lucie Co., alton Beach, E of US 
Kral 18366 (VDB). use Rosa Co., nea 


r Yellow River, 


Schallert 8145 (S). Wakulla Co., near Spring Creek, 
a ee (DUKE, a LL, NCU, SMU, USF, 
VDB). Walton Co., ca. 5 m Seagrove Beach along 
US pa Kral "19360 (VDB). "Washington Co., 3.5 mi. E 
of di Godfrey 5962 (DUKE, FLAS, CH, NCU 


a: Baker Co., SW corner of county, Thorne 


ooly Co., 3 mi. S-SE 
i. S Pinehurst, Kral 48130 (VDB). 
"8 er Claxton, Godfrey 72098 (FSU, 
1.3 mi. gie Duncan 
17021 (FLAS, NCU, SMU). Long Co., mi. SE of 
Ludowici on GA Rt. 99, Boole 1140 (B, is SMU). 
McIntosh Co. of graveyard on Wesley Lake Rd., 
Bozeman 2227 (FLAS, GA, NCU). Sumter Co., wet pine 
barrens, eal 1109 (E, GH, MO, NY, US). Wa re Co., 
Wayer ages, s.c. 4176a (US). MISSISSIPPI: Hancock 
o., . 90, 2.5 mi. W of jct. with State Hwy. 43, 
Raven 18581 (DS). Harrison Co., Gulfport, Lloyd & 
Tracy 225 (NY). Jackson Co., Hwy. 90, between Orange 
Grove and Pecan, Jones 14898 (FSU, GH, NY, VDB). 
NORTH CAROLINA: Brunswick Co., on Co. Rt. 1335, 2 mi. 
NW of jet. of Co. Rt. 1340, Leonard & Radford 2205 
(C, CM, E, FLAS, MISS, NO, NY, TENN, TEX, s 
UNA, USCH, VDB, WCUH). New Hanover Co., 
Fisher on the Lower Cape Fear Peninsula, Godfrey 6195 
(GH). SOUTH CAROLINA: Bamberg Co., on US 301, 0.3 
with SC 64, Bozeman et al. 114004 (NCU). 
Georgetown Co., 1,750 ft. due ENE of intersection of 
US 17 & Belle Isle Gardens Road, Rayner 1081(USCH). 
Hampton Co., 0.2 mi. N of Luray on US Rt. 321, Ahles 
& Bell 18288 (FSU, NCU, USF). Marion Co., just E of 
secondary road 49; 8-10 mi. N of intersection with Great 
Pee Dee River, Rayner 1109 (USCH). Orangeburg Co., 
2.7 mi. N of Branchville City on (just o l 
Rayner 1106 (USCH). Mexico, TABASCO: Huimanguillo, 
Cowan 2632 (MO), 2277 (NY), 3111 (MO). 


— 


» 


Ludwigia linifolia is a relatively uniform species. 
Its variability chiefly affects the size of the capsules 
and the lengths of the bracteoles, which are at- 
tached to the sides of the capsule. Its closest affinity 
is definitely with L. stricta, a Cuban endemic, from 
which L. linifolia differs by having narrower leaves, 
larger floral parts (including sepals, anthers, fila- 


ments, and style), and longer fruits and bracteoles. 
Characters of the two species and the only other 
petaliferous member of the section, L. linearis, are 
listed for comparison in Table 

Ludwigia linifolia has somewhat showy petals 
and the most prominent nectary disc lobes occur- 
ring in sect. Microcarpium. The nectary disc is 
bright yellow and produces copious nectar at an- 
thesis. Herbarium specimens often reveal empt 
anthers held away from the stigma (mechanical 
self-pollination has not occurred yet). This obser- 
vation suggests that L. linifolia is probably modally 
outcrossing, which is corroborated by limited field 
observations. 

Ludwigia linifolia occurs with L. alata, L. cur- 
tissii, L. glandulosa, L. linearis, L. microcarpa, 
and L. pilosa in the field. Natural hybrid popu- 
lations have, however, been found only between L. 
linifolia (n = 8) and L. i 


is contrary to what one would expect for the out- 


curtissii (n = 32). This 


crossing L. linifolia. However, experimental hy- 
bridizations (Peng, 1988) indicate that artificial 
hybrids between the diploid L. linifolia and plants 
of any other ploidy level in sect. Microcarpium 
are generally difficult to obtain. Repeated trials with 
different parental strains were successful in some 
cases, but the resulting hybrids were usually weak 
and needed special attention to reach flowering 
stage. Attempts to synthesize the natural hybrid L. 
linifolia X L. curtissii in the experimental green- 
house were unsuccessful. 


2. Ludwigia stricta (Wright ex Griseb.) Wright 
in Wright & Sauvalle, Fl. Cubana 54. 1873. 
Based on /snardia stricta Wright ex Griseb., 
Cat. Pl. Cub. 107. 1866. TYPE: eastern Cuba, 

Lo Wright 2555 (holotype, 

GOET; isotypes, C, GH, MO, NY, P, US). 

Figure 


Plants glabrous throughout. Stems erect or as- 
cending from stolons, simple or much branched, 
10-60(-85) cm tall. Stolons 5-10 cm long, l- 
2.5 mm thick, the leaves elliptic, 7.5-13 mm long, 
3-5 mm wide, glabrous, the base attenuate into 
winged petioles up to 2 mm long. Cauline leaves 
deciduous from below in late growing season, ob- 
lanceolate to narrowly oblanceolate or narrowly 
elliptic, 11-22 mm long, mm wide, gla- 
brous, the apex acute, the margin with remote, 
minute hydathodal glands, the base narrowly cu- 
neate into winged petiole 0-1.5 mm long. Stipules 
minute, ovate or lanceolate, succulent, dark brown- 
ish purple, 0.2-0.3 mm long, 0.1-0.2 mm wide. 


Flowers in upper leaf axils, their subtending leaves 


Volume 76, Number 1 243 
1989 


Peng 
Ludwigia sect. Microcarpium 


bi Ver. qae Nem e Do WANLING PENG 
DI. RES AN SS 
ON) == 
1 N a 
| 77 NEST 
7 Q = 


FIGURE 37. Ludwigia stricta. a’, c, d, d', and f from Cuba (Wright 2555, US); a, b, and e from plants grown 
from seeds from Cuba (1982, Leiva s.n., M 3304, MO).—a, a’. Habit, stem continues at arrows.—b. Flower.— c. 
Capsule.—d, d'. Adaxial and abaxial sides of leaf.—e. Petal.—f. Cross section of fruit. 


244 


Annals of the 
Missouri Botanical Garden 


not reduced. Sepals green, narrowly ovate-trian- 
gular, ascending, (1.8—)2—3 mm long, 0.9-1.5 mm 
wide, the apex narrowly acute or acuminate, the 
margin entire. Petals yellow, elliptic or narrowly 
obovate, 2.3-2.9 mm long, 1.2-1.3 mm wide, the 
apex obtuse, the base attenuate. Anthers 0.5-0.7 
mm long; filaments 0.75-1.1 mm long. Pollen grains 
shed as tetrads. Nectary disc yellow, elevated 0.25- 
0.3 mm on top of the ovary, 1-1.25 mm across, 
4-lobed, minutely papillose. Style 0.55-0.9 mm 
long, glabrous; stigma subglobose, 0.25-0.5 mm 
across. Capsules cylindrical, 5-6(-7) mm long, 
1.6-1.8 mm thick, minutely papillose; pedicels 0— 
0.9 mm long. Bracteoles attached at capsule base, 
narrowly lanceolate, 0.7-2.5(-3) mm long, 0.2- 
0.4(-0.6) mm wide, the margin entire. Seeds red- 
dish brown when mature, elliptic-oblong in outline, 
slightly curved on both ends, 0.5-0.6 mm long, 
0.2-0.25 mm thick, the surface cells nearly iso- 
diametric. Self-compatible. Gametic chromosome 
number, n = 8 


Distribution. Ludwigia stricta is a uniform 
species known only from Cuban lagoons and sa- 
vannas (Fig. 38). Flowering and fruiting from Au- 


gust through November. 


Representative specimens examined. CUBA. PINAR 
DEL RÍO: Herradura, Britton et al. 6613 (NY); Pinar del 
Río, Britton et al. 7233 (NY); 3 km S of Herradura, 
Ekman 17742 (S). ISLA DE JUVENTUD (ISLA DE PINOS): 
between Los Indios and La Siguanea, Ekman 12179 (Sy; 

otel Colony, Rostánski 2028 (MO); 1982, Leiva s.n. 
(cultivated, MO). 

Ludwigia stricta has a close affinity with L. 
linifolia, which is distributed in the southeastern 
United States and occurs on the Yucatán Peninsula 
in Tabasco, Mexico. Both are diploids; they are 
similar in stature, have four petals, shed their pollen 
as tight tetrads, have subcylindric capsules that 
dehisce irregularly from the sides, and have reddish 
brown seeds with subisodiametric surface cells. A 
comparison of these two species and the other 
petaliferous diploid species, L. linearis, is pre- 
sented in 

Viable seeds of [a 
only recently. Plants have been grown and crossed 
with all three other diploid species of sect. Micro- 
carpium in order to assess their genetic relation- 
ships. 


stricta have become available 


3. Ludwigia linearis Walter, Fl. Carol. 89. 
1788. Isnardia linearis (Walter) DC., Prodr. 
3: 60. 1828. TYPE: U.S.A. “the Carolinas”: 
T. Walter 665 (holotype, BM, Walter Her- 
barium, p. 66; photograph, MO). Figure 39. 


d+ ilie ian Michaux, Fl. Bor.-Am. 1: 88. 
03. LECTOTYPE: U.S.A. South Carolina: near wet 

renda T Aug.-Sept., 1785-1796, A. Michaux 

s.n. (P; photographs, GH, MO; isolectotypes, MO, 
sheets; lectotype here designated, based on 

Tem collection of authentic material at P 


.S.A. Texas: Harris Co., Hosnton. July 1843, F. 
J. Lindheimer 58 (holoty ype, GH; isotypes, BM, 
GOET, MO, P, PH, US). 


Plants glabrous to densely minutely strigillose 
or sometimes puberulent, the hairs 0.05-0.15 mm 
long. Stems erect, often well branched, (22-)50- 
100(-145) cm tall, often with distinct aerenchyma 
in lower part. Stolons up to 30 cm long, 0.8-0.25 
mm thick, seen in late fruiting season only, the 
leaves purplish, narrowly elliptic to very narrowly 
elliptic, 12-23 mm long, 2.8-8.5 mm wide, gla- 
brous to sparingly minutely strigillose, the base 
attenuate into petioles up to 5 mm long. Cauline 
leaves green, linear to elliptic linear, 16-60(-85) 
mm long, 0.9-3.7(-5.6) mm wide, glabrous to 
densely minutely strigillose or puberulent, the apex 
very narrowly acute, the margin entire, the hy- 
dathodal glands obscure, the base very narrowly 
acute or cuneate, subsessile. Stipules narrowly ovate 
or lanceolate, 0.15-0.3 mm long, 0.05-0.15 mm 
wide. Flowers many, in leaf axils, their flower- 
subtending leaves not reduced. Sepals green, tri- 
angularly ovate to narrowly triangular, the apex 
acuminate or elongate acuminate to cuspidate, as- 
cending, 2.3-5(-5.6) mm long, 1-3(-3.5) mm 
wide, glabrous, densely minutely scaly strigillose, 
strigillose, or puberulent without, glabrous within. 
Petals yellow, obovate to subrotund, 3-6 mm long, 
(2-)2.5-5 mm wide, the apex obtuse, the base 
attenuate. Anthers lanceolate oblong, 1-2 mm long, 
sporogenous tissue 4-8 packeted; filaments 1.1- 
2(-2.2) mm long. Packets in the sporogenous tissue 
opening and shedding tetrad pollen almost simul- 
taneously. Nectary disc yellow, elevated (0.25-)0.35- 
0.55(-0.7) mm on top of the ovary, 1.3-2.5 mm 
across, 4-lobed, glabrous to minutely strigillose or 
strigillose, especially around lobe margins. Style 
yellowish green, (0.4—)0.7-1.5 mm long, glabrous 
throughout to densely strigillose, especially in the 
lower part. Stigma yellowish green, elongated, shal- 
lowly 4-lobed on apex, (0.6-)1—1.9 mm long. Cap- 
sules elongate obpyramidal, often with a shallow, 
central, longitudinal groove on each side, 5-10 
(712) mm long, 2-5.5 mm thick, glabrous, densely 
scaly strigillose, strigillose, or puberulent, subsessile 
or with distinct pedicels up to 3.5(-5) mm long. 
Bracteoles subequal, attached near capsule base to 
4.5 mm above base or on the pedicel, linear, 0.4— 


Volume 76, Number 1 Peng 245 
1989 Ludwigia sect. Microcarpium 
S 
FIGURE 38. Distributions of Ludwigia alata (circles) and L. stricta (triangles). 


4(-7.5) mm long, 0.1-0.3 mm wide, deciduous. 
Seeds light brown, oblong-elliptic with slightly curved 
ends, 0.45-0.65 mm long, 0.15-0.25 mm wide, 
the surface cells oblong, elongate either parallel or 
transversely to the seed length. Self-compatible. 
Gametic chromosome number, n = 8 
Distribution. Ludwigia linearis is common 
in drainage ditches, along river or stream banks, 
swales, edges of pocosins, sandy soil in wet mead- 
ows, brackish marshes, and occasionally on dis- 


turbed ground. It is distributed along the Atlantic 
coast from southern New Jersey through Delaware, 
the eastern halves of North and South Carolina, 
southern Georgia, to its southern limit in central 
Florida. Westwards, this species extends through 
the panhandle of Florida and across Alabama, 
southern Mississippi, and Louisiana, to the central 
Gulf coast of Texas. To the north, L. linearis 
ranges through eastern Texas and across ron E 
Arkansas, and there is a northward 
from northern Alabama into Tennessee (Fi ig. 40). 


246 Annals of the 
Missouri Botanical Garden 


Y ZA 


SS 


NS NS d 


ES 
» 


JM 
E 


As 
E] 
y 


ni 
4 


AN Pe 
| fes | 
NF AN 


A 7 


FIGURE 39. Ludwigia linearis. All but a’ from South Carolina (Jasper Co., Boufford et al. 21633, MO); a' from 
de (Nansemond Co., Fernald et al. 15322, GH).—a. Habit, erect stem. —a'. Lower stem with basal stolons. — 
aves.—c. Flower bud.—d. Flower.— e. Petal.— f, f'. Adaxial and abaxial views of stamen.—g, g'. Immature 

uds mature capsules. —h. Cross section of capsule. 


Volume 76, Number 1 
1989 


Peng 247 
Ludwigia sect. Microcarpium 


FIGURE 40. 


glabrous morph; 4 = intermediate morp 


Incorrectly reported from Japan (see discussion). 
Flowering from late June through September; fruit- 
ing from late July through mid December. 


Representative specimens examined. (l: subgla- 
brous morph; 2: Pr io. morph; : glabrous 

morph; 4: intermediate m ) 
U.S.A. ALABAMA: Baldwin v m 5 mi. N of yt Sho. 
Kral 38264 (4, VDB); 13.5 mi. W of Foley on US Hwy 
98, Raven 18590 (1, DS). Barbour Co., ca. 5 mi. 
Eufaula by Lake Eufaula, Kral 33241 (4, SMU, a 
Butler Co., ca. 11 mi. N of Georgiana, by US Hwy. 65, 
Kral 40891 (1, VDB; mixed with 2). Cherokee Co., by 


Covington Co., ca. 6 mi. N of Florala on US Hwy. 331, 
Davenport 788 (1, UNA). Cullman Co., 1897, Eggert 
s.n. (1, MO). Dale Co., by paee her just N 
of Newton by AL Hwy. 123, Kral 54450 (1, VDB). 
Dallas Co., MoAr s.n. (2, 7 à Kalb d. near i 
Ruth 415 (1, NY, US). Elmore Co., 5 m 

tumpka, by AL Hwy. 9, a 136595 (1, VDB). Beaia 


Distribution of Ludwigia linearis. O = subglabrous morph; 6 = densely strigillose morph; A = 


Co., 4.6 mi. E of Boykin, just W of Conecuh River by 


.H of 
Clark 7342 (1, NCU). eue Co., N of Flat Rock, Kral 
36531 (1, FSU, VDB). Lee Co., near Auburn, 1897, 
Earle & Baker s.n. (1, F, MO 
near Society Hill, 1857, s.c. (1, NY). Mobile Co., 2 mi. 


Z 
® 
le] 
£ 
e 
La 
O 
o 
5 


Dera (4, NCU, ). Pick Co., S side of 
Aliceville, Kral 51383 (4, B Russell Co., 1.3 mi. 

Wo enix City, by US . 280, Kral 44243 (1, 
VDB). Sh us a mi. 5 2 AL Hwy 25 & Calera 
turnoff by U . 65, Kral 33584 (1, SMU, VDB). 

ashington Co. uas i. S of St. Stevens, Kral 37216 
(1, FLAS, VDB). ARKANSAS: Ashley Co., Lone Pine Prairie, 


, MO, NO, NY SMU, TENN, TEX). 

O mi. S of Hampton, Kral 24536 (2, 

VDB). Dallas Co., near Fordyce, Demaree 67870 (1, 
a o. 


S ar 10 
63154 (3, MO). Lonoke Co., near Carlisle, Demaree 


248 


Annals of the 
Missouri Botanical Garden 


37980 (2, SMU, VDB; 3, SMU). Prairie Co., near Hazen, 
Demaree 37904 (3, SMU, VDB); along Hwy. 70, E of 
i i . Union 
Co., near El Dorado, Hoiberg 591 (2, SMU). White Co., 
1.2 mi. S of McCrae, by US Hwys. 67, 167, & 64, Thomas 

& Smith 15187 (2, DS, SMU, TENN). DELAWARE: Kent 
Co., near Hamington, 1872, Canby s.n. ussex 
Co., 3 mi. S of Laurel, Hotchkiss 7919 (1, US). FLORIDA: 
Alachua Co. E of se sal 1940, Murrill s.n. (1, US). 
cClenny, Curtiss 6007 (1, DS, FLAS, 
Y, SMU, US). Bay Co., near Lynn 

Haven, Godfrey 59120 (1, FSU, SMU). Bradford Co. 
near Sampson City, Dennison & Arnold 171 (1, FLAS): 
W of Lawtey, Raven 18694 (4, DS). Calhoun Co., 5 mi. 
W of Blountstown, Godfrey & Kral 55035 (1, FSU, GA, 
GH, NY, S, SMU, TENN, UNA, USF). Citrus Co., near 
Chassahowitzka, Genelle & Fleming 1562 (1, GA, TENN, 
Co., near Doctors Inlet, 1939, Murrill s.n. 


. with FL Hwy. 17 on ride 
, FLAS). Escambia on 
ground e Center, Burkhalter x be FLAS). Flag. 
ler Co., i. E of Bunnell, 1937, t et al. s.n. (1, 
FLAS). Franklin Co., 41.7 mi. W of je US hwys. 98 
& 319, Peng et al. 4351 (1, MO). Gulf Co., Do 
Weewahitchka & Port St. Joe, Godfrey & Kral 54167 
(1, DUKE, FSU, GH, NY). Hamilton Co., 11 mi. E of 
Suwannee River by FL Hwy. 6, Godfrey 75386 (1, FSU). 
Hillsborough Co., 1.5-1.7 mi. S of Hwy. 674, on E side 
of Taylor Grill Dr., Peng 4325 (1, MO). Holmes Co., 
near Westville, s.c. 4175c (1, JE; 4, GH, US). Jackson 
Co., N of Sneads, Lake Seminole, Godfrey & Houk 61398 
(4, TEX); S of Marianna, along Hwy. 71, Hood 2642 
(1, TEX). Jefferson Co., ca. 15 mi. SSW of Monticello, 
Kral 5825 (1, FSU, GH, , USF). Lake Co., near 
Eustis, Nash 1490 (4, NY). Loos Co., N of Woodville, 
Perdue & Godfrey 1858 (1, C, GH, LL, NCU, SMU, 
TEX, US, USF). Liberty Co., 1 mi. E of Telogia, Morar 
8 (1, DUKE, FSU, MSU). Madison Co., 8 mi. W of 
Greenville, Kral 3742 (1, FSU, GH, NCU, SMU). Man- 
atee Co., NE of jet. hwys. 62 € 39, Shuey 1710 (1, 
USF). Okaloosa Co., E of Oak Grove by FL Hwy. 2, 
Yellow River, Godfrey 75428 (1, FSU). Pinellas Co., 
Brooker Creek, Poppleton 698 (1, USF). St. Johns Co., 
1941, West & Arnold s.n. (1, 

of Maxton, Yellow River, 

Godfrey & Houk 62565 i ree: 
WSW of Ma 


Vols Co., S of Seville. 1943, in 
FLAS). Wakulla Co., near St. Mar s Re fuge, Morar 17 
(1, FSU, NCU, SMU, US, VDB). reel Co., at Mossy 
Head, Godfrey 55244 (1, FLAS, FSU, GA, GH, N 


mi. 
(1, FSU, USF); S of Chipley along Hwy. za Hood 2836 
(4, FLAS). GEORGIA: Bartow Co., ca. 4.5 mi. SE of Adairs- 
ville on Cassville enden Greear 64 3491, GA, NCU). 
Berrien Co., ca. 3 mi. Alapaha, reg 23488 (1, 
GA, VDB). Charlton Co., es ng GA 4 between 
Moniac & St. George, Jones et al. 23045 "ü. i Chat- 
ham Co., along Dutchtown Road, 1959, Mellinger s.n. 
(1, GH); "pon N of Richmond Hill, Kral 18947 (4, VDB). 


Coffee Co., near Douglas, 1900, Harper s.n. (l, 
US). Coo , 7.6 mi. SE of Ade l, Jim Reeves p 
Faircloth 2928 (1, GA, NCU). Decatur Co., 1.5 mi. 


of Climax, Godfrey 75563 (4, FSU). Dooly Cm 0.75 mi. 


eorgia Pacific Plant, S of Vienna by US H l, 
Godfrey 76005 (4, FSU); 2.7 mi. S-SE of Unadilla, Har 
per et p 16817 S GA). Early Co., 2 mi. S o ilton, 


Crab Co., 3.5 mi. NW of Calvary, Faircloth & Faircloth 
1509(1, MO, NCU). Irwin Co., 7.3 a SW of Irwinville, 
at Tift-Irwin Co. line, Faireloih 5410 (1, GA, NCU). 
Lanier Co., 1 mi. E of Mimi land, vie 3605 (1, GA, 
NCU). Lee Co., 1.5 mi. SW of Smithville, Pyron & 
McVaugh 1948 TR GA). prm Co., Camp Stewart near 
Hinesville, 1943, Guimm s.n. (1, CM). Long Co., 4.2 mi. 


261, Bozeman & Radford 
NCU). Lowndes Co., 13 mi. E of Valdosta in 
Grand d Fairc loth 2080 (1, CA, NCU). McIntosh Co., 


of Meansville, Harper 2244 
ichmond Co., 1903, € s.n. 
,E e Lu Eyles 7637 (1, 

Co. i. E of Donaldsonville, 
Godfrey & Clewell 5021014, > FSU). Thomas Co., near 
Thomasville, 1903, Taylor s.n. (1, TEX). Tift Co. 42.5 
mi. W of Alapaha River jct. with Tift & Berrien counties, 
Plummer & Pullen 897 (1, GA 


Co., near Waycross, Eggleston 5078 (1, Ahi 
Co., near Jesup, Curtiss 921 (1, CM 
MO, ND, NY, US). LOUISIANA: Acasa Par 


ing, in 24488 (3, 
i. N of Oil City, Thieret 24467 (3, DS); along 
LA Hwy. 1, S of jet. LA Hwy. e ds oma. Js 
Overby 36294 (2, NCU). Calcasieu Parish, r 
W of Sulphur, near Stegall, Thieret 20580 (2, DS, DUKE, 
FSU, GA, LL, SMU). Evangeline Parish, 6.2 mi. SE of 
jct. of LA hwys. 106 € 107, Sullivan 1633 (2, USF). 
Grant Parish, 4.5 mi. N of Dry Prong, Shinners 24945 
(4, SMU); 1.5 mi. E m. US Hes 165 opposite LA Hwy. 
we Barrett Quail F , Tomas et al. 2995 (2, NCU, 
B). nation: Parish, mi. S of Montpelier, 
mos 24850 (2, DS). isis is Parish, ca. 4 mi. E 
of Bellwood, Kral 16192 (2, VDB). Ouachita Parish. 2 
mi. E of Dh rn Thomas et al. 30860 (2, FLAS, GH, 
SMU, TENN). ire Parish, 8 mi. N of M rede 
Kral o (2, VDB). St. Tammany Parish, 2.5 m 
of Tali iii Raven 22104 (1, DS, NCU); Hwy. 1.59 at 
P exit, Darwin & Sundell 578 (2, NO); 3.7 


.2 mi. E of Arcola, Tangipahoa 
River, Thieret 27823 (2, DUKE. FSU, GA); Ponchatoula, 
Cocks s.n. (4, NO). Union Parish, 4 mi. NE of Haile, 
Thieret 23691 (2, DS, SMU). Vernon Parish, 2 mi. E of 
aCamp, Kral 15805 (2, VDB). Washington Parish, 
Franklinton, dign 46898 (1, DS, SMU, VDB). W 
ster Parish, ca. 2 mi. W of Springhill, Allen 8474 y^ 


Volume 76, Number 1 
1989 


Peng 249 
Ludwigia sect. Microcarpium 


Vincent 1798 (3, FLAS, VDB). Winn Parish, 3 mi. N 
of Winnfield by LA Hwy. 1231-2 at jct. with US Hwy 
165, Thomas & Cicala 31931 (4, NCU, NY). MARYLAND: 
Wicomico Co., 6 mi. E of Salisbury, 1872, Canby s.n. 
(1, NY). Ms Co., near Ocean City n Carter 
s.n. (1, NY). Mississippi: Clarke Co. 2 nee o 
Pachuta, n 10750 (4, MISS). MM Co. „a ca. 3.5 
mi. N of Stone Co. line, W of Hwy. 49, Rogers 4664 
(1, MO, NCU); near Ragland Hills, Rogers 3807 (4, 


Demaree 36 311 (1, GH, SMU); Waveland & Hwy. 90, 
Jones & Jones 9435 (2, MISS, NCU); on MS Hwy. 43, 
ca. 5 mi. NE of MS 


SMU, TEX, VDB); on MS Hwy. 57, 2.8 mi. N 
Hwy. 90, Peng et al. 4355 (4, MO). Lamar Co., 5 mi. 
SW of Hattiesburg, Ray 5957 (MISS; 1, GA, GH, NCU, 
NY, USF; 4, NY). Lauderdale Co., S of Lauderdale, Jones 
14990 (1, GA, MISS); E of Chunky, US Hwy. 80 Fa 
of Chunky River, Kral 22036 (2, VDB). Marion Co., S 
of Hwy. 48/35 jct., ca. 3 mi. N of MS-LA border, 
Musselman 1238 (2, NCU). Monroe Co., near B 
1916, Bailey s.n. (4, MISS). Pearl River Co., 5.8 mi. S 
of Carriere, N of Picayune, Jones & Hudson 9748 ü. 
MISS); 1 mi. E of Picayune, Sargent 9879 (2, MO); 8 
mi. NW of Picayune, Sargent 8376 (4, CAS). Perry Co., 
5 mi. E of Runnelstown, Jones 9587 (1, MISS). Simpson 

., 5.8 mi. S of Puckett and N of Mendenhall, MS Hwy. 
13. gom 19257 (2, GA, MISS). Smith Co., near Tay- 
lorville, Tracy 8715 (2, F, GH, MO, a pom NY, 
TAES, US). Walthall CH along MS Hwy. 21, just over 
LA Bander, Kral 19393 (1, VDB). NEW JERSEY: Atlantic 


Martindale s.n. (1, F, NY, US). Camden Co., near Atco, 
1872, Commons s.n. (1, NY). NORTH C AROLINA: Bator 
Co., along NC Hwy. 33, 3 mi 
Duke 54- $280. NCU). Bertie Co. 
on NC Hwy. 17, Blomquist 16226 (1, DUKE). Bladen 
Co., 5.2 n hd of Dublin, Duke 5499 (1, NCU). Bruns- 
wick Co., 0.2 mi. W of Longwood on NC Hwy. 904, 
Peng et ^ du MO). Carteret Co., ca. 14 mi. NE 
of Beaufort on Rt. 70, Wilbur 9336 (1, DUKE). Chatham 
Co., of Wilsonville, on Co. Rd. 1008, Campbell 
1077 (1, NCU). Columbus Co., 2.1 mi. NW of Old Dock 
n NC Hwy. 130, Leonard & Radford 2192 (1, CM, 
FLAS, FSU. GA, NCU, NY, SMU, USF, VDB). Craven 
Co., 0.8 mi. N of US Hwy. 17 on Co. Rd. 1224, Peng 
et al. 3737 (1, MO); jet. of US Hwy. 70 & Street Ferry 
Rd., Duke 5468 (4, NCU). Cumberland Co., 8 mi. N of 
Fayetteville, on Bos 15A, Duke 54152 (1, NCU). Dare 
Co., along Hwy. 64 N of Manteo, Duke 54-236 (1, 
NCU). Duplin Co., 2.5 mi. SSE of Rose Hill on US Hwy. 
117, Ahles 35773 (1, NCU). Edgecombe Co., between 
er & Enfield, Godfrey & Fox 49642 (1, NCSC). 
Gates Co., : mi. NE of Chowan he js Hwy. 158, Beal 
2121 (1, NCSC). Greene Co., ESE of Jason, 
Radford 40379 (1, vae Prid Co.. near Weldon, 
1900, Williamson s.n. (1, NY). Erw Co, near Dunn, 
Godfrey 6149 (1, DUKE, GH, US). Hoke Co., 4 mi. 
SSW of Ashley Heights, Ahles 36379 a, cu Hyde 
Co., 8.5 mi. SW of Fairfield, Duke 54-233 (1, NCU); 


— 


mi. NW of Sladesville, Radford 39039 (4, e 


Iredell Co., near States ville, s.d., Hyanis 


W of Deep Run, Radford 31671 (1, NCU). Martin Co., 
2 mi. W of Hamilton on NC Hwy. 125, Duke 54182 (1, 
NCU). Nash Co., 0.8 mi. S of Taylors Cross Roads and 
1.3 mi. E of paved road, Ahles & Bell 16714 (1, CM, 
NCU). New Hanover Co., 7 mi. N of Sea Breeze, 2 
16055 (1, NCU). Northhampton Co., 4 mi. 

Jackson, des s Mill i Beal 2758 (1, NCSC). Ges. 
Co., 5.5 mi. S of Hwy. 41, on Co. Rd. 1003, S E 
Comfort, Peng et al. 37 90 (1, MO). Pamlico Co., 4 r 

W of Grantsboro, Duke 54-225 (1, NCU). Pender Co. 


o 


CA, NCU). Pitt Co., 3 mi. E of Grim 
Hwy. 264, Fox 3182 0, FSU, GH, MO, NC 
mond Co., 3.5 mi. of Hoffman at Camp Mackall 
Radford 14440 (1, at NCU). Robeson Co., 3 mi. 

of Lumberton, on N ; Hwy. 72, Duke 54157 (1, "NEU. 


Co., 1.5 mi. NW of Columbia, Radford 42484 (1, NCU). 
Washington Co., 4.2 mi. ESE of Skinnersville, Radford 
1, NCU). Wayne Co., Neuse State Park, Gillespie 


10) 

NCU). Beaufort Co., 
(1, NY); 2 mi. SSE WEN 
on Co. Rd. 43, Ahles & Bell 18018 (4, NCU). Berkeley 

o., 10 mi. NW of Bonneau, Godfrey & Tryon 1628 
(1, GH, NY); 0.7 mi. E of Interstate 26 and US Hwy. 
17-A, Peng 4390 (4, MO). Calhoun Co., 0.5 mi. NE of 
jet. of Co. rds. 26 & 27 on 26, Ahles 35400 (4, NCU). 
Charleston Co., 4.8 mi. E of SC Hwy. 174 on Hwy. I. 
E of Osborn, Peng et al. 3901 (1, MO); 24.5 mi. N of 
SC Hwy. 41, on US Hwy. 17, Peng et al. 3903 (4, MO). 
Chesterfield Co., Montrose near xb 52, Radford 15762 
(1, TENN). Clarendon Co., . NE of Turbeville, 
Radford 28107 (4, NCU). CE Co., 1.2 mi. W of 
Dorchester Co. line, on SC Hwy. 61, Raven 20462 (4, 
DS); 1.2 mi. S of Orangeburg- Colleton Co. line by E 
Hwy. 21, Bell 4542 (1, C, NCU). Darlington Co., nea 
Boggy Gully Bay, Sawyer 2569 (1, ban Dillon a ʻi 
1.7 mi. NE of Bingham on SC Hwy : 
(1, NCU). Dorchester Co., 0.3 mi. $ of 


Florence Co., NW 
(4, rh aede c po: Ait rene Radford 28679 
(4, NCU). Hampton i. NNW of Shirley on Co. 
Rd. 20, pow: bs 'NCU. ier Co., Myrtle 
Beach, Godfrey & Tryon 1161 (1, DUKE, F, GH, MO, 
NY, TENN, US); along Waccamaw River at the crossing 
of SC Hwy. 9, Raven 18721 (4, DS, NCU). Jasper Co., 
0.2 mi. N of Co. Rd. S 27-101 on US ph 17, just $ 
of Ridgeland, Peng et al. 3935 (1, MO). Lee Co., 3 m 

of Woodrow near SC Hwy. 441, Radford 29422 a, 
NCU). Lexington Co. i. dm ki Be 
Creek, Radford 29890 (1, NCU). ras Co. 4, CA. 


250 


Annals of the 
Missouri Botanical Garden 


mi. S of Britton Neck on Co. Rd. 49, Bell 11005 (1, 
NCU); 5 mi. S of Britton's Neck, Kral 19177 (4, VDB). 
Marlboro Co., 3.5 mi. E of Wallace, Radford 19085 " 
NCU). Newberry Co., Billy Dreher Island, Buff 80 ( 
USCH). Orangeburg Co., near Eutawville, Eggleston ate 3 
(1, GH, NY); od 1900, s.c., s.n. (4, US). Saluda 
Co., 1 mi. ard, Hwy. 23, Mass ey & Masse ey 3005 
(1, 'NCU); p mi. WSW of Ward, Radford 30358 (4, 
NCU). Sumter Co., 4 mi. S of Scottsville, near US Hw MA 
le Radford 29575 (1, NCU). wed Co., 2.3 
of ae aera near US Hwy. 521, Radford 
28280 (1, GH, NY, VDB). TENNESSEE: "Co Co., near 
Tullahoma, s.c. 4175 (1, GH, NY, US). Cumberland Co., 
NE of Crossville, Finger Lake, Norris & Sharp 16183 


(1, ). avi son Co., near Nashville, Cumberland 


s. 41A & 4420, Highland Rim 
Forest Experiment Station, = Qu et al. 50923 (1, 
Hang ay rren Co. of Morrison, by TENN 
Hwy. 5& power us Kral 53781 (1, VDB). TEXAS: 
recibas Co., ce. Barkley 13576 (2, F, MO, NY, 
Macs US). Angelina .9 mi. N of Bouton Miis 
mpgrounds, Fg 4003 (2, LL). Aransas Co., 
sas Wildlife Refuge, Fleetwood 9344 (2, 
3 


on TX Hwy. 6, Mase 311 (2, NCU, ai Cham 
Co., hare Tharp s.n. (2, X). Galveston Co., E side 
of Hwy. Dickinson. Waller 3030 (2, GH, isa 


Gregg T es , York s.n. (2, TEX). Hardin Co., nea 
Fletcher, Palmar 10673 (2, DS, MO); NW of Saratoga, 
Kral 21084 (4, VDB). Harris Co., near Sheldon, Rever 


(2, LL). Jasper Co., 1.5 mi. N of Kirbyville, dl il 26762 
(2, GH, LL). Dita Co., W edge of China, between 
Rt. 90 & R.R., Correll 36478 (2, LL). Johnson oi near 
Buchanan, 1898, Eggert s.n. (2, MO 

Manchester on Farm Rd. 195, Correll & Correll . 35909 
(4, LL). Lavaca Co., Borcher Ranch, 18 mi. SE of 


W of Wadsworth, Sharik $73. 1585 (4, MO). 
mi. W of Cleveland, TN 21039 
(2, VDB); Conroe, 2s Tharp s.n. (3, TEX). Morris 
i Naples, Correll 37471 (2, LL). 
Nacogdoches Co., 77 aller 258 (2, DS, TAES, TEX). 
Newton Co., along Rt. 87 N of jct. with Rt. 253, rs 
iioi MO, 1 TEX) 1939, Tharp s.n. (4, M 
Orange Co., mi. E of Bridge City, Correll ree 
(2, LL). Red River Co., N of Clarksville, Lundell 14011 
(3, NY, TEX). Rusk Co., 1879, s.c., s.n. (2, F, beh 
Travis Co., near Austin, 1929, Whitehouse s.n. (2, C 
Tyler Co as 1.5 m Mb li dus 69, Ani 
34015 LL). Upshu r Co., S of New Diana, 
e. 18977 (2, SMU). us Co; near Hempstead, 
Hall 221 (2, N 


x~ 


SE of Guinea, Fernald & Long 9108 (1, 
12 mi. E of McK 

Henrico Co., 
: à Suffolk, Kral 9508 (1, 
FSU, NCU). Norfolk Co., 0.5 mi. W of Bowers un 
Hubricht B2567 (1, MO). Prince George Co. 
SE of Petersburg, at head of Po , 

6652 (1, GH, NY). Princess Anne Co., near Cape Henry, 


dunes & Long 4961 (1, GH, NY, US). icing 
, SW of Corinth, Whitefield's Millpond, co. 
Ha 15320 (1, GH, NY, US). Sussex Co., 3 mi. W of 
Stony Creek, Howell 20491 (1, NCU), Warwick n 
near Newport News, 1907, Long & Long s.n. (1, US). 


The variation pattern of Ludwigia linearis is 
highly complicated as a result of a great deal of 
intergradation among slightly differentiated geo- 
graphical races. I have classified the varied pop- 
ulations into four categories: 1) the subglabrous 
morph, 2) the densely strigillose morph, 3) the 
completely glabrous morph, and 4) the interme- 
diate morph, which is discussed below. 


1. Subglabrous morph. Stems sparingly mi- 
nutely strigillose. Cauline leaves glabrous except 
for the minutely strigillose margin. Sepals 2.3-4 
(-4.5) mm long, 1.3-2.4(-3) mm wide, the apex 
acuminate, minutely scaly strigillose abaxially. An- 
thers 1.1- ong, the filaments 1.3-1.8 
(-2.1) mm long. Nectary disc minutely scaly strig- 
illose around margins of the lobes. Style (0.4—)0.7— 
0.9(-1.2) mm long, glabrous. Stigma (0.6-)1-1.5 
mm long. Capsules densely minutely scaly strigil- 
lose, 5-8.5(-10) mm long, 2-4(-5) mm thick, 
sessile or with a constricted, pedicel-like base. Brac- 
teoles 0.4-1.6(-2.5) mm long. Seed surface cells 


columnar, elongate parallel to the seed length. 


The type of Ludwigia linearis Walter belongs 
to this morph. Plants of this sort are distributed 
along the Atlantic coast from southern New Jersey 
through Delaware, eastern Maryland, eastern Vir- 
ginia, eastern halves of North Carolina and South 
Carolina, and Georgia, to their southern limit in 
the central Florida peninsula. They extend west- 
ward through the panhandle of Florida and across 
Alabama and southern Mississippi, reaching east- 
ern Louisiana (Washington and St. Tammany par- 
ishes only). Northward, they occur throughout the 
eastern half of Alabama and extend to central 
Tennessee. 


Stems and 
leaves densely minutely strigillose to puberulent in 
some cases. Sepals 3-5 mm long, 1.5-2.5(-3.5) 
mm wide, the apex cuspidate or elongate-acumi- 
nate, minutely strigillose or puberulent abaxially. 
Anthers (1.1-)1.3-2 mm long, the filaments 
(1.2-)1.5-2(-2.2) mm long. Nectary disc minutely 
scaly strigillose throughout or only surrounding 
margins of the lobes. Style (0.6-)0.9—1.5 mm long, 
glabrous to densely strigillose, especially on the 
lower part. Stigma (1-)1.3-1.9 mm long. Capsules 
densely minutely strigillose or puberulent, 5-10 
(-12) mm long, 3-5.5 mm thick, the base sessile 


2. Densely strigillose morph. 


Volume 76, Number 1 
1989 


Peng 251 
Ludwigia sect. Microcarpium 


or with distinct pedicels up to 5 mm long. Brac- 
teoles 0. .5) mm long. Seed surface cells 
columnar, pane transversely to the seed length. 


Plants of this morph were recognized at varietal 
rank, Ludwigia linearis var. puberula, by 
gelmann & Gray (1845). They occur most fre- 
quently in central and southern Arkansas, western 
Louisiana, and the eastern half of Texas. Scattered 
populations, however, occur east of the Mississippi 
River in eastern Louisiana and southern Mississip- 
pi. A single population was found in Dallas County 
in southcentral Alabama. 

In addition to being much more pubescent than 
plants of the subglabrous morph, plants of the 
densely strigillose morph are slightly larger in the 
size of most floral parts. Furthermore, their seed 
surface cell pattern is different from that of the 
subglabrous morph. The cells are elongate parallel 
to the seed length in the subglabrous morph but 
are elongate transversely to the seed length in the 
densely strigillose morph. 


3. Completely glabrous morph. Characters 
of this morph are exactly the same as those of the 
densely strigillose morph, except that the former 
is glabrous throughout. Indeed, many plants of the 
completely glabrous morph were collected along 
with those of the densely strigillose morph. Ex- 
amples are as follows: 


U.S.A. ARKANSAS: Prairie Co., low moist areas in rice 
paddies, P. O. Hazen, Demaree 37904 e AL gla- 
brous morph: TEX, VDB-22422, VDB-4 5; mixed 
collection of both the completely glabrous xe and the 
densely strigillose morph: e 3 

t 
“of completely glabrous morph 
and the densely erige morph: CAS). LOUISIANA: Bos- 
sier Parish, 10 mi. S of Plain Dealing, fire lane in young 
pine-hardwood stand, sandy loam, Shinners 24488 (com- 
pletely clips orph: NO; densely strigillose morph: 

SMU). TEXas: Waller Co., ditches, Hempstead, Hall 221 
(completely aa morph: GH, MO; densely strigillose 
orph: NY; mixed collection: F, NY — 2 sheets, US). 


In these cases floral and vegetative characters 
and even the seed surface pattern are exactly the 
same in the completely glabrous and the densely 
strigillose individuals. Plants of these two morphs 

y well differ only by a single allele that controls 
ub. expression of pubescence in plants. It is also 
worth noting that plants of the completely glabrous 
morph are only found scattered in Arkansas, Lou- 
isiana, and Texas, the main distribution range of 
the densely strigillose populations. Plants of the 
completely glabrous morph were previously in- 

d by Munz (1944, 1965) as a part of the 
population of the subglabrous morph (as Ludwigia 


linearis var. linearis), which is certainly not ap- 
propriate, as discussed above. 

imilarly, completely glabrous individuals that 
are we single-allele mutants in populations 
consisting mainly of densely pubescent plants but 
are otherwise similar morphologically occur in Lud- 
wigia hirtella Raf. (sect. Ludwigia; Peng, un- 
published). 


4. Intermediate morph. Any plant that is 
more pubescent than those of the subglabrous morph 
and less pubescent than the densely strigillose morph 
is considered “intermediate” here. The pubescence 
ranges from scattered minutely strigillose on abax- 
ial midvein and/or submarginal veins to sparingly 
strigillose on both surfaces of the leaves. The cap- 
sule wall is minutely strigillose and not scaly strig- 
illose. Floral characters generally fall in the in- 
termediate range of the subglabrous and the densely 
strigillose morphs. The seed surface cells, mostly 
elongate transversely to the seed length, are similar 
to those of the densely strigillose morph. In some 

ses, however, the seed surface does not have 
sell. defined cells and is difficult to study. At other 
times, it is clearly intermediate between that of the 
subglabrous and the densely strigillose morphs. 

Plants of the intermediate morph are widely 
scattered throughout the range of both the subgla- 
brous morph and the densely strigillose /complete- 
ly glabrous morphs. They are found in two counties 
in coastal North Carolina, very well represented in 
the Coastal Plain of South Carolina, scattered in a 
few counties in Georgia, Florida, and Alabama, and 
abundant in Mississippi, where the subglabrous and 
densely strigillose morphs meet. West of the Mis- 
sissippi River, they are also found as scattered 
populations in Arkansas, Louisiana, and Texas, the 
main distribution range of the densely strigillose 
morph. 
Plants designated as intermediate morph" here 
represent an artificial assemblage of plants of Lud- 
wigia linearis that may have originated from var- 
ious combinations of somewhat differentiated pop- 
ulations. For example, the intermediate plants east 
of the Mississippi River often co-occur with subgla- 
brous plants and may represent hybrids and their 
derivatives between plants of the subglabrous and 
of the densely strigillose morphs. On the other 
hand, intermediate plants that grow west of the 
Mississippi River may represent hybrids and their 
derivatives between the completely glabrous morph 
and the densely strigillose morph, as suggested by 
the following mixed collections: 


.S.A. ARKANSAS: Prairie Co., along 70, E of Carlisle, 
Buker in 1956 (mixed collection of one glabrous plant 


252 Annals of the 
Missouri Botanical Garden 
TABLE 4. Comparison of subglabrous and densely strigillose populations in Ludwigia linearis. 
Characters Subglabrous Populations Densely Strigillose Populations 
Leaves Completely glabrous on both sides but Densely minutely strigillose or puberulent 
with ME strigillose margins on both surfaces 
Bracteole length (mm)  0.4-1.6(-2 0.5-4(- 1.5) 
Sepal 
Apex Acuminate Elongate-acuminate or cuspidate 
Length (mm) 2.3- 3-5(-5.6) 
Vestiture Densely minutely scaly-papillose, the Densely minutely strigillose, the hairs 


hairs 0.02- 
Stigma length (mm) (0.6-)1-1.5 
Style Glabrous 
Seed surface cells 
Pedicel length (mm) 


0.05 mm long, appressed 


Elongate parallel to the seed length 
0-0.4 


0.06-1 mm long, appressed to ascending 
(1-)1.3-1.9 
Glabrous to densely minutely strigillose 
Elongate transversely to the seed length 
0-3.5(-5) 


and two intermediate plants). LOUISIANA: Winn Parish, 
NW 


pletely glabrous morph: USF; intermediate morph: NCU). 


Plants of the subglabrous populations are some- 
what distinguishable from the densely strigillose 
plants (Table 4) and are separable geographically 
(the boundary is roughly along the Mississippi Riv- 
er). Engelmann & Gray (1845) formally recognized 
the distinctiveness of the two morphs. However, 
with the abundant intergradation between the two 
races of L. linearis, as already noted by Engelmann 
& Gray (1845), “*... these characters (of L. lin- 
earis var. puberula) pd shading away into 
the ordinary L. linear a as the 
occurrence of the ddr i ious populations 
that have further complicated the character com- 
binations of L. linearis, I think a discussion of the 
variation pattern within this species is more useful 
than recognizing infraspecific taxa. 

Ludwigia linearis was reported as naturalized 
in marshy areas of Japan (Asai, 1970), and this 
report was taken up by Osada (1976) in his illus- 
trations of the naturalized plants of Japan. How- 
ever, an examination of Asai’s voucher specimen 
at TI (Japan: Honshu, Mie Pref., Suzuka-shi, Asa- 
higaoka-machi (Shirako-machi), October 9, 1966, 
H. Ohta s.n.) revealed that it actually represents 
L. perennis L. (sect. Caryophylloidea), which is 
widely distributed in the tropics and subtropics of 
the Old World and ranges north to Taiwan and 
southeastern China (Raven, 1963). 

wigia linearis can readily be recognized by 
its sublinear leaves, four yellow petals, and elon- 
gate-obpyramidal capsules with a shallow, central, 
longitudinal groove on each side. The only other 
species in sect. Microcarpium with which L. lin- 
earis might be confused is L. linifolia, which also 


occurs in the Coastal Plain and the Florida pen- 
insula, but much less commonly. These two species 
are similar in having very narrow leaves, petalifer- 
ous flowers, and elongate capsules. My biosyste- 
matic study (Peng, 1988) has shown that they 
differ by at least a reciprocal translocation. A com- 
parison of morphological characteristics in these 
two related species and L. stricta, which is endemic 
to Cuba, is petaliferous, and has elongate capsules, 
is summarized in Table 

Ludwigia linearis and the unrelated Latin 
American congener L. latifolia are unique in the 
genus in having conspicuous, transverse paren- 
chymatous septa in the anthers, which divide the 
sporogenous tissue into packets (Eyde, 1977; Tobe 
& Raven, 1986). Before the dehiscence of the 
anther, however, all of these packets fuse and form 
one continuous area of sporogenous tissue as a 
result of disintegration of the septa, leaving the 
conspicuous vestiges of the distintegrated septa on 
the inner surface of the anther wall (Tobe & Raven, 
1986). 

Ludwigia linearis is facultatively outcrossing, 
having four stamens held away from the stigma at 
early anthesis and arched over, tightly appressed 
against the unique, elongate stigma a few hours 
after anthesis. 

Ludwigia linearis is usually very floriferous and 
has the highest seed set per capsule in sect. Mi- 
crocarpium, about 600 seeds per mature capsule. 

Ludwigia linearis co-occurs with L. glandu- 
losa, L. lanceolata, L. linifolia, L. microcarpa, 
L. pilosa, L. sphaerocarpa, and L. suffruticosa 
in the field. Confirmed natural hybrids include L. 
linearis X L. glandulosa (two collections) and L. 
linearis X L. sphaerocarpa (one collection). In 
both cases, hybrids are triploid, fertility as to seed 
and pollen is nearly zero. Ovaries turn yellowish 


Volume 76, Number 1 
1989 


Peng 253 
Ludwigia sect. Microcarpium 


or pink after anthesis and then abort. One to four 
petals are often present in most flowers. 


4. Ludwigia glandulosa Walter, Fl. Carol. 88. 
1788. TYPE: U.S.A. “the Carolinas”: s.d., 7. 
Walter 612 (holotype, BM, Walter Herbar- 
ium, p. 66; photograph, MO). 


Plants subglabrous. Stems erect, usually well 
branched, 10-80(-100) cm tall, glabrous to spar- 
ingly minutely strigillose on the ridges formed by 
the decurrent leaf bases, the hairs ca. 0.05-0.1 
mm long. Stolons up to 20 cm long, the leaves 
purplish, narrowly elliptic, 15-35(-55) mm long, 
5-12.5(-20) mm wide, attenuate into petioles 3— 
10 mm long. Cauline leaves narrowly elliptic to 
sublinear, sometimes elliptic, those on the main 
stem 30-120 mm long, 3-25 mm wide, those on 
the branches usually reduced, 8-45 mm long, 2- 
10 mm wide; leaves glabrous to sparingly minutely 
strigillose on abaxial veins, the hairs ca. 0.025- 

.05 mm long, apex acute to very narrowly acute, 
margin entire in outline, densely fringed with mi- 
croscopic, papillose-strigillose hairs, hydathodal 
glands often visible, base attenuate, sessile or with 
petioles up to 15 mm long. Stipules UPS purple, 
ovate-triangular, succulent, ca. 0. .35 
long, 0.05-0.25 mm wide. Flowers Fic numer- 
ous, congested in leaf axils. Sepals ovate-deltoid, 
ascending, 1.1-2.3 mm long, 1-1.75 mm wide, 
glabrous, apex short acuminate or acute, margin 
entire, fringed with minute strigillose hairs. Petals 
0 s 0.3-0.5 mm long; filaments nearly 
translucent, 0.55-1 mm long. Pollen grains shed 
as tetrads. Nectary disc greenish, raised ca. 0.25- 
0.4 mm on top of ovary, 0.6-1.8 mm across, 
4-lobed, glabrous. Style pale green, 0.3-0.75 mm 
long, glabrous. Stigma greenish, subglobose, 0.25- 
0.5 mm across. Capsules subcylindrical with 4 shal- 
low grooves, each below the midvein of the sepal, 
2—7(-9) mm long, 1.3-2(-3) mm thick, glabrous 
to minutely papillose-strigillose, the hairs 0.02- 
0.05 mm long, sessile or with pedicels up to 0.35 
(-0.5) mm long, well spaced on main stems and 
primary branches, often congested on short sec- 
ondary or tertiary branches. Bracteoles flanking 
capsule base or attached up to 2.1 mm above 
capsule base, rarely on the short pedicel, lance- 
linear or linear, 0.35-1 mm long, 0.1-0.35 mm 
wide. Seeds light brown, kidney-shaped, the ends 
slightly pointed, ca. 0.5-0.75 mm long, 0.25-0.4 
mm thick, surface cells columnar, elongate either 
parallel or transversely to the seed length. 
compatible. Gametic chromosome number, n — 16. 


KEY TO THE SUBSPECIES OF LUDWIGIA GLANDULOSA 


Capsules (4-)5-8(-9) mm long; seed surface cells 
elongate parallel to seed length 


a sp. glandulosa 
Capsules 2-4 mm long; seed surface calls elongate 
transversely to seed length ... b. subsp. brachycarpa 


The two entities here regarded as subspecies of 
Ludwigia glandulosa were first recognized as dis- 
tinct by Torrey & Gray (1838-1840). Ludwigia 
glandulosa subsp. brachycarpa differs from subsp. 
glandulosa in its smaller stature, narrower leaves, 
and smaller flowers and capsules. Furthermore, the 
seed surface pattern of these two entities—a fea- 
ture that is often diagnostic in Ludwigia sect. 
Microcarpium —is sharply distinct, as given in the 


ey. 

Ludwigia glandulosa subsp. glandulosa is very 
common and widespread throughout the Atlantic 
and Gulf coastal plains and the Mississippi Em- 
bayment, westward to eastern Texas and south- 
eastern Oklahoma. Subspecies brachycarpa ranges 
just on the western edge of that of the subsp. 
glandulosa in northern Texas and Oklahoma. It 
grows in the same area as subsp. glandulosa 
throughout much of its range but extends farther 
south and west. The general distinctiveness of these 
two subspecies, without doubt, is maintained by 
their modal autogamy; vegetative reproduction by 
means of stolons may likewise play a role in pre- 
serving favored genotypes. 


4a. Ludwigia glandulosa Walter subsp. glan- 
dulosa. Figure 


Jussiaea brachycarpa Lam., Encycl. 3: 331. 1789. TYPE: 

U.S.A. South Carolina: s.d., J. Fraser s.n. (holotype, 
, photograph, 

Ludwigia heterophylla Poiret in Lam., En cycl. 
3:512. 1814. TYPE: U.S.A. * T e 
1800, L. A. G. Bosc s.n. (holo 

"e eytindelen Elliott, duce po t. S.C. & Ca. 

ins . South Carolina: her 

ar piera t, Aug.-Sept., s.d., 

CHARL; isotypes, GH, 

MO, NY; possible isotype, PH). 


Stems often reddish, (20-)40-80(-100) cm tall. 
Cauline leaves elliptic to very narrowly elliptic, 
those on main stem 32-120 mm long, 4-21 mm 
wide, those on the branches much smaller, 10-45 
mm long, 3-10 mm wide, petioles 1-15 mm long. 
Sepals ovate-deltoid, the apex acuminate, 1.25- 
2.3 mm long, 1.15-1.7 mm wide. Petals 0. An- 
thers 0.3-0.5 mm long; filaments 0.55-1 mm 
long. Pollen grains shed as tetrads. Nectary disc 
glabrous. Style 0.3-0.5 mm long; stigma 0.35- 
8(-9) mm long, 


Suppl. 
1798- 


5 mm across. Capsules (4—)5- 
1.6-2(-3) mm thick, sessile or with pedicels up to 


254 Annals of the 
Missouri Botanical Garden 


y p 


fi 
fi 


3 


4 
‘ P. 


f Y m i WA i à 
Hill Ji T WAN 
N AN | y wl MN IPN 4u 


FicURE 41. Ludwigia glandulosa subsp. glandulosa (Virginia: Southampton Co., Fernald & Long 9109 
GH). — a. Habit, upper stem. —a’. Lower stem. — b. Leaf. — c. Flower bud. —c’, Flower. —d. epal. —e. Partly dissected 
flower. —f, f'. Adaxial and abaxial views of stamen.— g. Capsule. —h. Cross section of capsule. 


Volume 76, Number 1 
1989 


Peng 255 
Ludwigia sect. Microcarpium 


0.35(-0.5) mm long. Bracteoles flanking capsule 
base or attached up to 2.1 mm above capsule base, 
rarely on the short pedicel, 0.5-1 mm long, 0.15- 
0.35 mm wide. Seeds 0.5-0.7 mm long, 0.25- 
0.35 mm thick, the surface cells oblong, elongate 
parallel to the seed length. Self-compatible. Ga- 


metic chromosome number, n = 16 


Distribution. Plants of L. glandulosa subsp. 
glandulosa are commonly found in roadside ditch- 
es, marshes, pond borders, wet meadows, swales, 
alluvial floodplains, peaty bogs, moist pinelands, 
swampy woodlands, and occasionally on waste 
ground. Ludwigia glandulosa subsp. glandulosa 
occurs throughout the Coastal Plain from extreme 
southeastern Virginia southward through eastern 
North Carolina, South Carolina, south and central 
Georgia to the panhandle of Florida and westward 
across southern Alabama, Mississippi, and Louisi- 
ana, reaching its western limit in eastern Texas. 
To the north, this subspecies extends through ex- 
treme southeastern Oklahoma, south and central 
Arkansas, and up the lower Mississippi basin to 
southeastern Missouri, central and eastern Ten- 
nessee, extreme western Kentucky, southernmost 
Illinois, and southwestern Indiana. Disjunct pop- 
ulations of L. glandulosa subsp. glandulosa are 
found in northcentral Missouri (Fig. 43). Flowering 
from June to September; fruiting from June to 
October. 


po specimens examined. U.S.A. ALA- 
, near Pratville, 1880, Mohr s.n. (US). 
i. N of Spanish a A AL Hwy. 
225, Kral 37447 (WDB). Barbour Co., mi. N of 
Eufaula, on AL Hwy. 165, Kral 33190 (C. FSU, VDB). 
Colbert Co., near Littleville, along US Hwy. 43, Kral 
44012 (VDB). Dale Co., near Clayhatchee, Kral 22078 
(VDB). Dallas Co., at mi. 161 of Alabama River, in Miller's 
Ferry Reservoir, Wiersema 401 (UNA). Escambia Co. - 


Kral 44816 (VDB). Geneva Co., NE s 
d River, Kral 31911 (FSU, VDB). 

, ca. 1 mi. E of Gainesville, Harper 4524 (GA, GH, 
MO, Nu UNA, WCUH). Houston Co., ca. 6 mi. S of 


o 
Ft. Deposit, Kral 40910 (VDB). Macon “Co. . 19 mi. W 
, Raven 18562 (DS). 


Haynes 6750 bob Mobile Co., Curtiss 6503 (DS, 
MO, NY, US). Monroe Co., 3 mi. E of Chrysler on AL 
Hwy. 59, Shome Creek, Kral 40948 (VDB). Montgomery 
Co., 2 mi. N of Montgomery near Tyler-Goodwin Bridge 
on Albanis River, Justice & bely 137 (US). Ma 
Co., E of Decatur on AL Hwy. 67. Kral 36436 (VDB). 
Perry Co., W of Sprott on Hwy. 183, Cahaba River, 


erford, Clark 16208 (NCU). 


sey s.n. (F, NY). Wilcox Co., 0.5 mi. E of Camden 1958, 


Williams s.n. (FSU). ARKANSAS: Ashley Co., near Bg x 
Demaree 70209 (MO). Calhoun Co., near Thor n, De- 

maree 36618 (DS). Clark my near ArcDelphia, Patter 
8069 (CAS, MO, NY). , near Corning, Palmer 
6082 (DS, POM, US). Gee Co., near Kingsland, 
Demaree 23304 (MO, SMU). Columbia Co., near Mag- 
nolia, Demaree 39191 (FSU, GH, NCU, SMU). Conway 
Co., near Petit Jean Mountain, Moore 811 (SMU, TENN). 
Craighead Co., near Lake City, St. Francis River, De- 
maree 7076 (F, GH, ND, SMU, TEX, US). Desha Co., 
near Dumas, Demaree 48844 (DS, NO, SMU). Drew Co., 

near Wilmar, Demaree 66828 (MO). Garland Co., on 
Big Island in Lake Hamilton, Demaree 39457 (CAS, 
SMU, VDB). Grant Co., near Sheridan, pena 19467 


reene Co., near Paragould, 


SMU). Hot Spring Co., 1896, Eggert s.n. ( ee 
SE of Haynes, S of Clay Hill, MeDaniel 837 (GA). Laike 
Co., near Carlisle, Denir ree 56843 (DS, SMU). Monroe 


nd Mine 
(DS). Prairie Co., near Hazen, Demaree 15457 (DS, MO, 
NY, SMU). Pulaski Co., near Little Rock, 1885, Hasse 
s.n. (CM, NY). St. Francis Co., 2.5 mi. NE of Madison, 
upper end of Keithley Lake, Mefaniel 9730 (FSU, MO). 
Sebastian Co., Massard Prairie, Armstrong 193 (TEX). 
Yell Co., near Plainview, ea 46645 (DS, NCU, 
VDB). FLORIDA: Escambia , near Hwy. 4 on Escambia 
River, Redfearn & Kral 2777 (FSU, CH). Gadsden Co., 
River cw Nash F, GH, MO, MSC, NCU, 


ND, NY Gulf Co., ps si 1896, s.c., s.n. (US 
Hamilton Co., a. 2 mi. S of Genoa, 1946, Arnold & 
pies s.n. (US). Holmes ay hi 0. 5 mi. E of Westville, 4 ie 


US). 
Redfearn 2836 (FSU, GH, NCU). Jackson Co., 
eie Lake Seminole, Godfrey & Houk 61400 (FSU, 


Raven 18617 (DS). Leon Co., 2 mi. E of Tallatianace, 

Gode 55604 (FSU, GA, GH, NY, ba dr "rne Co., 
W side of pae River, Morar 7 (SM 

VDB). Madison Co., 6.8 mi. 

e Raven 18628 We Okaloosa Co., 
L Hw 


& Dille 412 (MO). Taylor Co., N of 
hey & Kral 54954 (FSU, GH, NY en Wakulla Co., 

mi. N of Crawfordsville, Kral 3006 (FSU, GH). Walton 
a at Mossy Head, Godfrey 55243 (FLAS, GA, GH, 
NY, SMU, TENN, UNA, USF). ceorcia: Bibb Co., Oc- 
mulgee River, SE of Macon, Tel Welch s.n. (GA). 
Bryan Co., on US Hwy. 17, 0.5 mi. SW of GA Hwy. 
I-95, Dille & Dille 351 Tia dies n Co., 2 mi. 


US Hwy. 1, Peng 23 (MO). Evans Co., 1 mi. 
of GA Hwy. 16 
25 & 301, Peng 4103 (MO). Grady Co., 9.5 mi. 
Cairo, Kral 15174 (VDB). Jeff Davis Co., NW of AE 


256 


Annals of th 
Missouri Bond Garden 


4119 (MO). Lowndes Co., E side of Nevins Hall, Valdosta 
State College, Faircloth 4723 (NCU). McDuffie Co., Bow- 
den's Pond, 11.75 mi. S31%E of Thomson, 1.5 mi. SSE 
of Arrington Mill, Duncan 11522 (GA, MO, SMU) 
McIntosh Co., 3.2 mi. NW of ject. of Cox Rd. & ph 
251, on Cox Rd., Peng 4139 (MO). Screven Co. 
Murray Hill, 1907, Cuthbert s.n. (FLAS). Seminole Co. 
paved Co. rd. between Desser & Reynoldsville, Jones el 
al. 23645 (GA). Sumter Co., Muckalee Creek, Americus, 
, NY). Tattnall Co., Altamaha 
River, 2 mi. NE of Lane's Pur GA Hwy. 169, Boze- 


mi. W of dicen Thorne 6331 (GA). ILLINOIS: Massac 
Co., Mermet Lake, SW of Mermet, Evers 85808 (DS, 
MO). Pulaski Co., along Cache River, W of Pulaski, 
Bailey & Swayne 27 10 (NCU). INDIANA: Posey Co., SW 
corner of Sect. 32 of Point Tp., ca. 10 mi. SW of Mt. 
Vernon, Deam 37699 (GH, NY, US). KENTUCKY: Cal- 
loway Co., near Backusburg, on W fork of Clarks River, 
Athey 2471 (NCU, NY, VDB). Fulton Co., near Hick- 
man, along KY Hwy. 1282, 0.7 mi. E of jct. of KY Hwy. 
94, Athey 2060 (NCU, NY). Graves Co., along KY Hwy. 
339, Obion Creek bottoms, Browne et al. 73E7.8 (NCU, 
y. 1523, 0.5 mi. from 
wy. 62, Athey 1132 (NCU, NY). Mc- 
Cracken Co., Pool Road, Athey 1117 (NCU, NY). roui- 
SIANA: Acadia Parish, 5 mi. S of Eunice, Kral 17195 
(VDB). Allen Parish, 7.2 mi. W of Kinder, Shinners 
21504 (SMU). Beauregard Parish, 1-1.5 mi. S of Bayou 
Amacoco, Ewan 20106 (DS, NO). Bienville Parish, along 
US Hwy. 20 at Leaterman Creek, W of Gibsland, Sect. 
10, T18N, R7E, Thomas et al. 5209 (SMU, TENN). 
Bossier rr along Clear Lake, S of 2 Road, 
Balogh 552 (NCU). Calcasieu Parish, 2.5 mi. SW o 
Starks, rud 25084 (SMU). Caldwell Parish, 3 mi. 
Gra yson, Shinners 21917 (SMU). Catahoula Par- 


mi. 
W-SW of Mansfield, Shinners 20349 (SMU). East Felici- 
ana Parish, MacManus RR station, E of Jackson, Ewan 
18772 (NO). Franklin Parish, Winnsboro, Civitan Park, 
Thieret 30050 (A, B, FLAS, GA, NCU, SMU, TEX, 
VDB). Iberia Parish, old air base, New Iberia, Thieret 
KE, FSU, GA, SMU, TEX). Iberville Parish, 
Er a s.n. (NY). Jefferson Parish, 5 
e Road, Ewan 17757 (DS, GH, 
NY). Lafayette Parish, l mi. N of Lafayette, Fi 
Py Correll 9436 (DUKE, GH, NY). Lincoln Parish, 

mi. E of Ruston, Kral 15736 (VDB). Livingston Pa 

Denham Springs, 0.5 mi. N of Range Ave. & US Hw 
12, Montz 2253 (NO). Morehouse Parish, ca. 4 mi. NE 
of Beekman, Sect. 26, T23N, Sd Vincent 456 (DUKE, 
USF). Natchitoches Parish, 1 mi. W of Clarence, 
along Red River, Thieret 30247 (DS, DUKE, FLAS). 
Orleans Parish, near New Orleans, 1836, Arnott s.n. (P). 
Quachita Parish, 3 mi. SW of West Monroe, Kral 8094 
e Rapides Parish, vicinity of — Ball 640 
(F, GH, MO, NY 
Mangham a along LA Hwy. 15, Vincent 499 (FSU, USF). 
a Parish, near bridge across Sabine River to TX 
y. 21, Demaree 48220 (DS, NO, SMU, VDB). St. 
uie Parish, Bonnet Carre Spillway near river road at 
westguide levee, Montz 2929 (NO). St. Landry Parish, 


ichland Parish, ca. 1 mi. S of 


ca. 4 mi. NW of Grand Coteau, Thieret 16454 (DS, 
DUKE, FSU, GA, SMU, VDB). St. Tammany Parish, 
edge of Pearle River under Interstate Hwy., N of Pearle 
River, Raven 18576 (DS, NCU). Tangipahoe Parish, W 
limits of Hammond, Raven 22100 (DS). Tensas Parish, 
ca. 3.5 mi. of Newellton, along Hwy. to Newlight, 
Thieret 23962 (DS, SMU). Terrebonne Parish, near Hou- 
ma, 1913, Wurzlow s.n. (NY). Union Parish, ca. 2.5 mi. 
S of Lillie off US Hwy. 167, Vincent 397 (GA). Vermillion 
Parish, ca. 2 mi. SW of Abbeville, along Vermillion River, 
Thieret & Bernard 17863 (DUKE, FSU, SMU, TEX, 
USF). Vernon Parish, S limits of Leesville on Hwy. 171, 
Raven 22088 (DS, NCU). Washington Parish, Mitchell 
., ca. 3 mi. S of Bogalusa, Darwin et al. 763 (MO, 
NO). Webster Parish, 3.5 mi. E of Minden exit on US 
Hwy. 20, Kral ele (MSU, VDB). West Carroll Parish, 
1903, Moseley s. . West Feliciana Parish, 1 mi. 
id ates kefield, "Thieret 24068 (DS). Winn Parish, ca. 
N of Verda, S27, TON, R3W, Vincent 166 
DUKE FLAS, USF). MARYLAND: Somerset Co., Crisfield, 
: ee n. (MSC, mixed en L. palustris). MIS- 
ms Co. river floodplain, 9 mi. W-NW of 
Doloroso, Tons et al. 13529 (FSU, MISS, NCU). Amite 
Co., roadside of Longleaf Plantation, ca. 5 mi. S of Liberty, 
Jones et al. 19946 (GA, MISS). Bolivar Co., 3.5 mi. W 
of Duncan E e Hwy. 444, Pullen 70420 (MISS). 
Calhoun Co. i. E of Bruce on MS Hwy. 32, Pullen 
E a MIS, aue Co., 4.7 mi. E of Vardaman 
S Hwy. 8, Temple 3535 (GA, MISS, NCU). Clarke 
Co. , 3 mi. S of Pachuta, Ray 5072 (GA, GH, NCU, NY, 
TENN, USF). Clay C o., Tibbee Creek bottoms, ca. 3.8 


otto 
oto line, pido 70877 (MISS). Forrest Co., 
Ragland Hills, Leaf Riv 
Franklin Co., edge of Clea 
10089 (MISS). George Co., a 
cola P.O., Demaree 35758(GH, SMU), G 
Leaf River on MS Hwy. 98, Jones & 
(MISS). Hancock Co., along Jordan River S of Kiln, Jones 
& Jones 14145 (MISS, TEX). Harrison Co., near Biloxi, 
Tracy 6463 E (e MSU, NCU, NY, US, VINDOB). 
Holmes Co., i. S of Tchula near open R/ s of 
MS Hwy. 12, “Pullen 70402 (MISS). Humphreys Co., 

MS Hwy. 12, 2.6 mi. NW of Holmes-Humphreys n 
line, Pullen 69973 (MISS). Jackson Co., Pascagoula air- 
port rd., Jones & Jones 14941 (GH, MISS, NCU). Jasper 
Co., 6 mi. via rd. from Enterprise along Souinbuey Creek, 
Jones 15162 (MISS, TEX). Jefferson Co., 6 mi. N-NW 
of Church Hill at base of loess bluffs, Jones & Noble 
10171 (MISS). Jones Co., 6.6 mi. NE of Moselle on MS 


9m 
MISS, TENN). Lauderdale Co., 
a ie 14995 (GA, MISS, SMU, USF). Leake Co., 
pu line, ca. 1 mi. N of Pearl River, S of Wiggins 


=> 
z 


on arcis 3, Jones et al. 18127 (MISS). Lincoln Co., 
11 mi. W a Brookhaven, ai & Wilbur 3296 (GH, 
NY, SMU, US). Madison Co., mi. SE of Canton at 


Ross Barnett Reservoir, Jones D (MISS). Monroe 
Co., ca. 3.5 mi. W of Amory, vicinity of Tombigbee River 
bridge on US Es "s Pullen 67 1032 (MISS). Mont- 
gomery Co., 5. E of Duck Hill on MS Hwy. 404, 
Pullen 701013 (MISS). Neshoba Co., 5.1 mi. 

Choctaw Indian Reservation, W of Philadelphia on MS 


Volume 76, Number 1 
1989 


Peng 257 
Ludwigia sect. Microcarpium 


Hwy. 16, Temple 11996 (MISS). Panola Co., 2.4 m 
of ries River bridge on MS Hwy + 6, Templo 5 5893 


3.5 mi. N of Tallahatchie-Quitman county line on MS 
Hwy. 3, Pullen 641453 (GA, MISS). Rankin Co., ca. 3 
mi. SE of Johns on MS Hwy. 18, Temple 11960 (GA, 
MISS). Scott Co., edge of lake, Raworth Recreation Area, 
between Morton and E Jones et al. 17874 (FLAS, 
MISS). Sharkey Co., ca. 7 mi. W of Holly Bluff in Delta 
National Forest, Pullen 691196 (MISS). Simpson Co., 2 
mi. N of D’Lo on MS Hwy. 49, Jones 14065 (MISS, 
VDB). Smith Co., 5.1 mi. S of Pulaski and N of Trenton 
on MS Hwy. 481 at oe ke 19151 (MISS, SMU). 
Tishomingo Co., MS Hwy. 72 at bong Creek, Coleman 
48529 (TENN). Union "in E 7 mi. SE of Etta on county 
rd., around creek bridge, Pullen 70842 (MISS). Wash- 
ington Co., 1 mi. E of Longwood, Pullen 65302 (MISS, 
NCU). Wilkinson Co., 8.4 mi. SW of MS Hwy. 33 near 
Crosby on MS Hwy. 563, Temple 12148 (MISS). Winston 
., Nanih Waiya St. Pk., S-SE of Louisville, Jones et 
al. a (MISS, VDB). Yazoo Co., 3.5 mi. NW of jet. 
Hwy. 49E on MS Hwy. 3, Temple 10189(GA, MISS). 
oa. I: Butler Co., swamps in Blue Spring, 1892, Eg- 
i Php: Cape Giradeau Co., 3 mi. W of Arbor on 
9, Steyermark 64138 (F). Sin ome Co., near 

Can 1893, Bush s.n. (GH, MO, ND, NY). Pemiscot 
., 7 mi. S of Portageville, Steyermark 9131 (MO). 
Stoddard Co., woods bordering Swan Pond, T28N, R10E, 
Sect. 35, 4 mi. S of Advance, Steyermark 66146 (MO). 


— 


ar Paw 
E of Chaonia, T26N, R7E, Sect. 32 & part 
of 33, Steyermark 6197 (F, MO). NORTH CAROLINA: An- 
son Co., Brown's Creek on NC Hwy. 74, W of Wadesboro, 
Duke 54-111 (NCU). Beaufort Co., mad SE 300 
yds. N of Wynns Service Station on US . 17, Duke 
54-176 eph Bertie Co., under bridge of yond Creek 
. 70E of Wi Py iie Duke 54-261 (NCU). 
ton on road to Bluefield, 


1227 (NCU). p Co., dich a Belgrade, Duke 54- 
247 (NCU). Chatham Co., 7 mi. E of Pittsboro, RR R/ 
Hwy. 64 crossing, Campbell 1046 (NCU). 


Columbus Co., 2.3 mi. N of Tabor City on NC Hwy. 410, 
Duke 54-91 (NCU). Craven Co., 0.8 mi. N of Hwy. 17 
on Co. Rd. 1224 (rd. to Tuscarora), Peng 3739 (MO). 
Cumberland Co., 1 mi. N of Beard, the 0.4 mi. W-NW, 
Ahles & [d 33545 (NCU). Durham Co., jct. of Co. 
Rd. 1242 (Biscayne Rd.) and NC Hwy. 54, along New 
Hope Creek, Boufford & Wood 1886 1 (MO). Edgecombe 
Co., beside Swift Creek on Hwy. 85, Duke 54-7 1 (NCU). 
Gates Co., 1 mi. NE of Chowan gd on NC Hwy. 158, 
Beal 2132 (NCSC). Greene Co., 4 mi. W of Farmville 
on US Hwy. 70, Duke 54-168 (NCU). pies Co., Byrd's 
Mill Pond, S of Bunn Level on NC Hwy. 15a, Duke 54- 
149 (NCU). Hertford Co., 1.1 mi. NE of Como then 1.9 
mi. E-NE on dirt rd., Ahles & Duke 45982 (NCU). Hyde 

o., 0.5 mi. N of NE Creek, on US Hwy. 264, 
Duke 54-280 (NCU). Johnston Co., Neuse Ri ras 3 mi. 
N-NE of Cox Mill, Radford 27789 (NCU). Jones Co., 
0.4 mi. W of Co. Rd. 1105 (Catfish Rd.) on Forest Service 


Rd. 172 (Murey Branch Rd.), E-SE of "xn off NC 
Hwy. 58, Peng et al. 3801 (MO). Lee Co., 2 mi. NW 
of Sanford along Dig Buffalo Creek, Houck 359 (NCU). 
Martin Co., 2.3 m ha of Robersonville, Radford 39396 
(NCU). Nash Co., r Moccasin Creek and US Hwy. 
264 W of Middlesex, "Radford 44350 (NCU, US). North- 
is Co. mi. of Margarettsville, then 0.3 mi. 
S, Ahles & Misa, 52529 (NCU). Pamlico Co., 5 m 
W of Grantsboro on NC Hwy. 55, Duke 54-227 (NCU). 
Pasquotank Co., 1.5 mi. N of Nixonton by ypa Creek, 
Ahles & Duke 51239 (NCU). Pender Co., 4 mi. of 
Montague on Black River, S aA et al. Ae (NCU). 
Perquimans Co., 2 mi. E-NE of Chowan- oe 
county line, Ahles & Duke 48002 (NCU). Pitt Co., 
mi. E of Pactolus near NC Hwy. 33, Radford eee 
TEX). Robeson Co., 4.7 mi. S of Fairmont on NC Hwy. 
41, Duke 54-143 (NCU). Rowan Co., 0.1 mi. N of Stanly- 
Rowan county line, Ahles & Haesloop 57224 (NCU). 
Stanly Co., 3 mi. E of Richfield on NC Hwy. 49, Duke 
54-103 (NCU). Union Co., 3.5 mi. E of New Salem, 
Ahles 34111 (NCU). Wake Co., 6.5 mi. S of e ^ 
Yates Pond, Beal 1837 (NCSC). Washington Co., 
mi. NW of Creswell, Radford 38902 (NCU). Wilson d 
4.6 mi. W of Wilson, Radford 40791 (NCU). OKLAHOMA: 
Atoka Co., 3.5 mi. SW of Atoka, Cory 56815 (SMU). 
Choctaw Co., near Hugo, Xo radi 6348 (CAS, MO, NY, 
US). LeFlore Co., near Page at pond margin, Stevens 
2626 (DS, GH, MO, NY, US). Ma Co., 2.5 mi. SE 
of Enos, Happy Hollow, near Lake ‘Texoma, Goodman 
6696 (RSA, TEX). McCurtain Co., i. 
of Eagletown, reorum swamp, Waterfall y 0406 (SMU), 
Sequoyah Co., 0.5 mi. SE of Blackgum on OK Hwy. 
100, Wallis oe (SMU). SOUTH CAROLINA: Aiken Co. 
North Augusta, near Savannah River, Ahles & Crutch- 
oe d (NCU). Allendale Co., Coosawhatchie River, 
à . SE of Fairfax on County Hwy. 21, Bell 5064 
(NCU. USCH). Bamberg Co., Salkehatchie River on US 
, NE side of river, Ahles & Haesloop 
i. W of Co. Hwy. 111 
on Co. Hwy. re n 3759 (NCU, U). Berkeley Co., 
Santee River, 3 mi. NE of Pineville, N & Tryon 
644 (CAS, DUKE, F. GH, ae NY, TEN 
ton Co., 24.5 mi. N of SC 
Recreation Area, S of MeClellenvile. Ed et al. 3896 
(MO). Cherokee Co. . 29A on dirt 
road along Broad River les & por É 1925 (NCU). 
Chesterfield Co., 1 mi. W of McBee in the Sandhills, 
i S 8025 (F, CH. MO, NY, TENN, US). Clarendon 
, NE shore of Lake Marion, ca. 4.5 mi. SW of St. 
Fail off US Hwy. 15, Bradley & Sears 3561 (MEXU, 
MISS). Colleton Co., 1.2 mi. W of Dorchester county 
line on SC Hwy. 61, Raven 20461 (DS). Darlington Co., 
damp flats, Smith 2001 (NCU). Dillon Co., Sweat Swamp, 
ca. 3 mi. N of Little Rock on Co. Hwy. po ET & 
Leisner 32234 (NCU). Dorchester Co. . 0.4 mi. N of 
Reevesville on Co. Hwy. 16, Ahles & E 31 905 
(NCU). Georgetown Co., 2.8 mi. NE of Yauhannah, Pee 
Dee River near US Hwy. 701, Radford 28609 (NCU). 
Greenwood Co., 1.5 mi. N of Ninety Six, Wilson Creek 
near hes Hwy. 246, Radford 26581 (NCU). Hampton 
., 0.8 mi. N-NW of Shirley on Co. Hwy. 20, Ahles & 
Bell 18249 (NCU). Horry Co., along Waccamaw River 


— 


amden 
adfo NCU), Marlboro me 
9 mi. SW of Bennettsville, Pee Dee River, near US Hwy 


258 


Annals of th 
Missouri EH Garden 


52, Radford hp (CM, FLAS, NCU, SMU). - 
Cormick Co., i. SE of Clarks Hill, Savannah Rive 
near SC Hwy. 28, Radford 30647 (NCU, VDB). Ora iue 
burg Co., 3.9 mi. S-SE of Eutawville on Co. Hwy. 104 
along Sandy Run, Ahles 35003 (NCU). Richland Co. 15 
mi. below Columbia along Congaree River, 1965, Batson 
s.n. (USCH). Saluda Co., near US Hwy. 178 at Mayson, 
Radford 26906 (NCU). Sumter Co., 4 mi. E-NE of Shi 
loh, Jordon Swamp, Radford 27500 (FSU, NCU). 
TENNESSEE: Carroll Co., edge of cat-tail pool, Svenson 
4385 (GH). Crockett Co., Cypress Creek N of Bells, Sharp 
& Clebsch 6754 (TENN). Franklin Co., near Huntland, 
Svenson 10403 (GH). Grundy Co., N of Mt. View, God- 
frey 69774 (FSU, NCU). Hardeman Co., N of Middle- 
burg, Sharp & Clebsch 8208 (TENN). Henderson e 
Beech River SW of Chesterfield, Sharp 
(TENN). Lake Co., Reelfoot Lake, 1924, piu & 
Jennings s.n. (CM). Montgomery Co., 1.5 mi. NW Sad- 
lersville by US Hwy. 41, Kral 48384 (VDB). Obion Co., 
near Bayoudechien, Walnut Log, Sharp & Clebsch 6207 
(TENN). Shelby Co., McKellar Lake, Ensley a 
yard, Rogers 34826 (TENN). White Co., 9 mi. N of 
Sparta, swamp S of Cookeville, Norris E Sharp 16241 
TENN). rEXAs: Angelina Co., near and, Lewis 
Oliver 5245 (DH). Bastrop Co., near PES. Tharp 224 


leburr Slough above bridge on Shell Rd. 
Refuge, Fleetwood 10159 (SMU). Brazos Co., Fish Lake 
behind Easterwood oe College Station, Massey 309 
(NCU, TAES, TEX). Cass Co., ca. 2.5 mi. SW of McLeod 
along Frazier Creek, Conal 301 13 (TEX). Fayette Co., 
near Muldoon, Ripple 51-914 (TEX). Galveston Co., S 
edge of Alta Loma, Correll & Correll 38997 (LL). Con 
zales Co., , Bogusch 943 (TEX ). Gregg Co., ponds along 

i Hwy. 149, Correll 


Co., 8 mi. E of Edna, Navidad River Se Webster 
& Wilbur 3142 pow is Jasp , near Harrisburg, 
1875, Joor s.n. (F). Lamar Co. mi. NE of Paris 
on TX Hwy. 195, Correll 37518 (TEX). Liberty Co., 
5.7 mi. E of — n 19425 (DH, FLAS, NCU). 
Limestone a 4m Donie on Hwy. 80 & 0. 
mi. E on 39, b mb 3088 (SMU). Marion Co., 
9.6 mi. NW a Jefferson, Shinners 30257 (SMU). Mont- 
gomery Co., mi. E of Montgomery, 1950, Henry 
s.n. (CM). Nacogdoches = Waller 254 (DH, TAES, 
TEX). Newton Co., f Bon Wier, Kral 20731 
(VDB). Orange Co., iong chen River near bridge, E of 
Orange, Correll 26792 (TEX). Panola Co., 8 mi. NE of 
Carthage, Sabine River, Shinners 15656 (SMU, TAES). 
Polk Co., McCordell Lake, E of TX Hwy. 59, /Vixon et 
al. 5363 (NCU). Rains Co., a 2 p S of Point, Van Vleet 
678 (SMU). Red River Co. T 7.5 mi. N of Clarksville 
along TX Hwy. 37, Correll 57495 (NY, TEX). Titus Co., 
N of Winfield, McGregor 750 (SMU). Upshur Co., 5 mi. 
N of Ore City, along Cypress Creek, Correll 1317 1 (NY, 
eer TEX). Waller Co., near ae ir Hall 
n. (F, GH, NY, US). Wood Co. FEA 
Kral 1239 (FSU). VIRGINIA: e Ca, SW of 


& 


wy 


o o., ca i. E-NE o 
(FSU, VDB). Norfolk Co. S of North Landing, Fernald 


& Long 4076 (GH, NY). Prince George Co., near Gary 
Church, Fernald & Long 6307 (GH, NY). Princess Anne 


Co. 
Southam mp 

Drewryville, 1970, Churchill s.n. (MSU). Warwick Co., 
near Newport News, 1907, Long & Bartram s.n. (NY 
York Co., E of Tabb’s, Fernald & Long 8787 (GH). 


Ludwigia glandulosa is distinctive in being the 
only tetraploid species in sect. Microcarpium with 
cylindric capsules. The other three species with 
elongate capsules are the diploid, petaliferous, nar- 
row-leaved L. linearis, L. linifolia, and L. stricta. 
Raven & Tai (1979) suggested that L. linearis is 
similar to L. glandulosa and might be directly 
related to it. The biosystematic study of Peng 
(1988), however, demonstrated that the genetic 
relationship between the diploid L. linearis and the 
tetraploid L. glandulosa is not any closer than that 
between L. linearis and other tetraploids. Fur- 
thermore, it has been shown that no extant diploids 
are involved directly in the formation of any tet- 
raploid species in sect. Microcarpium. 

Ludwigia glandulosa subsp. glandulosa is 
known to occur together with subsp. brachycarpa, 
L. linearis, L. linifolia, L. microcarpa, L. pilosa, 
|. sphaerocarpa, and L. palustris (sect. Dantia) 
in the field. It has been known to form natural 
hybrids with all but L. microcarpa, an extreme 
selfer, and the facultatively outcrossing £. linifolia. 
Attempts to synthesize L. glandulosa X L. lini- 
folia in the experimental greenhouse also failed 


~ 


Peng, 1988), presumably owing to incompatible 
genomes. Artificial hybrids between L. glandulosa 
subsp. glandulosa and L. microcarpa, however, 
were healthy and floriferous. 

Duke (1955) was the first to record a natural 
hybrid population of Ludwigia glandulosa subsp. 
glandulosa x L. pilosa, although he did not fully 
describe the situation he observed. Tetraploid hy- 
brids like this and others are usually fertile and are 
intermediate in such diagnostic characters as ves- 
titure, sepal and bracteole length, capsule shape 
and size, seed surface pattern, and whether pollen 
is shed singly or as tetrads. Triploid natural hybrids 
involving L. glandulosa subsp. glandulosa and 
diploids like L. linearis and L. palustris may be 
floriferous but are highly sterile as to pollen and 
seeds. Their ovaries turn yellowish or pink and 
dehisce early. Ludwigia glandulosa subsp. glan- 
dulosa (stems erect, leaves alternate) X L. palus- 
tris (stems prostrate, leaves opposite) generally re- 
sembles L. glandulosa subsp. glandulosa but has 
slightly broader leaves and smaller flowers. The 
stems are erect and then prostrate and erect again 
in some cases; the leaves are alternate like L. 


Volume 76, Number 1 
1989 


Peng 259 
Ludwigia sect. Microcarpium 


glandulosa subsp. glandulosa. By contrast, hy- 
brids of L. glandulosa subsp. glandulosa and L. 
linearis are similar to L. linearis, having some 
yellow petals and narrow leaves. The number of 
petals on each flower varies from 0 to 4. 


4b. Ludwigia glandulosa Walter subsp. 
brachycarpa (Torrey & A. Gray) Peng, Ann. 
Missouri Bot. Gard. 73: 490. 1986. Based on 
L. cylindrica Elliott 8 brachycarpa Torrey 
& A. Gray, Fl. : 524. 1840, non 
Jussiaea brachycarpa Lam., Encycl. 3: 331. 
1789. L. glandulosa Walter var. torreyi 
Munz, Bull. Torrey Bot. Club 71: 164. 1944, 
illeg. subst. LECTOTYPE: U.S.A. Texas: Austin 
Co., San Felipe, “third collection," 1833- 
1834, T. Drummond 84 (GH; isolectotypes, 
GOET, K — 2 sheets, W; Peng, Ann. Missouri 
Bot. Gard. 73: 490. 1986). Figure 42. 


Stems sometimes reddish, 10-55(-90) cm tall. 
Cauline leaves linear-elliptic to linear, sometimes 
very narrowly elliptic, those on main stem 30- 
50(-70) mm long, 3-5(-10) mm wide, those on 
the branches 8-36 mm long, 2-3(-8) mm wide, 
sessile or with petioles up to 10 mm long. Sepals 
ovate-deltoid, the apex acute or short acuminate, 

1 mm long, 1-1.75 mm wide. Petals 0. 
Anthers 0.3-0.5 mm long; filaments 0.55-1 mm 
long. Pollen grains shed as tetrads. Nectary disc 
obscurely minutely papillose. Style 0.35-0.75 mm 
long; stigma mm across. Capsules 2-4 
mm long, 1.3-2 mm thick, sessile or with pedicels 
up to 0.15 mm long. Bracteoles flanking capsule 
base, 0.35-0.75 mm long, 0.1—0.2 mm wide. Seeds 
0.55-0.75 mm long, 0.3-0.4 mm thick, the sur- 
fa lls oblong, elongate parallel to the seed length. 


Self-compatible. Gametic chromosome number, n 


Distribution. 
subsp. brachycarpa are usually found along ditch- 
es, in low meadows, coastal prairies, seeps in sandy 

woods, moist sinkholes in granites, and old clay 
fields. This subspecies occurs along the Gulf coast, 
from extreme southwestern Louisiana to Nueces 
County, Texas, and northward through eastern 
Texas to southcentral Oklahoma (Fig. ow- 
ering and fruiting from April to November. 


Plants of Ludwigia glandulosa 


Representative specimens examined. U.S.A. LOUI- 
SIANA: Calcasieu Parish, 1 mi. S of Ti F Ewan 
21343 (NO). Cameron Parish, 2 mi. W of jct. of State 
hwys. 82 & 27, Peng et al. 4367 (MO). pii o 
Atoka Co., 6 mi. SE of Lane on OK Hwy. 3, Massey & 
Massey 2496 (MO, NCU, SMU). iis Co., N of Ard- 
more, Goodman 5961 (GH, RSA, SMU). Coal Co., 5 mi. 


i^ of bau on OK Hwy. 3, Massey 2484 (NCU, 

MU). L Co., 5 mi. SE of Ardmore in Lake Murray 
vw Park, ” Crutchfield 3449 (LL). TEXAS: Aransas Co., 
near Lenoir's Landing, Copano Bay, Correll & Correll 
18905 (GH, LL). Bastrop Co., Tharpe 870 (TEX). Bra- 
zoria Co., Columbia, 1980, Bush s.n. (MO, NY). Brazos 
Co., F ish Lake behind Easterwood oe Massey & 
Reaux 586 (LL, NCU). Burnet Co., 2 mi. N of Marble 
Falls, Granite Mt., 1946, Barkley & n nue s.n. (NY). 
Calhoun Co., Port O'Connor, 1930, Tharpe s.n. (TEX). 
Dallas Co., Dallas, 1877, Reverchon s.n. (NY). De 
Co., Lake Dallas Fish Hatchery, Sk e 1894 (NY, 
TEX). De Witt Co., western part of c 


veston Co., 1942, Nelson s.n. (G O, TEX). Gonzales 
Co., 7 mi. S of LT. Correll & Johnston 17481 
(LL). Harris - . 200 mi. S of TX H 59 near jet 
with Santa mi. N of Ov oor Co., 
Tranverse 1490 (F, SMU, TEX). Jasper Co., Herr 
75, Joor s.n. (LSU, US). Jefferson Co., on Hwy. 

12 mi. W of Beaumont, Munz & Gregory 23458 ME 

Co., ca. 18 mi. of Yoakum, Tharpe, Rogers 
& York 49177 (TEX). Limestone Co., ca. i. SE of 


T , MKT RR at Gapland crossing, 
Smith 863 (TEX). Montague Co., 2 mi. NW of Stoneburg, 
Skinners 26622 (SM Co., Mustang Island, 


"m 
7 5 


San Patricio he 3 mi. NE of Ingleside, Jones 932 (SMU). 
Travis Co., Austin, Tharpe 870 (TEX, US). Waller Co., 
Hempstead, Hall 219 (F, NY, US). 


Plants of Ludwigia glandulosa subsp. brachy- 
carpa occur in the western range of the TE 
and are frequently found in the same county a 
the typical subspecies, but mixed collections sug- 
gesting that they occur side by side are known only 
in the following: TEXAS: Fort Bend Co., 1 mi. NW 
of Arcola, water-filled roadside ditch of swampy 
brushlands, Raven 19405 (DS, mixed collection 
of two plants of subsp. glandulosa and one plant 
of subsp. brachycarpa; FLAS, subsp. brachycar- 
pa only). 

Natural hybrid populations between these two 
subspecies are known in two cases from OKLAHOMA: 
Atoka Co., ca. 0.25 mi. S of small community of 
Boehler and SE of Atoka, Taylor 20332 (VDB), 
and TEXAS: Galveston Co., ca. 2300 block of Broad- 
way, San Leon, Waller 3822 (GH). These hybrids 


are intermediate in stature, leaf shape and size, 


of columnar cells ‘ee are d transversely 
to the seed length). 


elongate, an 


5. Ludwigia polycarpa Short & Peter, Tran- 
sylv. J. Med. Assoc. Sci. 8: 581. 1835. /s- 


260 Annals of the 
Missouri Botanical Garden 


FIGURE 42. Ludwigia glandulosa subsp. brachycarpa (Louisiana: Cameron Co., Peng 4367, MO).—a. Habit, 
upper portion of erect stem.—a’. Lower stem with stolons.—b, b'. Adaxial and abaxial views of lower leaf. —c. 
Flower. —c'. Flower bud. —d, d'. Adaxial and abaxial views of stamens. —e. e'. Mature capsule. — f. Cross section of 
capsule. —g. Longitudinal section of capsule. 


Volume 76, Number 1 
1989 


Peng 
Ludwigia sect. Microcarpium 


RE 43. Distribution of Ludwigia glandulosa subsp. glandulosa (circles) and L. glandulosa subsp. brachy- 


carpa. (triangles). 


nardia polycarpa (Short & Peter) Kuntze, 
Rev. Gen. Pl. 1: 251. 1891. LECTOTYPE: U.S.A. 
Kentucky: marshes around Louisville, s.d., /7. 
A. Griswold s.n. (PH; isolectotypes, DWC, 
GH, NY, 2 sheets; lectotype here designated). 
Figure 44. 


Plants glabrous. Stems erect, well branched, 
(10-)25-60(-85) cm tall, the leaf base decurrent 
and forming ridges 0.15-0.25 mm broad, these 
sometimes minutely strigillose; aerenchyma well 
developed on lower stems when submersed. Stolons 
short, several from the erect stem base, 2.5-15 
(-22) cm long, 1-2.3 mm diam., usually seen only 
in late flowering season, frequently branching from 
the nodes; internodes short, leaves often clustered 
on stolon apex and overlapping, narrowly elliptic 
or oblanceolate, 7.8-20(-32) mm long, 2.4-8 


(-12) mm wide, glabrous, the margin entire or re- 


motely denticulate, papillose-serrulate; petioles 0— 
5 mm long. Cauline leaves very narrowly oblong- 
elliptic, 35-110 mm long, 4-10(-17) mm wide, 
the apex narrowly acute or acuminate, the margin 
entire, densely minutely papillose-serrulate except 
or the lowest Y, this usually smooth, hydathodal 
glands obscure, the base very narrowly cuneate or 
long attenuate into winged petiole 1-10 mm long. 
Stipules dark purplish, narrowly to broadly ovate, 
succulent, 0.15-0.35 mm long, 0.1-0.25 i 


Flowers many, sessile or nearly so in leaf axils, 


mm wide. 


sometimes borne almost to the base of stems; flow- 
er-subtending leaves not reduced. Sepals greenish, 
triangularly ovate, the apex elongate acuminate, 
spreading horizontally with reflexed tips, 2.5-4.5 
mm long, 1.5-3.2 mm wide, glabrous, the margin 
entire, minutely papillose-serrulate. Petals 0. An- 
thers 0.5-0.85 mm long; filaments yellowish green, 
0.7-1.5 mm long, the base dilated. Pollen grains 


262 Annals of the 
Missouri Botanical Garden 


FicunE 44. Ludwigia polycarpa (Massachusetts: Franklin Co., dit od s.n., DS).—a. Habit, upper 


portion of stem.—a’. Stem base with newly formed stolons.— b. Lea . Portion of nu are branch. —d. 
Portion of branch with mature capsules.—e. Flower bud. x Flower. F. Partly dissected flower to show stigma, 
style, and nectary disc.—g, g'. Adaxial and abaxial views of stamens.—h. Sepal.—i. Bracteole.—j. Capsule. —k. 


Cross section of capsule. 


Volume 76, Number 1 
1989 


Peng 263 
Ludwigia sect. Microcarpium 


shed as tetrads. Nectary disc yellowish green, raised 
ovary top, 1.8-3 m 
4- lobed, glabrous. Style yellowish green, 0.5-0.8 
mm long, glabrous; stigma greenish yellow, cuboid, 
0.4-0.8 mm long, distinctly 4-lobed on the apex. 
Capsules oblong-obovoid with constricted base, ob- 
scurely 4-sided with rounded corners, 4-7 mm 
long, 2.5-5 mm thick, glabrous to obscurely mi- 
nutely strigillose; pedicels 0.1-0.3 mm long. Brac- 
teoles flanking the capsules, attached 0.5-2.5(-3) 
mm from capsule base, linear-lanceolate with a 
swollen base, 3.5-6.5(-8) mm long, 0.4-1(-1.3) 


mm wide, the margin minutely papillose-serrulate. 


mm on mm across, 


Seeds light brown, narrowly oblong with curved 
ends, 0.5-0.6 mm long, 0.2-0.3 mm wide, the 
surface cells elongate parallel to the seed length. 
Self-compatible. Gametic chromosome number, n 
= 16 


Distribution. Plants of Ludwigia polycarpa 
grow in ditches; on moist prairie; on alluvial ground 
of ponds, lakes, and rivers; in marshes; in swales; 
along edges of lagoons; and in low fallow fields. 
Contrary to all other species in sect. Microcar- 
pium, L. polycarpa is centered primarily in the 
central Midwest, ranging from southcentral Mis- 
souri, eastern Kansas, eastern and northeastern 
Nebraska, northward and eastward across Iowa and 
southern Minnesota into central Wisconsin and the 
Upper Peninsula of Michigan. In the east, this 
species occurs in extreme southern Ontario and 
the eastern part of the Lower Peninsula of Mich- 
igan. Ludwigia polycarpa extends southward 
through western and southern Ohio, northcentral 
Kentucky, and western West Virginia. Scattered 
populations occur in central Virginia, southcentral 
Pennsylvania, Massachusetts, and Connecticut. A 
highly disjunct, presumably introduced population 
has been recorded from Kootenai County, Idaho 
(Fig. 45). Flowering from June to September; fruit- 
ing from July to October. 

Representative specimens exa 


ned. U.S.A 
CONNECTICUT: Hartford Co., Bissell & "Clark 245 (B, C, 
N 


n. (GH). Johnson Co., N of Homestead, 
1924, Shimek : s.n. (NY). Lee Co., ca. 1.25 mi. W-SW 
of Ft. Madison, Davidson et al. 684 (TEX). Louisa Co., 
Davidson 4035 (TEX). Palo Alto Co., 5 mi. E of Ruthven 
i S of the viaduct over Hwy. 18, Hayden 9416 
(GH, MO, NY, US). Polk Co., Van Bruggen 3152 (NY 
SMU). Poweshiek Co., rich slough beside US Hwy. 

about 1.5 mi. W of Victor, Russell 815258 (USF). Story 


“os 


Co., Ames, 1877, Arthur s.n. (P). Warren Co., 1 mi. 
NW " Indianola, Van Bruggen 3105 (?). Washinton 
Co., 3 mi. S of Crawfordsville, Wagenknecht 874 (F). 
Webster Co., Fort Dodge, 1904, Somes s.n. (NY). IDAHO: 
Kootenai Co., head of Fernan Lake, Coeur d’Alene, Rust 
1353 (NY, US). ILLINOIS: Adams Co., 


(NY, SMU). Cook Co., 
9384 (FSU, NCU, VDB). DuPage Co., W of Lacey road 
near the Milton—Lisle Township line NE of Lisle, Swink 
1828 (F). Gallatin Co., 1 mi. N of Shawneetown, Evers 
85336 (MO). Hancock Co., Augusta, 1847, Mead s.n. 
(F, MO). Henderson Co., Mississippi bottoms near Oquaw- 
(E, MO, US). Iroquois Co., 1888, 
aukegan, 

5W of 


Co. NW a Pleasant Hill, Eb 93796 (NCU). St. Clair 
Co., vicinity of Kingshighway, Neill 15378 (NCU, MO). 


1 838 
nebago Co., 1858, Bebb s.n. (GH, UPS). INDIANA: Black- 
ford Co., 2 mi. W of Mollie on S side of road, yd 325 
(MO, US). Delaware Co., along NYC RR, ca 
Yorktown, gc 18365 (SMU, uo Y Fayette 

Gardner 545 (NY). Howard Co., N 
ae EK 22 var Jackson IS abou ES 75 mi. 
Chestnut Ridge, Deam 14039 (NY, S). Lake Co., Whit- 
ing, 1893, Britton s.n. (NY). Martin Co., old river slough 
just N of White River and W of road 45, Tryon 416 
(DUKE). Newton Co., about 5.5 mi. SW of Lake Village, 
Deam 56715 (CM CH), Porter im Dune sd Lansing 
1539 (F). Posey o. in Scuffle pond, ca i. SW of 
Mt. Vernon, Kriebel 8378 (ND). Pulaski no dich on 
Jasper-Pulaski game reserve, . W Rd. 43, 1.5 mi. 


d. 10, Friesner 16176 CAS E FLAS, p Randolph 
U : 


Ro 24336 (GH 
KANSAS: Douglas Co., 4 mi. SE Lawrence, McGregor 
Ane (US, NCU, SMU). Jackson Co., marshes, Clothier 
1141 (GH, MO, NY, US). Linn Co., i 
Stephens 44334 (NCU). Miami Co., 
(CAS). KENTUCKY: Boone Co., pond e 1,000 ft. inland 
from Aurora ferry (just opposite Aurora, Indiana) along 
wy. ca. 2 mi. W of Petersburg, along Ohio River, Thieret 
50135 (VDB). Fayette Co., Lexington, Short s.n. (NY, 
) MAINE: Franklin Co., E side Pine Hill, Deerfield, 
Ahles 64607 (DS, NCU, SMU). Hampshire Co., Hock- 
anum Rd., Northampton, Ahles 86398 (GA, SP). Mid- 
dlesex Co., Winter pond, Winchester, Raven 16514 (MO). 
MICHIGAN: Genesee Co., Linden Park, 1903, Farwell s.n. 
(DS). Houghton Co., near Houghton, 1916, Dodge s.n. 
(LL). Macomb Co., wet thicket W edge of woods, frequent, 


Toledo RR between Stein and S Otter rds., 1972, 
ill & Thompson s.n. (MSU). Oakland Co., Hubbard marsh, 
1916, Chandler s.n. (MSU, US). St. E Co., near Port 
Huron, 1893, Dodge s.n. (SMU, T , TENN, S 
cola Co., Akron Tp., 1895, due s.n. (S). 

Co., Marsh Keeler, Pepoon 619 (MSU). Washtenaw Co., 

3 mi. W of Third Sister RES ee 8357 (DS, F. 
MO, NY, US ayne Co. out from Detroit, 
1851, Cooley s.n. (MSU). MINNESOTA: i Cgo Co., Wis 


264 


Annals of the 
Missouri Botanical Garden 


Distribution of Ludwigia polycarpa. 


FiGURE 45. 


Volume 76, Number 1 
1989 


Peng 265 
Ludwigia sect. Microcarpium 


lizenus 503 (MO). Hennepin Co., near Minneapolis, 1892, 
Burglehaus s.n. (MO, MSU). Houston Co., along back- 
water of the Mississippi River, 2 mi. S of La Crescent, 
Moore 26506 (CAS, EH, NCU). Mower Co., near Taopi, 
Rosendahl et al. 7261 (GH). Nicollet Co., edge of the 
Red Rock outcrops 3-4 mi. W of Courtland, 1947, Moore 
. (GH, US). Winona Co., river bank, 1897, 


mi. NW of Archie, Steyermark 60785 
(F, UPS). as Co., Little Chariton River, 4 m 
of Keytesville Steyermark 26456 (MO). Clark Co., 1892, 
Bush s.n. ( Daviess Co., Old Grand River, S of 
Packwood Lake, 4.5 mi. W of Lock Springs, Steyermark 
74203 (F). DeKalb Co., between Clarksdale and Bayfield, 
Steyermark 14943 (MO). Franklin Co., Shaw Arboretum, 
Pinetum Lake, Dille 436 (MO). Grundy Co., Oxbow Lake 
i i SW of 
Trenton, Steyermark 68498 (F). Henry Co., 
i. SE of Hartwell, Steyermark 7542 (F). Iron Co., 
Iron Mountain Lake, 1924, Kellogg s.n. (MO). Jackson 
Co., Sibley, Bush 4173 (GH, MO, NY, SMU, US). Jasper 
Co., near Waco, Palmer 18476 (NY). Johnson Co., din 
Creek, 3.5 mi. E of bip io along Hwy. 50, 5 
ermark 24543 (F, NY, TENN). Knox e bottom mcd 
along old channel of Salt River, 1.5 mi. SW of Locust 
Hill, Steyermark 70647 (F). Lewis Co., Mississippi River 
bottoms, E of Shiloh ppc along Doe Run, 7 mi. N of 
Canton, Steyermark 74081 (F). Lincoln Co., W Fork of 
Cuivre River, 5.5 mi. SW of Davis, Steyermark 72351 
(F). Linn Co., Locust Creek, Pershing State Park, 4 mi. 
SW of Laclede, Steyermark 40422 (F, MO). Livingston 
Co., Shoal Creek, 1 mi. NE of Dawn, Xo dre 6845 


(F). Macon Co., Chariton River, 5 mi. w Cam- 
bria, Steyermark 40580 (F, MO, US). Mon mtgomery Co. 
ca. 1 mi. W-SW of Wellsville, idis & Gu 11 90 (MO). 


Osage Co., along Gasconade River W side, 8 mi. SE 
of Linn, Steyermark 79318 (MO). "Rando olph Co., poe 
Spring Creek, 2 mi. S of Huntsville, AA dle mark 934 

(MO). Ray Co., 1892, Bush s.n. (MO, NY). St. eie 
Co., 3 mi. E of St. Peters, Bauer or (P). St. Louis Co., 

Mississippi River near Baden, Steyermark 8751 (MO). 
Saline Co., along road leading to bridge over Blackwater 
River, Sect. 32, 5 mi. of r 
21527 (F, MO). Schuyler Co., N side of Rt. 4, in 
of Chariton River, just inside ‘Schuyler County, just E of 


Livonia, Steyermark 70272 (F). Shelby Co., alluvial woods 
along NW side of Salt River, 3.5 mi. f Shelbina, 
Steyermark 66574 (MO). Vernon Co., Clear Chae 4- 


W of dies City, po sed. 9688 (F). 
, Ainsworth, Clements s.n. (GH, NY, US). 
, Valentine Lakes Refuge, Tolstead 411242 
. S of Shuler on Hwy. 15, ers 


ison Co.. 4 mi. 


o. 

Lucas H x^ a 1878, Sanford 
Co., ca . W of Mee Stuckey 6267 (SMU). 
PENNSYLVANIA: ere hin on . E-SE of Millers- 
burg, Berkheimer 1454 1 (CM). VIRGINIA: Buckingham 
Co., Sprouses Corner (near Dillwyn), Bartley & Pontius 


— 


539(NY). WISCONSIN: Brown Co., Fort Howard, Elmore's, 
1881, Schuette s.n. (F, 
1892, True s.n. (GH). Crawford Co., 
1861, Hale s.n.(?). Dane Co. Madison ann 

(NY). Grant Co., Wyalusing State Park, Musselman 3825 
(NCU). Kenosha Co., along Lake Michigan, just N of 
mus state line and E of RR, /ltis 20258 (MO). La 


1 (GH, MO). cue Co., 1 
nroe x 3 2] near Wyeville iab. 
» Hale s s.n. (NY, US). WEST VIR- 
EH. Bartley & Pontius 
476 (NY). CANADA. ae EL. Co., Sandwich, Ma- 
c — 44462 (CM, i. SW of Windsor, Rogers 
7 (MSU); near Amherst 1892, Macoun s.n. 
x NY, US). Kent Co., 11 W-SW of Cedar Springs, 
4 mi. S-SE of Merlin, al side 1 pe Erie, Fosberg 39627 
(C). Lambton Co., ard, Lake Huron, Macoun 
44461 (F, GH, NY}: : S.C. 188 cg Welland Co., Port 
Colborne, 1937, Allen s.n. (NCU) 


Ludwigia polycarpa is the only species in sect. - 
Microcarpium that occurs consistently north of 
37°N, in the central Midwest of the United States. 
The report of Kral (1976) on the range extension 

f L. polycarpa to Covington Co., Alabama (Kral 
40992, FLAS, GH, MO, NCU, NY, US, USF, 
VDB), was based on a pure F, stand of natural 
hybrids between L. pilosa and L. glandulosa. 

Ludwigia polycarpa is therefore effectively iso- 
lated geographically from all other species in the 
same section except L. sphaerocarpa, which has 

ide distribution throughout the Atlantic and Gulf 
coastal plains, also with scattered populations in 
the northeastern states as well as around the south- 
ern border of Lake Michigan (Fig. 45). Indeed, 
natural hybridization takes place where geograph- 
ical isolation is not maintained, as is evidenced by 
the presence of fertile hybrid populations of L. 
polycarpa X L. sphaerocarpa found in Starke 
Co., Indiana (Peng, 1988). Furthermore, sterile, 
intersectional hybrids between L. polycarpa and 
the prostrate, opposite-leaved L. palustris (sect. 
Dantia), which is very widespread throughout the 
eastern half of the United States, are found in 
Ballard Co., Kentucky, the southern border of the 
range of L. polycarpa (Peng, 1988) 

Herbarium specimens of Ludwigia polycarpa 


are rarely misidentified due partly to its geograph- 
ical isolation and partly to its distinctive morphol- 
ogy. Plants of L. polycarpa are generally uniform 
and are characterized by having densely minutely 
papillose-serrulate leaf margins, elongate-acumin- 
ate sepals with reflexed tips, long bracteoles (3.5— 
6.5(-8) mm) attached above the capsule base, and 
the oblong-obovoid capsules that frequently exhibit 
reticulate markings along the rounded corners. 
Plants of Ludwigia polycarpa produce stolons 


266 Annals of the 
Missouri Botanical Garden 


5 
D LA e,i,j S MENO 
ff li 


dd'ghk!, 3 , 


FIGURE 46. Ludwigia suffruticosa. All but a’ from Florida (Leon Co., Godfrey 53576, DUKE); a' from Florida 
(Manatee Co., Shuey 2024, FLAS). — a. Habit, erect stems arising from horizontal rhizomes. —a'. Stolon. — af = 
c. Inflorescence. —d. Flower.—d'. Flower, a sepal removed to show stamens and pistil. —e. Sepal.—f, f'. Adaxial 
and abaxial views of stamens.— g. Nectary disc and pistil. —h. Nectary disc.— i. Bracteole.— j. Flower-subtending 
leaf. —k. Capsule. —1. Cross section of capsule. 


Volume 76, Number 1 
1989 


Peng 267 
Ludwigia sect. Microcarpium 


from their lower nodes in late flowering season, 
like other species do. Their stolons, however, are 
much shorter, normally up to 15 cm, and their 
leaves, unlike those of any other species of sect. 
Microcarpium, are often densely congested and 
overlap at the apex of the sparsely leaved stolons. 
The modified stolon is comparable to the peren- 
nating turion of Epilobium (Keating et al., 1982), 
which apparently serves as an adaptation to the 
severe winter that plants of L. polycarpa have to 
endure in the northern states. 

The short stolons are occasionally found growing 
from the nodes of aerial stems that have fallen to 
the ground. Such specimens include: 


U.S.A. ILLINOIS: dde 1886, Ohlendorf s.n. 
Toe ae Smith 676 (F). MISSOURI: Saline 
Blackwater River, Sect. of Ridge Prairie. 
omar 21527 (MO). 


E 


Ludwigia suffruticosa Walter, Fl. Carol. 
90. 1788. Isnardia suffruticosa (Walter) 

Pl. 1: 251. 1891. Lud- 
wigia capitata Michaux, Fl. Bor.-Am. 1: 90. 
1803, nom. illegit., based on L. suffruticosa 
Walter. Isnardia capitata (Michaux) DC., 
Prodr. 3: 60. 1828. TYPE: U.S.A., “the Car- 
olinas," s.d., 7. Walter (type not seen, not 
located at BM). Figure 46. 


Kuntze, Rev. Gen. 


Plants bearing rhizomes and stolons. Stems erect, 
unbranched or slightly branched, (16-)30-90 cm 
tall, glabrous except for the rachis and stem base, 
these often puberulous to densely hirtellous, the 
hairs 0.2-0.6(-0.8) mm long, aerenchyma in the 
lower part very rarely seen. Rhizomes 1 -several, 
6-55 cm long, 1.25-4.5 mm thick, often branched, 
glabrous to densely hirtellous, the leaves dark pur- 
plish, minute, scalelike, oblate or suborbicular, ca. 
0.3-0.6 mm long, 0.6-0.9 mm wide, appressed. 
Stolons less commonly seen than rhizomes, 8-80 
cm long, 1.1-2.2 mm thick, branched, glabrous 
to densely hirtellous, the leaves purplish, oblong or 
oblanceolate-elliptic to spatulate, 4-35 mm long, 

.6-15 mm wide, glabrous to pilose, the petioles 
1-6 mm long. Cauline leaves green, lance-elliptic, 
lance-linear to linear, the lower ones shorter, ob- 
long-oblanceolate to oblong or oblong-lanceolate, 
25-95 mm long, (1-)3-9 mm wide, glabrous to 
+ pilose on those at stem base/peduncles, the apex 
manifestly elongate tapering into a sharp point or 
acute on lower leaves, the margin entire, hyda- 
thodal glands usually obscure, submarginal veins 
parallel to the midvein often prominent on the 
abaxial surface, the base rounded or obtuse, sessile. 


Stipules deltoid, 0.25-0.45 mm long, 0.15-0.4 


mm wide. Flowers in racemose inflorescence l- 
5(-12) cm long, 1-2 cm wide, terminal on the 
stems or branches; the subtending leaves much 
reduced and bractlike, lanceolate or lance-elliptic, 
4—12(-18) mm long, 1.35-3.5(-4) mm wide, gla- 
brous to hirtellous along margin and venation of 
the abaxial surface, especially on the lower half. 
Sepals whitish, broadly ovate or deltoid, ascending, 
2.3-3.5(-4) mm long, 2.3-3.2(-3.8) mm wide, 
glabrous, apex acuminate, margin entire, base con- 
nate 0.6-1 mm above the ovary. Petals 0. Anthers 
0.75-1(-1.3) mm long; filaments pale yellowish, 
1.2-2 mm long, distinctly dilated toward the base. 
Pollen grains shed as tetrads. Nectary disc yellow- 
ish, raised 0.5-0.6 mm on top of the ovary, 1.8- 
3.1 mm across, the lobes obscure, glabrous. Style 
pale yellowish, 0.9-1.7 mm long, glabrous; stigma 
whitish, 0.4-0.75 mm long, 0.5-0.75 mm thick, 
distinctly 4-lobed on the apex. Capsules broadly 
obpyramidal with rounded corners, sometimes sub- 
spherical, 2.5-4.25 mm long, 2.5-4.5(-5) mm 
thick, glabrous or in a few cases minutely strigillose, 
the hairs ca. 0.05-0.1 mm long, pedicels 0.5- 
1.5(-2) mm long. Bracteoles flanking capsule base 
or on the short pedicel, 3.5-5(-6) mm long, 
(1.2-)1.4-2 mm wide, glabrous to pilose without, 
glabrous within. Seeds brown, elliptic oblong, curved 
on both ends, 0.5-0.6 mm long, 0.2-0.25 
thick, surface cells nearly isodiametric. Self-com- 
patible. Gametic chromosome number, n — 16. 


mm 


Distribution. Plants of L. suffruticosa grow 
in sandy ditches, along river marshes, in meadows, 
in limestone sinks, in cypress swamps, and in moist 
pinelands. This species occurs along the Atlantic 
coast from southeastern North Carolina through 
eastern. South Carolina, southern Georgia, and 
throughout the northern three-quarters of the Flor- 
ida peninsula. To the west this species occurs 
through most of the Florida panhandle, extreme 
southeastern Alabama, and southern Georgia (Fig. 
47). Flowering from May to September; fruiting 
from June to October. 


Representative specimens examine ed. U.S.A. ALA- 
BAMA: Houston Co., onwood, 


MO ). Bay Co., ' just S of Ba ay- 
Adams 697 (DUKE, FLAS, FSU, CH. NC 
Bradford Co., Sampson City, 1946, [men dam & Arnold 
side of Cocoa, Kral 5484 


953 (C, FLAS). 
s.n. (USF). Columbia Co., Lake City, 1893, Rolfs s.n. 


268 


Annals of the 
Missouri Botanical Garden 


FIGURE 47. Distribution of Ludwigia suffruticosa. 


4 mi. I NW of Arcadia, duy 17965 
, 3.5 mi. N of Hines, , Pasture 
dl Co. near pes en Curtiss 

: 


5135 rs GH, JE. MSC, NY, US, ue adsden Co., 


(GA). De Soto Co., 
V 


n. (F LAS). uid p 
Hwy. "29 and x Hwy. 2 

Kavin. 18678 (DS). Hamilton Co., 5 mi. E of us 
1941, West & Arnold s.n. (FLAS). Hernando Co., Cooley 


et al. 7068 (NCU, USF). Highlands Co., 10 mi. S of 


Lake Placid, Brass 15203 (FLAS, GH, USF). Hillsbor- 
ough Co., 1.5 to 1.7 mi. S of FL Hwy. 674 on the E 
side of Taylor Grill Dr., Peng et al. 4327 (MO). Indian 
River Co., near Eau Gallie, Curtiss 927 (CM, F, FLAS, 
GA, MO, NCU, NY, US). Jackson Co., near Dellwood, 
along FL Hwy. 69, Mi 2759 (FLAS). Lake Co., vicinity 
of Eustis, Nash 950 (F, GH, MO, MSU, NY, US 
Co., Hitchcock DE GH, MO, NY, US). Leon Co., 
mi. S of Tallahassee, Godfrey 53576 (DUKE, FLAS, NY. 
SMU, USF, VDB). Levy Co., 5 mi. SE of Lebanon Station, 
Kral 7807 (GA). Madison Cos 4 mi. N of Greenville on 
221, Ramsey 104 (FSU, NCU, US). Manatee 
Co., 6.7 mi. W of jct. of FL Hwys. 64 and 675, E of 


Due Dress 
of 


Bradenton, Perdue 1779 (C, FSU, GA, GH, LL, NCU, 
SMU, Marion Co., Lake Kerr, 1935, West & 
Arnold s.n. (FLAS). Martin Co., Jonathan Dickinson State 
ark, w of Pinegrove Campground, Popenoe 351 (FTG). 
Nassau Co., ca. 7 mi. SE of Yulee, Kral 18629 (VDB). 
Obss lis Co., US Hwy. 441, 20 mi. E of Kenansville, 
l mi. S bj R Co. line, Clark 3477 (Duke). Orange 
Co., of Oakland, Kral 7735 (FLAS, GA, GH, 
US, USF). Osceola Co., side he main road leading S 
through Poinciana S of US h 17 & 92, Wunderlin 
& Shuey 5779 (TENN, USF), "Palm Beach Co., 2.0 m 
W of Lantana, Howard 12960 (C, DS, DUKE, FLAS. 
GA, GH, MO, MSC be AE 5. SMU, TENN, TEX, U, 
US, W). Pasco Co., B i. E of Zephyrhills, Kral 7264 
(GH, NY, USF ). dn Co. 23 L Power 
Hwy.19, 1 m 
(TENN, USF). Polk Co., SE p Hes 
of Lake Wales, off FL Hwy DS, 
USF). Putnam Co., Martin & ashe 110(FLAS, USF). 
St. Johns Co., Palm Valley, 1951, Buker s.n. (CM). 
Sarasota Co., 7 mi. N of S entrance 
region, Dodson 4515 (USF). ety Co., 
Schallert 15229 (AAU, JE, U, WAG). So 
mi. SW of Tarrytown, Kral 7925 (FLAS, CH, USF). 


Volume 76, Number 1 
1989 


Peng 269 
Ludwigia sect. Microcarpium 


wannee Co., 4 mi. E of Wellborn, Wiegand & Man- 

i. N of Shady Grove, 

, 1 mi. E of Worth 

Springs, Murill 775(?). Volusia io. 3 mi. N-NW of 

DeLeon Eo Kral 1699 (CA, GH, UsF. VDB). Wa- 
, 0.5 mi. 


kulla N of cT pe on US Hwy. 
98, Kral 3027(?). Washington Co., i. S of Wausau, 
Webster & Wilbur 3622 (GH, NY, SM U, US). GEORGIA: 


Baker Co., ca. 8 mi. SE of Milford, Kral 20493 (VDB). 
Broo y^ Co. Ns 5 mi. SE of Quitman, Faircloth 3338 
(GA, NCU). Charlton Co., between Folkston and 
T na Correll 5517 (DUKE, FSU, GA, TENN). Cook 
. SE of Adel, Faircloth 2927 (GA, NCU). 
hola 0 E 5 mi. S of Stockton on US Hwy. 129, 
Norsworthy 162 (MO, NCU). Effingham Co., near Lu- 
ciene Bay, Eyles 6376 (GA). Glynn Co., St. Simons Island, 
& Thorne s.n. (TENN). Grady Co., Susina 


Millen, Pyron & McVau h 919(GA). Lee Co., ca. 7 mi. 


O, 
NY, SMU, US). NE df Hinesville. 
a i. E 
SE of Long-Tattnall county line on GA Hwy. 
RJ with GÀ Hwy. 261, Faust & Jones 19981 (FSU, GA, 
VDB). Lowndes Co., 9.5 mi. SE of Valdosta, intersection 
of Southern RR and Howell Hwy., Faircloth 2887 (GA, 
NCU). McIntosh Co., 1.8 mi. S-SW of N tip of Sapelo 
Island, Duncan 20406 (CAS, DUKE, F, FLAS, GH, 
MISS, NCU, SMU, TENN, TEX, US, USF). Mitchell 
Co., Duncan 6682 (GA). Screven Co., W of Blue Springs, 
Duncan 5581 (GA, TENN). Sumter Co., S of Leslie, 
Harper 484 (F, GH, MO, MSC, NCU, NY, S, US). 
ayne Co., SE of Lindy's Bluff NE of Jesup, Duncan 
3759 (FLAS, GA, MISS, 2 US, USF). NORTH vaa 
LINA: Brunswick Co., ca. 2 mi. W of NC Hwy. 13 
Co. Rd. 1536, Boiling i. Bradley & Pind 
3308 (BOON, C, ECUH, MEX, MISS, S, TEX, UNA, 
U, US, USCH, WCUH). Bladen Co., 2.5 mi. E of Eliz- 
abethtown, Godfrey & Fox 49485 (DUKE, FSU, MO, 
NCSC, NY). New Hanover Co., Fort Fischer on the lower 
Cape Fear Peninsula, Godfrey 4698 (GH, NLSC, NY, 
US). Oran ge Co., Taft, Schallert 15229 (M). SOUTH CARO- 
Bay 


vannah River P 


Ahles & Bell 15855 (CM, NCU). Bamb 
of jct. of Co. Hwy. 59 and US Hwy. 601 o y. 
59, S.SE of Bamba, Ahles & Haesloop 30589 (NCU). 
Beaufort Co., Bluffton, 1887, Mellichamp s.n. (US). 
] mi. N of Russelville, Martin 48 (DUKE, 


Q 
z 
p 
t 
z 
z 
Es 
2 
o 
a 
ze 
lal 
a 
> 
9 
3 
O 
o 


frey & Tryon 1565 (GA, N 
W of Davis ues cid PRG LAS NCU, SMU). 
Colleton of Walterboro, Kral 19202 (VDB). 
Darlington a Hirovills 1908, Coker s.n. (NCU, NY). 

eorgetown Co., 9 mi. N of Georgetown, Godfrey & 


Jasper Co., NW of Tillman, Ahles & Williamson 54838 
(N E Lexington Co., Columbia, Godfrey & is 1241 
(GH, US). Sumter Co. . 1884, Smith s.n. (F, GH, US). 
Williamsburg Co., 6 mi. N of Kingstree, Wiegand & 
Manning 2200 (CH). 


Ludwigia suffruticosa is a very distinct species 
that has many features unique within sect. Micro- 
carpium. Its leaves are rounded to obtuse at the 


base and clearly sessile. It has a very compact, 
racemose inflorescence at the very top of each 
erect stem, as is suggested by the name “Ludwigia 
capitata," an illegitimate synonym of L. suffru- 
ticosa Walter, coined by Michaux (1803). The 
stems are usually single or occasionally with one 
or two or more branches arising directly beneath 
the main inflorescence. These branches always rise 
above the main inflorescence and are each ter- 
minated by a smaller compact raceme. Further- 
more, each flower is usually subtended by a re- 
duced, bractlike leaf in addition to the bracteoles 
that flank the ovary. It is also the only species in 
sect. Microcarpium that perennates mainly by un- 
derground rhizomes. Stolons that float or send roots 
into mud, prevalent in many other species, are not 
common in L. suffruticosa. It is especially peculiar 
in having variously pubescent inflorescences and 
lower stems, stolons, and underground rhizomes 
but is otherwise completely glabrous in the mid- 
dle—see below. 

The flowers of this species are characteristically 
very compactly arranged into a raceme up to 5 
cm long. The following specimens examined, how- 
ever, have less compact and longer inflorescences 
but are otherwise typical of Ludwigia suffruticosa: 


Caloosa Forest/Expt. 


U.S.A. FLORIDA: Charlotte Co., 
o q inflorescences 7 


no 
rescenced individuals (same oliva Dille 4 
GEORGIA: Wayne Co., in very sandy soil SE of T Lindy’ 
bluff, just NE of Jesup, Dunkan 3759 (GH, inflorescences 
8 cm long; NCU, inflorescences 10.5 cm long). 


The distribution pattern of pubescence in plants 
of Ludwigia suffruticosa is peculiar and variable. 
The plants range from being completely glabrous 
to densely, minutely hirtellous with multicellular 
hairs up to 0.6(—0.8) mm long on rhizomes, stolons, 
stolon leaves, stem base, and basal leaves of the 
lower plant body and peduncles, pedicels, flower- 
subtending leaves, and bracteoles of the upper part 
but are completely glabrous in the middle part of 
the plant body. The capsules are usually completely 
glabrous. In the following specimens, however, erect, 
microscopic protuberances ca. 0.02-0.1 mm long 
are found evenly but not densely distributed on the 
fruit wall: 


U.S.A. FLORIDA: Gilchrist Co., in swamps 4 mi. E of 
Trenton, 1940, West & Arnold s.n. (FLAS). Hillsborough 
Co., Tampa, Gates Dr. in sandy field W of Gates Apts., 
Dille & Dille 424 (MO); in wet sim pong SR-581, 
ca. 5 mi. N of Tampa, Funk 72; Gre Kuczynski 
48; Robbins 130; Rose 11; er 10 E USF). Leon 
Co., bordering Lake Jackson on NW side, just N of route 


270 


Annals of the 
Missouri Botanical Garden 


FicunE 48. 


Stem hs with stolons. — 


27, NW of Tallahassee, Anderson 4497 (GA, MO); on 
FL 363, 3 mi. N of Woodville, Dille & Dille 421 (MO); 
Lake Jackson, adjacent to US 27, Faircloth 246 (MO, 
NCU); shores of small pond, 6 mi. S of Tallahassee, 
Godfrey 53576 (DUKE, FLAS, NCU, NY, SMU, USF, 
VDB); wet sands, exposed shores of sinkhole pond, by 


Ludwigia lanc ns prs Highlands Co., Peng et al. 4193, MO).— a. Habit, erect stem.—a’. 
b, b'. Flower. — c. Cross section of capsule.— d. Dehiscent capsule. 


Spring Hill Rd., S of uses cua S icd 76066 on 
along US 319, ca. 5 mi. S of T assee, Hende 

63-1683 (TEX); margin e a al lake along FL 271, 
ca. 5 € S of Tallahassee, Henderson 67-1455 (VDB); 
near Lake Jackson, 8 mi. NW of Tallahassee, 1940, 
Hocking s.n. (FLAS). Orange Co., Godfrey 57330 (FSU). 


Volume 76, Number 1 
1989 


Peng 
Ludwigia sect. Microcarpium 


Polk Co., margin of a small lake, SE of Hesperides, ca. 
8 mi. E of Lake Wales, off number 60, Lakela 24806 
(DS). 

In a specimen collected from Florida (Leon Co., 
4 mi. S of Tallahassee, on Rt. 369, 1950, Sargent 
s.n. (GA)) the capsule wall is minutely pilose with 
hairs ca. 0.4 mm long. 

The sepals of Ludwigia suffruticosa are nor- 
mally glabrous, but in the following specimen the 
sepals are minutely pilose: 


U.S.A. FLORIDA: Duval Co., vicinity of Mayport & 
Jacksonville, 1870-1876, Keeler s.n. (NY). 


Ludwigia suffruticosa, L. alata, L. pilosa, and 
L. sphaerocarpa are the species in sect. Micro- 
carpium that do not undergo mechanical self-pol- 
lination. With its whitish creamy sepals, which are 
very showy in the dense flower aggregates, the 
cross-pollinating L. suffruticosa successfully at- 
tracts many insects, mostly bumblebees, honey- 
bees, and wasps. It occurs with at least four species 
of Ludwigia (L. alata, L. linearis, L. lanceolata, 
and L. pilosa) in the field, hybridizing with all but 
L. linearis. These results are mirrored in the bio- 
systematic study (Peng, 1988): artificial hybrids 
between L. linearis and L. suffruticosa are weak 
and inviable, whereas the other combinations that 
occur in nature have been synthesized successfully 
and were very vigorous in the experimental green- 
house. 

Remarks on natural hybrids between Ludwigia 
suffruticosa and either L. alata or L. lanceolata 
are provided in the accounts of L. alata and L. 
lanceolata. Natural hybrid populations of L. suf- 
fruticosa X L. pilosa have previously been given 
a varietal status within L. suffruticosa (as L. cap- 
itata Michaux 8 pubens) by Torrey & Gray (1840), 
which was subsequently and incorrectly considered 
to be a synonym of L. pilosa by Munz (1944, 
1965). The hybrids are densely hirtellous through- 
out, but the hairs are not as long as those of L. 
pilosa, and the sepals are neither cuspidate nor 
elongate-acuminate, but rather acuminate. Fur- 
ther, their stems are not well branched, their leaves 
are often long and narrow, their flowers are in 
more or less congested racemes on the upper part 
of the stem or branch, and the leaves that subtend 
their flowers are slightly reduced and bractlike. 
The combination of these characteristics clearly 
suggests that such plants represent hybrid popu- 
lations of L. suffruticosa X L. pilosa. The possi- 
bility that L. ravenii, the only species other than 
L. pilosa that is consistently densely hirtellous, was 
involved in their parentage is reduced because it 
is rare and not known to occur with L. suffruticosa. 


Furthermore, the seed surface of the hybrid plants 
consists of subisodiametric cells, a characteristic of 
both L. suffruticosa and L. pilosa, but not of L. 
ravenii. 


7. px lanceolata Elliott, Sketch Bot. S.C 
& Ga. 213. 1817. Isnardia ancla 
(Elliott) Bo Prodr. 3:61. 1828. TYPE: U.S.A. 
Georgia: in swamps, s.d., J. Leconte s.n. 
(holotype, CHARL; photographs, GH, MO; 
isotypes, P, PH). Figure 48. 


Plants glabrous. Stems often brownish purple, 
erect, leafless below, branched above, 45-100 cm 
tall; aerenchyma well developed in lower stems 
when submersed in water; leaf bases slightly de- 
current along the stems. Stolons up to 40 cm long, 
2 mm thick, the leaves remote, 1-3 cm apart, 
purplish, rotund or elliptic to broadly elliptic, 5— 
27 mm long, 6-12 mm wide, apex rounded, obtuse 
or acute, base attenuate into winged petioles 2— 
10 mm long. Cauline leaves dark green, the lower 
ones sometimes purplish, leaves on the main stem 
elliptic, oblanceolate or narrowly oblanceolate, those 
on the branches usually much reduced, ranging 
from narrowly elliptic to very narrowly elliptic or 
linear; in general, leaves 20-75 mm long, 2-7.5 
(-14) mm wide, apex acute to narrowly acute, 
margin entire with minute hydathodal glands, base 
narrowly cuneate or sometimes attenuate into a 
winged petiole up to 5 mm long. Stipules dark 
reddish purple, ovate to very widely ovate, suc- 
culent, 0.2-0.45 mm long, 0.15-0.25 mm wide. 
Flowers in upper leaf axils. Sepals greenish, shal- 
lowly triangular or deltate, ascending, 1 < 
long, 1.8-3.3 mm wide, glabrous, apex acute to 
acuminate, minutely papillose along the margins. 
Petals 0. Anthers 0.4-0.6(-0.75) mm long; fila- 
ments nearly translucent, 1-1.4 mm long, dilated 
toward the base. Pollen grains shed as tetrads. 
Nectary disc yellowish green, raised 0.4-0.6 mm 
on ovary apex, 1.75-2.6 mm across, 4-lobed, gla- 
brous. Style yellowish green, 0.5-0.7 mm long, 
glabrous; stigma yellowish, subglobose, 0.3-0.5 mm 
long, 0.7- Capsules obpyramidal, 
winged on the corners, 3. 5-5 mm long, 2.5-4.5 
mm thick (as measured from wing to wing), gla- 
brous, occasionally minutely strigillose, the wings 
0.3-0.7 mm wide, usually minutely strigillose; ped- 
icels up to 0.5 mm long. Bracteoles located at or 


m thic 


slightly above capsule base and on 2 sides, variable 
in shape and size even in the same plant, ovate- 
elliptic to very narrowly elliptic, 1.5-4.3 mm long, 
.4-1.4 mm wide, margin minutely papillose, often 
with a distinct, rounded, swollen tissue 0.5-1 mm 


o 


272 Annals of the 
esa Botanical Garden 
i ON bus LÀ. Vy 
Y A 
Y f . ( 
a. = E 
EE 
e 
= A dae EN e. d 
i 2 x 
o ES : 
à p ur 
$0 ^ - me 400 5004 m 
FIGURE 49. Distribution of Ludwigia lanceolata. 


across below the point of attachment. Seeds light 
brown, narrowly oblong with constricted ends, 0.65- 
0.75 mm long, 0.25-0.3 mm wide, the surface 
cells nearly isodiametric. Self-compatible. Gametic 
chromosome number, n — 16. 


Distribution. Plants of Ludwigia lanceolata 
grow in ditches, low meadows, cypress swamps, 
moist pinelands, edges of pocosins, and on sandy 
peaty soil. This is a fairly uncommon species, with 
scattered populations occurring along the Atlantic 
coast of southern North Carolina, South Carolina, 
eastern and southern Georgia, and the Florida pen- 
insula. This species reaches its western limit in the 
central panhandle of Florida (Fig. 49). Flowering 
from June to October; fruiting from June to No- 
vember. 


Representative specimens examined. U.S.A. FLORI- 
DA: Brevard Co., Okeechobee region, Fredholm 5987 
(GH, MO, NY, US). Columbia Co., Lake City, Vash 2502 
(E, F, GH, MO, MSU, NY, US). De Soto Co., 5 mi. N 


of Arcadia, 1938, West s.n. ri Franklin Co., Apa- 
lachicola, pp 4177 (GH, MO, NY, US, W). Gil- 
christ Co., . E of Trenton, ipn West & año 
s.n. (FLAS). Hamilton Co., Cypress Swamp, 
Jasper, 1941, West & Arnold s.n. (FLAS). Highlands 
Co., on Co. Rd. S-634, of US Hwys. 27 & pe 
Peng et al. 4183 (MO). Hillzborough Co., 1.5 to 1.7 m 
S of Hwy. 674, on E side of Taylor Gill Dr. ., Peng et al 
4326 (MO). Jefferson Co., vicinity of Lloyd, Godfrey & 
Houk 61411 (FSU). Madison Co., 8 mi. W of Greenville, 
Kral 3741 (FSU, NY, USF). Marion Co., W of Lake 
E Martin £ Cooper 877 (FLAS). Polk Co., 4.5 
. E of Haines City, on FL 580, 1963, Conard. s.n. 
(G i. Putnam Co., Fowlers Lake, 1935, Weber s.n. 
(FLAS). St. Johns Co., 5 mi. W of St. Augustine, Godfrey 
& Reinert 61 185(FSU). Taylor pu 1.2 mi. SE of Salem, 
pa gA 64738 (DS, FSU, LL, NCU). Volusia Co., ca. 
cae S jet. state 44 & 415, Kral 18435 (VDB). Wakulla 
, Cow mp, Apalachicola National Forest, NW o 
Crono, Lind 64876 (FSU). GEORGIA: Glynn 
Co., St. Sim ., Ford & Thorne 2788 (TENN). Long 
Co., 5 mi. W ol S 301, on Hughes River Rd., Bozeman 
& Radford 2042 (NCU). Lowndes Co., pine- -barren pond 
S of Melrose, Harper 1605 (E, F, GH, MO, MSU, NCU, 
NY, US). McIntosh Co., 1.25 mi. N of Fort Barrington, 


Volume 76, Number 1 
1989 


Peng 273 
Ludwigia sect. Microcarpium 


TABLE 5. Comparison of Ludwigia alata with L. lanceolata. 
Characters L. alata L. lanceolata 

Stems Often winged, sometimes obscurely so Nearly smooth 
Sepal 

Color Whitish adaxially Greenish adaxially 

Length Subequal to capsule Shorter than capsule 
Anther length (mm) 0.55-0.9 0.4-0.6(-0.75) 
Filament length (mm) EISIN 1-1.4 
Pollen grains shed Singly In tetrads 

ectary disc iis (mm) 2-3: 1.75-2.6 

Style length Rh 0.8-1.3 0.5-0.7 
Breeding system Obligate outcrossing Modally autogamous 
2 M sul bun the Bulging out longitudinally in the central part Smoot 
Seed length /width ratio a. Ca. 2.4-2.8 
Surface cells Columnar Suborbicular 
Chromosome number n= n — 16 


x Road, Bozeman 2176 ae GA, NCU). NORTH 
CAROLINA: New Hanover Co. Wikuington, 1867, 
Canby s.n. (NY). SOUTH CAROLINA: Raus Co., Bluffton, 
Mellichainp s.n. ( 


Ludwigia lanceolata is interfertile with all sev- 
en other tetraploid members of sect. Microcar- 
pium. However, it is most similar to the hexaploid 
L. alata in general aspects and fruit morphology, 
and is suspected to have been one of the parents 
of that species (Peng, 1988). For a detailed dis- 
cussion of these two species, see remarks under L. 
alata and Table 5. 

Ludwigia lanceolata has been found growing 
with L. pilosa, L. suffruticosa, both tetraploids, 
and with the diploid L. linearis. Ludwigia alata 
and L. lanceolata have not been found growing 
together, although they have overlapping ranges 
in the eastern and central Florida panhandle. It is 
worth noting that, although both species occur in 
generalized, wet habitats, such as the borders of 

itches, ponds, and swamps, L. lanceolata, unlike 
L. alata, is not found in the limestone prairies and 
brackish marshes that prevail in the Everglades 
region of southern Florida. No hybrids have been 
observed, but they would be very difficult to detect 
without measuring pollen stainability or determin- 
ing chromosome numbers. 

Ludwigia lanceolata intergrades with L. pilo- 
sa, and fertile hybrids between them are frequent 
in nature. Such hybrids were named L. simulata 
by Small (1903). The type at NY (West Florida, 
Biltmore Herbarium) shows a combination of 
densely strigillose hairs and four-angled or barely 
winged capsules; it apparently arose following hy- 
bridization between a parent with strongly winged 
capsules (L. alata or L. lanceolata) and one that 


was pubescent (L. pilosa, L. ravenii, or L. sphae- 

rocarpa). The isodiametric seed surface cells, com- 

bined with the fact that these plants shed their 

pollen grains as tight tetrads clearly suggest L. 

lanceolata and L. pilosa as the parents. Two other 

collections identified as this hybrid are also from 
lorida 


U.S.A. FLORIDA: Franklin Co., Apalachicola, Chapman 
s.n. (F, US). Highlands Co., Bar Point, Lake Childs, 
Brass 15532 (GH, US). 


Ludwigia lanceolata also intergrades with L. 
suffruticosa. For example, putative parents and 
introgressed hybrid populations were found in the 
same sandy ditch by a prairie in Hillsborough Co., 
Florida (Peng 4324, 4328, MO). The leaves of 
the hybrids were narrow like those of L. suffruti- 
cosa, but the stems were well branched above the 
middle. The flowers are produced in somewhat 
congested racemes at the top of each branch, and 
the capsules were obpyramidal and weakly four- 
angled. Peng 4324 had 85% stainable pollen and 


formed 16 bivalents in meiosis. 


8. Ludwigia alata Elliott, Sketch Bot. S. C. & 
Ga. 1: 212. 1817. Isnardia alata (Elliott) 
DC., Prodr. 3: 61. 1828. LECTOTYPE: U.S.A. 
South Carolina: Charleston Co., found in damp 
places on Sullivan's Island, in the wooded part 
of the island, July-Sep., s.d., S. Elliott s.n. 
(PH; no authentic material found in Elliott’s 
herbarium at CHARL, so sheet from PH se- 
lected as lectotype). Figure 50. 


Plants glabrous. Stems erect or somewhat 
sprawling, 40-1 20(-160) cm tall, branched above, 


aerenchyma distinct in lower parts when sub- 


Annals of the 
Missouri Botanical Garden 


Ficure 50. 
stolon. —b. Flower.—c. 
f. Cross section of stem. 


mersed, slightly to distinctly winged, the wings up 
to 1.8 mm broad. Stolons several from lower nodes, 
often seen in the flowering season, 8-65(-95) cm 
long, 0.7-2.5 mm diam., the leaves often purplish, 
especially on the abaxial surface, variable in shape, 
ranging from orbicular, rotund, oblanceolate to 


, 


Ludwigia alata (Florida: Martin Co., Peng et al. 4203, MO). —a. Habit, erect stem.—a'. 
Capsule. — d. Cross section of mature capsule. —e. Branch, showing decurrent leaf base. — 


Basal 


(broadly) elliptic, 4-26 mm long, 4-15 mm wide, 
base attenuate into petioles 1.5- 10 mm long. Cau- 
line leaves lance-elliptic, very narrowly elliptic to 
linear (those of lower stems sometimes oblanceolate 
or oblance-elliptic), 18-100 mm long, 2-12.5 
(720) mm wide, those on the branches that subtend 


Volume 76, Number 1 
1989 


Peng 275 
Ludwigia sect. Microcarpium 


flowers often 2-4 times smaller than those on the 
main stem, the apex acute to narrowly acute, mar- 
gin with remote but pronounced hydathodal glands, 
very occasionally minutely papillose-serrulate on 
the upper margin, base narrowly cuneate or nar- 
rowly acute, sessile or with petioles up to 3 mm 
long. Stipules dark brownish purple, triangular- 
ovate or lanceolate, succulent, 0.2-0 m long, 
0.1-0.3 mm wide. Flowers in upper leaf axils, 
sometimes rather congested and in an inflores- 
cencelike arrangement. Sepals creamy white with- 
in, the apex pale green, broadly ovate-deltate and 
boat-shaped, flaring outward, apex acute or acu- 
minate, 2-4 mm long, 1.6-4 mm wide, margin 
smooth or minutely papillose-serrulate. Petals O. 
Anthers 0.55-0.9 mm long; filaments nearly trans- 
lucent, 1.1-1.7 mm long, slightly dilated toward 
the base. Pollen grains shed singly. Nectary disc 
bright yellow, square with rounded corners, raised 
—0.8 mm on top of the ovary, 2-3.3 mm across, 
prominently 4-lobed, glabrous. Style pale green, 
-1.3 mm long, glabrous; stigma light yellowish, 
subglobose, 0.25-0.75 mm across, shallowly 
4-lobed on top. Capsules glabrous, obpyramidal, 
winged on the corners, 3-5 mm long, 2.8-4.5 mm 
wide (including both wings), the wings ca. 0.5-0.9 
mm broad, smooth or minutely papillose, central 
portion on each side of the wall often somewhat 
bulging longitudinally, sessile or with pedicels up 
.8 mm long. Bracteoles flanking capsule base, 
lance-elliptic or narrowly so, 2.4-4.7 mm long, 
-1.5 mm wide, glabrous, margins minutely pa- 
pillose or smooth. Seeds light brown, ellipsoid, 
slightly curved on both ends, 0.6-0.7 mm long, 
0.3-0.35 mm wide, the surface cells elongate 
transversely to the seed length. Self-compatible. 
Gametic chromosome number, n = 24. 


Distribution. The habitats of L. alata include 
ditches; shores of ponds and lagoons; peaty or 
sandy swales; open cypress swamps; sandy borrow 
pits in open pine woods; swampy, flat outcrops of 
oolitic rocks; wet savannas; tidal flats; brackish 
marshes; sandy beach strands; and hammocks. This 
species occurs along the eastern Coastal Plain from 
southeastern Virginia through eastern North Car- 
olina, with sporadic populations in eastern South 
Carolina and Georgia; it reaches its southern limit 
at localities scattered throughout the Florida pen- 
insula. To the west this species extends along the 
Gulf Coast through the Florida panhandle into 
southern Alabama, Mississippi, and extreme east- 
ern Louisiana (Fig. 38). Proctor (1982) reported 
disjunct populations from Jamaica. Flowering from 
June to October; fruiting from July to November. 


Representative specimens examined. U.S.A. ALA- 
BAMA: Baldwin Co., ca. 7 mi. W of Gulf LE Kral 
32612 (MO, SMU, VDB). FLORIDA: Bay Co., ca. 10 mi. 

-NW of Mexico, Kral 22985 (VDB). Bere. Das, SW 
shore of Lake Poinsett, 0.5 mi. N of St. Johns River ‘inlet. 
Kral 5335 (DS, FSU, NCU, NO, SMU). Broward Co., 
near E gate of Alligator Alley, Lakela & Meaghen 31895 
(NCU, NY, USF). Citrus Co., Chassahowitzka, Genelle 
& Fleming 1551 (GA, USF). Collier Co., 4.8 mi. W of 
Monroe Station on N side of US Hwy. 41, Peng et al. 
4242 (MO). Dade Co., 2.5 mi. W of Pinelands, on main 
road of Everglades National Park, Sot Finelands and 


Pa-Hay- S E ie ¡e et al. 5 (MO). Dixie 
Co., 3m Hines, Kral & Rd 3040 (FSU, 
NY, USF). 


Duval Co., near Jacksonville, Curtiss 929 (C, 
F, FI, FLAS, GA, GH, MO, U, , P, US). Escambia 
Co., inland from Gulf Beach, SW Pensacola, Kral 17602 
(VDB). Franklin Co., 38.8 mi. W of jct. (one that is closer 


i o 
Godfrey & Henderson 62980 (FSU, LL). Hendry Co., 
Atwater M-74 (FLAS). Hernando Co., Weekiwachee Riv- 
er, below the Springs, Godfrey & Hawk 63101 (FSU). 
Hillsborough Co., Tampa, 1876, Garber s.n. (F, US). Lee 
Co., Myers, Hitchcock 113 (F, GH, MO, NY, US). Levy 
Co., near Yankeetown, Cooley et al. 6971 (FSU, NCU, 
USF, VDB). Manatee Co., north fork of Manatee River, 
S of State Rd. 62 and SW of Duette, Shuey 1768 (USF). 
Marion Co., Rainbow River, between Dunellon and Rain- 
bow Springs, Godfrey 63112 (FSU, MO). Martin Co., 


mi. 
Largo, Ferborgh & Brockman 287 (FSU). Okaloosa Co., 
Santa Rosa Island, W of Destin, McDaniel 4978 (FSU). 

, Killarney, 1889, Veslulund s.n. (S). Palm 
, SW of der. E of Loxahatchee National 
Wildlife Refuge, ca. 10 mi. W of Gulf Stream, 1970, 
Poltevint s.n. (FLAS). Pinellas Co., Clearwater, Rolfs 
592 (F). Polk Co., near N end of Lake Pierce, 1963, 
Conard s.n. (FLAS). Putnam Co., 0.5 mi. N of Welaka, 
1940, Laessle s.n. (FLAS). Santa Rosa Co., Blackwater 
River, vicinity of Milton, Godfrey 7547 1 (FSU). a 

Co., 2 mi. E of G Geneva, Kral 50914A (FSU, SMU). Tayl 
Co., ca. 20 mi. NW of Cross City, Godfrey & Houl 
aa (DUKE, FSU, LL, MSU, NCU, SMU). Volusia 

4 mi. S of jet. FL Hwys. 44 and 415, Kral 18436 

(VDB). Wakulla el St. Marks Wildlife Refuge, Godfrey 

bi FSU, CA, GH, NY, S, SMU, TENN, UNA, 
USF). cEoRGIA: Camden Co., White Oak River, 1902, 
Harper s. n. (E, 


- 
p 
3 
Sed 
"PP 
e 
B. 
e 
e, 


.25 m 

and Mill Bayou, Darwin & Sundell 

(NO). wisis Hancock Co., Kiln off MS 

95 (NCU). Harrison a Ship Island, 
US). 


Long Beach, Godfrey 48436 (FSU). Camden Co, E side 
of Pasquotank River on US Hwy. 158, E of Elizabeth 
City, Bozeman & Radford 11531 (C, CM, FLAS, NO, 
SMU, TENN, TEX, U, UNA, USCH, WCUH, East Car- 
olina Univ. herbarium). Carteret Co., SE of Mullet Pond 
on Shackleford Banks, Anderson 408 (DUKE, SMU). 
Craven Co., New Bern, Kearney 1961 (US). Currituck 
Co., Northwest River, near Moyock, Blomquist 14379 
(DUKE). Dare Co., 6 mi. S of Buxton, Radford et al. 


276 


Annals of the 
Missouri Botanical Garden 


7593 (GH, NCU). Gates Co., near Sunbury, s.d., Le Conte 
s.n. (NY). Hyde Co., Ocracoke Island, Kearney 2325 
(US). New Hanover Co., Wrightsville Sound, 1963, Ahles 
& McCrary s.n. (NCU). Onslow Co., Bear Island or 
Hammock Beach State Park, W quarter of island, Wilbur 
9576 (DUKE). SOUTH CAROLINA: Charleston Co., Santee 
River, N of McClellanville, Godfrey & Tryon 718 (GH). 
A N & US Hwy. 17, 
Raven 18719 (DS, NCU). Jasper Co., 0.4 mi. SW of 


VIRGINIA: Norfolk Co., Northwest River below Northwest, 
Fernald & Long 14965 (C, GH, S). Princess Anne Co., 
along Blackwater River, SW of Pungo Ferry, Fernald & 
Long 13981 (GA, GH, MO, NY, SMU, TENN, TEX, 
US). Jamaica: Hanover Co., in the Great Morass, ca. 1.9 
mi. SW of Logwood, Proctor d (FTG, GH, IJ, MO). 
Westmoreland Co. . N-NE of Negril, inland 
from milepost 23, Proctor 37733 (F, FTG, IJ). 


Ludwigia alata has been confused with L. lan- 
ceolata (Long & Lakela, 1976; Raven & Tai, 
1979; Godfrey & Wooten, 1981; Wunderlin, 
1982; Clewell, 1985), owing to their similarity in 
being glabrous, apetalous, and most notably, in 
having winged, obpyramidal fruits, a character not 
shared by other species of sect. Microcarpium. 
Raven & Tai (1979) reported the chromosome 
numbers of L. alata. They reported tetraploid 
counts of n — 16 and hexaploid counts of n — 2 
as a result of not differentiating L. lanceolata from 
L. alata. It has now become clear that L. alata 
is a hexaploid and L. lanceolata is a tetraploid 
(Peng, 1988). In my biosystematic study (1988), I 
reported that reciprocal artificial hybrids between 
the two species exhibit a modal meiotic configu- 
ration of 16 bivalents and 8 univalents, which 
suggests that they have two genomes in common. 
Similar results were obtained when the hexaploid 
L. alata was crossed with any of the other tetra- 
ploid species of sect. Microcarpium. Cytogenetic 
and morphological data suggest that L. alata may 
have originated following hybridization between the 
tetraploid L. lanceolata and the diploid L. micro- 
carpa or populations ancestral to them (Peng, 
1988). 

Despite their superficial resemblance, Ludwigia 
alata differs sharply from L. lanceolata in that 
the pollen grains are shed singly in L. alata and 
as tetrads in L. lanceolata (Praglowski et al., 1983), 
and the seed surface cells are columnar in £. alata 
vs. suborbicular in L. lanceolata. Other characters 
used by monographers to differentiate the two 
species include seed shape (ovoid in L. alata vs. 
cylindric in L. lanceolata) and sepal shape and 
size (slightly longer in length than width and nearly 
as long as the capsule in L. alata vs. about as long 
as wide, shorter than the capsule in L. lanceolata) 
(Small, 1933; Munz, 1944, 1965). These char- 


acters, however, are not always clear and reliable. 


The leaf base is usually decurrent in Ludwigia 
alata, resulting in the winged stem. The wings are 
up to 1.8 mm wide in extreme cases. In L. lan- 
ceolata, by contrast, the stems are nearly smooth. 
However, the character of the winged stem in L. 
alata is too variable to be diagnostic. 

Ludwigia alata has, like all other tetraploids of 
the section, lost its petals in the course of evolution. 
In a specimen collected from Duval Co., Florida 
(Creager 639, NCU), one vestigial petal is present 
on a single flower. Progeny of Peng 4205 collected 
from Dade Co., Florida, exhibited minute, vestigial 
petals on some flowers, too, in greenhouse culti- 
vation. Although normally lacking petals, L. alata 
has whitish sepals that flare outward. This char- 
acter, along with the fact that the distance between 
the androecium and the gynoecium is such that 
mechanical self-pollination cannot usually occur, 
indicates that L. alata is an obligate cross-polli- 
nator. By contrast, L. lanceolata has greenish, 
ascending sepals and is modally self-pollinating. 
Ludwigia alata and L. lanceolata are compared 
in Table 5. 

Ludwigia alata occurs with L. curtissii, L. 
linifolia, L. microcarpa, L. pilosa, L. simpsonii, 
and L. suffruticosa. Natural hybrids between L. 
alata and L. pilosa, and between L. alata and L. 
suffruticosa, all of which require insect vectors to 
effect pollination, are abundant. (See Peng [1988] 
for voucher information of such natural hybrids.) 
The hybridity of field populations of L. alata x 
L. pilosa is initially suggested by the combination 
of the winged ovaries/capsules and minute, uni- 
form pubescence found in these plants. Of the 15 
taxa in sect. Microcarpium, only L. alata and L. 
lanceolata have winged capsules, and they are 
glabrous. Ludwigia pilosa and L. ravenii are con- 
sistently pubescent, and L. sphaerocarpa is vari- 
able in vestiture; none of them has winged capsules. 
The parentage of the hybrids between L. alata (n 
— 24) and L. pilosa (n — 16) is further verified 
by (1) their reduced level of stainable pollen (ca. 
45-70%, which is indicative of a heteroploid cross), 
and (2) a mixture of columnar and isodiametric 
cells and some other irregular cells on their seed 
surface (among the pubescent putative parental 
species, only L. pilosa differs significantly in the 
seed surface pattern from L. alata). 

Natural hybrids between Ludwigia suffruticosa 
(n = 16) and L. alata (n = 24) are difficult to 
recognize in the field. They are very similar to L. 
alata in general aspect, having winged capsules 
and slightly winged stems, as well as pollen grains 


are borne only in the upper leaf axils of each 


Volume 76, Number 1 
1989 


Peng 277 
Ludwigia sect. Microcarpium 


branch; their capsules are smaller, presumably as 
a result of reduced seed set; and their seed surface 
consists of a mixture of many columnar but rela- 
tively shorter 
metric cells (typical of L. suffruticosa), and is ir- 
regular 


cells, as well as somewhat isodia- 


9. Ludwigia sphaerocarpa Elliott, Sketch Bot. 
S. C. & Ga. 1: 213. 1817. Isnardia sphaer- 
ocarpa (Elliott) DC., Prodr. 3: 61. 1828. 
Ludwigia sphaerocarpa Elliott var. typica 
Fern. & Griscom, Rhodora 37: 174. 1935. 
TYPE: U.S.A. South duum. TE EARN Co., 
in swampy grounds near Orangeburg, flowers 
in August, s.d., S. Elliott s.n. (holotype, 
CHARL; photographs, GH, MO; possible is- 
otype, PH). Figure 51. 


Ludwigia sphaerocarpa Elliott var. deamii Fern. & 
37: 174. 1935. TYPE: U.S.A 
, low border of Lake Walker, 
W of Baile eytown, 23 Aug., 1925, C. C. Dea 
42350 (holotype, GH). 
a eee kii 'arpa Elliott var. jungens Fern. & 
Griscom, Rhodora 37: 174. 1935. TYPE: U.S.A. 
Vir p nia: : Princess Anne Co., Cape Henry, pool in 
sandy barrens, 23 Sep., 1933, M. L. Fernald & L. 
Griscom 2862 (holotype, GH). 
Ludwigia uer c da Eliott var. macrocarpa Fern. 
Griscom, Rhodora 37: 174. 1935. TYPE: U.S.A. 
Masse Plymouth Co., Lakeville, stony shore 
of Quitacas Pond, 27 Aug., 1899, P. Rich 
s.n. (holotype, GH). 


Stems erect, well-branched, (40-)60-110 cm 
tall, glabrous to densely strigillose; aerenchyma 
often very prominent in the lower part. Stolons 
many, up to 90 cm long, 2-3.5 mm thick, floating, 
sometimes branched, the leaves purplish, narrowly 
elliptic or oblanceolate, sometimes spatulate, 9-30 
mm long, 4-8(-13) mm wide, glabrous to densely 
strigillose, apex acute or obtuse, margin with dis- 
tinct hydathodal teeth or sometimes subentire, base 
attenuate into + winged petiole 1.5-3 mm long. 
Cauline leaves narrowly elliptic or lanceolate to 
sublinear or linear, those on the main stem 
100 mm long, 4.5-11(-16) mm wide, those of the 
branch 20-50(-60) mm long, 2.5-5(-6) mm wide, 
glabrous to densely strigillose, apex acute to very 
narrowly acute, margin entire, hydathodal glands 
often visible in main cauline leaves, base attenuate 
or narrowly cuneate into petioles 1-4(-10) mm 
long. Stipules reddish purple, lanceolate or deltoid, 
0.15-0.4 mm long, 0.1-0.2 mm wide. Flowers in 
upper leaf axils, lax to very crowded on branches. 
Sepals green without, yellow within, ovate-deltoid, 
ascending, 2-3.5(-4) mm long, 1.6-3(-3.3) mm 
wide, glabrous to densely strigillose on both sur- 
faces, apex acuminate, margin entire. Petals 0. 


Anthers 0.5-0.8 mm long, filaments yellow, 1- 
1.7 mm long, slightly dilated toward base. Pollen 
grains shed as tetrads. Nectary disc bright yellow, 
raised 0.4-0.6 mm on ovary apex, 1.5-3 mm 

across, 4-lobed, glabrous to short hirtellous between 
the lobes. Style yellow, 0.55-1(-1.25) mm long, 
glabrous to strigillose, especially below. Stigma yel- 
low, subglobose, 0.35- mm long, 0.4-0.7 mm 
thick. Capsules sometimes pinkish, subglobose, 
(1.8-)2-4(-4.5) mm long, 2-4 mm thick, glabrous 
to densely strigillose, pedicels 0.5-1.2(-2.3) mm 
long. Bracteoles subopposite on short pedicel, rare- 
ly flanking capsule base, linear to very narrowly 
lanceolate, rarely lanceolate, 0.5-1.5 mm long. 
Seeds brown to light brown, elliptic, 0.45-0.7 mm 
long, 0.25-0.35 mm wide, the surface of a mixture 
of columnar cells, these elongate and transversely 
elongate to the seed length, sometimes oblique, the 
orientation of cells often variable from seed to seed. 
Self-compatible. Gametic chromosome number, n 

16 


Distribution. Plants of L. sphaerocarpa grow 
in drainage ditches, on stream or pond shores, and 
in river marshes, swales, swamp forests, edges of 
limestone sinks, peaty bogs in pastures, and inter- 
dunal marshes. This species has an extensive dis- 
tribution along the Atlantic coast, ranging from 
Massachusetts to northcentral Florida. To the west 
it extends through the panhandle of Florida, south- 
western Georgia, southeastern Alabama, central 
and southwestern Louisiana, and reaches eastern 
Texas. Disjunct populations occur in southcentral 
Tennessee, extreme southwestern Indiana, along 
Lake Michigan in northeastern Illinois and north- 
western Indiana, and in westcentral New York (Fig. 
52). Flowering from June to September; fruiting 
from August to November. 


Representative specimens examined. U.S.A. ALA 
BAMA: Covington Co., Blue Pond, Conecuh National For- 
est, SW of A Kral 44739 ri. GA, GH, MO, 
SMU, US, VDB). Elmore Co., ca. mi. E-SE of jct. 
with US Hwy. uds & Co. Rd. 4, ix; 325 (UNA). 
Geneva Co., SW of Hartford, McDaniel 7654. 
CONNECTICUT: Middlesex Co., 


Ellendale, Commons s.n. 18 
Co., 4 mi. N of Suwannee, Godfrey 56178 (FSU, GA, 
GH, NY, TENN, ide Flagler Co., near Korona, 1944, 
Butts s.n. (GH). Franklin Co., 0.5 mi S o 
the Ochlockonee River, Godfrey 54110 (FSU, GH, NY). 
de Co., 4 mi. E of Trenton, 1940, West & Arnold 
n. (FLAS). Holmes Co., just E of a E Godfrey 
590 13 (FSU, in 2 sheets). Jackson 
Wildlife Refuge Area, Monoson 55 (CH 
ca. 4 mi. E of Wacissa, Godfrey 631 73 (FSU). Leon 


278 Annals of the 
Missouri Botanical Garden 


A q LZ 
SE] / y 
5 p 
Ww 
A / 
f | a 
e. 
A - 
/ 
"e 


FIGURE 51. Ludwigia sphaerocarpa. All but a’ from South Carolina (Jasper Co., Peng et al. 3986, MO); a’ 
from South Carolina (Jasper Co., Boufford et al. 21650, MO).—a. Habit, erect stem. —a'. Basal stolon. — b. Flower. — 
c, c’. Adaxial and abaxial views of stamen.— d. Fruit. — e. Cross section of fruit. 


Volume 76, Number 1 
1989 


Peng 279 
Ludwigia sect. Microcarpium 


FIGURE 52. 


Co., 2 mi. SE of Tallahassee, McDaniel 3708 (FSU). 
Madison Co., 6 mi. E of "da wel Wiegand & Manning 
2199 (GH). St. Johns Co., .N En jet. of FL 201 
and C-305, on FL 207, nz et Fut 4159 (MO). Taylor 
Co., 1.2 mi. SE of Salem, Godfrey 64726 (DS, LL, NCU). 
Te Co., 2 mi. N of ado dine 1943, West & Arnold 

n. (FLAS). Wakulla Co., 6 mi. E of Sopchoppy along 
Odhlockones River, Morar 28 (DUKE, FLAS, FSU, MSC, 
US). Washington Co., Curtiss a (C, F, a D 
S, UPS, US). GEORGIA: Baker , Dunca 
4055 (GA). Burke oe near Shell Blu, Eyles 64 78 (CA). 
Decatur Co., along y. 97, 1.3 [o Eo 
Thorne & Davidson 12147 (FSU, "GA. Dooly Co., 
N of Pinehurst, Kral 51611 (VDB). Emanuel Co. 

Oh 


nd 
Altamaha River, Peng et al. 4115 (MO). Lowndes Co., 
near based Curtiss 67 10 (GÀ, GH, MO, NY). Mitchell 
a , 5 mi. N of Camilla, Thorne 5792 (GA). Seminole 
s Lake, lots 99 and 102, Dist. 21, Thorne 
Davidson 16764 (FLAS). Sumter Co., Harper 553 (F, 
GH, MO, NY, US). ILLINOIS: Cook Co., Chicago, 1860, 


Distribution of Ludwigia sphaerocarpa. 


1.8 mi. S of 
and 0.8 mi. W of Co. line. Friesner 22974 
(FLAS, XR S SMU). Lake Co., Pine, 1896, Unbach s.n. 
(DS, SC, NY, S, US). Porter M Wilson (Dune 
E 1958, Hennit a. n. (F, NY, SMU, W). Posey Co., 
Scuffle Pond in the Nolta Erwin ds ca. 8 mi. SW 
of Mt. Vernon, Deam 59451 (NY). Starke Co., SW 
corner of Bass Lake, Friesner 16306 (GH, MO, SMU). 
LOUISIANA: Cameron Parish, Lacassine Wildlife Refuge, 
Thieret 16410 (DUKE, GA, MO, SMU). Rapides jee à 
Red River, 1840, Hale s.n. (US). MARYLAND: Wicomi 
Co., Salisbury, 1889, Canby s.n. (NY). Worcester Co. 
15 mi. N of Berlin, O'Neill 7489 (CAS, DS, F, ND, NY). 
MASSACHUSETTS: Middlesex Co., Concord River, Bedford, 
1908, Fernald s.n. (GH). Plymouth Co., Middleboro Ponds, 
Lakeville, Churchill 595 (MO). Suffo Ik Co., near Boston, 
1830, Greene s.n. (NY). NEW JERSEY: Atlético Co., Ham 
monton, Gershay 505 (GH). Burlington Co., Atsion, Foie 
4697 (GH, TENN). Cape May Co., Bennett, 1913, Stone 
NY). Cumberland Co., 1902 5 


ug eg s.n. (F). INDIANA: Jasper Co. 


$ 
8 


Middlesex Co., Spotswood, Taylor 2 
Co., Denmark Pond, Mackenzie 4758 (MO, , S). 
Sussex Co., Hopkins Corners, 3 mi. NE of Lafayette, 


280 


Annals of the 
Missouri Botanical Garden 


1935, Edwards s.n. (NY). NEW YORK: Queens Co., N of 
I: Long Island, 1900, Bicknell s.n. (NY). Rich- 
., Staten Island, 1879, ise s.n. (GH). Suffolk 
bn rbor, Muenscher & 

ut i Esopus Pond, 
). Ween Co., Lake 
Mohegan, ade Martens s.n. (C, DUKE, F, GH, NY, 

). Ya ., s.d., c ell s.n. (US IN 

Dare Co., s Hwy. 345, 3.75 mi. ON of Wanchess, 
la Island, Radford & Stewart 895 (NCU). Hoke 
o., 4 mi. rrr Heights, Ahles 36372 (NCU). 
Johnston Co., P of Princeton, Radford 27780 
(CM, GH, FLAS, NCU, NY, SMU, TENN, VDB). Wash 
ington Co., 3.6 mi 


. RHODE ISLAND: 
uth Weg iar T. 
OUTH CAROLINA: Aiken Co., 1.4 m 

of Monetta on Sc ghé and 0.2 mi. N on SC Huy. 
266 (near ld mui Cemetery) Massey & Massey 2995 
(MO, NCU). Bamberg Co., 4 mi. SW of Denmark on Co. 
Rd. 26, Ahles E (NCU). Barnwell Co., Savannah 
River Operations Area of the 
1964, Kelley ss s.n. (USCH). Dalo: Co. 
Pineville on SC Hwy. 45, Ahles & Baird EAT (NCU). 
Calhoun Co. Riley, along US Hwy , Ahles 35324 
(NCU). Clareidan Co., 13 m i SW. Bi A Mi Godfrey 
& x 1457 (DUKE, F, CH, MO, NY, US). Colleton 

o., mi. E of Smoaks on SC Hwy. 217, Ahles & 
Bell a (NCU). Dorchester Co., along US Hwy. 15, 
N of SC Interstate 15, Leonard & Radford 1988 (NCU). 
Hampton Co., 2.1 mi. S of Gifford En I Hwy. 321, 
Bell 3948 (NCU, USF). Jasper Co., 6.6 mi. N of SC 
Hwy. 46 on US Hwy. 17, N of Hardeeville, Peng et al. 
3986 (MO). Kershaw Co., Adams ey 2.5 mi. NE 
of US Hwy. 1 at Camden, on SC Hwy. 15, Radford et 
al. 11388 (B, BOON, DS, East vos Univ. Herb., 
— Pis NCU, S, TEX, UNA, UH). Lee Co., 

s Creek, near US Hwy. 15 : Epis 

Radford 27252 (FSU, NCU). TENNESSEE: Coffee Co., 


.W 
of ‘Winchester, 1976, Nini s n. (VDB). VIRGINIA: Creen 
ville Co., Cephalanthus swamp, ca. 1 mi. N of Skipper's, 
Fernald & Long 8788 (GH). Isle of Wight Co., m 
Pond, S of Berns Church, apio & Long 7547 (GH 

). Nansemond Co., ca. 7 mi. E-NE of Suffolk, Kral 
13766 (FSU, VDB). NL Co., Lake Drummond, Great 
Dismal Swamp, W of Walleceton, Fernald & Long 13402 
(GH, MO, NY, SMU, TENN, US). Princess Anne Co., 
E of T-1, of RR, near picnic — Egler 40281 
(NY). Sussex Co., N of Stony Creek, nald & Long 
8790 (GH). York Co., NW of Grafton, Fernald & Long 
7548 (GH, NY, US). 


Ludwigia sphaerocarpa is highly polymorphic. 
Its variability is largely the result of its wide geo 


action of differential adaptation and genetic drift 
made possible by its pattern of distribution, and its 
possible hybrid nature (see below). Hybridization 
with L. pilosa in regions where their ranges overlap 
apparently provides an important source of vari- 
ability in L. sphaerocarpa. 


Populations of this species consist of individuals 
that vary in overall pubescence, leaf shape and 
size, fruit size, and density of fruits on branches. 
Fernald & Griscom (1935) recognized var. jun- 
gens, var. macrocarpa, and var. deamii based on 
various combinations of these characters. In their 
treatment, the varieties were separated as follows: 


Rameal leaves strongly reduced, glabrous or pubes- 
cent, lanceolate. 

Mature hypanthium small, 2.5-3.2 mm long, 

m broad, averaging broader than long. 

Hypanthium pubescent; leaves narrowly line- 

ar-lanceolate and attenu ate Lees 
COMIS var. "typica" [var. TM 

Hypanthium, branches, and leaves pubescent; 
eaves more broadly lanceolate, not a 


Haire penu rum larger, 3.5-4.6 mm pe ng, 
3.2-4 mm broad, averaging longer than broad 
var. macrocarpa 
Rameal leaves scarcely smaller than the primary 
nes, pubescent, narrowly oblong; stems pubes- 
cent; hypanthium eE 3 mm long, about 
as wide var. deamii 


In his revisions of the New World species of 
pad Munz (1944, 1965) reduced var. deamii 

o synonymy under var. jungens and presented 
ds following key to the varieties: 


Fruit crowded on branches, 3.5-4.5 mm long, not 
as wide. Massachusetts to New Jersey ..... 
var. macrocarpa 
Fruit not ue on branches, 2-3.2 mm lon 
usually 
Main stems id leaves usually glabrous; principal 
leaves linear-lanceo ate and over 7 cm lon 
Rhode Island to Florida .......... var. sphaerocarpa 
Main stems and leaves ae io to strigose, lan- 
ceolate, usually less than 7 cm long. New Jersey 
to North Carolina, Michigan, and Indiana ..... 
var. jungens 


A study of numerous herbarium specimens not 
available to Fernald & Griscom (1935) and Munz 
(1944, 1965) has revealed that correlations be- 
tween these characters are not consistent; the 
often intergrade. Indeed Munz (1944) noted that 

r. typica (the typical variety) and var. jungens 
Maot freely.” Under Ludwigia sphaerocar- 
pa var. macrocarpa, he also wrote, “New Jersey 
material is sometimes difficult to distinguish from 
var. typica (the typical variety)." In my opinion, 
it is best to treat L. sphaerocarpa as representing 
an assemblage of variable populations without rec- 
ognizing the infraspecific taxa. These populations 
share the following diagnostic characters: subglo- 
bose capsules, diminutive bracteoles (shorter than 
1.5 mm long), and ovate-deltoid, yellowish, con- 
spicuous sepals. 


Volume 76, Number 1 
1989 


Peng 281 
Ludwigia sect. Microcarpium 


It is also of interest to note that, although the 
seed surface cell pattern is generally very regular 
within populations of members of Ludwigia sect. 
Microcarpium (Figs. 19-28, 30-34), this is not 
the case for L. sphaerocarpa (Fig. 29). The seed 
surface is arranged in columnar cells both trans- 
versely elongate and parallel to the seed length, 
the former alignment often predominating in the 
central part of the seeds. Seeds with variously 
oriented surface cells are also seen in some pop- 
ulations. Comparison of the seed surface pattern 
in populations of L. sphaerocarpa with that of 
various artificial hybrids strongly suggests that ear- 
lier hybridizations within the interfertile tetraploid 
group of sect. Microcarpium may have resulted 
in this widespread series of populations that are 
more or less stabilized in some of their character- 
istics, and such hybridizations may still be contrib- 
uting to the variability of this series of populations. 

The biosystematic study of Ludwigia sect. Mi- 
crocarpium (Peng, 1988) demonstrated that hy- 
brids between species that differ in their seed sur- 
face cell pattern always show a mixture of cell 
types or cells of intermediate shapes and sometimes 
of randomly oriented cells. Similarly, hybrids re- 
sulting from crossing a densely hirtellous species 
with a glabrous one always exhibit minutely villous 
or strigillose pubescence. Of the tetraploid Lud- 
wigia taxa that are interfertile with L. sphaero- 
carpa, L. ravenii and L. glandulosa subsp. 
"pehyearpe are the only ones with columnar seed 

lls elongate transversely to the seed length, 
= mw L. polycarpa and L. glandulosa subsp. 
glandulosa are the only taxa with columnar seed 
surface cells elongate parallel to the seed length. 
Both subspecies of L. glandulosa, having narrow, 
subcylindrical capsules, are unlikely to be parents 
of L. sphaerocarpa. In seed surface pattern and 
overall pubescence, L. ravenii (densely hirtellous) 
and L. polycarpa (glabrous) seem to be suitable 
parental candidates. However, that they both have 
oblong-obovoid capsules and are highly autoga- 
mous, in contrast to subglobose capsules and out- 
crossing habit of L. sphaerocarpa, renders it un- 
likely that they have been involved directly in the 
formation of L. sphaerocarpa. One or possibly 
both parents of L. sphaerocarpa are likely extinct. 

The morphological variation in Ludwigia 
sphaerocarpa is further complicated by its fre- 
quent natural hybridization with L. pilosa, which 
has apparently resulted in many hybrid swarms or 
introgressed populations. These plants generally 
exhibit varying degrees of intermediacy in diag- 
nostic characters such as seed surface pattern, 
overall pubescence, bracteole size, sepal shape and 


size, and leaf shape. The following specimens show 
intermediacy in at least two of the above characters 
or a combination of some of the diagnostic features 
of L. pilosa with some of those of L. sphaero- 
carpa: 


U.S.A. ALABAMA: Escambia Co., 2.5 mi. N of Atmore 
dapes off Hw wy. 21, Harvey 275 (UNA). FLORIDA: Citrus 
i. S of Homosassa, Kral 7772 (FLAS, GH, NCU, 
Us. DE VDB). Columbia Co., 2.5 mi. N of Ellisville, 
along U.S. Hwy. 41, Raven 18634 (DS). Dade Co., near 
Lake Okeechobee, Fredholm 6182 (GH, NY, US). Duval 
Co., near Jacksonville, Curtiss 4324 (DS, ND, NY, US), 
Christ 5214 (GH). Highlands Co., S end of Lake Istok- 
poga, Raven 18683 (DS, NCU); E of Sebring, Carter 
p, 1948, Garrett s.n. (FLAS); 
Parker Islands, Brass 15325 (US). Hillsborough Co., 
River State Park, along U.S. Hwy. 301, 1978, Rochow 
& Lopez s.n. (USF); 1 mi. N of bridge on Tarpon Springs- 
Lake Fern Road, Poppleton 699 (DS). Lake Co., in the 
ip of Eustis, Nash 1041 (F, GH, MO, MSC, NY, 
PH, S, US). Madison Co., 6.8 mi. W of Greenville, along 
U.S. Hw . 90, Raven 18629 (DS); 10 mi. S of Greenville, 
Kral 3769a (FSU, GA, GH, NY, SMU, TENN, UNA, 
USF). , NE of intersection of state roads 62 
& 39, Shuey 1712 (USF). Polk Co., ca. 3.3 mi. SW of 
Polk City, along FL Hwy. 33, Smith & Myint 586 (FLAS, 
FSU, NCU). St. Johns Co., St. Augustine, 1875, Reyn- 
olds s.n. (F, FLAS, MO, NY —2 sheets, PH); 1876, 
arber s.n. (FLAS). Wakulla Co., vicinity of Panacea, 
Godfrey 60276 (FSU). GEORGIA: Colquitt Co., 8 mi. NE 
of Moultrie, Kral 15008 di uie iss Co., 0.8 mi. 
S of Interstate 16 on U.S. Hwy. 1, Peng et al. 4025 
(MO), Peng et al. 4026a (MO) Richmond Co., Augusta, 
Sanitary Dairy Ys 1924, Hildebrand s.n. (DUKE). 
Wilcox Co., 10.9 mi. NNW of Abbeville, Hardin & 
icio 14246 (CA). NORTH CAROLINA: Robeson Co., ca 
S of Luberton, Kral 19030 (VDB, E with 
L. ra SOUTH CAROLINA: Aiken Co., 1869, s.c., s.n. 
, St. Helena Island, 1884, Cuthbert 
E of Smoaks, Raven 


18718 (DS, E shouts, NCU). TEXAS: Freestone Co., 5 m 


SE of Dew, on Hwy. 75, Coeli 35249 (LL); 15 mi. SE 
of Fairbald, Kral 111 (FSU). Gonzales Co., Ottine, T 
Tharp s.n. (MO, SMU, TEX). Harris Co., Gosby, 1923, 
Fischer s.n. (S, US); Hamble, Tharp 4518 (MO), M 


don, Reverchon 2301 (MO). Hardin Co., 1937, Thar 

s.n. ne br sm ta Nacogdoches, Waller 273 

(SMU, TAES). ARE D. cm. de Mauriceville, 

Shinners perce “Wood i. N of Mineola, 

Correll 35021 (LL); Lake Ellis, ended & Lundell 11748 
U). 


Some of the above putative hybrid populations 
are from central and southern Florida, where nei- 
ther typical Ludwigia pilosa nor L. sphaerocarpa 
occur (Peng, 1988, fig. 22). This suggests that 
their physiological characteristics, and thus their 
ecological tolerances, may differ from those of the 
parents in such a way as to allow them to occur 
in novel habitats and areas where neither of their 
parents can survive 

The following collections are unusual in being 
densely hirtellous (having erect hairs ca. 0.4-0.8 


282 


Annals of the 
Missouri Botanical Garden 


mm long), but they are otherwise typical of Lud- 
wigia sphaerocarpa. They may represent intro- 
gressants between L. sphaerocarpa and either 
pilosa or L. ravenii. 


U.S.A. FLORIDA: Flagler Co., vicinity of Andalusia, 
Godfrey & Reinert 61149a (FSU). SOUTH CAROLINA: 
Beaufort Co., 0.8 mi S of Yemassee on Co. Rd. 3, Bell 
4703 aia Raven 18716 (DS, FLAS, TEX). Hampton 
Co., 0.8 mi. NNW of Shirley on Co. Rd. 20, Bell & 
Ahles 1 8254 (NCU). Jasper Co., 5 mi. S of Ridgeland, 

U.S. Hwy. 17, Dille & Dille 346 (MO); 7 mi. N of 
Hardeeville, on U.S. Hwy. 17, Dille & Dille 349 (MO 
TEXAS: Angelina Co., N of Boykin Springs, Angelina Na- 
tional Forest, Correll 34910 (LL). Hardin Co., between 
Silsbee and Kountze, S of Hwy. 418, SW of county dump 
grounds, Amerson & Watson 228 (LL). E. Co., Hum- 
ble, Tharp 4518 (US). Montgomery Co., mi. W of 
Cleveland, Kral 21041 (DS, VDB—2 ctor 


— 


Plants of the following collections were found to 
have one to three vestigial, yellow petals in some 
of the flowers: 


U.S.A. FLORIDA: Flagler Co., near prey 1944, Butts 
s.n. (GH). Wakulla Co., on U.S. 98, 5 mi. W of FL 59, 
Dille & Dille 401 (MO). NEW YORK: nd Co., Long 
Pond, Sag Harbor, Muenscher & Curtis 6357 (NY). 


Judged by their morphology as well as their full 
seed set, these plants are more likely to be mutants 
showing an ancestral trait rather than hybrids be- 
tween Ludwigia sphaerocarpa and one of the 
petaliferous species of sect. Microcarpium. 

Almost all populations of Ludwigia sphaero- 
carpa. are at least sparingly strigillose, especially 
on the ovary /capsule walls. The following two gla- 
brous collections from Louisiana are exceptional: 


.S.A. LOUISIANA: Cameron Parish, in “The Pool," 
Lacassine Wildlife Refuge, Thieret 16410 (DUKE, GA, 
MO, SMU). Rapides Parish, Red River, 1840, Hale s.n. 
(US). 


They seem to parallel the glabrous variants of 
Ludwigia linearis (see above) in which a few gla- 
brous plants may grow intermixed locally with 
densely strigillose individuals. 

Ludwigia sphaerocarpa is one of the few species 
in sect. Microcarpium in which the distance be- 
tween the androecium and the gynoecium is such 
that self-pollination is impossible without the inter- 
vention of pollen vectors. Although the flowers of 
L. sphaerocarpa are apetalous, their sepals are 
adaxially yellowish. This, along with the bright yel- 
low nectary disc, which produces copious nectar, 
attracts many insects to the flowers of this appar- 
ently largely outcrossing species. 

n the field, Ludwigia sphaerocarpa grows with 
L. glandulosa, L. linearis, L. microcarpa, L. pi- 
losa, and L. polycarpa and hybridizes with all but 


the small-flowered, modally selfing L. microcarpa. 
The natural hybrids generally exhibit intermediate 
morphology, most notably in fruit shape and size, 
bracteole length, overall pubescence, and seed sur- 
face cell pattern. The fruits are plump and contain 
abundant seeds in homoploid (tetraploid) hybrids. 
In the heteroploid (triploid) L. sphaerocarpa (n = 
16) x L. linearis (n — 8), one to four petals were 
present in most flowers, and seed set was negligible. 

In the experimental greenhouse, artificial hy- 
brids have been synthesized between Ludwigia 
sphaerocarpa and the diploid L. linearis, L. mi- 
crocarpa, the tetraploid L. glandulosa, L. poly- 
carpa, L. suffruticosa, L. lanceolata, L. pilosa, 
the hexaploid L. alata, and the octoploid L. cur- 
tissii (Peng, 1988). 


10. Ludwigia pilosa Walter, Fl. Carol. 89. 
1788. Isnardia pilosa man Kuntze, Rev. 
Gen. Pl. 1: 251. 1891. TYPE: U.S.A., “the 
Carolinas,” s.d., T. Walter 658 Wido BM, 
Walter Herbarium p. 66; photograph, MO). 
Figure 53 


Ludwigia rudis Walter, Fl. Carol. 89. 1788. TYPE: U.S.A. 
Carolinas,” s.d., T. Walter s.n. (type not found 
" BM, Walter Herbaria: possible isotype, P (“No. 
3”), photograph, MO). 

Ludwigia hirsuta Lam., Encycl. 3: 614. 1792. TYPE: 
U.S.A. South Carolina: 1785-1792, J. Fraser s.n. 
(holotype, P; Pih m GH, MO). 

Ludwigia mollis Michaux, Fl. Bor.-Am. 1: 90. 1803. 


Isnardia mollis (Michaux) Poiret in Lam., Encycl. 


wet places, 1785-1796, A. Michaux s.n. (holotype, 
P; photographs, GH, MO; possible isotype, PH). 

Plants hirtellous throughout. Stems much 
branched, 40-120 cm tall, often with prominent 
aerenchyma below when submerged in water, the 
hairs 0.25-0.95 mm long, translucent or spotted 
with light brown pigments. Stolons up to 2.5 m 
long, creeping in mud or floating in water, occa- 
sionally bearing flowers and fruits, the leaves ob- 
ovate or elliptic to orbicular, 6.5-20 mm long, 5- 
11 mm wide, densely hirtellous to nearly glabrous, 
margin with distinct hydathodal teeth, petioles 2- 
7 mm long. Cauline leaves (upon drying) usually 
brownish green adaxially, brown abaxially, elliptic 
or lance-elliptic to very narrowly so, 15-80(-100) 
mm long, 3-12(-14) mm wide, those on the main 
stem usually 1.5-4 times longer than those on the 
branches, the veins reddish, the hairs 0.3-0.5 mm 
long, apex acute or narrowly acute, margin entire, 
hydathodal glands obscure, base acute to narrowly 
acute or attenuate, sessile or with petioles up to 
2(-10) mm long. Stipules brownish purple, ovate 
to lanceolate, 0.2-0.25 mm long, up to 1 mm 


Volume 76, Number 1 Peng 283 
1989 Ludwigia sect. Microcarpium 


/ 


FIGURE 53. Ludwigia pilosa (South I inet Co., Peng 4397, MO). —a. Habit, erect stem. — a'. Basal 
stolons. —b. Cauline leaf. — tolon leaf. — wering branch. —d. Flower bud.—e. Flower.—f. Flower, partly 
dissected to show stigma, style, and disc. —g. Sep al. —h, h'. Adaxial and abaxial views of stamen. — i. Fruit. —j. Cross 
section of capsule. 


284 


Annals of the 
Missouri Botanical Garden 


wide, usually obscured by hairs on the stem. Flow- 
ers usually congested in upper leaf axils. Sepals 
very conspicuous, creamy white within, usually also 
tinged with pink along the main veins and edges, 
occasionally reddish when grown in exposed sites, 
ovate-triangular or triangular, with 3 parallel main 
veins that are usually raised and prominent on the 
adaxial surface, apex elongate acuminate to sub- 
cuspidate, ascending with reflexed tips, 3.5-5.5 
(76) mm long, 2-4 mm wide, densely hirtellous 
within. Petals O. Anthers 0.6-0.9(-1.3) mm 
filaments yellowish, 1.5-2.5 mm long, distinctly 
ilated below. Pollen grains shed as tetrads. Nec- 
tary disc bright yellow, turning black upon drying, 
raised 0.3-0.75 mm on top of the ovary, 2-3.8 
mm across, indistinctly 4-lobed, densely hirtellous 
around the style base and often sparsely hirtellous 
between the lobes. Style 1 -2 mm long, sparsely to 
densely hirtellous, especially below; stigma 0.3- 
0.6 mm thick. Capsules subglobose or sometimes 
oblong-obovoid with rounded corners, 3-5 mm lon 
3-4.5 mm thick, densely hirtellous, sessile or 
with pedicels up to 1 mm long. Bracteoles flanking 
the capsules, attached 1-2.25 mm above capsule 
base, linear-lanceolate or narrowly elliptic, hirtel- 
lous, 3-6.5(- 7.2) mm long, 0.25-1.5(-1.7) mm 
wide. Seeds brown, elliptic oblong or oblong ovoid, 
slightly curved on both ends, 0.5-0.7 mm long, 
0.25-0.35 mm thick, the surface cells nearly iso- 
diametric. Self-compatible. Gametic chromosome 
number, 16. 


long; 


gu 


Distribution. 
frequently found in roadside ditches, 
marshes, in swales in sandy pine flats, at edges of 
pocosins, in peaty bogs, in low grassy savannahs, 
and in swamp forests. This species is distributed 
along the Atlantic coast of extreme southeastern 
Virginia, eastern and central North and South Car- 
olina, southern Georgia, and northern Florida. It 
extends westward through the Florida panhandle, 
southern Alabama, Mississippi, and Louisiana, 
reaching eastern Texas. A disjunct population oc- 
curs in northern Alabama (Fig. 54). Flowering from 
late June through September; fruiting from late 
July through November. 


Plants of Ludwigia pilosa are 
in river 


Representative specimens examined. U.S.A. ALA- 
BAMA: Baldwin Co., Fort Morgan Peninsula, Demaree 
35917 (FSU, GH, SMU, USF, VDB). Barbour Co., ca. 
5 mi. N of Eufaula on AL 1-65, Cowickee Creek camp 
by Lake Eufaula, Kral 33185 (VDB). Butler Co., ca. 11 

mi. N of Georgiana by AL 1-65, Kral 40892 (VDB). 
Covington Co., ca. 8 mi. S of Opp, by US nn 331, 
Kral 32063 (SMU, VDB). Crenshaw Co., near Dozier 
Reed 2052 (TEX). Dale Co., near Cla E Kral 
22079 (DS, VDB). Dallas Co., just W of Selma, by Hwy. 


22, Perdue 4155 (FSU, VS). Elmore Co., W s 
Elmore, by Mortar Creek, Kral 40808 (VDB). Escambia 
Co., 1.3 mi. E of ba wbeek, Kral 33831 (MO, VDB). 
Geneva Co., 3.2 mi. S of Hartford on AL Hwy. 167, 
Kral 36728 (VDB). Houston Co., just E of jct. S of 
Spring Hill Church, pe Creek Swamp, Clark 7343 
(NCU). Lowndes Co., 10 mi. E of Minter, near Fostoria, 
Kral 48927 ee! Macon Co., Earle & Earle 49 (GH, 
O, ND-G, US). Madison Co., Earle & Earle 49 
(GH, MO, ue US). Mobile Co., on Hollinger's Island, 
Harper 4087 (GA, MO, NCU, NY, UNA, US, VDB). 
Pike Co., Spring Hill, Bush 339 (S). Washington Co., 
3.1 mi. S of St. Stevens, Kral 37234 (FSU, VDB). Arcs 
DA: Alachua Co., near Waldo, 1936, Survey s.n. (FLAS). 
Bay Co., near Long Beach, Godfrey & Kral 55075 (FSU, 
NCU), Bradford Co., near US Hwy. 301, Radford 8245 
NCU). Calhoun Cb. 10 mi. S M oo Adams 
700 (FLAS, FSU, GH, NY, SMU, VDB). Clay Co., 2.7 
mi. S of Cove Springs along US Hwy. 17, Raven 1 8686 
(DS). Columbia Co., 3.6 mi. NE of Lake City, Osceola 
National Forest, Pirdus 1824 (C, FSU, GA, GH, NCU, 
SMU, TEX, USF). Duval Co., near Jacksonville, Curtiss 
924 (CM, F, FLAS, GA, GHM, MO, MSU, NY, US). 
Escambia Co., near Pensacola, Burkhalter 2919 (FLAS). 
Franklin Co., 38. 8 mi. W of jct. of US Hwys. 98 & 319, 
Peng et al. 4345 (MO). Gadsden Co., ca. 3 mi. NW of 
Quincy along US Hwy. 90, Webb & Ross 1011 (TENN). 
Gulf Co., between Weewahitchka & Port St. Joe, Godfrey 
& Kral 54168 (DUKE, FSU, GH, NY, TEX, USF, VDB). 
Hamilton Co., 3 mi. E of Ae 1941, West & Arnold 
s.n. (FLAS). Hole Co., 4.1 m 
mo 5206 (FSU). Jakt Co.. 5m 
. 90, Hood 2593 (FLAS). cdm Co., 
Wacissa ‘along Tram Rd., along St. Marks River, Godfrey 
70908 (FSU). Leon Co., 6 mi. N- aires, Kral 
3687 (FLAS, GA, GH, NY, S, SMU, TENN, eon piah 
Liberty Co., just S of Gadsden Co. line on Rt. 26 ar 
jet. with Rt. 20, Anderson 4477 (FSU, MO). d sed 
.8 mi. of Greenville on US Hwy. 90, Raven 
1 8625 (DS). es Co., Spes near Yellow 
. 2, Godfrey 7542 7 (FSU). Santa Rosa 
n Godfrey 73798 (FSU). Wa- 
, 6.1 of Sopchoppy n FL Hwy. 375, 
Wooten 2270 (FSU), Walton Co. . W of jet. US 
Hwy 1 on US Hwy. 98, near ii Rosa Beach, joan 
es DS NCU). Washington Co., 3.5 mi. E of Car 
ville, Kral & Godfrey 5961 (FSU, USF). GEORGIA: na 
pling "E . near Baxley, s.d., s.c. 923h (US). Atkinson 
o., ca. 5 mi. S of Pierson, sian Plain Province, Dun- 
can 3015 (GA, GH, US). Baker Co., ca. Big Cypress 
area, ate Ford & e: s.n. (TENN). Brooks Co., 
17 mi. N of Madison, FL on GA Hwy. 33, Ramsey & 
Stripling 153 (FSU, SMU). Bryan Co., S side of Rich- 
mond Hill, Kral 18806 (VDB). Bulloch es. 3.5 mi. from 
jet. of GA Rt. 67 


~ 


wy. 17, Raven 18701 (DS, 
FLAS). Charlton Co., GA Hwy. 94, between Moniac & 
St. George, Jones et ul 23027 (GA). Chatham Co., near 
Savannah, s.d., s.c. 4169C (S, US). Colquitt Co., 4.8 mi. 
NW of Borlin, Coastal Plain Province, Faircloth 2676 
(NCU). Cook Co., 6 mi. W of Adel at the Porter Purvis 
Farm, Coastal Plain Province, Faircloth 2401 (GA, NCU). 
Decatur Co., 1.5 mi. E of Climax, by US Rt. 84, Godfrey 
77562 (FSU). Dooly Co., 2 mi. N of Pinehurst turnoff, 
by I-75, Kral 51610 (VDB). Early Co., W central bound- 
ary of Co. in Moccasin Pond, Duncan 4025 (GA, MO, 


Volume 76, Number 1 


Peng 285 
Ludwigia sect. Microcarpium 


1989 
D 
á "n 
D di sE L ( 
e PE 4 PEN a" y SÉ 
pou MÀ G9 i LA, AS 
S Ml oe 1 ES 
he ( c» b 
V) Vi ^ id | 
Y f le ( y p. ke 
a UW E A 
/ NY ^ Moy» > 
| e AN 
) E fa ag 
LATER Or 
e 
e 0900s 
e. 
. * eee ee 
007o e* ee t 
e o e (Fe) q j 
4 — Cf 
\? aA m P 
e SY Ys 
ii 
Mm 
A 
"n 50 i 200 z 400 500km 
\ 
i 
Y 
m 
FIGURE 54. Distribution of Ludwigia pilosa. 


SMU). Echols Co., 2.6 mi. E of Statenville, Kral 51645 
(VDB). Effingham Co., near Eden, 1875, Curtiss s.n. 
(US). Emanuel Co., 0.8 mi. S of I-16 on US 1 along the 
Ohoopee River, Peng et al. 4024 (MO). Evans Co., 1 
mi. S of jct. GA 169 & US 25 & 301, nuo 5& 
301, Peng et al. 4107 (MO). Grady Co., 3.5 mi. NW 
of Calvary on the J. R. qi. property, OMA 1510 
(GA, MO, NCU). Irwin Co., 7.3 mi. SW of Irwinville at 
Tift-Irwin county line, Pai 5408 (GA). Lanier Co., 

2 mi. S of Lakeland, along GA Hwy. 31, Thigpen et al. 
226 (NCU). Liberty Co., near Hinesville, Camp Stewart, 
1943, Gumm s.n. (C) Long Co., 4.2 mi. SE of Tattnall 
county line on US 301 & 25, Peng et Hm 4110 (MO). 
Lowndes Co., 7.3 mi. n of Mi um Faircloth 2634 (GA, 
NCU). McIntosh Co., i. E of power line on Wesley 
Lake Rd., Bozeman an (GA, NCU). Richmond Co., 
Sanitary Dairy Pond, Augusta, 1924, Hildebrand s.n. 
(DUKE mixed with L. pilosa x L. sphaerocarpa). Screv- 
en Co., 7 mi. S of Sylvania, Kral 22363 (VDB). Seminole 
Co., 1.5 mi. E of Donaldsonville, Godfrey 63209 (FSU). 
W) 


(NCU). Wayne Co., -NW o 
Odum, Duncan 23859 (FLAS, CA). os Co., 4.5 


mi. W of Alamo, Kral 22330 (VDB). Wilcox Co., 10.9 
mi. N-NW of Abbeville, Duncan 


MISS, NCU, SMU, , USF). LOUISIANA: Beauregard 
Parish, 6 mi. o , near Zion Eu , Thieret 
28115 (FSU). Calcasieu B Pa , 2.5 mi. f Sulphur, 
just S of St lr iig d 20591 (DS, AM lies de 
Parish, near Kai in 2415-4 (NCU, SMU). Ra E 
ides Parish, near andria, s.d., Hale s.n. (GH). S 
Tammany Parish, of St. Tammany along bos 


J 
a Thieret 21648 (DS, FSU). Tangipahoa Parish, 
l = W of Robert, oe. & in 9288 (DUKE, 
U, ND). Vernon Parish, c i. E of Rosepine off 
3 1146, Allen & Vincent 651 (CA). Washington Parish, 
near Franklinton, Demaree 46896 (MO, NO, SMU). mis- 
=A Clarke Co., 1896, Schuchert s.n. (NY). Coving- 
on Co., S of Collins, Hwy. 49, Jones 10908 (GA, MISS, 
NCU) F orrest Co., near Rayland Hills, Rogers 4096 
George Co. ., near Agricola, Demaree 32797 

VD 


7 


dsboro, Demaree 29713 (DUKE, NCU, SMU, 
TEX, VDB). Jackson Co., 2 mi. W of US 10 & MS i 
on MS 90, Peng et al. 4354 (MO). Jones Co., 6.6 m 
NE of Moselle on MS Hwy. 11, Raven 18568 (DS). 


286 


Annals of the 
Missouri Botanical Garden 


Lawrence Co., i. N of Monticello on Hwy. >El Jones 
9976 (MISS). M Co., 0.8 mi. S of San ook on 
wy. 35, Jones et al. 20280 (FSU, GA, MISS, VDB). 
Pearl River Co., 6 mi. NW of Poplarville, Darwin et al. 
729 (MO, NO). Perry Co., 5 mi. E of Runnelstown, Jones 
9599 (MISS). Stone Co., near McHenry, U. of Miss. 
Forest Lands, Demaree 34056 (DS, SMU, USF). Wal- 
thall Co., N of Varnado, just m T line on LA 21, 
Kral 17394 (VDB). Wayne Co., 2.7 mi. E of county 
line, Hwy. 84, Jones 9724 (MISS). NoRTH a 
Beaufort Co., near Washington, Sargent 54 (US). Bruns- 
wick Co., 0.2 mi. W of ~ m n NC 904, Peng et 
al. 3886 (MO). Carteret Co., 0.25 mi. from Hwy. 70 o 
rd. to Harker's Island, Preig 15572 (DUKE, VDB). 
Columbus Co., 2.7 mi. N of Wananish, Fox & Whitford 
1889 (FSU, NCSC, SMU). Craven Co., 6.3 mi. NE of 
Jones Co. line on county rd. D Peng et al. 3832 
(MO). Cumberland Co., 1.5 m of Godwin, just E of 
Damat grab cd Fox eR NCSC, NY, SMU). 
Dare Co., jus of Manteo city pups nig US 64, 
Duke 54237 yee Duplin Co., (i of Pin 
Hook, Ahles 35840 (FLAS). e Co.. 8 mi 
of Lillington on taroen Sige at NC 210, Beal 2702 
(NSC). Hyde of Swan Quarter, Oosting 
30 (DUKE, B Cu Co., near Statesville, s.d., 
Hyanus s.n. (MO). Johnston Co., E of Clayton, near jct. 
of NC Hwys. 96 & 42, Radford 44388 (NCU). Jones 
Co., 0.4 mi. W of county rd. 1105 (Catfish f on 
Fo rest Service Rd. 172 (Mirey Branch Rd.) E-S E of 


64, icd d 39337 
anover Co., ` 


of Comford, di ng e 
i of Gra ib n ipd Creek, Beal 2521 DUK. 
NCSC). Pender Co., 1 mi. S of Ward's Creek on US 42 
Duke 6472 (NCU). Pitt i 400 yds. E of jct. of NC 
zm 11 & 903, Duke 54173 (NCU). Robeson Co., 
mi. E of Daystrom on rd. to dispar Ahles & 
Lese: 32962 (NY, VDB). Sampson Co., 0.9 mi. S of 
Halls Store, vicinity of Mill Pond, Ahles & Pone 33640 
(NCU). Scotland Co., 4.7 mi. E-NE of Johns on rd. to 
Maxton, Ahles 3700 (NCU). Tyrell Co., 1.5 mi. SE of 
Cross Landing, Pu pd kie (NCU, USF). SOUTH 
CAROLINA: Aiken Co., , HWR s.n. (US). Allendale 
, ca. 2 mi. W of Barton « on rid Rd. 23, Bell 5106 
(NCU). Pese Co., 0.7 mi. W of Olar on SC 64, then 
0.8 mi. S on dirt rd. (SW of Olar), Ahles 37683 (NCU). 
Beaufort Co., 5 mi. S of Hardeeville, Eyles 4400 (GA). 
Berkeley Co. 10 mi. N 


17, E of Osborn, Peng et al. 3908 (MO). Chesterfield 
Co., 3 mi. NW of Pati near SC 102, Radford 15873 
(NC U). Clarendon Co., i. S of Sandinia, Black River 
Swamp, Godfrey & leon 1486 (F, GH, MO, NY, TENN, 
US). Colleton Co., 4.2 mi. E of Smoaks, Raven 
ua reia ie Co., across lake from Hartsville, Smith 

NCU). Dorchester Co., 1.5 mi. N-NE of St George 
on US 15, Ahles & Haesloop 37776 (NCU). Horry Co., 
on SC Hwy. 9, 3 mi. NW of jet. US Hwy. 17, Dille & 
Dille 342 (MO). Jasper Co., 2.5 mi. N of SC 46 on US 

1, N of Hardeeville, Peng et al. 3973a (MO). Richland 
Co.,.7 mi. SE of Columbia, Smith 860 (GH). Florence 
Co., say at N‘limits of Scranton, Raven 20456 (DS). 
Ham .8 mi. NW of Yemassee on SC 28, Ahles 
& BU. 18351 (NCU). Horry Co., 2 mi. N of Little River, 


yí 


beside Hwy. 905, Duke 54115 (NCU). Jasper Co., 2.5 
mi. N of SC Hwy. 46 on US Hwy. 17, N of Hardeeville, 
Peng et al. 3973-a (MO). Kershaw Co., 4.5 mi. SE of 
Bethune, near Lynches River, Radford 27737 (NCU). 
Lee Co., 6 mi. W of Bishopville, near SC 34, € > 
Swamp, Radford 29437 (NCU). Marlboro Co., 
E of Wallace, near SC 90, Radford 15626 TOES 
Orangeburg Co., Eggleston 5025 (GH, MO, NY). Sumter 
Co., 15 mi. E of Mayeville, near SC 53 €: Lynches n 
Radford 27561 (NCU). Williamsburg Co., 3.5 m o 
Andrews, Kral 19168 (VDB). TEXAS: Hardin Co., : 
N of Gist, Cory 50923 (GH, NY, SMU). Jasper Jd 20 
mi. NW of Jasper, Correll 28573 (TEX). Montgomery 
= just W of Fostoria, Correll 33994 (GH, TEX). New- 
n Co., 1.5 mi. E of Kirbyville, State Forest No. 1, Cory 
19813 (DS, GH, NY, SMU, US). Tyler Co., ca. 3.5 mi. 
pe Warren, Hyatt Lake Estates, Coret & Correll 
6729 (TEX). VIRGINIA: Nansemond Co., S of Baines Hill 
SE Adams Swamp, Fernald & Long 13705 (C, GA, 
GH, MO, TENN). 


Ludwigia pilosa is a distinctive species in sect. 
Microcarpium characterized by being densely hir- 
tellous throughout; reddish brown leaf midveins 
especially abaxially; sepals elongate-triangular, 
creamy, and often tinged with pink, reflexed at the 
tips; and capsules subglobose to somewhat oblong- 
obovoid, with a pair of long bracteoles attached 
slightly above the base. The only species that has 
consistently been confused with L. pilosa is the 
recently named L. ravenii (Peng, 1984). The sim- 
ilarity in these two species is in aspect and in the 
vestiture. They differ in many other characters, 
notably floral parts and breeding systems. Lud- 
wigia ravenii is modally autogamous, with obscure 
greenish sepals, and all of its floral parts are sig- 
nificantly smaller than those of L. pilosa. Further, 
the seed surface cells are transversely elongate 
rather than subisodiametric. 

The flowers of Ludwigia pilosa, in contrast, 
are unable to undergo mechanical self-pollination. 
Their showy sepals and copious nectar, which is 
produced by the nectary disc, unfailingly attract 
various insect visitors such as ants, wasps, bum- 
blebees, honeybees, and moths to effect pollination. 
Ludwigia pilosa grows with diploid L. linifolia, 
L. microcarpa, tetraploid L. glandulosa, L. lan- 
ceolata, L. ravenii, L. sphaerocarpa, L. suffru- 
ticosa, and the hexaploid L. alata, which is closely 
allied to the tetraploid species group. It forms fertile 
hybrids with probably all tetraploid species and with 
L. alata when they grow together. Natural hybrids 
between L. pilosa and L. ravenii are presumably 
rare (L. ravenii is rare itself) and difficult to detect. 
An intersectional hybrid between L. pilosa and L. 
arcuata (n = 16; sect. Dantia) was found in Mobile 
Co., Alabama (Peng, 1988). All natural hybrids 
are intermediate morphologically, having vestiture 
similar to that of L. pilosa but shorter and other 


Volume 76, Number 1 
1989 


Peng 287 
Ludwigia sect. Microcarpium 


features diagnostic of the other putative parents 
(e.g., the winged fruits of L. lanceolata, elongate 
capsules of L. glandulosa, and additional features). 
Two such hybrids have previously been named: L. 
capitata B pubens Torrey & A. Gray (= L. pilosa 
x L. suffruticosa) and L. simulata Small (= L 
PE X L. lanceolata). Kral (1976) reported a 

tension of L. polycarpa to Alabama, based, 
in fact; on a natural hybrid population of L. pilosa 
x L. glandulosa. The only species that hybridizes 
extensively with L. pilosa and produces many hy- 
brid swarms or introgressed populations is L. sphae- 
rocarpa. Plants of these populations exhibit a vary- 
ing degree of intermediacy between the two species. 
They are discussed further under L. sphaerocar- 


pa. 

Hybrids between Ludwigia pilosa and the other 
tetraploid species or the hexaploid L. alata are 
readily synthesized in the experimental greenhouse 
and are quite fertile. It is generally difficult to obtain 
hybrids between L. pilosa and diploid species or 
the L. curtissii complex. Floriferous and vigorous 
hybrids of L. pilosa X L. linifolia and L. pilosa 
X L. curtissii, however, were obtained. Seed set 
was nonexistent or negligible in these hybrids. 


11. Ludwigia ravenii Peng, Syst. Bot. 9: 129. 
1984. TYPE: U.S.A. South Carolina: Berkeley 
Co., 0.1-0.2 mi. SW of jct. of Co. 16 and 
Co. 6, on Co. 16 W of Moncks Corner, scat- 
tered in roadside sedgy, sandy ditch, on loam 
densely covered by Sphagnum subsecunda- 
tum Nees, associated with Cyperus, Xyris, 
Eriocaulon, Paspalum, Panicum, Hyperi- 
cum, Polygonum, Rhexia, Liatris, Eupato- 
rium, Ludwigia linearis Walter, L. hirtella 
Raf., 9 Sep. 1982, Ching-I Peng 4402 (ho- 
lotype, MO-3041593; isotypes, B, BM, DAO, 
DUKE, F, FLAS, FSU, FTG, GA, GH, HAST, 
KYO, LE, MISS, MISSA, MO, NCSC, NCU, 
NY, P, PE, TAI, TAIF, TENN, TI, US, USF, 
VDB). Figure 55. 


Plants densely hirtellous throughout. Stems erect, 
usually well branched and leafy, (15-)35-90 cm 
tall, densely hirtellous, the hairs 0.3-0.6(-0.75) 
mm long; aerenchyma (if any) slightly developed, 
in lower stems. Stolons slender, several from the 
base, to 18 cm long, 0.6-1.5 mm thick, glabrous 
or hirtellous, occasionally bearing flowers and fruits, 
the leaves purplish, elliptic to rotund, 10-18 mm 
long, 6-14 mm wide, apex rotund to acute, base 
attenuate into a petiole 1-3 mm long. Cauline 
leaves pale green on both surfaces, narrowly lance- 


elliptic, 13-65 mm long, 4-15 mm wide, densely 


hirtellous on both surfaces, the hairs 0.2-0.5 mm 
long, apex acute, marem entire a à minute a 
dathodal glands, b ly winged 
petiole 1-8 mm long. S rue reddish purple, lan- 
ceolate to widely deltate, succulent, 0.25-0.45 mm 
long, 0.15-0.25 mm wide. Flowers many, in leaf 
axils, not congested. Sepals green on both surfaces, 
broadly ovate-deltate, 1.5-3 mm long, 1.4-2.1 
mm wide, densely hirtellous without, glabrous with- 
in, apex acuminate, margin entire. Petals 0. An- 
thers 0.3-0.35 mm long, 0.4-0.5 mm wide; fil- 
aments 0.7-1.1 mm long. Pollen grains shed as 
tetrads. Nectary disc greenish, raised 0.3-0.4 mm 
on ovary apex, 1.4-2.5 mm across, 4-lobed, gla- 
brous. Style 0.25-0.5 mm long, glabrous; stigma 
subglobose, 0.25-0.45 mm across. Mature cap- 
sules oblong-obovoid, obconical when young, (3-) 
4—5(-5.3) mm long, 2-3.5(-4) mm thick, densely 
hirtellous, pedicels 0.2-0.5 mm long. Bracteoles 
attached near capsule base, lanceolate or elliptic 
to very narrowly so, (1.5-)2-4.3 mm long, 0.25- 
0.9 mm wide, hirtellous. Seeds light brown, elliptic- 
oblong with slightly curved ends, 0.5-0.7 mm long, 
0.25-0.35 mm thick, pluriseriate in each locule, 
free, the surface cells elongate transversely to the 
seed length. Self-compatible and modally autoga- 
mous. Gametic chromosome number, n = 16. 


Distribution. Ludwigia ravenii occurs in wet, 
peaty habitats, such as ditches or margins of ponds, 
bogs, and swamps. It is found as scattered popu- 
lations in coastal southeastern Virginia, eastern 
North Carolina, and southeastern South Carolina 
and with somewhat disjunct populations in north- 
eastern Florida (Fig. 56). Flowering from July to 
September; fruiting from August to October. 


Specimens examined are cited in full in Peng 


(1984). 


This recently discovered species consists of scat- 
tered, morphologically very uniform populations. 
Owing to its densely hirtellous vestiture and relative 
rareness, it was previously not well understood and 
was confused with Ludwigia pilosa, the only other 
species of sect. Microcarpium that is consistently 
densely hirtellous; that species, however, is wide- 
spread and common. As was shown in the table 
comparing L. ravenii with related species (Peng, 
1984), L. ravenii is very distinct from L. pilosa. 

Ludwigia ravenii has small inconspicuous flow- 
ers with greenish sepals and short stamens and 
styles. Its anthers are firmly appressed to the stig- 
ma shortly after anthesis. These characteristics 
strongly suggest that it is modally autogamous. This 
was corroborated by my field study at the type 


288 Annals of the 
Missouri Botanical Garden 
M 
NI LÁ 
(3 
Pd 
— 


(ye 
MN Y n 


CEY. 
FEAS, SS 
Sy | 


ge 
r 


E 


FIGURE 55. Ludwigia ravenii (North Carolina, Sampson Co., Boufford & Wood 18898, CM).—a. Habit. — b. 
lower.— c. Fruit. —d. Seed. (Reprinted with permission from Systematic Botany.) 


locality (Peng, 1984): for two hours by the ditch 
where L. ravenii grew, I did not observe any insect 
visitors, whereas 0.5 mi. away in the same ditch, 
I witnessed a swarm of honeybees visiting and 
hovering over flowers of a population of L. pilosa. 

Ludwigia ravenii does not appear to grow ag- 
gressively. It starts to form stolons only late in its 
fruiting season (September to October). Unlike most 
other tetraploid members of sect. Microcarpium, 


these stolons are usually slender and short, resem- 
bling those of L. microcarpa. Consequently, the 
populations consist of scattered individuals instead 
of large colonies connected by stolons, as is typical 
in L. pilosa and other species. 

Live plants of Ludwigia ravenii were not col- 
lected until after the completion of my biosyste- 
matic study of Ludwigia sect. Microcarpium, so 
its genetic relationships with the other tetraploid 


Volume 76, Number 1 
1989 


Peng 289 
Ludwigia sect. Microcarpium 


FIGURE 56. 


Distribution of Ludwigia ravenii. 


members of the section remain obscure. The ex- 
tensive crossing data already available for other 
species clearly demonstrate, however, that pollen 
stainability and seed set are high in hybrids between 
any two tetraploid species, no matter how different 
morphologically they are (Peng, 1988). Circum- 
stantial evidence thus suggests that L. ravenii is 
almost certainly interfertile with other species of 
the tetraploid group of sect. Microcarpium. 


12. Ludwigia microcarpa Michaux, Fl. Bor.- 
Am. 1: 88. 1803. /snardia microcarpa (Mi- 
chaux) Poiret in Lam., Encycl. Suppl. 3: 188. 
1813. LECTOTYPE: U.S.A. South Carolina: 
Berkeley Co., Goose Creek, wet places, 1785- 
1796, A. Michaux s.n. (lectotype, P; pho- 
tograph, MO; here selected). Figure 57. 

Plants glabrous, varied in habit, ranging from 
erect, unbranched, slender herbs to being profusely 
branched with somewhat woody bases. Stems erect 


or ascending, rarely prostrate and rooting at the 
nodes, 5-60 cm tall, 0.4-1.5(-4) mm thick, the 
leaf bases distinctly decurrent, with or without 
aerenchyma at stem base. Stolons pinkish, several 
from base, slender, 4-15(-25) cm long, 0.4-0.75 
mm thick, the leaves broadly elliptic to suborbicu- 
lar, 2.2-6.6 mm long, 2.2-5.2 mm wide, petioles 
1.5-5 mm long. Cauline leaves obovate-spatulate 
or oblanceolate, sometimes narrowly oblance-ellip- 
tic, 4-17 mm long, 1.5-10 mm wide, glabrous, 
apex acute or mucronate, margin subentire, with 
prominent hydathodal glands forming minute teeth, 
minutely papillose-strigillose, the hairs ca. 0.05 mm 

ong, occasionally smooth, base attenuate into 
winged petioles 0.5-5 mm long; main veins on 
each side of the midrib 3-4, obscure adaxially, 
conspicuous and slightly raised abaxially; branch 
leaves sometimes remarkably smaller than main 
cauline leaves. Stipules reddish purple, lanceolate- 
deltate, succulent, ca. 0.13-0.15 mm long, 0.1- 

0.13 mm wide. Flowers axillary, usually not crowd- 


290 Annals of the 
Missouri Botanical Garden 


6 ] 
T ; QR $^ " i] 
«3 BA N " " 1 
3 DD t 
f aq Y "NO 2) 1 
Q 3 | 
M a ‘ S t 
(3 > Q AN 
Q Y i 
Eu un TU i x 
p d s. ww 
x of wa 
XS s 
D he 


FIGUR Ludwigia microcarpa. 
Carolina (Brunswick Co., Leon 
stolons. — b. 


All but a' from Florida (Monroe Co., Godfrey 63525, MO); a' from North 
ard & Moore 7579, .—a. Habit, showing erect stem. —a”. Erect plant with basal 
ortion of branch with flower and fruit.—c. 2 iew of fruit. —d, d'. Adaxial and abaxial views of 
stamen. —e. Stigma and style with pollen on stigma. — f. 


Volume 76, Number 1 
1989 


Pen 291 


Ludwigia sect. Microcarpium 


ed. Sepa is green with whitish, thickened base, 
ascending in flower and spreading in fruit, ovate- 
deltate, 0.9-2 mm long, 1-1.85 mm wide, gla- 
brous, apex acuminate, margin remotely minutely 
j| JM ETC. occasionally smooth. Petals 0. 
Ant .1-0.2 mm long; filaments translucent, 
0.4— o. 55 mm long. Pollen grains shed singly. Nec- 
tary disc greenish, nearly flat on ovary apex, 0.5- 
1.2 mm across, 4-lobed, these represented by 4 
greenish rhomboid spots, alternating with 4 smaller, 
rounded reddish scars of anther base on top of the 
mature capsule, glabrous. Style pale green, 0.25- 
0.4 mm long, glabrous; stigma green, subglobose, 
ca. 0.05-0.1 mm thick. Capsules obconical, 1- 
1.5 mm long, 1.4-1.9 mm thick, minutely puber- 
ulent, the hairs ca. 0.05 mm long, thin-walled, 
containing 6-20 seeds, these sometimes recogniz- 
able from outside as bumps, sessile or with pedicels 
up to 0.15 mm long. Bracteoles flanking capsule 
base, sublinear or narrowly oblong, 0.35-1.2(-1.5) 
mm long, 0.1-0.4 mm wide, usually with a swollen 
base, glabrous. Seeds dark reddish, ovate-oblong, 
0.5-0.55 mm long, 0.3-0.4 mm thick, the surface 
cells transversely elongate, glabrous or sometimes 
densely covered by strigillose hairs (waxes?). Self- 
compatible and modally autogamous. Gametic 
8. 


chromosome number, n = 


Distribution. Plants of Ludwigia microcarpa 
grow in roadside ditches, ponds and 
streams at the borders, low meadows, low areas in 
open woods, edges of swamp forests, brackish 


marshes, 


marshes, hammocks, and solution pits of limestone 
on marl prairies. This species occurs along the 
Atlantic coast of North and South Carolina and 
Georgia, and throughout the Florida peninsula. It 
extends westward across the panhandle of Florida, 
southern Alabama, reaching southern Mississippi, 
with a single population in southwestern Louisiana. 
To the north, L. microcarpa occurs through west- 
ern Georgia and central and western Alabama to 
southcentral Tennessee. Scattered populations are 
found in northcentral North Carolina and south- 
central Missouri (Fig. 58). Furthermore, L. mi- 
crocarpa occurs on several Caribbean islands, in- 
cluding the Bahamas, Cuba, and Jamaica. Flowering 
and fruiting throughout the year. 


Representative specimens examined. U.S.A 
E of Mobile on US ‘00, 
b Co., ca. 7 mi. en- 


). Bib 
terville by Co. 27, Kral 47928 (VDB). Calhoun Co., 5.5 


mi. W Piedmont io US 278, Kral 37736 de Cher- 
okee Co., 4 2 mi. = H n pp Eo 31769 (GA, 
NCU, SMU, VD. Chilto . 9 mi. N of 


Clanton and AL 22, Kral d d "Clarke Co., on 
Co. 19, N of Choctaw Bluff, Kral 35473 (NY, VDB). 


Conecuh Co., ca. 4 mi. NE of Lenox, Kral 40954 (VDB). 
Covington Co., 2 mi. W of Opp, Kral 20622 (VDB). 
Geneva Co., W side of Geneva, Kral 32026 (FSU, SMU, 
VDB). Houston Co, Chattahoochee State Park, Godfrey 
75953 (FSU). Morgan Co., ca. 4 mi. S of poma Kral 
33465 (C, SMU, VDB). St. Clair Co. . ca. 3 mi. NW of 
Asheville, by US 231, Kral 32361 (NCU, SUA VDB). 
FLORIDA: KE ane Co., pu 1938, Arnold & West 
s.n. (DUKE, FLAS). Bay Co., 1 mi. N of Panama City, 
FL 77, Godfrey d (MO). Broward Co., S of Wiles 
Road, 1.4 mi f US 441, ca. 3 mi. N of Margate, 
1964, Will s.n. vee FSU). Calhoun Co., banks of 
Chipola River, Wofaltha, Godfrey 57758 (FSU, GH). 


Charlotte Co., Punta Gorda city, on US Hwy. 41, ca. 1 
mi. S of jc US Hwys. 17 & 41, Peng et al. 4294 
(MO). Citrus Co., 5m mi. S BL Kral 7770 s 
US). Clay Co., Me arms on FL 


.Wo 

16, Peng et al. 4161 dor Colier Co., on FL Hwy. 82, 
1.2 mi. N of C-890; 5.7 mi. E of 
Peng et al. 4281 (MO). Columbia Co., Camp O'Leno, 
Beck 243 (FLAS). Dade Co., 25 mi. W of FL Hwy 

on US Hwy. 41, Peng et al. 421 7 (MO). Dixie Co., 4 
mi. NW of ¿Cros City, 1 s 56021 d DS, DUKE, 
> LL, pug I ES ENN). pi 

, FL H N n bud. 295, Peng e 

al "147 (MO). rub rons Pensacola, Godfrey 74408 
(FSU, NCU, MO). Franklin Co. 41.7 mi. W 


of La Belle, 
ertiction 63-1611 (FSU). aris Un N , 3 mi. W of 
Weeki Wachee Springs, Kral & Kral 7029 (FLAS, GA, 
GH, NY, USF). PEERS Co., 2.8 mi. N of FL Hwy. 
582, on E S of FL Hwy. 581, Peng et al. 4332 (MO). 
Holmes Co., 1 mi. of Wentville, Kral & Redfearn 
2885 (FSU, cH. NCU). Indian River Co., canal margins 
d mi. W of Vero Beach, Kral 4942 (FSU, VDB). Jack- 
on Co., Lake Seminole, N of Sneads, Godfrey & Houk 
61399 (FSU, NCU, SMU). Lake Co., 3.7 mi. S of Mas- 
cotte city limit, on county rd. 33 , Peng et al. 4166 (MO). 
ee Co., 3.8 mi. S of Gladiolus Dr. ER on San con 
Blvd. (865), Peng et al. 4288 (MO). Leon Co., 
lockonee River W of Tallahassee, Godfrey 73131 (FSU, 
MO, NCU, TENN). Levy Co., 0.5 mi. S of Gulf Hammock, 
Kral & Kral 6963 (FLAS, GA, CH, USF). Liberty Co., 
3.3 mi. E of FL 65 on FL 20, Boufford et al. 18407 
(MO). Madison Co., 2.6 mi. W of the Aucilla River on 
US 90, Boufford & Ahles ae (CM, MO). Manatee 
Co., US 301 2 mi. N of Sarasota, Raven 18657 (DS). 
Marion Co., Eureka, Ne cet 60141 (FSU, MSU). Martin 
Co., 4.3 mi. E of Okeechobee and Martin Co. line, on 
FL Hwy. 710, d Brady Ranch, between RR and Hwy., 
Peng et aL 4200 (MO). Monroe Co., along rd. to Watson 
ae Big Pine Key, Killip 44291 (bs, FLAS, GA, 
H, NY). Osceola Co., N of FL 532, ca. 1.5 mi. E of 
erat 4, Wunderlin & Sh huey 5829 (USF). Palm 
Co., NW of Loxahatchee in the Corbitt Wildlife 
CREW po Kral 5634 (FSU, SMU, VDB). Pasco Co., 
Aripeka, Godfrey 57244 (FSU). Pinellas Co., along FL 
, ca. 1 mi. E of its p. me 1 enda erson 63- 
en (FSU, JE, TEX). Polk Co., along US 192, just 
nside the county line, z^ e edge of : inti Hen- 
BL abe 63-1632 (FSU). P m Co. i. S of San 
Me Godfrey & Reinert ns (FSU). St. Johns Co., 
1.9 mi. N of jet. of FL . 207 & C-305, on FL Hwy 
207, ee et al. 4156 (MO). Sarasota Co., on FL Bw. 


292 


Annals of the 
Missouri Botanical Garden 


FIGURE 58. 


Distribution of Ludwigia microcarpa. 


72, 0.5 mi. E of W boundary of Myakka River State 
Park, Peng et al. 4313 (MO). Sumter Co., Bushnell, 14 
mi. S of Rt. 301, Sargent 6728 (SMU). Suwannee Co., 
Mayhaw ponds 5 mi. W of O'Brien, 1946, West & Arnold 
s.n. (FLAS). Taylor Co., on US Hwy. 98, 3.9 mi. W of 
jet. of US 27, 19 & 98 (Perry), Peng et al. 336 (MO). 
Wakulla Co., 2 mi. N of St. Marks, ss 18601 (DS, 
NCU, USF). Washington Co., Alligator Creek, Rt. 90, 
W of Chipley, Hood 2927 (F LAS) e G MEE Baker Co. 

Ivey's Mill Pond, Eyles 7183 (DUKE, CA). Bartow p. W 
at edge of broad ven runway from large spring S of 
Aubrey Lake, 5.6 mi., N12' E of Cartersville, Harris 

Duncan 12797 (DUKE, RU GA, U, SMU, 
TENN, US, USF, VDB). Camden Co., 6.6 mi S of Wood- 


bine on US 17, Raven 18703 (DS, NCU). Crisp Co., 
pond near Flint River, W of Cordele, Eyles 6612 (GA). 
Decatur Co., along Spring Creek below dam, lots 30 and 
31, District 21, Thorne & Davidson 17267 (GA). Early 
Co., Cohelee Creek, 2 mi. NW of Hilton, Duncan 6808 
(GA). Floyd Co., 4.7 mi. N5' W of Cave Springs, Venard 
& Dou 13114 (GA). Gordon Co., 1 mi. N of Fairmo 


f 
2 Co. line, S of Hinesville, Kral 18859 (VDB). Long 
Co., mi. SW of jct. of US Hwys. 301 & 25, and GA 
Hwy. 00 a on US Hwys. 301 & 26, along Altamaha River, 
Peng et al. 4127 (MO). McIntosh Co., 0.4 mi. 


Volume 76, Number 1 
1989 


Peng 293 
Ludwigia sect. Microcarpium 


jet. of Cox Rd. and GA Hwy. 251, on Cox Rd., Peng et 
al. 4133 (MO). Sumter Co., in pine barrens, Harper 471 
(F, GH, MO, NY, US). Walker Co., Chickamauga, s.d., 
s.c. 267 la (US). LOUISIANA: Calcasieu Parish, vicinity of 
Lake Charles, Md 26 (F, MO, NCU, NY). mis- 
, Hatcher's Spring in valley of Greer 
i. N of Greer, Steyermark 27987 s 
GH). MISSISSIPPI: "Clarke Co., Jackson Prairie, 1.5 m 

S.SW of Pachuta, Jones & Jones 14614 (FSU, CA. 
MISS, NCU, USF, VDB). Harrison Co., ca. 4 mi. N o 
Biloxi, Rogers 6780 (GH, NCU, NY). Jackson Co., 1 
mi. N of Franklin Creek Church, ca. 5 mi. NE of Orange 
Grove, Jones & Catchot 17421 (GA, MISS, TENN, 
USF). Jasper Co., 9 mi. N of Montrose, McDaniel 2631 
(NY, UNA). Smith Co., ca. 4.5 mi. of Bay Springs, 
Rogers 46028 (TENN). Wayne Co., Waynesboro, Pol- 
lard 1215 (F, GH, MO, NY, US). NORTH CAROLINA: 
Brunswick Co., roadside ditch along SR 1335, 3.3 mi. 
SE of SR 1333, Leonard & Moore 7579 (MO). Carteret 
Co., SE of Mullet Pond on Shackleford Banks, Anderson 


mh s 


Co. line; on 17, N of Hampstead, Peng et al 
4877 (MO) o., Wheeley's Ch., Batson 1074 
(DUKE). SOUTH CAROLINA: Allendale Co., pond 

26, 6.l mi. S of jct. with y ct. wi 


along railroad tracks, Moncks C 
dell 3406 (BOON, Charleston Co., 24.5 mi. N of 


wy. 41 on 7, vicinity of Bu ck Hill Rec- 
reation Area, S of MeCiellanvile, Peng et al. hus 897 (MO). 
Clarendon Co., Santee Reservoir, 2.8 m of Jo 


rdan 

| xe 31050 (CH, NCU, NY, VDB). Du Čo., 

W of Canadys on SC 61, Ahles & Bell 17888 
ey. "CA, NCU, USF). Dorchester Co., 2.9 mi. N of 
Rosinville on US 15, Ahles & Baird 54898 em 
Georgetow mi. N of Georgetown, Godfrey & 
Tryon 1043 (DUKE, F, GH, MO, NY, TENN, US). Horry 
Co., cart road across sand plain behind dunes, Myrtle 
Beach, Godfrey & Tryon 1163 (CAS, DUKE, F, GH, 
MO, NY, TENN, US). Orangeburg Co., Branchville, s.d., 
s.c. 2671c (US). TENNESSEE: Coffee Co., 3.5 mi. S 
Manchester, on US 41, Kral 26038 (FSU, VDB). Roane 
Co., shoreline of the Tennessee River, E of Thief Neck 
Isl., ca. 8 mi. SW of Kingston, Wofford & Dennis 51043 
(TENN, VDB). Union Co., stream bed of Little Barnes 
Creek, Underwood 385 (TENN). BAHAMAs: Great Abaco 
Co., along Abaco camp road, S of Marsh Harbour, Correll 
& aia 50684 er d cy E sl., along 
Forest Drive, about Marsh Harbour, 
Correll £ Meyer ji E FIG, NY ‘SMU, US). Grand 
Bahama Isl., Dead Man's Reef, 9 mi. E-SE of Mes End, 
Lewis 7110 (FTG, NY). CUBA. HABANA PROV.: Laguna 
x darn León 8724 (GH, NY). ISLE OF PINES: San 
Juan, Britton et al. 5527 (CM, F, GH, MO, NY). Las 
VILLAS: Santa Clara, Caibatten, at Delores, Ekman 16316 
S) 


dalena, Cayamas, Baber 4648 (POM). Cuba, Wright 


2554 (G). JAMAICA. SAINT ANN PARISH: Moneague, Prior 
902 (US); E side of the Black River T ca. l mi. 
N of Mountainside, Proctor 19694 (A, NY); a 
District, fac 33464 (IJ, RSA); s dn 27748 (IJ). 


Ludwigia microcarpa is the only diploid species 
of sect. Microcarpium that is apetalous and has 
more or less spatulate leaves. It has the smallest 
stature, leaves, flowers, and fruits and has the 
fewest seeds (ca. 10—20) per capsule of any species 
in the section. Occasionally it may be confused 
with plants of the “L. curtissi complex” (including 
a hexaploid, L. simpsonii, and an sopla, E 
curtissii), which have similarly spatulate leaves a 
turbinate fruits. It is, however, Lon maet A 
the “L. curtissii complex" by its minute capsules 
(1-1.5 mm long), which are shorter than or sub- 
equal to the sepal, its nectary disc of four flat 
greenish spots, and its reddish seeds. For detailed 
comparison, see Table 6 

Ludwigia microcarpa is uniform in reproduc- 
tive structures but variable in vegetative habit. The 


stems are, as a rule, erect and usually well branched 
but are sometimes single. The stems and branches 
are straight or slightly zigzag. In outline, the plants 


subequal, ascending branches throughout the nodes; 
or they are triangular or ovate when the branches 
are gradually shorter from lower nodes up. The 
stems are normally annual and herbaceous; the 
plants may perennate either by forming stolons or 
by sending new shoots up directly from the spent 
stem base of the previous year. Woodiness is seen 
in some Florida specimens, presumably as a result 
of a favorable climate. The following specimen from 
Leon County, Florida, has an exceptionally woody 
base about 8 mm thick: sandy alluvium bordering 
wet woodland, Ochlockonee River, W of Tallahas- 
see, Godfrey 73131 (MO). 

Most plants of Ludwigia microcarpa start to 
flower when young; it is common to find plants 5- 
10 cm high with flowers/fruits. The apetalous flow- 
ers are very small, with a tiny, greenish nectary 
disc from which a small amount of nectar is pro- 
duced. Mechanical self-pollination apparently pre- 
dominates in this species. In the experimental 
greenhouse, however, it 
crossed to the diploid L. linifolia and L. linearis, 
tetraploid L. glandulosa, L. lanceolata, L. poly- 
carpa, L. sphaerocarpa, and hexaploid L. alata 
and L. simpsonii; these combinations have resulted 


been successfully 


in the production of vigorous, floriferous hybrids 
(Peng, 1988). None set any seed. The F, hybrids 
between L. microcarpa and other petaliferous dip- 


294 


Annals of the 


Missouri Botanical Garden 


TABLE 6. 


Comparison of Ludwigia microcarpa with related species. 


Characters 


L. microcarpa 


L. simpsonii 


L. curtissii 


Chromosome number 


Habit 


Leaf 
Phyllotaxy 


Lateral veins (abaxial) 


Sepal length (mm) 
etals 


Anther length (mm) 
Filament length (mm) 


Nectary disc 
Width (mm) 
Elevated or flat 


Capsule 


Dehiscence 
Bracteole 


Length (mm) 
Width (mm) 


Seed 
r 
Surface cells 


Number / capsule 


n=8 


Erect 


Alternate 


Slightly raised, 
conspicuous 

0.9-2 

Absent 


0.1-0.2 
0.4-0.55 


0.5-1.2 
Nearly flat 


1-1.5 
1.4-1.9 
Ring-dehiscent 


0.35-1.2(-1.5) 
0.1-0.4 


Reddish brown 


n= 24 

Erect, ascending, or 
decumbent, occasionally 
prostrate and rooting 
at the nodes 


Alternate except often 
opposite or subopposite 
in seedlings or at lower 


nodes 
Not raised, usually obscure 


1.2-1.8 
Absent 


0.25-0.35 
0.55-0.8 


0.9-1.3 


Elevated as conspicuous 
bumps 


1.5- F4 2.5) 


1.5 
P oou dehiscent 


0.9-1.5(-2.5) 
0.35-0.85 


Light brown 
Transversely elongate 


Ca. 40-80 


n = 32 


Usually erect 


Alternate 


Not raised, usually obscure 


1-3 vestigial petals 
occasionally present 
0.3-0.55 


0.75-1(-1.25) 


0.9-1.6 


Elevated as conspicuous 
bumps 


(2-)2.5-4(-4.7) 
2-3(-3.5) 


Loculicidal-dehiscent 


1.5-3.5(-4) 
0.4-0.8 


Light brown 
Tranversely elongate 


Ca. 50-150 


loids were consistently petaliferous, suggesting that 
petal formation is controlled by a dominant gene. 
In other aspects, these F, hybrids were intermediate 
between their parents. Hybrids between L. micro 
arpa and species of higher ploidy level generally 
resembled the latter but tended to have smaller 
leaves and flowers. 
Ludwigia microcarpa grows with L. alata, L. 
curtissii, L. glandulosa, L. linearis, L. linifolia, 
. pilosa, L. simpsonii, and L. sphaerocarpa. 
Natural hybrids have been found between L. mi- 
crocarpa (n — 8) and both L. simpsonii (n — 24) 
and L. curtissii (n = 32), both of which resemble 
L. microcarpa morphologically and share a com- 
mon genome with that species (Peng, 1988). Since 
L. simpsonii and L. curtissii differ only in quan- 


titative characteristics, which often overlap, it is 
impossible to distinguish between L. microcarpa 
X L. curtissii and L. dae di X L. simpsonii 
without knowing the chromosome number, or with- 
out seeing the populations in m field. These hy- 
brids are usually readily recognizable by their erect 
stems; small and oblanceolate to spatulate leaves; 
apetalous flowers; elevated nectary discs; and mi- 
nute, obpyramidal ovaries that abort after anthesis. 
he corners of the ovary/capsule often appear 
winged to some extent, apparently as a result of 
the aborted ovaries, which fail to develop into seeds 
in the locules of such hybrid individuals. 
Hybrid populations between Ludwigia micro- 
carpa and L. simpsonii are apparently common 
in Florida, where the two species grow in the same 


Volume 76, Number 1 
1989 


Peng 295 
Ludwigia sect. Microcarpium 


habitats, often intermixed. These plants have a 
modal meiotic configuration of 8 bivalents and 16 
univalents (Peng, 1 

Confirmed hybrid populations of Ludwigia mi- 
crocarpa X L. curtissii were found in Martin Co., 
Florida, 4.3 mi. E of Okeechobee and Martin coun- 
ty line, on FL 710, at Brady Ranch (Peng 4202, 
MO). Here they occurred along with putative par- 
ents by a wide depression between the highway 
and a railroad. Plants of L. microcarpa grew along 
the swamp border, whereas plants of L. curtissii 
stood in deeper water but were not as abundant as 
the former. Hybrid plants were scattered between 
them. The hybrids formed 8 bivalents and 24 uni- 
valents at meiosis (Peng, 1988). 

Intersectional hybrid populations of Ludwigia 
microcarpa with L. palustris (sect. Dantia) were 
found in several counties of North Carolina, Geor- 


gia, and Florida (Peng, 1988). Their leaves were 


bopposite or opposite; their fl 
apetalous, with minute ovaries, and aborting after 
anthesis; and the stems were prostrate and rooting 
at the nodes to ascending, suberect, or both. The 
combination of alternate leaves with a more or less 
prostrate habit in North American Ludwigia is 
usually indicative of hybridization between plants 
of sect. Microcarpium and sect. Dantia. 


13. Ludwigia simpsonii Chapman, Fl. South. 
Florida: Manatee Co., 
ground, s.d., C. T. 
US-1384069; isotypes, GH, MO in part, US— 
2 sheets). Figure 


Ludwigia gn Helwig, Repert. Sp. Nov. 25: 53. 
28. TYPE: Cuba. Pinar del Rio: Remates, Ciénaga 
La Tumba, wet places, 16 June 1920, E. kman 
11304 (holotype, B, destroyed; isotypes, S—2 
sheets). 
Plants glabrous, erect, ascending, decumbent, 
or often prostrate and rooting at the nodes. Stems 
10-60(-75) cm tall, well branched, leaf base slight- 


ly decurrent. Stolons rarely seen, new shoots arising 


from erect stem base or nodes of the trailing main 
stem, ascending at first, soon erect. Leaves alter- 
nate, those on lower nodes often opposite or sub- 
opposite, shape varied: spatulate, oblanceolate to 
very narrowly oblanceolate, or pesi in extreme 
cases, Ho 20) mm long, (1-)3- 7(-11) mm wide, 

e or mucronate, margin subentire, with 
ida glands, base attenuate into narrowly 
winged petioles 2-10 mm long. Stipules reddish 
purple, narrowly ovate-deltate, succulent, 0.15- 


0.25 mm long, 0.1-0.15 mm wide. Flowers in leaf 
axils, rarely congested. Sepals green with whitish, 
accrescent base, ovate-deltate, ascending, 1.2-1.8 
mm long, 1-2 mm wide, apex narrowly acute or 
acuminate, margin entire. Petals 0. Anthers 0.25- 
0.35 mm long; filaments nearly translucent, 0.55- 
0.8 mm long. Pollen grains shed singly. Nectary 
disc green, raised mm on ovary apex, 
0.9-1.3 mm across, distinctly 4-lobed, glabrous. 
Style greenish, 0.25-0.4 mm long; stigma yellow- 
ish, subglobose, 0.15-0.25 mm thick. Capsules 
obconical, 1.5-2.5 mm long, 1.5-3 mm thick, 
comprising ca. 30-70(-100) seeds, glabrous or 
occasionally with minute, puberulent hairs ca. 0.05 
mm long, sessile or with pedicels up to 0.4 mm 
long. Bracteoles attached on opposite sides near 
the base of capsules, lance-elliptic, 0.9-1.5(-2.5) 
mm long, 0.35-0.85 mm wide, swollen at base. 
Seeds light brown or brown, ellipsoid, 0.5-0.6 mm 
long, 0.3-0.35 mm thick, the surface cells trans- 
versely elongate, glabrous, occasionally covered by 
wax (?) taking the form of minute, appressed or 
ascending hairs. Self-compatible. Gametic chro- 
mosome number, n — 24. 


Distribution. 
sandy, peaty ditches, open pineland swamps, edges 
of cypress swamps, tidal flats or nearby marshes, 
and limestone sinks, sometimes growing as weeds, 
throughout most of eastern and southern Florida, 
with somewhat disjunct populations in the eastern 
part of the Florida panhandle and southernmost 
Mississippi (Fig. 60). This species also occurs in 
Cuba and Jamaica. Flowering and fruiting all year. 


Ludwigia simpsonii occurs in 


Representative specimens examined. U.S.A. FLORI- 
DA: Bradford Co., New River, 1896, Hitchcock s.n. (F). 
Brevard Co., North Merritts Island, E side of Rt. 3, N 
of Wilsons Cimet. Shuey M1391 (USF). Broward Co., 
Fort Lauderdale, Small & Carter 1171 (FLAS, GH, NY, 
, Punta Gorda, Curtiss 6757 (E, GA, 
, Big Cypress, 6 mi. W of 
Miles City, Brass 15832 (FLAS, GH, both mixed with 
L. microcarpa). Dade , 4 mi. SW of Florida City, 
Thorne 14899 (USF, ised with L. microcarpa). Duval 
Co., near Jacksonville, Curtiss 930 (GA). Highlands Co., 
l mi. E of Highlands Hammock State Park, along 
364, Popenoe 1520 (FTG). Hillsborough Co., 4 mi. E of 
Dale Mabry on West Waters Ave., Raven 18650 (DS, 
mixed with L. microcarpa). Lake Co., Eustis, Nash 2136 
(E, F, FLAS, GH, MO, MSC, NCU, ND, NY, US). Lee 
Co., Myers, Hitchcock 117 (F, GH, NY, US). Manatee 
D Manatee, Tracy 7602 (CM, F, GH, MO, MSC, NY, 

S, W). Monroe Co., Watson Hammock, Big Pine Key, 
Killip 44454 (S, mixed with L. 
Beach Co., near Jupiter, Curtiss 5545 (C, E, FLAS, GA, 
GH, MO, MSC, NY, US). Pinellas Co., Fort DeSoto Co. 
Park, S of Gulfport, P m Key, Gulf of Mexico, Lakela 
et al. 26366 ( ns Co., St. Augustine, 1877, 
Reynolds s.n. (F, mixed in L. lance olata; GH, mixed 


296 Annals of the 
Missouri Botanical Garden 


Ie Y 

INS N V4 

N NY 
wW \ | Q 
y Y 

WY Y A 

ES > A 

Ls N l L= Y 


Ber E yi ‘ !) : Z 


FIGURE 59. Ludwigia simpsonii. All but a' from Florida (Collier Co., Peng et al. 4272, MO); a' from Florida 
(Collier Co., Peng et al. 4273, MO).—a. Habit, erect stems.—a’. Basal ascending branches with some opposite 


Volume 76, Number 1 
1989 


297 


Peng 
Ludwigia sect. Microcarpium 


e Ludwigia simpsonii 
O Ludwigia curtissii 


FIGURE 60. 


th L. microcarpa; NY). Sarasota Co., 0.5 mi. E of W 
boundary of Myakka River State Park, on FL Hwy. 72, 
Peng et al. 4313 (MO). Seminole Co., Sanford, Nash 
2278 A FLAS, GH, d NCU, ND, NY, US). Volusia 
Co., 6 mi. E of Geneva River on E side of St. John's 
River, Kral 5171 (SMU) Wakulla Co., St. Marks, 1843, 
Rugel s.n. (NY, mixed with L. el MISSISSIPPI: 
Harrison Co. Mississippi City, 1900, Tracy s.n. (MISS) 
CUBA. PINAR DEL RÍO: Remates, Ciénaga La Tumba, Ek 


Distributions of Ludwigia simpsonii (closed circles) and L. curtissii (open circles) 


man 11304 (S). Cuba, Wright 2554 (G). JAMAICA: near 
Black River, Harris 9935 (BM, C, F, NY, P, RSA, US, 
Z). St. Elizabeth Parish, along the Broad River, 1 mi. 
below the Slipe Road Bridge, Proctor 24836 (IJ). 


Ludwigia simpsonii is very variable in habit 
and leaf shape. It, L. curtissii (including L. spa- 
thulifolia), and L. microcarpa form a closely knit 


E 
—d. Flower. —d'. Partly dissected flower.—e, e'. 


—c. Flower bud. 


leaves.—b. Leaf.— 
—f. Capsule. —g. Dehiscent capsule. 


stamen. 


Adaxial and abaxial views of 


298 


Annals of the 
Missouri Botanical Garden 


polyploid complex, with the diploid L. microcarpa 
evidently having played a role in the formation of 
the other species (Peng, 1988). Its distinctiveness 
from the diploid L. microcarpa is readily perceived 
in capsule size, shape of nectary disc, seed color, 
and seed number per capsule. The characters sep- 
arating L. simpsonii and L. curtissii, however, 
are nearly always quantitative and often overlap. 
Raven & Tai (1979) indicated that the diagnostic 
characters separating L. curtissii and L. simpson- 
ii, such as capsule shape and size and leaf shape, 
are not correlated with chromosome numbers, and 
that the two species tend to converge in the ex- 
perimental greenhouse. For reasons of this sort, 
these two species have frequently been merged 
under L. curtissii in the recent literature (e.g., 
Long & Lakela, 1976; Raven & Tai, 1979; God- 
frey & Wooten, 1981; Wunderlin, 1982). By con- 
trast, basing conclusions on abundant fieldwork and 
greenhouse and chromosome studies, I (1988) ob- 
served that the difference in chromosome number 
is correlated with mature capsule size. All plants 
with n — 24 have capsules 1.5-2(-2.5) mm long, 
whereas those with n — 32 have mature capsules 
(2-)2.5-4(-4.7) mm long. The morphology of 
plants with n — 24 fits closely the description of 
L. simpsonii, and the morphology of n — 32 plants 
clearly corresponds to that of L. curtissii (including 
L. spathulifolia). Comparison of these two species 
and the diploid L. microcarpa is in Table 6. 

Although Ludwigia simpsonii and L. curtissit 
frequently are found in the same locality, they 
appear to be ecologically distinct and seldom occur 
side by side. Ludwigia simpsonii tends to grow 
along roadsides with other weeds in moist sandy 
soil. Ludwigia curtissii usually grows farther from 
roadsides, sometimes mixed with tall grasses or 
sedges, in black muck and often in deep standing 
water. 

Ludwigia simpsonii sometimes grows with L. 
alata, L. curtissii, and L. microcarpa. Proven 
natural hybrids are known only of L. simpsonii X 
L. microcarpa; they occur in several scattered 
counties in Florida (Peng, 1988). These hybrids 
showed a modal meiotic configuration of 8 bivalents 
and 16 univalents and 4-24% stainable pollen. 
Their ovaries aborted after anthesis. Ludwigia 
simpsonii may form natural hybrids with L. cur- 
tissii, but such hybrids would be virtually impos- 
sible to pick out without studying their chromo- 
somes and would probably be identified as one of 
their putative parents. Artificial hybrids between 
these two closely related species have been syn- 
thesized; they produced moderate numbers of seeds 


and had ca. 80-90% stainable pollen. They con- 


sistently showed a meiotic configuration of 24 bi- 
valents and 8 univalents, which indicated that chro- 
mosomes of the three genomes in L. simpsonii pair 
with homologues in L. curtissii, leaving the eight 
additional chromosomes in this species unpaired. 
Based on morphological features and crossing re- 
lationships, the octoploid L. curtissii was probably 
derived following hybridization between a petalif- 
erous, narrow-leaved diploid similar to L. linearis 
or L. linifolia and the hexaploid L. simpsonii 
(Peng, 1988). 

Intersectional hybrids of Ludwigia simpsonii 
and L. repens (n — 24; sect. Dantia) have been 
found growing intermixed with both putative par- 
ents. These plants exhibited few, if any (0-1), 
associations of chromosomes at meiosis (Peng, 
1988), which corroborated an earlier report by 
Schmidt (1967). The lack of chromosome pairing 
in hybrids strongly suggests that the three genomes 
of L. simpsonii are distinct from one another and 
are not homologous with any of the genomes pres- 
ent in L. repens. Judged by the fact that L. simp- 
sonii often has opposite or subopposite leaves near 
the base of the stem or at the seedling stage, with 
a somewhat ascending, decumbent, or occasionally 
prostrate habit, it seems likely that it originated 
from hybridization between the diploid L. micro- 
carpa, to which it is similar morphologically and 
with which it shares one genome, and a possibly 
extinct member of sect. Dantia with prostrate stems 
and opposite leaves. 

Plants from the following collections are excep- 
tionally fully prostrate and root at the nodes: 


U.S.A. FLORIDA: Collier Co., 13 mi. N of U.S. 41, on 
U.S. 29, in vast grassy field behind roadside ditches (Big 
Cypress Swamp), Peng 4268 (MO), 4273 (in part, MO). 
They grow side by side with erect plants but are 
otherwise normal, small-fruited L. simpsonii, ex- 
hibiting 24 pairs of chromosomes in meiosis and 
setting abundant fruits with viable seeds. 

Artificial hybrids between Ludwigia simpsonii 
and the diploids L. linearis, L. linifolia, L. mi- 
crocarpa, the tetraploids L. glandulosa, L. lan- 
ceolata, the hexaploid L. alata, and the closely 
related octoploid L. curtissii have been synthe- 
sized, although with difficulty in some cases. Except 
or L. simpsonii X L. curtissii, which is highly 
fertile, all other combinations had 3-20% stainable 
pollen and produced very few, if any, seeds. 


14. Ludwigia curtissii Chapman, Fl. South. 
U.S., 2nd edition, suppl. 621. 1883. TYPE: 
U.S.A. Florida: Brevard Co., ponds near Cape 
Malabar, July 1879, 4. H. Curtiss 922 (ho- 


299 


Volume 76, Number 1 Peng 
1989 Ludwigia sect. Microcarpium 
M y 
VÁ W 
vA Y , 
Y, Wy W / b b 
WZ, U/ "| 
EN Y N Uy , 
| | | Mf 
NN y N Z N Wp | 
Y 7, - 
E y UE | 4 
WN " | 
uA. 
Y $e) n 


NS v 
W Ye | L 
À fi Y Va 


Ñ 


fi 2 
> W Y VY" 
cà M Y 
EN j WZ 
<= X YW > 
7a 
Ag 
“W 


mm 
a (20 _ |, \ 
b,b’ 10 IN 


cde L2. 1 
ff ,1 | 
gh L3 ., 
FIGURE 61. bye igia curtissii (Florida: Martin Co., Peng et al. 4199, MO).— a. Habit. —b. Adaxial and abaxial 

views of leaf. —c. Cross section of stem.— d. Flower bud.—e. Flower, partly dissected. —f, f'. Adaxial and abaxial 


views of Eu — g. Capsule. —h. Cross section of capsule. 


300 


Annals 
rene EE Garden 


lotype, US-2630449; isotypes, BM, F, GH, 
M, MO—2 sheets, NY —3 sheets, PH, US). 
Figure 61. 


Ludwigia Qr ad Small, Man. Southeast. S es 
1506. 19 dtd ue ade Co., 
ieri aa iden of Perrine, 16 Jan. 1909, J. K. 
& J. J. Carter 2990 (holotype, NY). 


Plants glabrous. Stems erect or ascending at 
base, very rarely creeping, 15-75 cm tall, single 
to well branched, the branches sometimes very 
slender; leaf bases slightly decurrent. Stolons not 
seen, new shoots ascending from base of the per- 
sistent, defoliated main stem of the previous year, 
very rarely prostrate and rooting at the nodes. 
Leaves variable in shape, most frequently oblan- 
ceolate-spatulate, sometimes spatulate or oblan- 
ceolate to very narrowly so, sublinear in extreme 
cases, 10-25(-30) mm long, 1.3-8 mm wide, gla- 
brous, apex acute or mucronate, margin subentire 
with hydathodal glands, base attenuate into winged 
petioles 3-12 mm long. Stipules reddish purple, 
narrowly ovate, succulent, 0.2-0.3 mm long, 0.15- 
0.25 mm wide. Flowers in leaf axils, usually not 
congested. Sepals green with whitish, accrescent 
base, widely deltate or triangular, ascending, 1.5— 
3 mm long, 1.2-2 mm wide, apex narrowly acute 
or acuminate, margin entire. Petal usually lacking, 
1-3 vestigial petals occasionally seen in some 
ers, these yellow, narrowly elliptic, 1-2.5 mm long, 

-] mm wide, apex obtuse, base attenuate. An- 
thers 0.3-0.55 mm long; filaments yellowish, 0.75- 
1(-1.25) mm long. Pollen grains shed singly. Nec- 
tary disc green, raised 0.3-0.4 mm on the ovary 
apex, 0.9-1.6 mm across, prominently 4-lobed, 
glabrous. Style greenish, 0.35-0.65 mm long; stig- 
ma yellowish, subglobose, 0.25-0.4 mm thick. 
Capsules obconical, (2-)2.5-4(-4.7) mm long, 2- 
3(-3.5) mm thick, glabrous, occasionally remotely 
minutely puberulent, the hairs 0.05-0.07 mm, 
pedicels 0.1—0.5 mm long. Bracteoles attached on 
opposite sides near base of capsule, narrowly lan- 
ceolate, lance-elliptic or oblong-linear, 1.5-3.5 
(-4) mm long, 0.4-0.8 mm wide, swollen at base. 
Seeds light brown, ellipsoid, 0.45-0.6 mm long, 
0.3-0.4 mm thick, the surface cells transversely 
elongate, glabrous, occasionally covered by waxe 
(?) that appear also like appressed or ascending 
hairs. Self-compatible. Gametic chromosome num- 


un 


ber, n — 
Distribution. Ludwigia curtissii grows in pine 

savannas and flatwoods, marshes, ponds, stream 

banks, sandy, peaty swales, limestone prairies, and 


in solution pits in limestone; it is restricted to the 
Bahamas and the peninsula of Florida (Fig. 60). 
Flowering and fruiting all year. 


Representative specimens examined. U.S.A. FLORI- 
DA: Brevard Co., near Eau Gallie, Indian River, Curtiss 
928 (FLAS). Charlotte Co., 3.5 mi. NW of Port Charlotte, 
Godfrey 65356 (DS, DUKE, FLAS, FSU, LL, MSC, 
NCU, US, USF, VDB). Collier Co., Alligator Alley, picnic 
area ca. 5 mi. W of Miles City, Avery 2039 (FLAS, 
mixed with L. simpsonii, FTG). Dade Co., Everglades, near 
Royal Palm Hammock, Small et al. 6644 (FLAS, GH, 
NY, US). DeSoto Co., near Punta Gorda, Bright 4541 
(CM). Franklin Co., St. George Island, Godfrey 71148 
(FSU). Glades Co., 4.4 mi. SE of jet. of FL Hwy. 29 with 
US Hwy. 27, Raven 18679 (DS, NCU). Hendry Co., 6 
mi. S of La Belle, Godfrey et al. biet (FSU, GH, SMU). 
Hernando Co., Choocochee Hammock, S of Br eo 
1921, Small et S s.n. rs NY). Hillsborough Co., 
Memorial Highwa 1.5 mi. from FL Rt. 580, NW 
of Tampa, Lakela 35261 (FSU, GH, NCU, SMU, USF). 
Lee Co., 5 mi. N-NW of Ft. Myers, Kral 7574 (FLAS, 
GH, US, USP). Manatee Co., 

(MO, US). Martin Co., rd. be 
aikin Dickinson State ades Correll et al. 49914 (FTG, 
NCU). Monroe Co., 3.8 m of Dade-Monroe county 
line, near FL Rt. 94, Godfrey et al. 63519 (FSU, MO). 
, W side of Big Pine Key, Killip 41573 (F, 


5636 (FSU, d. Pinellas Co., 4 mi. E of encajes 
Kral 7452 (FLAS, FSU, es ith L. palustris, 6A, 
GH, US qe oti Co., of V — Kral 749 
(FLAS, FSU, GH, VDB). a Co., 5 mi. W of New 


Smyrna, Kral 18457 (VDB). BAHAMAS: South Andros, 
ca. mi. SW of Congo Town airstrip, Correll et al. 
50247 (FTG, IJ, MO) Grand Bahama, Pelican Lake 
Area, Correll & Popenoe 51315 (MO) 


Ludwigia curtissii and L. simpsonii are both 
highly variable in habit and leaf shape and are 
often difficult to distinguish; the only consistent 
distinguishing character is the size of mature cap- 
sules. These two species are unique in sect. Mi- 
crocarpium in having loculicidal capsules and in 
having, along with the diploid L. microcarpa, more 
or less spatulate cauline leaves. Their capsules de- 
hisce by four longitudinally lenticular slits opposite 
the loculi. When dehiscence is complete, the cap- 
sule is split into four basally united parts topped 
with persistent but slightly shriveled sepals. This 
type of capsule dehiscence is highly specialized and 
has an anatomical basis (Peng & Tobe, 1987). 
Ludwigia curtissii and L. simpsonii are also dis- 
tinct in that they are the only species in sect. 
Microcarpium that rarely produce true stolons. 
From the perennial base of the previous-year stem 
they form new shoots that ascend at first and soon 
become erect. 


Volume 76, Number 1 
1989 


Peng 
Ludwigia sect. Microcarpium 


Some populations of Ludwigia curtissii from 
the Everglades of southern Florida and the Ba- 
hamas tend to have larger capsules (3.5-4.7 mm 
long) and were named L. spathulifolia by Small 
(1933). As their variation pattern closely parallels 
that of L. curtissii s. str., and they are octoploids 
with n — 32, they are here considered as variants 
of the slightly rdg fruited L. curtissii that do 
not warrant taxonomic recognit lon, as was sug- 
gested by Long & Lakela (1976), Raven & Tai 
(1979), Godfrey & Wooten (1981), and Wunder- 
lin (1982). 

Ludwigia curtissii co-occurs with L. alata, L. 
linifolia, L. microcarpa, and L. simpsonii and 
hybridizes with the diploid L. linifolia and L. mi- 
crocarpa, forming F,s that form flowers but not 
seeds. Ludwigia curtissii X L. microcarpa ex- 
hibited 8 bivalents and 24 univalents at meiosis. 
The pollen stainability was 1676 (Peng, 1988). As 
discussed above, natural hybrids between L. cur- 
tissii and L. simpsonii probably occur but would 
be difficult to recognize. Like L. simpsonii, L. 
curtissii forms intersectional hybrids with L. re- 
pens (n — 24, sect. Dantia) in the field. Such 
plants are vigorous and somewhat prostrate or sub- 
erect; they have alternate and opposite or subop- 
posite leaves, but do not form viable seeds. 

n the experimental greenhouse, artificial hy- 
brids have been synthesized between Ludwigia 
curtissii and the diploid L. linearis and the tet- 
raploids L. glandulosa, L. lanceolata, L. pilosa, 
L. polycarpa, L. sphaerocarpa, and L. suffruti- 
cosa. A few seeds were occasionally produced by 
some of the octoploid X tetraploid hybrids. 

Compared with Ludwigia simpsonii (n — 24), 
L. curtissii (n = 32) is generally taller, more 
robust, more often erect, tends to have narrower 
and longer leaves, and nearly always has larger 
flowers and capsules. Further, the leaves of L. 
curtissii are nearly always alternate throughout, 
and the flowers occasionally have one to three 
vestigial petals. As these two species have three 
genomes in common (Peng, 1988), it is apparent 
that L. curtissii originated following hybridization 
between L. simpsonii and a diploid, petaliferous, 
erect species with narrow and alternate leaves, 
larger flowers, and longer capsules. The likely can- 
didate for such a plant is only to be found in L. 
linearis, L. linifolia, the Cub 
or their progenitors. Indeed it is even possible that 
more than one of the above species hybridized with 
L. simpsonii in the past to form the present-day 
L. curtissii. If this is true, the morphological vari- 
ability observed in L. curtissii is expected 


LITERATURE CITED 


ANDERSON, E. 1948. Hybridization of the habitat. Evo- 
lution 2: 1-9. 

Asal, Y. 1970. On Ludwigia linearis Walter c6 
introduced to Japan as an alien weed. J. Jap. Bot 
45: 371-318. 

BRATSEVA, G. M. 1969. Palinologicheskie issledovaniya 
verkhnego mela i paleogena Dal'nego Vostoka. Akad. 


A 
CARLQUIST, S. agraceae, 
with notes on alternative modes of photosynthate 
movement in are’ ase woods. Ann. Missouri Bot. 
Gard. r 
— 1982. Te anatomy of Onagraceae: further 
ce root anatomy; significance of vestured pits 
and allied MORES in dicotyledons. Ann. Missouri 

Bot. Gard. 69: 755-769. 
CasTELLS, A. C. DE, W. T. ORMOND, M. C. PINHEIRO & 
M. T. DA Sia 1979. nds dos hidatodios e 
sua importancia no complexo Ludwigia L. Pin (3 
ae). Arch. Jard. Bot. Ri : 5- 

Cu A. W. 18 
States. Ivison, Blakeman, Taylor o., New k. 
1892 ora of the Southern United States. 
Revised reprinting of 2nd edition. American Book 
Co., New York. 
CLEWELL, A. F. 1985. Guide to the Vascular Plants of 
the Florida Panhandle. 


Florida State Univ. Press, 
wwe 
Duke, J. A. 1955. Distribution and speciation of the 
pu "S = in North Carolina. J. Elisha Mitchell 
oc. 71: -269. 
abor S. 1816-1821. A Sketch of the Botany of 


South ibi and Georgia, Volume I. J. R. Schenck, 
Charle 
ENGELMANN, E & A. Gray. 1845. Plantae Lindhei- 
rianae. Boston JI Nat. Hist. 5: 210-264. 

Eype, R. H. 19 Reproductive structures and evo- 
lution in Ludwigia (Onagraceae). I. Androecium, 
placentation, merism. Ann. Missouri Bot. Gard. 64 

4-655. 


1978. Reproductive structures and evolution 
in Ludwi ia (Onagraceae). II. Fruit and seed. Ann. 
Missouri Bot. Gard. 65: 656-675 

1981. Reproductive structures and evolution 
in Ludwigia (Onagraceae). III. Vasculature, nectar- 
ies, conclusions. Ann. Missouri Bot. Gard. 68: 379- 
412 


FERNALD, M. L. & L. GRISCOM. 
rs in southeastern Virginia. Rhodora 37: 
9. 


1935. Three days of 
7: 167- 


s R. K. & J. W. WoorEN. 1981. Aquatic and 
wetland plants of southeastern United States. The 
University of Georgia Press, Athens, Georgia. 

GONZALES GUZMÁN, A. E. 1967. A epi ew acu i xd 
on the Upper Los Cuervos and Mira ormatio. 

pes and Middle Eocene; Tibü area, ‘Colombi a). E. 

n. 


— 


W. M. Kein. 1960. d x 
of meiotic chromosomes of six genera in the 
graceae. Aliso 4: 505-521. 

66. Catalogus Plantarum Cubensium 
Ex hibens Collectionem Wrightianam Aliasque Mi- 
nores ex Insula Cuba Missas. G. Engelmann, kaikis. 


302 Annals of the 
Missouri Botanical Garden 
HELwic, B. PRAGLOWSKI, J., J. J. SkvarLa, P. H. m EN & J. W. 


1928. Wi N nonnullae cubenses. 

Repert. Spec. Nov. Regni 53 

A R.C. 1 The Ma due and systematics 
of Onagraceae: leaf anatomy. Ann. Missouri Bot. 
Gard. 69: 770-803. 

Raven & P. C. Hocu. 1982. Peren- 
nation in Epilobium (Onagraceae) and its relation to 
classification and ecology. Syst. Bot. 7:379-404 

KraL, R. 1976. Additions to some notes on the flora 
of the southern states, particularly Alabama and mid- 

e Tennessee. Rhodora 78: 438-456. 

Ku RABAYASHI, M., » H. LE wis & P. H. Raven. 


J. Bot. a 1003-1026. 
Lamarck, J. B. A. P. M. DE. 1789. Bg Se Mé- 
odique, es 3, Pt. 1. Panckoucke, Paris 


——. | Vp e Méthodique, Volume 3, 
Pt. 2. Panckoucke, 

LinnaEus, C. 1753. edo Soie lst editi 

Lonc, R. W. € O. LAKELA lora of Tropical 


orida. E of Miami Press, cin Gables, Florida. 


MicHAUX, A. 03. Flora ae ore Americana. Caroli 
Cr de O Pari and Stras 
Mu UNZ, P. 9 a in  Onagracea XII. A re 


es of Jussiaea. Dar- 


Studies in oun eae— XIII. The 
American bep of Ludwigia. Bull. Torrey Bot. 
Club 71: 152-16 
cd N. Amer. Fl. II. 5: 1- 


OSADA, T. 1976 Coloured illustrations of und 
plants of Japan. Hoik bl. Co., Os 
Penc, C. I 984. Lu 


s ravenii i (Onagraceae) a a 

new species from the Coas n of the sout 

eastern ae = Sys Bo t. 9: '129- 132. 

1986. ew o in Ludwigia sect. 

Mic Pec ds Dn Ann. Missouri Bot. 

Gard. 73: 490. 

. 1988. The biosystematics of Ludwigia sect. 

Microcarpium (Onagraceae). Ann. Missouri Bot. 

Gard. 75: 970-1009. 

& H. Tose. 1987. Capsule wall anatomy in 

relation to capsular dehiscence in Ludwigia sect. 
nue (Onagraceae). Amer. J. Bot. 74 
1102-111 

ib E L. M. 1814. Encyclopédie Méthodique. Suppl. 

. 2. H. Agasse, Paris. 


NowICKE. 1983. Onagraceae Juss.: Fuchsieae L./ 
2 e L. World Pollen and oe Flora 12: 1- 


Proctor G. R. 1982. More additions to the flora of 
naica. J. Arnold Arb. 63: 199-315 

Bd T. P. & E. Zarpini. 1987. The sys- 
tematics and evolution of Ludwigia sect. Myrto- 
carpus sensu lato Mises Monogr. Syst. Bot. 
Missouri Bot. Gard. 19: 1-12 

e Old World species. of Lud 

wigia t (including al with a synopsis of the 

genus TE eae). Reinwardtia 6: 327-427. 

^ survey of y dapes biology in 

M en New Zealand J. Bot. 17: 575-593. 
& AVEN. 1976. The genus Epilobium 

in Australasia: a systematic and evolutionary su. 

New Zealand Dept. Sci. Industr. Res. Bull. 216: 

321. 


& W. Tal. 1979. Observations of chro 
somes in Ludwigia (Onagraceae). Ann. Missouri Bot. 
Gard. 66: 862-879. 

Rouse, G. E. 1962. Plant microfossils from the Burrard 
Formation of — British Columbia. Micropa- 


[1868- 11873: Flora ee Imp. 
“La Antilla,” deCacho-Negrete, Hav 

ScuMipT, C. L. 1967. biosystematic pua of Lud- 
wigia sect. Dantia dip (rA Ph.D. Thesis. 
Stanford Univ., Stanford, Californ 

SHORT, C. W. € R. PETER. 1836. A second supple- 
end catalogue of the plants of Kentucky. Tran- 
sylvania J. Med. Assoc. Sci. 8: 575-582. 

SMALL, J. K. 903. Flora of the Southeastern United 
States. Published by the author, New Yor 

1933. Manual of the S t 
lished by the author, New York. 

Tose, H. € P. H. Raven. 1986. Evolution of p 

J. Bot 


En anthers in Onagraceae. Amer. 
488. 


Flora Pub- 


1838- 1840. A Flora of North 
y & Putnam, New York. 

AL 88. Flora Caroliniana. J. Fraser, London. 
WUNDERLIN, R. P. suide to the Vascular Plants 
of Central Florida. Univ. Press of Florida, Tampa 
and i. cities, Florida. 


73 
TORREY, 1 "n A. GRAY. 


TWO NEW SPECIES OF THE 
LUPINUS LANATUS COMPLEX! 


Ana Maria Planchuelo? and 
David B. Dunn? 


ABSTRACT 


The Lupinus lanatus complex is a group of closely related species that grow in southeastern Brazil and northeastern 


herbaceous, perenn 
long or longer, and lar 


Argentina. They are 
or three-foliate, stipules 3-7 cm 


ial or occasionally biennial plants with basal leaves in the juvenile plant simple 


e flowers 15-20 mm 


long having the banner glabrous and 


well reflexed from the wings. Two species, L. magnistipulatus and L. setifolius, are described for the first time. 


This group of species appears to be closely re- 
lated and can be recognized by the large stipules 
3-7 cm or more long and the presence of some 
simple or trifoliate leaves at the base of the juvenile 
plants. Completely developed leaves from adult 
plants have five or more leaflets. They are all 
herbaceous perennials or short-lived perennials, or 
occasionally function as biennials. The flowers are 
commonly 15-20 mm long and glabrous, with the 
banner well reflexed from the wings and the sides 
turned back. The calyx cup tapers into the pedicel 
without the gibbous enlargement that is commonly 
present in other lupines. 

We are suggesting that this group of species 
forms a second lineage derived from L. para- 
guariensis, which is thought to be the origin of 
the group, since it possesses morphological char- 
acteristics reminiscent of the group, including that 
the juvenile leaves and the first leaf produced on 
the lateral branches are simple. Planchuelo & Dunn 
(1984) described L. paraguariensis as showing a 
transitional stage between simple- and compound- 
leaved species. 


Lupinus lanatus has floccose lanate indument 
on both sides of the leaflets (see Fig. 1A). Vege- 
tatively it has been described as forming perennial 
mats up to four feet in diameter in sand dunes, 
with new growth developing seasonally from the 
tips of the stems which survive under the sand. 
Lupinus magnistipulatus, described in this paper, 
is similar to L. lanatus in the perennial condition 
but has less lanate hair coating (see Fig. 1B), and 
as its name suggests, the former develops a much 
larger free tip on the stipules. Lupinus setifolius, 
also described in this paper, appears to be biennial. 
The large free tip of the stipules and the loss of 
the lanate indument (see Fig. 1C) suggest the latter 
as a derivative from L. magnistipulatus. 

The species L. multiflorus, L. albescens, and 
L. aureonitens, treated by Planchuelo & Dunn 
(1984), are relatives of this complex. They have 
caespitose perennial habit and a narrow free tip of 
the stipules like L. lanatus but develop a much 
denser indument that grades from densely short- 
lanate to densely sericeous, completely hiding the 
leaf surface. 


KEY TO THE SPECIES OF LUPINUS LANATUS COMPLEX AND RELATIVES 


P 


p 


nts. 
2 Wing tips with an upturned toothlike tip. 


P 
3a. Leaves primarily cauline; stems erect, branching above 


3b. Leaves primarily basal; stems with branches primarily basal 


Basal leaves and first leaf of each lateral branch simple, the others palmately compound .... L. p 


paraguariensis 
escription in Planchuelo & Dunn, 1984). 


im leaves of mature plants palmately compound, exceptionally with few basal simple leaves on juvenile 


L. albescens 
(description in Planchuelo & gens 1984). 
ureonitens 
(description in Planchuelo & ela 1984). 


' The authors wish to express their appreciation to Dr. John Dwyer for reviewing the Latin descriptions, to Mr. 
John Myers for helping with the illustrations, and to the curators of DS, K, LIL, MO, NY, SI, UC, UMO, and US 


ie Es loan of the materials. 


nsejo Nacional de Investigaciones Sisi R Técnicas (CONICET), Argentina, Facultad de Ciencias Agro- 


oscar Casilla de Correo 509, Córdoba, Argen 


3 University of Missouri-Columbia, 202 Tucker Hall Columbia, Missouri 65211, U.S.A. 
ANN. Missouni Bor. GARD. 76: 303-309. 1989. 


304 


Annals of the 
Missouri Botanical Garden 


E l. Indument of: —A. L. lanatus, leaflet (x100).— 
aoe leaflet (x 100). 


2b. Wing tips rounded, without a toothlike tip. 


B. L. magnistipulatus, petiole (x 200).— 


eaflets mostly broadly obovate or na Ne with the tips rounded. Stipules foliaceous, 


with free portion ovate-lanceolate, 8-15 m 


a. Leaflets thinly floccose-lanate on both dee Plan lived perennials ............. 
a sides except for setaceous hairs on 

asi ng some areas of the 

Leaflets all narrowly ellipt ic to narrowly oblanceolate his tips acute or em Stipules not 

foliaceous, with free portion lance-attenuate, 

6a. flet 


NN 
a 


. L. magnistipulatus 
ios midrib, pins veins, an 
argins. Bienn L. setifolius 


= ceptionally 6 mm 


ets abundantly floccose-lanate or thinly lanate, má larger 7-11 cm pil Stipules 3-8.5 


- Stipules 1 


1. Lupinus lanatus Benth. in Mart., Fl. Brasil. 
15(1): 16. 1859. TYPE: Brazil. Rio Grande do 
Sul: Herb. Imp. Bras. 1511, not seen. Fig- 


ure 


Plants perennial, herbaceous, 25-60 cm tall, in 
clumps to 1.5 m wide, possibly rhizomatous or the 
stems arising from underground branches; stems 
fistulose, 8-12 mm diam., first internode of 
branches 4-9 cm long, others 1-2 cm long, cov- 
ered with soft tangled lanate hairs 4-5 mm long; 
stipules 3-8.5 cm long, commonly green like the 
leaflets, the free portion lance-attenuate, 3-5(6) 


m long, lanate as stems; 
leaflets 5-7, oblanceolate, the tips obtuse or round- 
ed, densely or thinly lanate on both sides, thinner 
above, the largest 7-11 cm long, 10-20 mm wide. 
Peduncles 2.5-8 cm long, fistulose, angular and 
lanate as the stems; racemes 15-50 cm long, flow- 
ers scattered, the rachis fistulose and lanate as 
stems; bracts caducous, lance-attenuate, 12-22 
mm long, lanate dorsally, glabrous ventrally; brac- 
teoles 3-5 mm long, lanceolate to lance-attenuate, 
attached just below the lips of the lateral sinuses 


.5-2.6 cm long. Calyx not enlarging as fruit deve 
(de 


. lanatus 
cm long, rarely to 7 cm long. 

ops mul oa 
escription in Planchuelo & Dui 1984). 


oliaceous as fruit develops ... 1. 
arger 4-5 


of the calyx; pedicels 1-2 mm long at anthesis, 

mm long in fruit, lanate. Calyces lanate out- 
udo. glabrous within, the base tapering into the 
pedicel, lower lip 13-17 mm long, 3.2-4.5 mm 
wide, often enlarging and foliaceous as fruit de- 
velops, trifid, central tooth 2.5-4 mm long, lateral 
teeth 2-3 mm long, arching outward, upper lip 8- 
13 mm long, bifid, the slit 5.6-9 mm deep, the 
lobes 2.3-2.5 mm wide, the lips connate 3 mm; 
banner glabrous, oval-ovate, 15-17.5 mm long, 
including the 2-2.5-mm-long claw, 12-13.5 mm 
wide, appressed 4-6 mm, reflexed 12-13 mm, 
reflexed/appressed ratio 2.4, the angle 130°, the 
tip bluntly rounded; wings arcuate, 15.5-18.5 mm 
long, 6.5-7.5 mm wide, the claw 3.5 mm long, 
lobe above the claw 1.6-2.2 mm wide; keel arcuate 
glabrous, 4.5 mm wide in the middle, the tip arcing 
back in a continuous curve, no definite angle, 80— 
85°; ovules 8. Legumes ascending to erect, 5-9 
cm long, 1-1.5 cm wide, shaggy lanate. 


The floral parts, especially the calyx and the 
banner, are remarkably similar to those of L. para- 
guariensis. The Argentinian specimens and Ram- 


bo 28195 from Rio Grande do Sul, Brazil, had 


Volume 76, Number 1 Planchuelo & Dunn 305 
1989 Lupinus lanatus 


FiGURE 2. Illustration of typical structures of Lupinus lanatus. — A. Upper part of the plant with inflorescence. — 
B. Stipules detached at the node. — C. Lateral view of the left side of the flower. —D. Calyx, cut at the left lateral 
sinus and opened so that the inside surface shows. —E. Banner petal flattened, dorsal view. —F. Wing petal.— C. 
Keel petals, enclosing the staminal tube and the pistil, with the mean number of ovules drawn. 


306 


Annals of the 
hp Botanical Garden 


unusually large stipules, some of them 6 mm wide. 
All the others have most of the stipules under 5 
mm wide. Although all the morphological charac- 
teristics of the large-stipuled specimens are those 
of the species, the enlargement of the free tip of 
the stipules suggests relationship with L. magni- 
stipulatus and even crossing between the two 
species. 

Lupinus lanatus occurs in the southernmost 
states of Brazil and in Misiones, Argentina. The 
plants are reported growing in open places and 


dunes. 


Representative specimens examined. ARGENTINA. 
MISIONES: Campo Grande, Bertoni 638 (LIL); Dep. Con- 
ce idis CUL dE Schulz 6965 (LIL); Dep. San Ja- 

Par collector's name illegible, Inst. Miguel 
Lillo ¡68108 (LID, BRazIL: locality unknown, Sello 3032 
(NY, UC). RIO GRANDE DO SUL: Viera, near Rio Grande, 
Archer 4303 (DS, K, NY, RB); S. Leopoldo, in dunes, 
Henz 33899 (NY); Jacaré, Rambo 38545 (MO); Apar- 
ados da Serra, próximo Capela das Ausentes, Pereira 
6444 (NY); Nonvaí, Municipio de Sarandi, at flumen Alto 
Uruguai, Rambo 28195 (LIL); Pareci Velho p. Cai, Ram- 
bo 43841 (LIL, US). 


2. Lupinus magnistipulatus Planchuelo & 
Dunn, sp. nov. TYPE: Brazil. Santa Catarina: 
Campos dos Padres, 1,900 m, R. Reitz 2377 
(holotype, SI; isotype, RB; photo, UMO). Fig- 
ure 3. 


Plantae perennes, 17-35 cm altae; caules erecti, fis- 
tulosi, spicata multis pilis patentibus, 4-7 mm longis or- 
tipulae virides foliaceae, 4-7 cm longae, parte 

.2-2 cm longa, parte libera divergenti et ascen- 
denti, 2-5 cm longa, 8-15 mm lata, lanata; petiolis 4- 
8.5(11) cm longi, lanati folia (3)5- t foliolis 
lanatis, oblanceolatis, apice rotundatis, 5-7 cm longis, 
15-20 mm latis, complanatis; racemus major 20-3 
longus, floribus ir bractae foliolaceae, lanceolatae, 
apice acuminatae, 12-23 mm longae, 2.5-4 mm latae, 
in dorso Testa bracteolae 3.5-6 mm longae, 0.5-1.2 
mm latae; flores calycibus in dorso lanatis, ventraliter 
glabris, labio inferiore triangulare, 10-16.6 mm longo, 

-4 mm lato, tridentato, denti bus 1.7-4.5 mm longis, 
lateralibus brevibus, labium superiore 7-10.6 mm longo, 
bifido, incisura 5-7.8 mm profunda, lobis 2.5-3 mm latis; 
vexilum glabrum, ovatum, 15.5-20.6 mm longum, 9.7- 
4-4.5 mm longo, veli reflexa 
m latae, 

e m longo; carina glabra, 3. 1 5 mm 
lata; ovula 4-8; legumina — immatura, 5 cm longa, 
mm lata; semina non v 


Plants short-lived perennials, 17-35 cm tall; 
stem erect, simple or with some basal branches, 
more often branching from the upper nodes after 
the first inflorescence, hollow, angular, to some- 
what fistulose, covered with soft, spreading, tangled 
hairs 4-7 mm long; stipules 4-7 cm long, mem- 
branaceous, encircling ca. 4076 of the stem cir- 


cumference, 1.2-2 cm fused to the petioles, the 
free portion green, foliaceous, ovate-lanceolate, di- 
verging away and ascending, 2-5 cm long, 8-15 
mm wide, thinly lanate on both sides; petioles 4— 
8.5(11) cm long, lanate as the stems; basal juvenile 
leaves simple, the rest compound; leaflets (3)5—9, 
broadly oblanceolate, the tips generally rounded, 
lanate on both sides, the largest 5-7 cm long, 15- 
20 mm wide, mostly complanate at maturity. Pe- 
duncles 4—6 cm long on the primary inflorescence; 
racemes all immature on the primary inflores- 
cences, 20-30 cm long, flowers scattered to sub- 
verticillate, the rachis lanate; bracts green, folia- 
ceous, caducous, lanceolate, the tips acuminate, 
largest below, smaller above, 12-23 mm long, 2.5- 
4 mm wide, lanate dorsally; bracteoles lanceolate, 
3.5-6 mm long, 0.5-1.2 mm wide, attached near 
or slightly below the lips of the lateral sinuses of 
the calyx; pedicels 2.6-5 mm long, ascending 
lanate. Calyces lanate outside, glabrous within, the 
lower lip triangular, 10-16.6 mm long, 3.5- 
wide, the tip trifid, the teeth 1.7-4.5 mm long, 
the laterals only slightly shorter, the upper lip 7- 
6 mm long, deeply bifid, the notch 5-7.8 mm 
deep, the lobes 2.5-3 mm wide, the lips connate 
2. mm laterally; banners glabrous, dins 
15.5-20.6 mm e including the 3.4-4.5-m 
long claw, 9.7-15.6 mm wide, reflexed 9-13. 5 
mm, appressed 5. 8- e 5 mm, reflexed/ berets 
ratio 1.45-1.6, the angle 127-144"; s 
21 mm long, 6.4-11 mm wide, the Pin 3 3-4.5 
mm long; keels glabrous, 3.4-5.5 mm wide in the 
middle, the angle 96-1035; ovules 4-8. Legumes 
shaggy lanate, immature, 5 cm long, 7 mm wide; 
seeds not seen. 


The name Lupinus magnistipulatus was sug- 
gested several years ago by Prof. A. Burkart on 
herbarium specimens. Some of the specimens cited 
here were labeled by Burkart as a new species, L. 
reitzii. Our interpretation of the material is that 
these specimens are juvenile plants and do not differ 
otherwise from the others. Therefore we have not 
recognized L. reitzii. 

The new species apparently grows only in south- 
eastern Brazil in the states of Santa Catarina and 
Paraná at altitudes from 700 to 1,650 m. Flow- 
ering occurs from late October to late January. 


ne Aedes ud specimens examined. BRAZIL. 
un. Guarapuana, Guara, Hatchback & Gui- 

maraes 20492 (NY, US). SANTA CATARINA: Mun. Bom 
etiro, Campo dos Padres, Smith, Reitz & Klein 7703 
(NY, as Smith & ca 1 0326 (US); Campo dos Padres, 
,90 R. Reitz 2377 (holotype, SI; isotype, RB; 
photo, UMO); Man. gum Campo de Palmas, 52 km 
W of Caqador, Smith & Reitz 9155 (SI, US); Mun. 


Volume 76, Number 1 
1989 


Planchuelo & Dunn 


307 
Lupinus lanatus 


18mm 


7 
SÍ 
% 


FIGURE 3. 
inflorescence. — B. Stipules detached at the node. — 
the left lateral sinus an 


Illustration of typical structures of Lupinus magnistipulatus. 
C. Lateral view of the 
nd opened so that the inside surface shows 


—A. Upper part of the plant with 
left side of the flower. — D. Calyx, cut at 
nner petal flattened, dorsal view. — F. Win 


—E. Ban 
petal. —G. Keel petals, enclosing the staminal tube and the pistil, with the mean number of ovules drawn. 


Ponte Serrada, by road to Xanxere, 700-900 m, ruderal, 
4 ith & = 13054 (US, SI, fragments); Mun. Sao 

Joaquin, Serr ratorio, Reitz & Klein 7417 (SI, 
US); Smith & Reitz 10135 (US) 


3. Lupinus setifolius Planchuelo & Dunn, sp. 
nov. TYPE: Brazil. Rio Grande do Sul: locality 
unknown, Rambo 36208 US SI; pho- 
tos, NY, UMO, US). Figure 


Plantae probabiliter biennes, 25-30 cm altae; caules 
erecti, fistulos si, pilis paucis patentibus ornatis; stipulae 
[nee foliaceae, 5-6 cm longae, parte adnata, 1.5-2 

m longa, parte libera E ovata, fpem 2.5-4 


cm longa, 10-15 mm lata; petioli 6-7.5 cm longi; folia 
5- "ds palmata, folia juvenalia basi plantae locata et folia 


har rotundatis ubique glabris praeter marginem et cos- 
infra pilis patulis setaceis ornatis; racemus pe 
floribus diffusis; bractae 13-20 mm longae, 3-5 m 
latae, EEES setaceo; bracteolae 3.7-4.8 m f y 
0.8-1 mm latae; flores calycibus i d dorso laxe subsericeis, 
ventraliter glabris, labio inferiore 10-11.3 mm ongo, 3- 


vexillum alabrum: ovatum, .9 mm longum, 11.5- 
12.5 mm latum, ungue 3-3. 5 mm longo, parte reflexa 


308 


Annals of the 
Missouri Botanical Garden 


Al 
| 


15mm 


FIGURE 4. 
B. Stipules detached at the node. — C. 
sinus and opened so that the inside surface shows. — E. 


Illustration of typical structures of Lupinus setifolius. — A. Upper part of the plant with inflorescence. — 
Lateral view of the left side of the flower. — D. 
Banner petal flattened, dorsal view. — F. 


Calyx, cut at the left lateral 
Wing petal. —G. 


Keel petals, enclosing the staminal tube and the pistil, with the mean number of ovules drawn. 


11-12 mm; alae 16-16.5 mm longae, 5.8-6.5 mm latae; 
carina glabra, 3.5-4.5 mm lata; ovula 5-6; legumina 

n visa 

Plants probably biennial, ca. 25-30 cm tall; 
stems erect, fistulose, 6—7 mm diam., angular from 
the ridges by veins from the petioles, with scattered 
spreading hairs 3-4 mm long, these more numer- 
ous above; stipules foliaceous, 5-6 cm long, the 


adnate portion 1.5-2 cm long, the free portion 
ovate-lanceolate, the tip acute or obtuse, 2.5-4 
cm long, 10-15 mm wide, glabrous on both sides 
except a few hairs on the veins and margins; pet- 
ioles of larger leaves 6-7.5 cm long; juvenile leaves 
at the base of the plant and first leaves of lateral 
branches with 3 leaflets, the upper leaves with 5— 
7 leaflets, these broadly oblanceolate, the tips 


Volume 76, Number 1 
1989 


Planchuelo & Dunn 309 


Lupinus lanatus 


rounded, glabrous on both sides, except a few 
spreading hairs on the veins below or on the margin, 
the largest 6-7 cm long, 20-25 mm wide, mostly 
complanate. Peduncle 5 cm long, with more nu- 
merous spreading hairs than the stem; raceme (im- 
mature) dense, the flowers scattered; rachis with 
denser spreading hairs; bracts lanceolate, the tips 
acuminate, 13-20 mm long, 3-5 mm wide, dor- 
sally pubescent on the veins, glabrous ventrally on 
the lower half; bracteoles 3.7-4.8 mm long, 0.8- 
l mm wide, glabrous with margin setaceous, nar- 
rowly lanceolate, attached just below the lips of 
the lateral sinuses of the calyx; pedicels 3 mm 
long at anthesis, spreading pilose. Calyces covered 
with long soft hairs outside, glabrous within, the 
base tapering but slightly gibbous above, lower lip 
10-11.3 mm long, 3-4 mm wide, the tip trifid, 
all teeth 2.3-3 mm long, 0.5-0.8 mm wide, upper 
lip 7.5-8 mm long, 4 mm wide, bifid, the notch 
4.5-5 mm deep, the lobes 1.2-1.5 mm wide; ban- 
ner glabrous, ovate, the base tapering into a claw, 
15.5-16.5 mm long including the 3-3.5-mm-long 
claw, 11.5-12.5 mm wide, reflexed 11-12 mm, 
appressed 4-5 mm, reflexed/appressed ratio 2.2- 
3, the angle double, the lower 132-1465; wings 
16-16.5 mm long, -6.5 mm wide, the tip 
rounded, the claw 3.5-3.8 mm long; keel glabrous, 
3.5-4.5 mm wide in the middle, the angle 92- 
109°, the tip not curled back; ovules 5-6. Legumes 
not seen. 
The description was based solely on the holotype. 
The taxon appears to be closely related to L. mag- 
nistipulatus. The flowers are smaller and the base 


of the calyx has a ridge above and below at the 
pedicel, while that of L. magnistipulatus tapers 
more gradually into the pedicel. 


Representative specimen examined. BRAZIL. RIO 
GRANDE DO SUL: locality unknown, Rambo 36208 (ho- 
lotype, SI; photos, NY, UMO, US). 


CONCLUSIONS 


The three species, L. paraguariensis, L. al- 
bescens, and L. aureonitens, not treated in this 
paper are included in the key to facilitate identi- 
fication of presumably closely related species. In 
seedlings of L. albescens grown from an herbarium 
specimen, the first leaf of the plant was simple and 
the second three-foliate. This particular charac- 
teristic should be common to the group of species 
treated in the key, although it is difficult to find in 
herbarium specimens. Considering the taxonomic 
studies of the genus Lupinus for South America 
(Planchuelo-Ravelo, 1984), this complex and its 
related species are representatives of the lupines 
of the “Atlantic Region" and are the only group 
of perennial plants with large compound leaves 
growing in the area. 


LITERATURE CITED 
PLANCHUELO, A. M. & D. B. DuNN. 1984. The simple 


leaved lupines and their p in Argentina. Ann. 
Missouri Bot. Gard. 71: 92-103. 

PLANCHUELO-RAVELO, A. M. 984. Taxonomic studies 
of Lupinus in South America. Pp. 39-53 in Pro- 
ceeding of the 3rd International Lupine Congress, 


La Rochelle, France 


AUGUSTUS FENDLER'S 
VENEZUELAN PLANT 
COLLECTIONS: 


Carol A. Todzia? 


ABSTRACT 


Between 1854 and 1858, Augustus Fendler resided in Colonia Tovar, a German village in the Coastal epa 


of Venezuela, where he assembled on 


map and list of Fendler 


e of the finest collections of plant specimens for his time. Fendler's life in Colon 


aton, A Engelmann, and Josep man 
e 


i A 
ed. An dii list dei il collection dus extracted rut 


his notebook is given for the angiosperm types that li collected. 


In the mid nineteenth century South America 
was poorly known botanically compared with most 
other parts of the world. At this time, plants gen- 
erally were collected by people who traveled through 
one or several countries, making plant specimens 
along their way (e.g., Ruiz & Pavón; Humboldt & 
Bonpland). Few collectors lived in an area long 
enough to sample a region thoroughly and collect 
the relatively rare species. Augustus Fendler was 
a notable exception. He resided four years in Co- 
lonia Tovar,’ a small German colony 35 miles west 
of Caracas, Venezuela (Rasmussen, 194 
collected over 3,000 ferns and flowering plants. 
These collections are noteworthy, not only for their 
excellence, but also because a great number of new 
species were based on his material. To this day, 
new species are being described from Fendler's 
Venezuelan specimens (e.g., Croat & Lambert, 


Although his Venezuelan collections are ex- 
tremely important, Augustus Fendler is perhaps 
best known for his collections of New Mexican 
plants between 1846 and 1847. Gray (1849) pub- 
lished an annotated list of Fendler's New Mexican 


plants, which Shaw (1982) discussed more re- 
cently. Fendler's other collecting trips to Panama, 
Venezuela, and Trinidad are not as well known as 
those he took in the United States. In Panama, 
Fendler (Fig. 1) collected near the Chagres River 
for four months in 1849 and 1850. He resided in 
Trinidad from 1877 until his death in 1883. His 
most valuable collections were made in Venezuela 
between January 1854, and September 1858. 

This paper is an account of Fendler's life in 
Venezuela constructed from his letters to D. 
Eaton, G. Engelmann, and A. Gray in the Yale 
University Library, Missouri Botanical Garden, and 
Gray Herbarium archives, respectively. This cor- 
respondence provides insight not only into Fendler, 
but also into the difficulties of life as a plant col- 
lector in the s. Fendler corresponded with 
Engelmann in German and with Eaton and Gray 
in English. These lucid and detailed letters are a 
chronicle of plant collecting in South America in 
the mid nineteenth century and are even more 
remarkable since German was Fendler's native lan- 
guage 

According to various biographical notes (Canby, 


! This work was supported by a National Museum Act Curatorial Trainee Grant to the Missouri Botanical Garden. 


I am grateful to M. Riley (MO), M. Hill (YU), and the librarians at the Gra 
nd J. Ewan pro 


assistance with archives. A. Sa e J. dise ermar 
rry, C. Blaney, J. E. 


y Herbarium and Arnold Arboretum for 
iie mulating ye ee on Fendler. I ocn 
, A. R. Sm 


oran, N. Mor . D. Stev 
and H. van der Werff for helpful Duel on the manuscript; R. Magill and M. ‘cae for quo Misi 


and J. Myers fi 


or preparation of the 
? Missouri Botanica 


map. 
en, P.O. Box 


the western part of Venezuela rines 1981) 


299, St. Louis, a 63166, U.S.A. Current address: Plant Resources 
U 


e of Aragua ae is not to be confused with Tovar in the state of Merida, in 


ANN. Missouni Bor. GARD. 76: 310-329. 1989. 


Volume 76, Number 1 
1989 


Todzia 
Fendler's Venezuelan 


Plant Collections 


FIGURE 1. 


1885; Gray, 1885; Rohl, 1944), Fendler was born 
in Gumbinnen, eastern Prussia, in January 1813. 
At the age of 12 he was sent to a preparatory 
school where he mastered the fine penmanship that 
was as delightful to his correspondents as it is to 
his present-day historians (Fig. 2). Fascinated with 
traveling, Fendler sailed from Bremen to Baltimore 
in 1836 and within two years moved to St. Louis. 
During a visit to Europe in 1844, Fendler met 
Professor Ernst Meyer of the University at Koe- 
nigsberg in Prussia, who encouraged him to collect 
plants in the western United States and send them 
to him for sale. On his return to St. Louis, Fendler 


Photograph of Augustus Fendler probably taken in St. Louis, ca. 1860. 


brought his specimens for identification to George 
Engelmann, the botanist who advised Henry Shaw, 
founder of the Missouri Botanical Garden (Stieber 
& Lange, 1986). Fendler and Engelmann struck 
up a friendship that spanned nearly 40 years. En- 
gelmann later introduced Fendler to Asa Gray, who 
described Fendler as “a close, accurate observer, 
a capital collector and specimen-maker" (J. L. 
Gray, 1893: 341). 

After a year of plant collecting in New Mexico, 
a failed attempt to collect plants near the Great 
Salt Lake, and two sojourns in Arkansas and Lou- 
isiana, Fendler settled in Memphis, Tennessee in 


312 


Annals of the 
Missouri Botanical Garden 


m ^ 1 AL 
t P 


/ 
eres aa Poa GU 


p Le p 
COP OAPAMA 


¿Hna APP Qd A E ASA 3, 
Ld, c er 
y 2 
Pm y r 
Zan Coe 
u^ uU ergi rone e 


dM. 


a 2» ¿ue em rf GA > Agi" Lor Sa p^ gs 


> 2 e ac 00 66 a d Las IA 4 ae ARO TA 


ira PP ai ite FA 


a 


PP oe MMC 


2 MEE olari AC 


> A Agr eS wag vM >) Las 47 The ame 


ITEE M e ALPP 2274 


E 
p Wee gZIECE. DIVI 
e: An Pg UL 


— 5 


P ¿CAS p^ 


7 ae an . aa Vasil Bre cw LZ reo A: reg fF 


a, ye REL 
A FA ote aio ^ ctc XM I A p d 


a GIRE PPP OPO fore core y dor a" ait. 


Puro co. 277 


AFD A p 
d a dz; e. EM s rato 
ra DAI M E ar A 


AA See Tte, 


ord 
diffe 


P d 


gt Gita A NS ve our p pd 


e p? AA OA v. p 


A k A ALT ae LF” A IAEA IAN 2 p gy» 


"bte AP en o wx MET to i al tL. 


: cp T bare Pr? Sac 


p en Iu 


UE He avum 
Per ESTE LAO IET LH 


"M Ara Teror wr of 7 A 


of Ae « ALAS PE ya aX ATO Vw 


p S y 22 


— P Faut or pom LA n Ee de SL 


fav 
Lm a 


LIL 


JO mme PEAD a E SETA governg lore 


FIGURE 2. 
Garden archives. 


1850, and prospered there for four years in the 
gas-lamp business (Stieber & Lange, 1986). In 
1853, when the lamp-lighting business in Memphis 
faltered, he was eager for a change of pace and 
wanted to resume collecting plants. He wrote to 
Gray on 11 August 1853 about his collecting 
plans: “My principal object in going to Caracas is: 
to go into a business again, by which I can support 
myself. But I hope I shall have plenty of leisure 


5:3 . aa dac ——Ó— = 


Letter by Augustus Fendler to Asa Gray (Memphis, Tennessee, July 13, 1853). Missouri Botanical 


to make botanical collections besides. If I collect 
again I shall aim to make the sets as large and 
complete as possible. Do you think 15 sets, each 
containing from 1,000-1,500 species of plants 
collected about Caracas could meet with a ready 
sale?" On 15 August 1853, he wrote Engelmann 
that he wished to move to a town near Caracas 
because “In a city with a population of 70,000 I 
believe to be able to find more security than here, 


Volume 76, Number 1 


1989 


Todzia 
Fendler's Venezuelan 
Plant Collections 


313 


Boo 77 SB A = Co 272 Aero 
PA ; Mum 777 B77 P ctp por, ov dq nee 7 SAC, Lami 
P Au E A Fe Tarro a 
e A ran P ai tr eri T wee PA a e 
VATES eee AA “SÍ 72 ha. ATA xt wf 

a ares 

Filtra, ea KEEFE. ina cl 


yd ra 
m MAE QS ZA TTA 


OÍ Lame Ll c are For 


Stem 5 


cae (Lp PLEO 
MIS on A 


Fer 


IN AE 


Fred 


3 : me cae a 
4 e c EXT or a PR d AREA 22 IE 
Lr D t 4 cw 


^ 
Pura E. “sf e T 


her SAT 


ME, AO 
p 


= 
"mm Fhe 4.2 


Pragara 


EC 


E O a j > Zane 
ride arcam párr 
D^ PL i “A i 
ER, DL 15 Se E MESA uu c Ame > 
Rr A 


C 


a: Gy LAII? LII 


: toe 


ino" Lez Ivo 


C d AT Ar? 


ne S pt eee P TW MA 4 » a E SD a 
Dee a ae gour mn Ena well Aime x y Gere 


rue AI E v P adatta PG 
p he RARE 


Lune er. 


Y aI ra ^ d p 


M pt Of, or Comente Ta 
LL 


DA ato 


Terra . 


-P7 LE PPT cn 


e eran 
PED cd ae IZ or LE 


A int iin Yo Pew died ozo. e P ss: A ooh aruit: 


FIGURE 2. Continued. 


and secondly, because I believe I shall have a better 
climate there. Then also, the wish to live again in 
a mountain valley and to be in the vicinity of a 
rich mountain flora! without having the difficulties 
of long and difficult communication as I found in 
Santa Fe." 

On the 24th of December 1853, Fendler sailed 
for Venezuela from New York harbor, 
panied by his chronically ill brother. They arrived 


accom- 


in La Guaira, Venezuela, on 21 January 1854. 
In his day, it took five hours to reach Caracas from 
the coast. Upon arriving, he commenced collecting 
plants in the vicinity. Prompted by the high cost 
of living in Caracas, he went up to Colonia Tovar 
after only six weeks. After reaching the settlement, 
he wrote to Gray on 16 December 1854: “The 
base of the valley is 5,600 ft. above the sea, and 
some of the surrounding mountains about 2,000 


314 


Annals of th 
Missouri B nd Garden 


ALAN ed, du eo 


aa rene . 


ESA 


(Ala 227 


p EU TES P iios xe oa di n d Lm Amo 
n d OPP PIEL, ^d , 


p—— à "c Lr 
" 
py c pm Sor AG c 


v onm 
A or, LOCA. 


Ji L2: €— Hepe A Arce 


- P24 oP? 


CA 


Ire I, an bat 
y sad rr 
e Mare edad 
g 
MARS memet eve js» 
Pe Loread 


Ce eor m rer AL 
P4 


rre, 


e 


^ ae Cee 


oo : e »: " an 
seo AS LAO Imre er oF, 


€ EPP? He 7? 


P os nr 2 P d ha nani dapes 


FIGURE 2. Continued. 


feet higher .... The road from Caracas leads at 
first along a small river, which is very rapid and 
has to be crossed 42 times. The valley in which 
this river flows becomes at last very narrow and 
the road turns off to the left ascending a high 
mountain, from the tip of which it leads on in a 
westerly course along the very summit of the prin- 
cipal mountain range, through the deep shades of 
the primitive forest and down the valley of the 
Colony." 

While at Colonia Tovar, Fendler also corre- 
sponded with Joseph Henry of the Smithsonian 
Institution, who published Fendler's letters and me- 
teorological observations in the Annual Report of 
the Board of Regents of the Smithsonian Insti- 
tution. Fendler was an ardent amateur meteorol- 
ogist and in Venezuela traveled with a barometer, 
rain gauge, and thermometer. From his corre- 
spondence with Gray, Fendler provided a very rich 
description of Colonia Tovar (Fendler, 1857): “The 
valley in which Colonia Tovar is situated was, so 
late as December, 1841, a perfect wilderness, cov- 
ered with primitive forest. Not even the existence 
of this valley was known fifteen years ago, neither 
to the government nor to its owner, although it is 
only thirty-five miles west of Caracas, the capital 
of Venezuela, and in a straight line cannot be more 
than twelve miles from the sea. And when an at- 
tempt was made to explore this region not even a 
guide could be found for the small exploring party 
of fifteen men, headed by Colonel Codazzi, a skillful 


ns A, 2 UR A 


officer and compiler of the new map of Venezuela. 
When this party at last succeeded in crossing this 
region and reaching the sea-shore, they thought 
they had achieved a most extraordinary thing, (to 
cross a distance of twelve miles in six days;) and 
after they had returned to their homes none of 
them had a desire to do the feat over again. This 
was a party of natives. And when, at a later period, 
after the establishment of the colony, another skill- 
ful engineer found, with a party of colonists, his 
way to the opposite port of the sea-shore, the party 
did not venture to go back the same route, but 
rather chose the way by sea to Laguayra, from 
there to Caracas and back to the colony, a very 
circuitous route certainly. Such is the nature of 
this mountain region, with its precipices, waterfalls, 
deep ravines, and its dense, almost impenetrable 
primeval forests. 

Shortly after arriving in Colonia Tovar, Fendler 
bought a small farm in 1854 for 47 dollars. He 
described his new home in his letter to Gray of 16 
December 1854: “My cottage stands on a small 
hill which is projecting from the sloping side of a 
mountain overlooking the greater part of the Col- 
ony. On the slopes of this little hill we have made 
terraces planted with Musa sapientum, with apple 
trees, Palms (Oenocarpus utilis), and stately tree 
ferns 14 feet high. Near the brown polished stem 
of the Palm the clear arch of a fountain (9 feet 
high) glitters in the tropical sun sending unceasingly 
its sparkling little stream to the top of the young 


Volume 76, Number 1 
1989 


Todzia 
Fendler's Venezuelan 


315 


Plant Collections 


Palm. This fountain I made in a few days without 
outlay of money. Two pieces of the trunk of Cerox- 
ylon andicola served as pipes." Fendler clearly 
enjoyed Colonia Tovar: “A visitor who never before 
lived in a valley like this finds here many pecu- 
liarities of vegetation, surface and climate which 
make him feel that he is not far off the land of 
perpetual peace; for he has entered the happy 
region of the ferns, the 'tierra templada de los 
helechos.' There is no scorching summer's heat, 
no fearful winter's cold, neither tornados to dev- 
astate the country nor gales to blind the inhabitants 
with sand and dust or penetrate their clothes and 
flesh with piercing frost" [Fendler to Gray, 16 
December 1854]. His years in the village were 
the most enjoyable time of his life, and later letters 
indicate that he very much would have liked to 
return there [Fendler to Engelmann, 20 July 1882, 
Trinidad ]. 

During his first year at Colonia Tovar, Fendler 
devoted much of his time to plant collecting. He 
wrote to Professor Henry: “In collecting botanical 
specimens, I have penetrated, without companion, 
the wilderness around in different directions, also 
that on the other side of the principal mountain 
range towards the sea, and can testify to the dif- 
ficulties and hardships which are met with in ex- 
ploring such a country . . . . In these woods, where 
the rays of the sun never touch the ground, there 
it is where moisture and a cool temperature reign 
forever. The trunk of every tree and its branches 
are covered with Ferns, Lycopodiaceae, Mosses, 
Hepaticae, Lichens, Orchids, Bromeliads, Araceae 
and besides Piperacae with many exogenous plants 
too numerous to mention” (Fendler, 1857). 

Fendler wrote in detail to Gray, who had offered 
to enlist subscribers for his sets of plants. “I have 
already 8 boxes full of dried plants. The most 
attention I have hitherto paid to ferns of which I 
have 255 species, most of them complete; other 
plants I have not yet enumerated. For the last four 
or five weeks I have been employed with arranging 
my collections but have not yet finished. The fine 
dry weather which is now approaching urges me 
again to renewed activity in the field. The woods 
here are very dense and impenetrable without a 
sabre. The neighboring country is much diversi- 
fied" [Fendler to Gray, 16 December 1854]. 

“Of Palms I have collected 6 species, of which 
5 grow in the colony. Tree ferns 9 or 10 species. 
Cruciferae and Umbelliferae are represented only 
by 2 or 3 species. 1 do not intend to send plants 
until I can send something complete, and until I 
can ascertain the safest way to forward them" 


[Fendler to Gray, 16 December 1854]. Less than 


a year later, Fendler wrote: **Of flowering plants 
I have about 1,850 species, although my search 
after these may be called but a superficial one. I 
think from 800-1,000 more species might be col- 
lected and of ferns perhaps from 80-100 more" 
[Fendler to Gray, 25 November 1855]. In addi- 
tion to ferns and angiosperms, Fendler collected 
fungi for M. A. Curtis, a mycologist in the Caro- 
linas, lichens for Professor E. Tuckerman, a li- 
chenologist at Amherst College, and mosses for 
William Starling Sullivant, a distinguished Ameri- 
can bryologist in Ohio. 

In 1857, after collecting in Colonia Tovar for 
over two years, Fendler found it increasingly dif- 
ficult to find novel species. “In coming to a new 
country it is at first comparatively easy to make 
a living by collecting plants; for the collector can 
make use of all the different species that meet his 
eye, and can carry them home without the aid of 
other hired persons or beasts of burden. But after 
he has explored the neighborhood and is obliged 
to make extensive excursions of from 8-14 days 
or 3 weeks, the case is different, and he soon finds 
that his earnings do not come up to his expenses" 
[Fendler to Gray, 1 July 1857]. 

After collecting thoroughly around Colonia To- 
var, Fendler took longer excursions crossing *'the 
principal mountain chain in three different places. 
One from Maracai to Columbia [Puerto Colombia ] 
to the very margin of the sea; the other from 
Valencia to San Estevan within 3 miles of Puerto 
Cabello; the third from Petaquire to within a few 
miles of the sea-shore. The two first regions are 
W est, the third one East of Colonia Tovar. Besides 
these I have made a number of minor excursions 
into the neighboring valleys” [Fendler to Gray, 1 
July 1857]. During these long excursions he re- 
flected: “The more I attempt to explore these re- 
gions the more am I convinced, that a great deal 
remains yet to be done with regard to discovering 
and making known the plants of this country. These 
extensive ranges of mountains with their innumer- 
able recesses and deep ravines, covered with the 
densest growth of primeval forest uninterrupted for 
more than 150 miles from East to West, have been 
in most places never been traversed by civilized 
men. When standing on some high mountain-peak 
and looking over a part of this territory, it needs 
no great power of imagination to be filled with a 
longing and an ardent desire to embark in enter- 
prises even of the most perilous kinds, in order to 
penetrate the unknown abysses and to bring to 
light the hidden treasures of vegetation. And when 
in such moments I compare my strength and m 
fast waning pecuniary means, which forbid to hire 


316 


Annals of the 
Missouri Botanical Garden 


a companion to help me carry the burden, then I 
feel somewhat akin to depression of mind which 
prompts me to leave the spot and retrace my steps 
homewards.” 

Because Fendler viewed plant collecting as a 
business, he was very concerned with the market- 
ability of his specimens, which were valued ac- 
cording to their origin, rarity, and completeness of 
identifications. Thus Fendler wished to have his 
plant collections be as rare as possible, as well as 
to have them determined. He was concerned that 
Karl Moritz, who had been living in Colonia Tovar 
for over ten years, would begin again to collect 
plants in his territory. “The present letter is chiefly 
written to make you acquainted with my intentions 
of sending the collections as soon as possible, that 
you may be enabled to correspond or to publish 
something to that account and secure me from 10- 
13 subscribers in time, because Mr. Moritz, who 
is still here has sometime ago commenced again to 
collect ferns, which he intends to send at an early 
date to Germany (I think to Berlin). I have seen 
Moritz's own set of his former collections when he 
used to travel about the greater part of Venezuela 
viz: the Orinoco, the mountains of Merida, Cu- 
mana, Caripe and a great many other regions; but 
the number of all his species of ferns will hardly 
exceed 250 or 260, while I have already 314, of 
which at least 290 are collected at the Colony or 
from 4-8 miles around it. There are several regions 
not far-off, where I never yet have been and which 
promise a very rich harvest of new species. I think 
I could augment the number of species considerably 
and discover many more new ones if | am sure 
that they find a market on which I may rely. I am 
therefore anxious to secure subscribers for them; 
so that I need not fear competition and then I shall 
go at it with renewed vigour; and if my life and 
health be spared I think of bringing together a 
rather complete representation of the vegetation 
of this and other parts of Venezuela. When I came 
here, Moritz seemed rather surprised that I should 
go at collecting plants in a place which he thought 
of being thoroughly ransacked, and he said that it 
would be a hard matter to find any new species. 
He seems however to have altered his opinion in 
this respect, after I had been in the field for some 
time; for I showed him many a plant which was 
new to him” [Fendler to Gray, 25 November 
1855]. Fendler’s Venezuelan collections surpass 
Moritz’s in quantity and quality (J. A. Steyermark, 
pers. comm.; A. R. Smith, pers. comm.). 

Plant collecting, however, was not profitable 
enough for Fendler to support himself and his in- 


valid brother. “Whenever I was engaged in sci- 
entific pursuits for any length of time I was soon 
obliged to go to work at some other more profitable 
business, although that business might not be in 
accordance with my inclinations. And now I find 
again that the little capital, which I had earned at 
Memphis during three years by distilling cam- 
phene, is rapidly vanishing, notwithstanding the 
most frugal mode of living. I am therefore under 
necessity to try my hand for a while at something 
else beside collecting plants, and have already com- 
menced to make experiments in brewing beer and 
making brandy; and shall be obliged (with regret 
do I say it), to spend most of my time behind the 
still . . . . By no means do I wish by the preceeding 
lines to convey to you the idea, that I am dis- 
couraged or that I have given up collecting plants. 
As long as strength and life and a few spare dollars 
remain with me I shall continue to do all I can for 
my favorite pursuit; and even now, whenever time 
and circumstances allow me to do so, I shall do it 
with the same zeal as I have done heretofore" 


[Fendler to Gray, 1 July 1857]. 


FENDLER'S COLLECTIONS 


In Venezuela Fendler collected under two num- 
ber series, one for the ferns beginning at no. 1 and 
ending at no. 500, and one for flowering plants, 
numbers 1-2,630. Sets of Fendler's Venezuela 
plants were purchased by the following people and 
institutions (the number before the name indicates 
which set went to that recipient). (2) Hooker (K), 
(3) Grisebach (GOET), (4) A. Gray (GH), (5) En- 
gelmann (MO), (6) Short (PH), (7) Charles Wyville 
Thomson (E), (8) Boissier (G), (9) De Candolle (G), 
(10) Oxford herbarium (OXF), (11) Harvey (TCD), 
(12) Delessert (C), (13) Martius (BR), (17) Eaton 
(YU), (18) Miss Elizabeth Morriss (?PHA), (21) 
Eaton (YU). It is not known whether sets 14-16 
were sold at a later date, or whether they remained 
at GH. Sets 2-13 included ferns and phanerogams; 
sets 17-23 consisted only of ferns. The first and 
most complete set was retained by Fendler at first. 
Later he sold this first set of angiosperms to Gray. 
"My full set of Venezuelan plants contained over 
twice as many species as set No. 4, and therefore 
I hope that by the addition of those plants your 
herbarium would gain something. 1 would rather 
see them incorporated with your herbarium than 
with that of anyone else. The naming of the price 
of the lot I leave to you as you can judge best of 
their value, and I know that I am in good hands 
when I am in yours" [Fendler to Gray, 28 October 


Volume 76, Number 1 
1989 


Todzia 
Fendler's Venezuelan 


317 


Plant Collections 


m 
Choroni 


e 
bn Esteban 


Esmeralda 


i la Guaira 
Caraypca 
Petaquire 


= e dE 
Punta Maya ip Urbina petant 
R. Maya NN DF M / 
Y Colonia a Tovar .— E es baa ) 
b A ——á Prid 
R. San Carlos: ES eee Adiuntas 


a 


NC 
a | Mp 4 
ES / VO 
Valencia OS - 4 
yo E Jj 
re 2» i 
EN ^ 4 m. j S 
f so | Gjigúe d US TENES EN 
"Oo lira NC = = 10° 


104 
68 


FIGURE 3. 


1863, St. Louis]. Ten cents per sheet was the usual 
asking price, although it was slightly higher for the 
orchids since they were often unicates. 

At times Fendler felt he was not justly compen- 
sated for his specimens. For example, Lindley 
thought the price too high. “If Dr. Lindley was 
advised to pay but £2 per 100 for my Orchidae, 
then I will ask nothing more, and he shall be very 
welcome to them if he finds them to be worth that 
much. But if on the contrary he finds that the 
greater part of them, from being common things, 
are of no use to him, which I am inclined to infer 
from his letter, then I should be very sorry indeed 
that I ever offered them to him. To me that would 
be at least objects of remembrance of the amount 
of labor, bestowed upon, and perhaps of an over- 
animated enthusiasm in searching for them” [Fen- 


dler to Gray, 27 March 1859, St. Louis ]. 


FENDLER's FIELD NOTEBOOK 


In 1859, Fendler sent Gray the field notes for 
his Venezuelan plants, retaining the information 
only on his personal set of specimens (set No. 1) 
[Fendler to Gray, 15 December 1859, St. Louis]. 
The field notes contain all dates and localities from 
which a particular species was collected, as well as 
elevation and notes about habit. Fendler numbered 
his plant collections according to the classification 
schemes presented in Gray's Botany of the North- 


Map of the localities in northern Venezuela where A. Fendler collected in 1854-1858. 


ern United States and Lindley's Vegetable King- 
dom rather than the order in which they were 
collected, as is usually done today. 

o obtain more complete sets of specimens, which 
could then be sold at a higher price, Fendler often 
applied the same number to specimens he believed 
to be the same species but were from widely distant 
localities. Fendler also attributed to the same col- 
lection number specimens collected on as many as 
five different dates. This was a common practice 
among professional plant collectors, particularly 
those working with Gray (Ewan, 1969), and a 
source of problems for plant systematists. The /n- 
ternational Code of Botanical Nomenclature 
(Burdet et al., 1983) states that an “isotype is any 
duplicate (part of a single gathering made by a 
collector at one time) of the holotype.” Since Fen- 
dler did not indicate on his specimens what specific 
date a particular sheet was collected, technically 

o isotypes can be designated. Of the 225 names 
in this list, 66 were collected on more than one 
date. Few taxonomists have previously been aware 
of this problem with the Fendler Venezuelan col- 
lections. 


LOCALITIES 


Almost all of Fendler's Venezuelan specimens 
bear blue labels imprinted at the top with “Plantae 


Venezuelanae” and “Prope coloniam Tovar legit 


318 


Annals of the 
Missouri Botanical Garden 


TABLE 1. Localities used in Fendler’s angiosperm list. 


Approx- 
imate 
Elevation 
State Latitude Longitude in Meters 
Agua Blanca = Quebrada Agua Blanca Aragua 9954'N 69°22'W 
Biscaina Aragua 10°20'N 67°15'W 1,000 
Campanero Carabobo 10°26'N 68°01'W 150 
Caraca D.F. 10°30'N 66°55'W 914 
Cariaca = Carayaca D.F. 10%32'N 67°07'W 800 
Chichiribichi = Chichiriviche D.F. 10°33'N 67?14"W 0 
zhoroni Aragua 10°29'N 67°37'W 0 
Colombia = Puerto Colombia Aragua 10°30'N 67°36'W 0 
Colonia Tovar Aragua 10°25'N 67°17'W 2,400 
Consejo = El Consejo Aragua 10°14’N 67°16'W 200 
Cumbote = Cumboto Aragua 10%24'N 66?26'W 70 
Esmeralda Aragua 10%25'N 67?*36'W 200 
Guacara Carabobo 10?14'N 67°53'W 438 
Güigüe Carabobo 10%05'N 67%47"W 420 
Hacienda Curisal (not located) 
La Guayra = La Guaira D.F 10%36'N 66°56'W 0 
Lagunassa = Lagunazo D.F. 10°25'N 67°10'W 2,000 
Las Aguntas = Las Adjuntas D.F. 10°26'N 67°01'W 900 
La Victoria Aragua 1014'N 67°20'W 585 
Macar .F. 10*26'N 67*02'"W 1,400 
Maracai = Maracay Aragua 10%15'N 67°30'W 445 
aya, Punta Aragua 10°32'N 67°24'W 1,400 
Petaquire E, 10°32’N 67°05'W 1,600 
Piedernales = Pedernales Carabobo 10°03'N 67°43'W 500 
Puerto Cabello Carabobo 10°28'N 68°01'W 0 
San Carlos River Aragua 10?19'N 67°19 W 
San Estévan = San Esteban Carabobo 10°26'N 68°01'W 200 
San Joaquin Carabobo 10?16'N 67°47'W 450 
Turmero Aragua 10?1 4/N 67°29'W 466 
Tuy River Aragua, Miranda 
Urvina = La Urbina .F. 10°27'N 67°14'W 1,400 
alencia Carabobo 10°11'N 72°12'W 478 


A. Fendler 1845-5” at the bottom. The collection 
number is usually on the label, although it occa- 
sionally is on a slip of paper attached to the plant. 
Apparently Gray had these labels printed as he did 
for Wright’s Cuban collections. Only the first set 
of Fendler’s specimens, now at GH, has any col- 
lection data on them. Although the labels indicate 
that the sets were collected near Colonia Tovar in 
the state of Aragua, they were in fact also collected 
in the state of Carabobo and in the Distrito Federal. 
Figure 3 maps the Fendler localities in northern 
Venezuela, and Table 1 presents a list of these 
localities. 


TYPE LIST 


Of Fendler’s 2,630. Venezuelan angiosperm col- 
lections, this study located records of 225 that 


have been designated as types. Since only the first 
of the 20 sets has collection data, full localities 
from the notebook are given for these types. In 
the following annotated list of angiosperm types 
based on Fendler Venezuelan collections, the num- 
ber in the parentheses refers to the number of 
dates listed in the notebook. The orchids, numbers 
1,359-1,489 and 2,124-2,155, were not includ- 
ed in the notebook. 

The species names listed in Appendix I were 
gleaned mostly from the Flora de Venezuela and 
the Catálogo de La Flora Venezolana (Pittier et 
al., 1945-1947), as well as with the help of several 
specialists (see Appendix I). The current accepted 
names are given when known to me. Herbaria data 
were gathered from the original descriptions, the 
TROPICOS data base at MO, the specialists listed 
in Appendix I, and the author's search through the 


Volume 76, Number 1 
1989 


Todzia 319 
Fendler's Venezuelan 
Plant Collections 


MO herbarium. Unfortunately, there is no record 
of the number of duplicates made of each collection 
number (although the number may be on the labels 
of set No. 1 at GH, as with the ferns (Smith & 
Todzia, 1989), and therefore there is no way to 
ascertain which specific specimens are at any one 
herbarium. Since GH received Fendler's first set 
of angiosperms, excluding the orchids which went 
to Lindley, there are probably specimens for all 
numbers there. However, only herbaria where a 
specimen is confirmed to be deposited are included 
in the list. Overall, Fendler's Venezuelan specimens 
are widely distributed in the United States and 
Europe. The following 18 herbaria are known to 
have some Fendler Venezuelan collections (Lan- 
jouw & Stafleu, 1957; pers. obs.): AMES, BR, 
BUF, C, CM, F, G, G-DC, GH, GOET, K, MO, 
NY, OXF, P, PH, S, and US. 


LITERATURE CITED 


BunpET, H. M. AL. (editors) 1983. International 
"à de of Botanical Nomenclature. Regnum Veg. 111: 
cor hus M. ia An autobiography and some rem- 
es e late August Fendler. I, Bot. Gaz. 

(Cr sra rt. 10: 285-290; II, Bot. Gaz. (Craw- 
seres, 10: 301-304; III, Bot. Gaz. (Crawfords- 


. & N. LAMBERT. 
Venezuela. Aroideana 9: 3-2 

69. Historical problems for the working 
m omist. an 18: 194-203. 

FENDLER, A. (Communication from A. Fendler.) 
Ann. Report Vire. Inst. 1858: 179-282. 
Gray, A. 1849. Plantae Fendlerianae Novi-Mexicanae. 

Mem. Amer. Acad., n. ser. 4: 1-1 
885. Amer. J. Sci. Arts, ser. 3, 29: 169- 
171. Reprinted in Scientific Papers of Asa Gray 1: 
465-467. Houghton, Miflin and Company, Bs 
[Obituary.] 
Gray, J. L. yen 1893. Letters of Asa Gray. 2 
volumes. Houghton, Mifflin and Company, Boston. 
Kitup, E. P. 1938. The American species of Passi- 
floraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 
1-613 


1986. The Araceae of 
213. 


Lanjouw, J. . A. STAFLEU. 1957. Index Herba- 
riorum a it (2); Collectors. Regnum Veg., Volume 


9. Utrecht, Netherlands. 
Pirrier, H., T. Lasser, L. SCHNIE, Z. LUCES DE FEBRES 
& ADILLO. 1945- 1947. ' Catálago de La Flora 


zolana. 2 volumes. Lit. Y. Tip. Vargas, Caracas. 
Rasmussen, W. 19 Colonia Tovar, Venezuela. 
ee History 17: 156-166. 

ROHL, m 944. August Fendler. Bot. Acad. Ci. Fis. 


SHAW, E. A 1982. Augustus Fendler's collection list: 
New ae 1846-1847. Contr. Gray Herb. 212: 


l- 
SMITH, A. R. & C. A. Topzia. 1989. Augustus Fendler's 
Venezuelan collections ue a and fern allies. Ann. 
0-349. 


Missouri Bot. Gard. 76: 


STEYERMARK, J. A. 1981. Erroneous citations of Ven- 
ezuelan localities. Taxon 30: 816-817. 

STIEBER, M. T. & C. Lance. 1986. Augustus Fendler 
(1813-1883), professional plant collector: selected 
correspondence with George Engelmann. Ann. Mis- 


souri Bot. Gard. 73: 520- SL 


APPENDIX I 


List of species with type specimens collected by A. 
Fendler in Venezuela (1854-1858). The collection dates 


eae), B. Hammel (Cyclanthaceae), J. Kuijt 
uer 


a erff (Laurace 
(Melastomataceae, Pllexiacene), and J. Zarucchi (Apo- 
cynaceae, Fabaceae). 


ANNONACEAE 
Rollinia fendleri R. E. Fries, Acta Horti Berg. 12: 


2, tab. 15. 19 


1 M LS f 
[Biscaina, 3,000 ft., 5/6/55], Fendler 196 (holotype, 
K). 


APIACEAE 


pirra venezuelensis Rose ex Mathias, Britto- 
: 226. 1936. 


[Co se Tovar, 6,000 ft., 13/4/54], Fendler 524 
(holotype, M: isotypes, G, K). 


APOCYNACEAE 
Amblyanthera fendleri Muell. Arg., Linnaea 30: 417. 
1859-1860. 


[A few mi. SE of Colonia Tovar, 5,000 ft., 12/4/54, 
6/1/54, nis i Fendler 1032 (G- DC, photo at 
- Mandevilla fendleri (Muell. Arg.) Woodson 
cera Pis was Muell. Arg., Linnaea 30: 417. 
1859- 


[A few mi. of Colonia Tovar, 7,000 ft., 12/5/54], 
Fendler 1 030 DES G-DC, photo at F; isotypes, 
B, BR, F, K, MO). 


ARACEAE 


Anthurium bredemeyeri Schott var. elongata Engl., 
Fl. Bras. 3: 80. 1878. 

[Colonia pc 6,000-7,000 ft., 25/5/54, 15/9/ 
54, 15/10/54], Fendler sae (lectotype, MO; iso- 
lectotype, 

= Anthurium bredemeyeri Schott 


320 


Annals of the 
Missouri Botanical Garden 


A fendleri Schott, Prodr. Syst. Aroid. 468. 
860. 


Dian, 3,000 ft., 
type, K; isotype, 
Anthurium nymphaeifolium K. Koch & C. Bouche 
Monogr. Phan. 2: 177. 1879. 
Co lonia Tovar, 5,000-6,000 ft., 12/ 
5/54, 28/7/54, 15/10/54], Fendler 1336 (syn- 
type, K). 
hurium ornatum Schott 
Philodendron deviatum Schott, Bonplandia 7: 29. 


10/5/54], Fendler 1343 (holo- 


[Biscaina, 3,000 ft., 10/5/55], Fendler 1329 (holo- 


type, 
Philodendron effusilobum Croat, Aroideana 9: 99, 
1986 


[Near Sen Estevan, 1,000 ft., 9/3/57], Fendler 2573 
(holotype, 
Phi ¡rie desata cera Schott, Bonplandia 7: 29. 
1859. 


Dre El 6,000-6,500 ft., 16/8/54, 19/6/ 
55, /55], Fendler 1327 (holotype, K; isotypes, 
18), 


ARALIACEAE 


—— fendleri Seemann, J. Bot. 2: 301. 1914 
A js i. S of Colonia Tovar, TV puer 
9 (MO). 


ASCLEPIADACEAE 
M cro A. Gray, Proc. Amer. Acad. Arts 


[La Son 20/11/56] Fendler 2119 (GH, MO). 
= Sarcostemma bilobum Hook. & Arn. subsp. lin- 


deniana (Decne.) Holm 


ASTERACEAE 


Clibadium parviceps S. F. Blake, Contr. U.S. Natl. 
Herb. 22: 598. 1924. 

[Between Maracai and Choroni, 4,000 ft., 12/12/56; 
also between Petaquire and Cariaca, 4,000 ft., 12/ 
2/58], Fendler 1967 (holotype, GH; isotype, US). 

Eupatorium PAS m B. L. Robinson, Contr. Gray 


Herb. 65: 
[Colonia Tovar, ES e ft, 23/6/55], Fendler 634 
(GH, photo a 
Peon steetzii B. L. Robinson, Proc. Amer. Acad. 
Arts 55: 36. 1919. 
[Colonia Tovar, 6,500 ft., 27/8/54], Fendler 647 
(holotype, GH). 
— tovarense B. L. Robinson, Proc. Amer. 
Acad. Arts 54: id 1918. 
[Between Pecan and the sea, 4,000 ft., 22/4/ 


58], Fendler 1947 eel ae GH; isotype, MO). 
= Ayapana tovarensis (B. L. Robinson) R. King & 
H. Ro vidé 
Eupatorium xestolepis B. L. Robinson m um 
lepis, Proc. Amer. Acad. Arts 54: 261. ed 
[Between Caracas and La Guayra; also iun 
of Colonia Tovar, 7/5/55, 10/5/55. Mov. 
Fendler 638 (holotype, GH). 
Hieracium tovarense Fries, T hod 152. O 
[Around Colonia Tovar, 6,000- O ft., 26/7/5 
7/5/55, 28/6/55, 23/8/55, UA Fender 
718 (G, photo at F). 
= Hieracium avilae H.B.K. 


Mikania aes ad B. L. Robinson, Contr. Gray Herb. 
61: 15. 


[Between ose and Campanero, 5,000 ft., 27/2/ 
51], ~ 2348 (holotype, GH; isotypes, MO, G, 
photo a 

Mikania bos B. L. Robinson, Proc. Amer. Acad. 
Arts 47: 196. 1911. 

[Tuy River below Colonia, 4,000 ft., 21/10/54], Fen- 
dler 626 (holotype, GH, photo at F). 

Mikania racemulosa Klatt, Abh. Naturf. Ges. Halle 
15: 325. 1882. 

[Colonia Tovar, 6,000- 7,000 ft., 31/3/54, 28/6/ 
55], Fendler 625 (GH, MO, photo at F) syntype 
with Schomburgk 480 (G). 

Mikania vitrea B. L. Robinson, Contr. Gray Herb. 61: 

[Colonia Tovar, 6,500 ft., eet ao 2349 

. Var. crassic Lm Steyerm. 
Piptocarpha sprucei Baker, Fl. Bras. 6(2): 129. 1873. 

[Between Maracai and Choroni, 2,500 ft., 28/1/57], 
Fendler 1957 (syntype). 

Senecio cucullatus Klatt, Abh. Naturf. Ges. Halle 15: 


. 1882. 
[Colonia Tovar, 6,500 ft., 24/8/55], Fendler 702 (P, 
MO, photo at F) 
Trichogonia rhadinocarpa eA L. Robinson, Proc. 
cad. Arts 42: 36. 
Mesina: 3.200 ft., 2/0/53) TUE 651 (GH). 


BEGONIACEAE 


Begonia eei dica Kunth & Bouché var. fen- 
dleriana A. DC., Prodr. 15(1): 289. 1864. 
[A few mi. i SE of Co m nia Tovar, in ravines, 3,500 ft., 
9/3/55, 1/5/55], Fendler 513 (G-DC, K). 


BROMELIACEAE 


Aechmea cymoso-paniculata Baker, J. Bot. 17: 165. 
1879. 


[Between Petaquire and the sea, 3,000 ft., 9/2/57], 
Fendler 2453 (holotype, K; isotype, MO). 
Aechmea fendleri André ex Mez, Bromel. Andreanae 
13. 1889 


[Between Petaquire and Colonia Tovar, 7,000 ft., 11/ 
2/51], Fendler 2454 (holotype, K; e GH). 
Bilbergia filicaulis Griseb., ms igl. Ges. Wis. 
Georg-Augusts-Univ. 1864: 
‘Between Valencia and Campanero, 500 ft. 7/3/57), 
Fendler 2452 (holotype, GOET, photo US). 
= Aechmea filicaulis (Griseb.) Mez 
Caraguata coriostachya Griseb., Nachr. Kónigl. Ges. 
Wis. Georg-Augusts-Univ. 1864: 
[Between Maracai and Choroni, 4,000 ft., 
Fendler 2167. 
— Guzmania coriostachya (Griseb.) Mez 
Nidularium albo-r m Griseb., pr Tore Ges. 
Wis. Georg-Augusts-Univ. 1864: 865. 
[High mountains E of Colonia Tovar, 24/6/55) Fen- 
1521 (holotype, GOET; isotype, G 
= Greigia albo-rosea (Griseb.) Mez 
Pitcairnia fendleri Mez, Monogr. Phan. 9: 387. 1896. 


13/12/56], 


[Between Caracas and La Guayra, 1,500 16/8/ 
55], Fendler 1550 alne, GOET, photo US). 
Tillandsia acorifolia Griseb., Nachr. Königl. Ges. Wis. 


eorg-Augusts-Univ. 1864: 19. 1865. 
[Seaside of mountains N of Colonia Tovar, 6,000 ft., 


Volume 76, Number 1 
1989 


Todzia 
Fendler's Venezuelan 
Plant Collections 


321 


22/1/56], Fendler 1771 Vip Ld GOET). 
— Guzmania acorifolia (Grise 
Tillandsia aurantiaca Griseb., Nachr. Kónigl. Ges. 
Wis. Georg-Augusts-Univ. 1864: 16. 1865. 
[Between Petaquire and Cariaca, 4, a a 12/2/58], 
Fendler 2575 (holotype, GOET, p o US). 
— Tillandsia tetrantha Ruiz Lopez n E var. au- 
rantiaca (Griseb.) Lyman B. Smit h 
h were caribaea Lyman B. Smith, Proc. Amer. 
. Arts 70: 155. 1935. 
[ n. mountains near Colonia Tovar, 6,000 ft., 14/ 
2/55, 17/6/55], Fendler 1523 (holotype, ed 


c 
N 


photo US). 
Tillandsia compacta Griseb., dw Konigl. Ges. Wis. 
Georg-Augusts- -Univ. 1864: 1865. 
[A few mi. SE of Colonia a 6, 000 ft., 25/5/54, 
21/6/54], Fendler 1508 (holotype, GOET; isotype, 
K). 


= Tillandsia compacta Griseb. var. compac 
oe excelsa Griseb. var. latifolia Crise. Suche 
igl. Ges. Wis. Georg-Augusts-Univ. 1864: 17. 


1863. 

[A few mi. SE of A zm 6,000 ft., 7/5/55], 
Fendler 1516 (GH, G 

= Tillandsia fendleri hei var. fendle 

Tillandsia fendleri Griseb., We at Königl. Ges. Wis. 
Georg-Augusts-Univ. 1864: 1865. 

[Between Caracas and Colonia Sh 6,000 ft., 7/5/ 
55], Fendler 1515 (holotype, GOET, photo US). 
= Tillandsia fendleri Griseb. var. fendleri 

Baker, J. Bot. 26: 143. 1888. 
25/5/54, 21/6/54], Fen- 


B olonia Tovar, 6,500 ft., 
dler 1509 (GH). 
Tillandsis i incurva Griseb., iw Kónigl. Ges. Wis. 
Georg- Augusts-Univ. 1864: y? 
[6 mi. SE of Colonia Tovar, Age , 28/8/54 
5/55], Fendler 1524 (holotype, GOET; isotype, CH). 
— Vriesea incurva (Griseb.) Read 
Tillandsia laxa Griseb., An Kónigl. Ges. Wis. Georg- 
Augusts-Univ. 1864: 1865. 
[Between Maracai and irn 3,000 ft., 29/1/57], 
— 2166 p.p. (holotype, GOET, phot o US). 
= Vriesia laxa (Griseb.) Mez 
Tillandsia lineatifolia Mez, Monogr. Phan. 9: 686. 
896. 
ERA Petaquire and the sea, 3,500 ft., 2/9/57], 
Fendler 2447 (GOET, a o GH). 
= Tillandsia anceps 


Tillandsia longibra racteata Baker, J. Bot. 26: 81. 1888. 


etween Petaquire and ne sea, 5,000 ft., 9/2/57], 

Fendler 2449 (synt 

pron splendens M ) Lemaire var. formosa 
itte 


Tillandsia longifolia Baker, Handb. Bromel. 185. 
1889. 


[A few mi. SE of Colonia Tovar, between rocks, 6,000- 
6,700 m, 7/5/55, 14/5/55], Fendler 1522. 
chr. Kónigl. Ges. 


[Between Maracai and Choroni, 6,500 fe, 29/1/51], 
Fendler 2159 (holotype, GOET, photo US; isotype, 
K). 


mania mucronata (Griseb.) Mez 
Tillandsia myriantha Baker, J. Bot. 25: 242. 1887. 
[7 mi. SE of Colonia Tovar, 5,000 ft., 14/5/55], 
Fendler 1530 (holotype, K; isotype, GOET). 
Tillandsia pleiosticha Griseb., Nachr. Kónigl. Ges. 
Wis. Georg-Augusts-Univ. 1864: 19. 1865. 


[High mountains E of Colonia Tovar, 7,000 ft., 11/ 
7/54, pf Fendler 1514 (holotype, GOET, 
photo US; i e, K). 

— Guzmania pleiosticha (Griseb.) Mez 

, Nachr. SE Ges. Wis. 

ug ‘Univ. (on 865. 


= . Sm ith 
Tillands sia tovarensis Mez, Monogr. Phan. 9: 769. 
[B Between Petaquire and Colonia Tovar, 26/10/57], 


Tillandsia ve 
eorg- bin s-Univ. 1864: 19. 1865. 

[High mountains E of Colonia Tovar, 7,000 ft., 7/5/ 

p? Fendler 1517 (holotype, GOET; isotypes, GH, 


= paa ventricosa (Griseb.) Mez 


BURSERACEAE 


Protium laxiflorum p var. fendleri Engl., Mono- 
g an. 4: 
[Large tree Misa a resin called “Taca mahac,” flow- 
ers gree a Tovar, 6,400 ft., 22/4/54, 19/ 
9/55], Fendler 177 (holotype, G, photo F). 


CAPPARACEAE 


Cleome moritziana Klotzsch ex Eichler, Fl. Bras. 13(1): 
250. n 
A few mi Colonia Tovar, 4,000 ft., 
Fendler Es ie enm F?). 


18/5/55], 


CELASTRACEAE 

mu fendleri Briq., Annuaire Conserv. Jard. Bot. 
Genéve 20: 360. 1919. 

[A few mi. SW of Colonia Tovar, 6,500 ft., 28/6/ 

55], Fendler 215 (F, G). 


CHLORANTHACEAE 
Vs PTET pew Solms-Laub., Prodr. 
16(1): 4 1869. 
[Colonia recta 6,500 ft., 26/7/56, horon C 4/9/ 
56], Fendler 1127 (lectotype G, photos , MO, 
NY; isolectotypes, BR, G, GH, MÓ, NY, S, US). 


COMMELINACEAE 
Phaeosphaerion efoveolatum C. B. Clarke, Monogr. 
2 . 1881. 
[Between Gar and Colonia Tovar, 6,000 ft., 23/ 
8/55], Fendler 1555 (holotype, G). 
CONVOLVULACEAE 


Ipomoea alatipes Hook., Bot. Mag. pl. 5330. 1862. 
[Between Biscaina and La Victoria, 2,000 ft., 8/9/ 


iana Kuntze, Rev. Gen. Pl. 2: 444. 


Ipomoea fendler 
1891. 


E Victoria, 2,000 ft., 
omoea calantha D 
Mohsen fruticosa Kuntze, Revis. Gen. Pl. 2: 444. 
1891 


21/11/56], Fendler 2083. 


[A few im W of Colonia Tovar, 7,000 ft., 17/4/55, 


3/2/58], Fendler 941. 


322 


Annals of the 
Missouri Botanical Garden 


[Between Turmero and Agua Blanca, 1,800 ft., 30/ 


1/57, 30/1/58], Fendler 2082 
CUCURBITACEAE 


Echinocystis polycarpa Cogn., Mém. Couronnés Autres 
Mém. Acad. Roy. Sci. Belgique 28: 90. 1878. 
gei 3, 000 ft., 23/2/54], Fendler 503 (G, photo 


Edmondia spectabilis Cogn., Monogr. Phan. 3: 420. 
1881. 
[Colonia Tovar, 6,500 ft., 5/5/54, 6/3/55], Fendler 
Mém. Couronnés Autres 
877 


4 no 
date], Fendler 490 (G, photo at F). 
Sicyos macrocarpus Cogn., Monogr. Phan. 3: 893. 
l 
[Near Citigiie, 2,000 ft., 25/1/55], Fendler 502 (syn- 
type, 
= Anomalosicyos macrocarpus (Cogn.) H. Gentry 
CYCLANTHACEAE 
Asplundia fendleri Harling, Acta Horti Berg. 18: 221. 


[In and near Colonia Tovar, 6,000-6,500 ft., 7/7/ 
54, 23/8/56], Fendler 1348 (holotype, K; e 


Evodianthus funifer (Poit.) Lindman subsp. fendler- 
uiid Acta Horti Berg. 18: 274. 1958. 

i s Hag sti, 
Fendler 2617 ae K; isotypes, GH, GOET) 


CYPERACEAE 

peine aig rudis C. B. Clarke, Kew Bull. 8: 39. 
el 

[A few mi. W of Colonia Tovar, 7,000 ft., 28/8/54, 

19/9/55, 19/7/56], Fendler 1591 (holotype, K). 

DIOSCOREACEAE 

Dioscorea fendleri R. Knuth, Pflanzenr. 4, Fam. 43: 
65. 1924. 


No information in notebook. Fendler 217 1. 


ELAEOCARPACEAE 


pas | 31 


Benth., J. Linn. Soc. Bot 5, suppl 


2: 70. 1861. 
[Seaside of mountains between Maracai and Chor 
3,000-4,000 ft., 26/10/56, 12/12/56], Fendler 


2489 (holotype, K). 


EUPHORBIACEAE 


Acalypha villosa bey var. intermedia Muell. Arg., 
Linnaea 34: 8. 1865. 
[La Victoria, 2, 500 bs 
(syntype, G-DC; isotype, 
[Near La Victoria, 2: 500 ft., 
Bos , 
nocarpus villosus (Jacq.) Kuntze var. inter- 
edius (Ma ell. Arg.) Kuntze 
Alchornea a Muell. Arg., Linnaea 34: 170. 
1865. 


21/12/56], Fendler 1825 


MO). 
14/11/56], Fendler 2408 


— Tovar, 6,500 ft., 7/4/54, 13/4/54, Fendler 
2 (G-DC, MO, photo F). 


- diera i villosus (Jacq.) Kuntze var. inter- 
medius (Muell. Arg.) Kuntze 
Euphorbia tovarensis Boiss., Prodr. 15(2): 59. 1862. 
[Maya, x 000 ft., 12/12/54], Fendler 1191 7 ain 
G; isotype, MO). 
Excoecaria obtusiloba Muell. Arg., Linnaea 32: 125. 
finata, 3,000 ft., 5/6/55], Fendler 1230 (G-DC, 
hoto F). 
Mabea rubrivenium Poeppig & Endl. 
[Colonia Tovar, 6,500 ft. 7/5/54, 16/6/55], Fendler 
1232. 


hice 15(2): 5. 1862. 
ayra, 1,000 ft., 16/8/ 


Pedilanthus fendleri Boiss., 
[Between Caracas and La 
55], Fendler 1202 (G-DC, as ind F). 
grin penninervia Muell. Arg., Linnaea 34: 158. 
865. 
ed Biscaina, 3,000 ft., 1/2/57], Fendler 2412 
(holotype, G-DC, photo 
Bs fendleri Muell. Arg., Linnaea 34: 179. 1865- 
18 


E 3,000 ft., 5/6/55], Fendler 1208 (holotype, 
G-DC, photo F). 


FABACEAE 


Calliandra laxa var. parvifolia Benth., Trans. Linn. 
Soc. London 30(3): 551. 1875. 
[Between La Victoria and Biscaina, 2,500 ft., 
51], Fendler 2255 (K). 
Eti fendleri Piper, Contr. U.S. Natl. Herb. 
20(14): 570. 1875. 
[Biscaina, 7/9/55], Fendler 248 (holotype, K; isotype, 
MO). 


1/2/ 


Cassia viciifolia Benth., Trans. Linn. Soc. London 27: 
544. 1871. 
[Between Turmero and La Victoria, 2,000 ft., 14/3/ 
57], Fendler 2230 (lectotype, K). 
= Senna viciifolia (Benth.) Irwin € Barneby 
Benth., Trans. Linn. Soc. London 


0. 1875. 
[Valley e Macarao, 3,500 ft., 18/1/57], Fendler 2259 
K). 


Machaerium oe Benth., J. Proc. Linn. Soc., 
Bot. 4(suppl.): 58. 1860. 
ies ad Macarao, 4, 000 ft., 4/7/56], Fendler 1750 
(syntype, K). 
[Near La Victoria, 2,200 ft., 21/11/56], Fendler 1866 
(syntype, 
Machaerium Me Pittier, Contr. U.S. Natl. Herb. 
20(3) 121. 1917. 
[La Victoria, E 000 ft., 14/11/56], Fendler 1865 
(holotype, GH; pie MO). 


imosa tov enth., Trans. Linn. Soc. London 
30(3): 407. 1875. 

m da p ft., 13/2/54], Fendler 344 (holotype, 
K; iso 


Moldenhawera mollis Benth., Fl. Bras. 15(2): 77 
[La Victoria, 2,000 ft., 21/11/56], Fendler 1864 (K). 
GENTIANACEAE 
iiri m magnificus Gilg, Engl. Bot. 22: 343. 
896. 


[10 mi. SE of Colonia Tovar, 5,000-6,000 ft., 8/4/ 
54, 19/6/54, 4/9/54], Fendler 830 (B, MO, photo 
F). 


Volume 76, Number 1 
1989 


Todzia 323 
Fendler's Venezuelan 
Plant Collections 


— Symbolanthus calygonus (Ruíz Lopez & Pavón) 


GESNERIACEAE 

Columnea fendleri Sprague, Kew Bull. 1912: 41. 
1912. 

[Between Petaquire and the sea, 5,000 ft., 


Fendler 2031 (probably K). 
— Columnea scandens L. var. fendleri (Sprague) Wieh- 
ler 


9/2/51], 


Monopyle leucantha Moritz ex Benth., Icon. Pl. 12: 
87. 1876. 


[Colonia Tovar, 5,800-6,200 ft., 31/2/54, 27/3/ 
54, 10/5/55, 17/6/54, 29/8/54], Fendler 794 


LAMIACEAE 
Salvia tovariensis m Annuaire Conserv. Jard. Bot. 
Gen 898. 
[Between ee oa Maya, 3,200 ft., 12/2/54, 12/ 
54], Fendler 876 (G). 
Suchys — Briq., Annuaire Conserv. Jard. Bot. 
115. 1898. 


[Near iid Tovar, 6,500 ft., 31/2/54, 19/12/54, 
20/6/55, 21/9/55], Fendler 879 (G, MO, photo 
F). 


LAURACEAE 
Gymnobalanus fendleri Meissn., Prodr. 15(1): 142. 
1864. 
Ex Valencia and Campanero, 5,500 ft., 7/3/ 
51], Fendler 2395 Coe ? PI 
= Ocotea fendleri (Meissn.) R 
Gymnobalanus latifolia uu Prodr. 15(1): 141. 
1864. 
[Between Colonia Tovar and Petaquire, 5,000 ft., 19/ 
2/55], Fendler 1095 (G-DC, photo F). 
Mespilodaphne vaginans Meissn., Prodr. 15(1): 104. 
1864. 


[Colonia Tovar, 6,000 ft., 21/6/54, PN Fen- 
dler 1098 (types, G-DC, photo F, G, K, MO 
US). 


— Ocotea vaginans (Meissn.) Mez 
Nectandra leucantha Nees & o var. attenuata 
Meissn., Prodr. 15(1): 151. 
[La Vistos. 2,000 ft., ee Fendler 2393 
(syntypes G, K, MO). 


LECYTHIDACEAE 


gx fendleriana Miers, Trans. Linn. Soc. 30: 
. 60, fig. 6. 1874. Fendler s.n. (K). 
- i Leg fendleriana (Miers) Kunth 


LILIACEAE 


Smilax ET H.B. a e fendleri A. DC., 
Mono. . 1: 143. 
[Near Colonia Tovar, ipu E m ft., 19/6/54, 3/ 
12/54, 16/9/55], Fendler 1546 (G, MO, photo at 
F). 


LOGANIACEAE 
à etw fendleri Sprague & Sandw., Kew Bull. 1927: 
29, 1927. 


b. Victoria, 2,500 ft., 21/11/56], Fendler 2314 (K). 


LORANTHACEAE 


Dendrophthora basiandra Kuijt, Wentia 6: 32. 1961. 
[Colonia Tovar, 6,800 ft., 10/8/54], Fendler 1104 
(holotype, GH; isotypes, GH, K). 
Dendropthora eichleriana Urban, Ber. Deutsch. Bot. 
Ges. 14: 287. 1896. 
[Colonia Tovar, 6,800 ft., 14/2/55], Fendler 1101 
(holotype, K; isotypes, GH, 
Phoradendron a Da. Bot. Jahrh. Syst. 
23, Beibl. 57: 97. 
[La Victoria, 2, 000 * , 21/1/56], Fendler 1811. 
Phoradendron condicion Urban, Bot. Jahrb. Syst. 
23, Beibl. 57: 5. 1897. 
[Biscaina, 3,200 ft., 5/6/55], Fendler 11 
Phoradendron i fendicrianum Eichler, Fl. E 5(2): 
129. 1868. 
[Colonia Tovar, 8/4/54, 10/5/54, 9/3/55], Fendler 
102 (MO). 


grep i. longipetiolatum Urban, Bot. Jahrb. 
Syst. 23, Beibl. 57: 6. 1897. 
[Colonia Tovar, 6,500 ft., 10/7/56], Fendler 1762. 
Phoradendron ovalifolium Urban, Bot. Jahrb. Syst. 
: 97. 


[Biscaina, 3,500 ft., 10/5/55], Fendler 1108. 
Phoradendron rigidum Urban, Bot. Jahrb. Syst. 23, 
Beibl. 57: 7. 1897. 
[Colonia Tovar, 6,500 ft., 
cin tovarense Urban, Bot. Jahrb. Sy 
Beibl. 5 8 
[A few mi. se: at Coliinia Tovar, 6,000 ft., 
Fendler 1 
= E. dipterum Eichler 
Phoradendron tubulosum Urban, Eng. Bot. Jahrb. 
Bei . 1897 


1/6/55], Fendler 1105. 
st. 23, 


26/1/56], 


7/9/55), Fendler 1106 (syntype). 


23, le 
[Biscaina, 3, ue ft., 
ndro e Trel., Genus Phoraden- 


Phorade venezuelens 
dron, 
[La Victori; 2; 000 ft., 
olotype, K; isotype, G 
wa dendron quandrangulare (H.B.K.) Krug & 


21/11/56], Fendler 1810 


Stachyphyllum fendleri van Tiegh., Bull. Soc. France 
565. 1895. 


Cun Tovar, 6,500 ft., 15/8/54, 24/10/55, 8/ 
11/56], Fendler 1125 ‘(holotype, P; isotypes, GH, 
MO). 


= Antidaphne viscoidea Poeppig & Endl. 


MELASTOMATACEAE 


Arthrostemma alatum Triana, Trans. Linn. Soc. Lon- 
don 28: 35. 1871. 


[Near La Victoria, 2,000 ft., 21/11/56], Fendler 1832 
(G, MO, photo F). 
Blakea grisebachii Cogn., Monogr. Phan. 7: 1071. 
1891. 


[Seaside of mountains, N of Agua Blanca, 4,000 ft., 
30/1/58], Fendler 2547 (syntypes, GH, GOET). 
Blakea longibracteata Cogn., Monogr. Phan. 7: 1074. 
1891. 
[Between Petaquire and the seashore, 5,500 ft., 22/ 
4/58], Fendler 2593 (holotype, GOET). 
Calophysa ? flexuosa Triana, Trans. Linn. Soc. London 
28: 141 
[Between ee and the sea, 4,000 ft 
Fendler 


= Clidemia d (Triana) Cogn. 


., 9/2/51], 


324 


Annals of the 
Missouri Botanical Garden 


Clidemia ampla Cogn., Monogr. Phan. 7: 1023. 1891. 
[Between Petaquire and the seashore, 5,500 ft., 22/ 
4/58, Fendler 2594 (syntypes, F, G, GH, GOET, 


K, MO). 
—— ciliata D. Don var. grandifolia Cogn., 
Monogr. Phan. 7: 1020. 1891. 
iiia prr and Choroni, 3,000 ft., 
Fendler 
= ads x D. Don var. ciliata 
~~ fendleri Cogn., Monogr. Phan. 7: 1016. 
891. 


24/1/57], 


[Mountains near Caracas, a few mi. SE of Colonia 
Tovar, 17/2/54, 7/5/55, 19/6/55, 3/1/55], 
Fendler 438 (G, MO, photo 

— grandifolia Cogn., Monogr. Phan. 7: 1018. 
91. 

[Bet etween Maracai and Choroni, 4,000 ft., 28/1/57], 
Fendler 2263 (syntypes, G, MO). 

Henriettella tovarensis Cogn., Monogr. Phan. 7: 1044. 
1891. 


[Near Colonia Tovar, 5,600 ft., 6/4/54, 22/9/55, 
3/10/55], Fendler 444 p.p. (G, photo F, MO). 
Henriettella tovarensis Cogn. var. angustifolia Cogn., 

Monogr. Phan. 7: 1044 l; 
[Near Colonia Tovar. 5, 600 ft., 6/4/54, 22/9/55, 
3/10/55], Fendler 444 p.p. (syntype, G, photo F). 
Heterotrichum glandulosum Cogn., Monogr. Phan. 
7: 955. 18 


[A few mi. SE a Colonia Tovar, 5,100 ft., 
3/1/55], Fendler 432 (G, photo 
= Micronia araguensis Wurdac 
Heterotrichum lucidum Triana, Trans. Linn. Soc. 
London 28: 134. 1871. 
[Colonia Tovar, 6,500 ft., 


4/9/54, 


2/9/56], Fendler 1845 


( 
Leandra fendleri Cogn., Monogr. Phan. 7: 653. 1891. 
[Near Colonia Tovar, 7,000 ft., 12/6/54, 3/12/54, 
12/5/55, 27/8/55], Fendler 439 (G, MO, photo 

F). 


— Leandra lindeniana (Naud.) Cogn. 
Meriania subumbellata Cogn., Monogr. Phan. 7: 430. 
1891. 


[Between Maracai and € 4,000 ft., 
29/1/57], Fendler 18: 
Miconia fendleriana n Monogr. Phan. 7: 822. 
1891. 


12/12/56, 


[Between Maracai and Choroni, 4,000 ft., 
Fendler 1835 (syntype, GOET). 
[Between Maracai and Choroni, 6,000 ft., 
Fendler 1836 (syntype, G 
Miconia multinervulosa Cogn., Monogr. Phan. 7: 926. 


12/12/56], 


12/12/56], 


[Near Colonia Tovar, 6,500 ft., 15/5/55, 23/8/56], 
Fendler 415 (syntypes, pho to i 
— Miconia theaezans (Bonpl.) 
Miconia resimoides oa. ae Phan. 7: 926. 
1891. 
[Near Colonia Tovar, 6, Ped S 15/5/55, 23/8/56], 
Fendler 414 (G, MO, 
Miconia rubens Naudin var. "latifolia Cogn., Monogr. 
Phan. 7: 921. 1891. 
[Between poi and Choroni, 4,000 ft., 
Fendler 1 
Miconia euam Cogn., Monogr. Phan. 7: 917. 
1891. 


12/12/56], 


[Colonia Tovar, 6,500 ft., 29/4/54], Fendler 419 (G, 
photo F, MO). 
— Miconia rubens Naudin var. rubens 


Tibouchina moritziana Cogn., Monogr. Phan. 7: 259. 
1891. Fendler 404 p.p. (MO). 
— T. geitneriana (Schltr.) Cogn. 


MELIACEAE 
— montana H.B.K. var. fendleriana C. DC., 
Monogr. Phan. 1: 654. 1878. 
[Colonia Torat. 6,300 ft., 17/5/54, 21/6/55, 24/ 
10/55], Fendler 139 (G, photo F). 
= Trichilia pallida Sw. 


MONIMIACEAE 


Siparuna venezuelensis Perkins, Bot. Jahrb. Syst. 28: 
679. 1901 


[Between Valencia and Campanero, 7/4/57], Fendler 
2358. 


MYRSINACEAE 


Ardisia robinsonii Mez, Pflanzenr. 4, Fam. 236: 77. 
1902. 
[Between Caracas and Colonia a 6,000 ft., 30/ 
4/57], Fendler 2357 (GH, o F). 
Conomorpha caracasana Mez, ‘neal 4, Fam. 236: 
260. 1902. 
[Colonia Tovar, 6,500 ft.], Fendler 751 (MO, photo 


Conomorpha glabra Mez, Pflanzenr. 4, Fam. 236: 
254. 1902. 


[High mountains around Colonia Tovar, 7,000 ft., 19/ 
3/55, 23/6/55], Fendler 752 (paratype, MO, photo 
F). 


r laurifolia Mez, Pflanzenr. 4, Fam. 236: 
254. 19 


[Crest of mountains near Colonia Tovar, 7,000 ft., 9/ 
11/54, 14/2/55], Fendler 756 (G, MO, photo s 
barer n Mez, Pflanzenr. 4, Fam 
8. 19 


[Between as and Colonia Tovar, 6,800 ft., 11/ 
2/57], Fendler 2015 (G, GH, MO, photo F). 
= klotzschii Mez, Pflanzenr. 4, Fam. 236: 

1902 


co. Tovar, 7,000 ft., 31/8/54, 1/9/54], Fendler 
3 sheeets, MO). 
Parathesis moritziana Mez, Pflanzenr. 4, Fam. 236: 
180. 1902. 
[Colonia Tovar, 6,500 ft., 6/4/54], Fendler 754 (ho- 
lotype, MO). 


ORCHIDACEAE 


Repert. Spec. Nov. Regni 
eih. 6: 30. 
No information in notebook. Fendler 
iia oam psilosepala Renz, Candles 11: 245. 


Cranichis fendleri Schltr., 
Veg. Be 1919. 


No [i PNE in notebook. Fendler 1402 (isotype, 
MO). 
Liparis fendleri Schltr., Repert. Spec. Nov. Regni Veg. 
Beih. 6: 32. 191 
No information in nobel Fendler 1422 (MO). 
— Liparis brachystalix Reichb. 
diem MAI rhynchus Reichb. f., Xenia Orch. 


Fendler s 
= praia brachyrrhyncha (Reichb. f.) Garay 


Volume 76, Number 1 
1989 


Todzia 
Fendler's Venezuelan 
Plant Collections 


Physurus hyphaematicus Reichb. f., Xenia Orch. 2: 
184. 1873. 


Fendler s.n 
— Aspidogne hyphaematica (Reichb. f.) Garay 
Physurus xystophyllus Reichb. f., Xenia Orch. 2: 183. 
1873. 


n. 
Pleurothallis apiculata Lindley, Fol. Orch. Pleuroth. 
17. 1859. 


o information in notebook. Fendler 2155 (K). 
Pleurothallis clandestina Lindley, Fol. Orch. Pleu- 
oth. 43. 1859. 


No information in notebook. Fendler 2148 (K). 
Pleurothallis a Lindley, Fol. Orch. Pleuroth. 
15. 1859. 


evasnerat 


No information in notebook. Fendler 1490 (K). 
— Myoxanthus exasperatus (Lindley) Luer 
Pleurothallis viridula Lindley, Fol. Orch. Pleuroth. 
19. 1859 


No information in notebook. Fendler 1486 (K). 
— Restrepiopsis viridula (Lindley) Luer 
Stelis alata Lindley, Fol. Orch. Stelis, 18. 1858. 

o information in Y natem Fendler 2154 (K). 
Stelis coriifolia Lindley, Fol. Orch. Stelis, 18. 1858. 
o information in notebook. Fendler 2444 (K) 
Stelis fendleri Lindley, Fol. Orch. Stelis, 18. 1858. 

o information in notebook. Fendler 1470 (K, MO). 
Stelis humulis Lindley, Fol. Orch. Stelis, 10. 1858. 
No information in notebook. Fendler 1467 (K). 
Stelis lutea Lindley, Fol. Orch. Stelis, 7. 1858. 


K). 
Stelis tenuilabris Lindley, Fol. Orch. Stelis, 18: 1858. 
No information in notebook. Fendler 1469 (K 
No information in notebook. Fendler 1471 (K, MO). 


PASSIFLORACEAE 


Passiflora velata a: Fl. ee e 560. 1872. 
[Between Turmero La Victoria, 2,000 ft., 14/3/ 
57], Fendler 2329 a bed by Killip, 
1938, K 


). 
= Passiflora serrulata Jacq. 


PIPERACEAE 


Peperomia adscendens C. DC., J. Bot. 4: 140. 1866. 
[Colonia Tovar, 6,500 ft., 1/4/54, 20/5/54, 3/6/ 
55], Fendler 1153 (holotype, G-DC; isotypes, GH, 
K, MO, NY, PH; also the type of Peperomia glabra 

C. DC 


Peperomia blanda H.B.K. var. dissimilis C. DC., 
Pe 16(1): 459. 1869. 
mi. W of perd es 7,000 ft., 
Fendler 1170 (GH, MO, NY). 
Peperomia carlosiana C. DC., J. 
[Valley of San Carlos River, 3,500 ft., 
Fendler 1148 (holotype, C. photo 


). 
= Peperomia lanceolata-peltata var. carlosiana (C. 


28/6/55], 


Bot. 4: 140. 1866. 
25/10/54], 


er 
C. DC., J. Bot. 4: 141. 1866 
12/12/57], 
Fendler 2402 (holotype, G, photo . 

, J. Bot. 4: 143. 1866. 
Co bos Tovar, 7,000 ft., 28/ 
9/55], Fendler 1169 (G, photo F). 


Peperomia diffusa C. DC., J. Bot. 4: 133. 1866. 
[A few mi. S of Colonia Tovar: Biscaina, 3,000-4,000 
ft., CE 5/6/55], Fendler 1178 (holotype, 
G-DC; isotypes, GH, K, MO, N 
Peperomia ellipticifolia C. DC., An 16(1): 463. 
1866. 


[Colonia Tovar, 6,500 ft., 10/5/54, 14/5/55], Fen- 
dler 1171 (holotype, e i isotype, GH). 

— Peperomia Mente hg (Ja .K. 

Peperomia ernstiana C. DC. Prodr. 16(1): 397. 1869. 

[Near the River Tuy, 5,000-5,500 ft., 2/6/54, 29/ 
7/54, 20/6/55, 3/7/55], Fendler 1163 (holotype, 
G-DC; isotypes, GH, MO, NY, PH). 

= Peperomia venezueliana 


C. DC. 
Peperomia fendleriana C. DC., Prodr. 16(1): 460. 
1869. 


[Colonia Tovar, 6,500-7,000 ft., 20/5/54, 16/6/ 
54], Fendler 1174 (syntype, G- DC; isotypes, GH, 
MO, NY, PH). 

Peperomia fragrans C. DC., J. Bot. 4: 140. 1866. 

[Maya, banks of River Tuy, y 000 ft., 20/6/55], Fen- 
dler 1156 (holotype, G-DC; VAR GH, K). 

Peperomia glabra C. J. Bot 143. 1866. 

[Colonia Tovar, 6,500-7, 000 ft., Py m Fendler 
1153 (holotype, K; isotype, CH: also the type col- 
lection of Peperomia indian DC. 


: . Bot. 4: 139. 1866. 
E of Colonia Tovar, 6,000 ft., 7/5/55], Fen- 
dle: 1164 (G, photo F; isotype, CH). 
— Peperomia alata Ruiz Lopez & P 
Peperomia lanceolata-peltata C. DC., F Bot. 4: 136. 
1866. 


aes 3i se ar 13/8/55], Fendler 1149 (syntype 
h Háffnann 414 (at B); isosyntypes, 
CH, K, MO, PH 
Peperomia linca. C. DC., J. Bot. 4: 145. 1866. 
[Colonia Tovar, 6,500-7,000 ft., 17/6/54, 3/11/ 
56], Fendler 1176 (syntype with Jameson s.n., both 
G-DC; isosyntypes, GH, Ha NY, P 
Dor var. emargi- 
u 
ear Colonia Tovar, 
Fendler 1152 (also 
ides. mds A. G-DC (per Trel. & Yuncker, 
Piperac. No. S. Amer. 685. 1950), isolectotypes, 
GH, MO, NY, PH). 
= Ce gri; n (L.) A. Dietr. var. emarginata 
(C. DC) T. 
Peperomia ce C. DC., J. Bot. 4: 133. 1866. 
[Near Colonia Tovar, 6, 500 ft., 16/5/54, 26/11/ 
56], Fendler 1166 (holotype, [3 photo F). 
Peperomia pseudodependens C. DC., J. Bot. 4: 137. 
1866 


[La Victoria, 2,000 ft., 20/11/56], Fendler 1817 
(holotype, G, photo F; isotype, G 
Peperomia pseudopeltoidea C. DC., Prodr. 16(1): 
434. 1869. 


[Colonia Tovar, 6,500 ft., 6/6/54, 16/5/55], Fendler 
11506 (G, photo F). 
Peperomia purpurella C. DC., Prodr. 16(1): 414. 
1869. 
[River Tuy below E im 5,000 ft., 20/8/54], 
Fendler 1161 (G, pho 
— Peperomia patula C. DC. 
Peperomia reflexa (L.) A. Dietr. var. coriacea C. DC., 
Prodr. 16(1): 452. 1869. 


326 


Annals of the 
Missouri Botanical Garden 


[Colonia Tovar, 6,500 ft., 3/4/54, 2/5/54, 5/8/54], 
Fendler 1183 (holotype, G-DC, isotypes, GH, MO, 
= Pe pe romia reflexa (L.) A. Dietr. 

Peperomia san- Kir prie C. DC., J. Bot. 4: 138. 


[San Carlos River valley, 3,500 ft., 
1151 (holotype, G, da 
Peperomia succu len C. DC., J. Bot. 4: 142. 1866. 
[10 mi. S of Colonia cite 3, 500 ft., 10 mi. S of 

Colonia Tovar, 18/5/55], Fendler 1157 (holotype, 


7/9/54], Fendler 


e, K). 
A. Diet. var. glabra C. DC., Prodr. 
16(1): 3 vi 
[Between ila and Choroni, 3,000 ft., 12/12/56], 
Fendler 1818 (holotype, G-DC; isotypes, GH, K, 
, jJ 
Pe mia tovari C. DC., Prodr. 16(1): 404. 1869. 
[Between Colonia "To ovar and Petaquire, 19/4/ 55], 
Fendler 1168 (syntype at G-DC with Moritz 1941 
(at BM); isosyntypes, GH, MO, NY, PH). 
Peperomia trinervula C. DC., Prodr. 16(1): 420. EE 
[High mountains E of Colonia Tovar, 7,000 ft., 
9/ 55], Fendler 1167 (lectotype, G-DC; Aut 


Peperomia venezueliana C. DC., J. Bot. (Hooker) 4: 
139. 1866. 


[Between Colonia Tovar and Petaquire, 6,500 ft., 22/ 
4/58], Fendler 2618 (holotype, G, photo F). 
ink var. firmius C. 


869. 
[Colonia Tovar, 6, 000 ft., 28/4/54, 7/4/54], Fendler 
a (syntype at G- DC with Bredemeyer 679 (at 
Willd.); 2 sheets at GH; isotype, GH). 
= Paer caracasanum Bredem. ex Link 
Piper cumbotianum C. DC., Prodr. 16(1): 268. 1869. 
[Between Agua Blanca and Cumbote, 5,000 ft., 30/ 
1/58], Fendler 2571 (G, photo F). 
Piper fendlerianum C. DC., J. Bot. 4: 213. 1866. 
[10 mi. E of Colonia Tovar, 1. 500 ft., 26/5/55, 17/ 
6/55], Fendler 1140 (G, photo F; isotypes, GH, K, 
PH). 


Piper glanduligerum C. DC., J. Bot. 4: 215. 
[Colonia Tovar, 6,500 ft., 6/4/54, 27/6/56], cid 
1137 ui Ee G, photo F; isosyntypes, GH, 
NY, U 


Piper dera C. DC., Prodr. 16(1): 289. 1869. 
[Colonia Tovar, 6,500 ft., 16/6/55], Fendler 1144 
(holotype, G-DC; €— GH, , PH) 
Piper ovale C. e . Bot. 4: 166. 16 
i Colonia Tovar, 4,000 ft., 
Fendler 2398 (syntype, G-DC; isosyntype, 
crassinervium 


iper 
Piper tovarense Trel. - Yuncker, Piperac. N. South 
Amer 


27/2/57], 
GH). 


: 27, fig. 
[10 mi. E of Colonia Tort, 7,000 ft., 17/6/55], 
,G 
Piper tristemon C. DC., Prodr. 16(1): 328. 1869. 
[1 of Co lonia Tovar, 7,500 ft., 26/5/55, 17/ 
6/55], Fendler 1141 (holotype, G-DC, photo F; is- 
oty (0) 
Piper (aii o = DC., J. Bot. 4: 216. 1866. 
[Between Agua Blanca and Cumbote, 5,000 ft., 30/ 
1/58], Fendler 2572 Ub. G-DC, photo F; iso- 
types, GH, MO, P 
— Piper glabrescens Va ) C. DC. var. venezuelense 
(C. DC.) Trel. 8 Yuncker 


Piper victorianum C. DC., J. Bot. 4: 218. 1866 

La Victoria, 2,000 ft., 21/11/56], Fendler 1139 
(holotype, G-DC, photo F; isotypes, GH, K, MO, 
PH). 


= Piper amalgo var. medium (Jacq.) Yuncker 


— 


POACEAE 


Chusquea fendleri Munro, Trans. Linn. Soc. 26: 61. 


[In and ‘about Colonia Tovar, common, 6,000- 7,900 
ft., 17/4/55], Fendler 1627 (holotype, K; isotype, 
MO). 


PODOCARPACEAE 
Podocarpus — Pilger, Pflanzenr. IV. 5(Heft 
18): 69. 1903. 
[Colonia E 6,500 ft., 6/4/54], Fendler 1289. 
= Prumnopitys harmsiana (Pilger) Laubenf. 


POLYGALACEAE 


Polygala fendleri Chodat, Mém. Soc. Phys. Genéve 
1, pt. 2, no. 2: 167. 1893. 

i. SW of Colonia Tovar, 5,000 ft., 13/9/55, 

8/9/56], Fendler 238 (G). 
Polygala tovarensis Chodat, Mém. Soc. Phys. Genéve 

2, no. 2: 32. 1893 

[Caracas: RUM ew mi. of Colonia Tovar, 500- 
6,500 ft., 31/1/54, 13/9/55, 19/7/56], Fendler 


240 (G). 
rg fendleri Chodat, Bull. Herb. Boissier 3: 
1895. 


Pendler s.n. 
RUBIACEAE 
Cephaelis aneurophylla Standley, Publ. Field Colum- 
bi us., - Ser. 8: 63. 
[Between Maracai and Choroni, 5, one ft., 12/12/56], 
Fendler 1979 Gas ype, GH; isoty dni 


— Psychotria pta (Standl. ) Sieva 
Cephaelis macrocarpa Standley, Publ. Field Colum- 
= ed ot. Ser, : 64. 1930. 
Bet ai and Choroni, 5,000 ft., 12/12/56], 
Fendler 1990 (holotype, GH; isotype, K). 
= Psychotria costanensis Steyerm. subsp. costanensis 
Elaeagia karstenii oe Field Columbian Mus., 
Bot. Ser 1930 
[Colonia Tovar, 13/8/54], Fendler 594 (holotype, GH; 
isotypes, F, MO). 
Malanea fendleri rd Publ. Field Columbian Mus., 
1930 


[Between Petaquire and Cariaca, 6,000 ft., 12/2/58], 
Fendler 2555 (holotype, GH). 
Malanea hirsuta ger Publ. Field Columbian Mus., 
Bot. Ser. 8: 65. 1930. 
[A few mi. S of Colonia Tovar, 3,500 ft., 16/5/55], 
Fendler 569 (holotype, GH; isotypes, K, M, MO, 
Y). 


Manettia schumanniana Sprague, Bull. Herb. Boiss. 
2(5): 834. 1905. 

[A few mi. E of Colonia Tovar, 7,500 ft., 
Fendler 589. 

Palicourea fendleri Standley, Publ. Field Columbian 
iris ot. 

[Between Maracai and Chor oni, E^ 000 ft., 29/1/57], 

Fendler 1983 na GH, photo F; isotype, K). 


5/1/57), 


Volume 76, Number 1 
1989 


Todzia 327 
Fendler's Venezuelan 
Plant Collections 


Rudgea ES Standley, Publ. Field Columbian 
Mus., : 72. 1930. 
[Between Pide A dd the sea, 4,000 ft., 9/2/57], 
Fendler 1993 (holotype, GH, photo F; isotype, K). 


SAPINDACEAE 


Matayba longipes Radlkofer, Sitzungsber. Math-Phys. 
Cl. Kónigl. Bayer. Akad. Wiss. München 9: 626. 
1879. 


[Colonia Tovar, 6,500 ft., RON Fendler 1748 
(holotype, G, photo F; isotype, M 


SAPOTACEAE 


Crepindodendron crotonoides Pierre, Not. Bot. Sap. 


A SO dnd a 100 ft., 26/1/57], Fendler 
749 (holot 


= Pouteria E (Purre) Baehni 


SOLANACEAE 


Cestrum laetum Francey, Candollea 6: 378. 1935. 
[Colonia Tovar, 6,500 ft., 13/4/54, 7/5/54], Fendler 
955 (holotype, NY 
Cestrum tovarense Francey, Candollea 6: 388. 1935. 
(Colonia Tovar, el ft., 31/8/55], Fendler 962 
(holotype, NY; isotype, MO). 
Saracha nitida Bitter, Repert. Spec. Nov. Regni Veg. 
4. 


[10 mi. E r Colonia Tovar, 
Mer 1007 (G, photo 
Solanum tovarense Bitter, Repert. Spec. Nov. Regni 
Veg. 18: 65. 1922. 
[Colonia Tovar, 6,000 ft., 27/4/54, 16/8/55, 22/ 
9/55], Fendler 980 (G, photo F) 


17/6/55, 21/9/55], 


STYRACACEAE 


qq macrotrichus Perkins, Repert. Spec. Nov. Reg- 
1906. 
[A ue mi. SE of Colonia Tovar, 4,500 ft., 9/3/55], 
Fendler 746 (G, MO, photo F). 


TURNERACEAE 


Piriqueta Hoe Urban, Jahrb. Konigl. Bot. Gart. 
Berlin 2: 69. 1883 
pou s of Calania Tovar, 4,000 ft., 
Fendler 


13/9/55], 


URTICACEAE 

Hemistylis macrostachya Wedd., Ann. Sci. Nat. (Paris) 
qn 

n Caraca La Guayra, 1,000 ft., 


and 
55] Fendler 1259 (G, photo 
Hemistylis odontophylla Wedd., Prodr. 16(1): 235. 


16/8/ 


l 
E Vitara, 2,000 ft., 21/11/56], Fendler 1827 (G, 
oto 
Phenax uliginosus Wedd., Ann. Sci. Nat. (Paris) 4(1): 
192. 
[Colonia px 31/4/54, 21/8/55], Fendler 1268 
(C, MO, photo F). 
ey cg e eu Ann. Sci. Nat. (Paris) ser. 


* h mountains d Colonia Tovar, 7,000 ft., 10/ 
5/54, 24/6/55], Fendler 1246 (G, MO, photo F). 


Pilea mollis Wedd., Arch. Mus. Hist. Nat. 8: 251. 
1855. 
[River Tuy below Colonia Tovar, 4,000-5,500 ft., 2/ 


7/54, 29/8/54, 22/4/58], Fendler 1241 (G, MO, 


hoto F). 
Bilge on urticella Wedd., Prodr. 16(1): 157. 1869. 
[Between Valencia and Campanero, 4,000 ft., 7/3/ 
57], Fendler 2429 (G, MO, photo F). 
Pouzolzia acalyphioides Wedd., Prodr. 16(1): 234. 


(Bicis, 3,000 ft., 10/5/55], Fendler 1264 (G, pho- 
to F). 


VERBENACEAE 


Aegiphila fendleri Moldenke, Repert. Spec. Nov. Reg- 
ni ; 33. 
[Between Agua Blanca and Cumbote, 4,000 ft., 30/ 
/58], Fendler 2032 (holotype, GH; isotypes, G, 


K). 
Aegiphila floribunda pu Moldenke, Repert. Spec. 
Nov. Regni Veg. 33: 122. 1933. 
diera Tovar, 6,000 ft., ioo 16/6/55], Fen- 
dler 845 (MO, W, photo F). 
Aegiphila lewisiana e Repert. Spec. Nov. 
Regni Veg. 33: 131. 
Troes Caracas and La Bx. 2,000 ft., 
55], Fendler 844. 
Citharexylum venezuelense Moldenke, Repert. Spec. 
Nov. Regni Veg. 37: 237. 1934. 
ira Caracas and La Guayra, 1,000 ft., 
55], Fendler 1298. 


16/8/ 


16/8/ 


APPENDIX II 


List of Fendler pp A cited in Appendix 


I by number and their basio 
27 Cleome moritziana Ei ex Eichler (Cappara- 
ae) 


ce 

115 Piriqueta den Pear Urban (Turneraceae) 

139 deir montana H.B.K. var. fendleriana C. 
C. (Meliac 

177 peo jur Engl. var. fendleri Engl. 


i Briq. 
Polygala fendleri Chodat (Polygalaceae) 
is Chodat (Po 


varensis Benth. (F 
404 Tibouchina moritziana Cogn. (Melastomataceae) 
Henriettella tovarensis Cogn. var. angustifolia 
. Cogn. (Melastomataceae) 
Miconia resimoides Cogn. (Melastomataceae) 
Miconia multinervulosa Cogn. (Melastomata- 
ceae 
419 Miconia tovarensis Cogn. (Melastomataceae) 
Heterotrichum glandulosum Cogn. (Melastoma- 
taceae 
Clidemia E D. Don var. grandifolia Cogn. 
(Melastomatace 
Clidemia fendleri Cogn. (Melastomataceae) 


Henriettella tovarensis Cogn. var. angustifolia 
Cogn. (Melastomataceae) 
Edmondia spectabilis Cogn. (Cucurbitaceae) 


328 Annals of the 
Missouri Botanical Garden 
490 Gurania wagneriana Cogn. (Cucurbitaceae) 1144 Piper nobile C. DC. (Piperaceae) 
502 Sicyos macrocarpus Cogn. (Cucurbitaceae) 1145 Piper tovarense Trel. (Piperaceae) 
503 Echinocystis polycarpa Cogn. (Cucurbitaceae) 1148 Peperomia carlosiana C. DC. De A 
513 Begonia fagopyroides Kunth & Bouché var. fen- 149 Peperomia lanceolata-peltata C. DC. (Pipera- 
dleriana A. DC. (Begoniaceae) 
524 OA venezuelensis Rose ex Mathias (Api- Mrs Peperomia pseudopeltoidea C. DC. (Piperaceae) 
Peperomia san-carlosiana C. DC. (Piperaceae) 
529 os NM fendleri Seemann (Araliaceae) Dos Peperomia magnoliifolia A. Dietr. var. 
569 Malanea hirsuta Standley (Rubiaceae) inulata C. DC. (Piperaceae) 
589 Manettia schumanniana Sprague (Rubiaceae) 1153 Peperomia adscendens C. DC. (Piperaceae) 
594 Elaeagia karstenii Standley (Rubiaceae) 1153 Peperomia glabra C. DC. (Pip e 
625 Mikania pa Klatt (Asteraceae) 1156 Peperomia fragrans C. DC. (Piperaceae) 
626 Mikania hexag . L. Robinson (Asteraceae) 1157 Peperomia succulenta C. DC. (Piperaceae) 
634 Eupatorium neritfolibm B. L. Robinson (Aster- 1161 Peperomia purpurella C. DC. (Piperaceae) 
1163 Peperomia ernstiana C. DC. (Piperaceae) 
638 Eupatorium xestolepis B. L. Robinson var. xes- 1164 Peperomia laevis C. DC. (Piperaceae) 
tolepis (Asteraceae) 1166 Peperomia patula C. DC. (Piperaceae) 
647 Eupatorium steetzii B. L. Robinson (Asteraceae) 1167 Peperomia trinervula C. DC. (Piperaceae) 
651 Trichogonia rhadinocarpa B. L. Robinson (As- 1168 Peperomia tovariana (Piperaceae) 
teraceae) 1169 Peperomia decurrens C. DC. (Piperaceae) 
702 Senecio cucullatus Klatt (Asteraceae) 1170 Peperomia bl H.B.K. var. dissimilis C. DC 
718 Hieracium tovarense Fries (Asteraceae) (Piperaceae) 
746 Styrax macrotrichus Perkins (Styracaceae) 1171 ti elli ipticifolia C. DC. (Piperaceae) 
749 Crepindodendron crotonoides Pierre (Sapota- 1174 Peperomia fendleriana C. DC. (Piperaceae) 
ceae 1176 Peperomia linearis C. DC. (Pip AE 
751 Conomorpha caracasana Mez (Myrsinaceae) 1178 Peperomia diffusa C. DC (Piperaceae) 
754 Parathesis moritziana Mez (Myrsinaceae) 1183 Peperomia reflexa A. Dietr. var. coriacea C. DC. 
756 Conomorpha laurifolia Mez (Myrsinaceae) (Piperaceae) 
757 Cybianthus klotzschii Mez (Myrsinaceae) 1191 Euphorbia tovarensis Boiss. (Euphorbiaceae) 
794 NE leucantha Moritz ex Benth. (Gesneria- 1202 Pedilanthus fendleri Boiss. (Euphorbiaceae) 
ae) 1208 Tragia fendleri Muell. Arg. (Euphorbiaceae) 
830 miens magnificus Gilg (Gentianaceae) 1230 Excoecaria obtusiloba Muell. Arg. (Euphorbi- 
844 Aegiphila lewisiana Moldenke (Verbenaceae) aceae) 
845 Aegiphila floribunda Moritz & Moldenke (Ver- 1241 Pilea mollis Wedd. (Urticaceae) 
benaceae) 1246 Pilea dauciodora Wedd. (Urticaceae) 
876 Salvia tovariensis Briq. (Lamiaceae) 1259 Hemistylis macrostachya Wedd. (Urticaceae) 
879 Stachys fendleri Briq. (Lamiaceae) 1264 Pouzolzia acalyphioides Wedd. (Urticaceae) 
941 Ipomoea fruticosa Kuntze (Convolvulaceae) 1272 Alchornea grandiflora Muell. Arg. (Euphorbi- 
955 Cestrum laetum Francey (Solanaceae) ceae 
962 Cestrum tovarense Francey (Solanaceae) 1289 Podocarpus harmsianum Pilger (Podocarpaceae) 
980 Solanum tovarense Bitter (Solanaceae) 1298 Citharexylum venezuelense Moldenke (Verbena- 
1007 Saracha nitida Bitter (Solanaceae 
1030 Dipladenia fendleri Muell. Arg. ERE a 1327 Philodendron fraternum Schott (Araceae) 
1032  Amblyanthera fendleri Muell. Arg. (Apocyn 1329 Philodendron deviatum Schott (Araceae) 
ceae) 1336 Anthurium nymphaeifolium K. Koch € € 
1095 Gymnobalanus latifolia Meissn. (Lauraceae) é var. ovatum Engl. (Araceae) 
1098  Mespilodaphne vaginans Meissn. (Lauraceae) 1343 Anthurium fendleri Schott (Araceae) 
1101 uiri la eichleriana Urban (Lorantha- 1346 Anthurium bredemeyeri Schott var. elongata 
Engl. (Araceae) 
1102 horda fendlerianum Eichler (Lorantha- 1348 Asplundia fendleri Harling (Cyclanthaceae) 
1422 Liparis fendleri Schltr. (Orchidaceae) 
1104 Dendrophthora d Kuijt ashe mds 1460 Stelis muscifera Lindley (Orchidaceae) 
1105 Phoradendron ri m Urban (Loranthaceae) 1461 Stelis lutea Lindley (Orchidaceae 
1106 Phoradendron ae Urban (Lora aos ae) 1467 Stelis humulis Lindley (Orchidaceae) 
1108 Ph ovalifolium Urban (Loranthaceae) 1468 Stelis m dley (Orchidaceae) 
1115 oe cuneifolium Urban (Lorantha- 1469 Stelis tenuilabris Lindley (Orchidaceae) 
1470 Stelis le i Lindley ved oca 
1125 A fendleri van Tiegh. (Lorantha- 1471 Stelis tenuilabris Lindley (Orchidaceae) 
ceae) 1486 Pleurothallis viridula Lindley (Orchidaceae) 
1127 mum pseudoandromeda Solms-Laub. 1490 Pleurothallis exasperata Lindley (Orchidaceae) 
(Chloranthaceae) 1508 Tillandsia compacta Griseb. (Bromeliaceae 
1137 Piper glanduligerum C. DC. (Piperaceae) 1509 Tillandsia grisebachiana Baker (Bromeliaceae) 
1139 Piper victorianum C. DC. (Piperaceae) 1514 Tillandsia pleiosticha Griseb d ipee 
1140 Piper fendlerianum C. DC. (Piperaceae) 1515 Tillandsia fendleri Griseb. (Bromeliac 
1141 Piper tristemon C. DC. (Piperaceae) 1516 Tillandsia excelsa Griseb. var. latifolia Griseb. 
1143 Piper caracasanum Bredem. ex Link var. firmius Bromeliace 


C. DC. (Piperaceae) 


( ae) 
Tillandsia ventricosa Griseb. (Bromeliaceae) 


Volume 76, Number 1 
1989 


Todzia 


329 


Fendler's Venezuelan 
Plant Collections 


Nidularium albo-roseum Griseb. (Bromeliaceae) 
Tillandsia caribaea Lyman B. Smith (Bromeli- 
aceae 

Tillandsia incurva Griseb. (Bromeliaceae) 
Tillandsia robusta Griseb. (Bromeliaceae) 
Tillandsia myriantha Baker (Bromeliaceae) 
Smilax scabriuscula Humb. & Bonpl. ex Willd. 
var. fendleri A. DC. (Liliaceae) 

Pitcairnia fendleri Mez (Bromeliaceae) 
Tillandsia longifolia Baker (Bromeliaceae) 
Phaeosphaerion efoveolatum C. B. Clarke (Com- 
melinaceae 

Rhynchospora rudis C. B. Clarke (Cyperaceae) 
Chusquea fendleri Munro (Poaceae) 

Matayba longipes Radlkofer (Sapindaceae) 


Phoradendron longipetiolatum Urban (Loran- 
thaceae) 

Tillandsia acorifolia Griseb. (Bromeliaceae) 
d ndron venezuelense Trel. (Lorantha- 


pc bilineatum Urban (Loranthaceae) 
Peperomia pseudodependens C. DC. (Pipera- 
ceae 

Peperomia tenella A. Dietr. var. glabra C. DC. 

(Piperaceae 

Acalypha villosa Jacq. var. intermedia Muell. 
Arg. (Euphorbiaceae) 

Hemistylis odontophylla Wedd. (Urticaceae) 

Meriania subumbellata Cogn. (Melastomataceae) 

Arthrostemma alatum Triana (Melastomataceae 

Miconia rubens Naudin var. latifolia Cogn. (Me 

lastomataceae) 

Miconia fendleriana Cogn. (Melastomataceae) 

Miconia fendleriana Cogn. (Melastomataceae) 

Heterotrichum lucidum Triana (Melastomata- 
ceae 

Machaerium tovarense Pittier (Fabaceae d 

Machaerium mortizianum Benth. (Fabaceae) 

Eupatorium tovarense B. L. Robinson Jem 
ceae 

Piptocarpha sprucei Baker (Asteraceae) 
Clibadium parviceps S. F. Blake (Asteraceae) 
Cephaelis aneurophylla Standley ise ed 
Palicourea fendleri Standley (Rubiaceae) 


ro — 


Colum p 

Aegiphila fendleri Moldenke (Verbenaceae) 
Ipomoea fruticosa Kuntze (Convolvulaceae) 
Ipomoea fendleriana Kuntze (Convolvulaceae) 


Ipomoea alatipes Hook. (Convolvulaceae) 


Pleurothallis clandestina Lindley (Orchidaceae) 
Stelis alata Lindley (Orchidaceae) 
Pleurothallis apiculata Lindley diaries 


Griseb. (Bromeliaceae) 
Dioscorea fendleri R. Kunth (Dioscoreaceae) 
Cassia viciifolia Benth. (Fabaceae) 

Calliandra laxa Benth. var. parvifolia Benth. 
(Fabaceae) 

Inga fendleriana Benth. (Fabaceae) 


Strychnos fendleri Sprague & Sandw. (Logani- 
aceae 
Passiflora velata Mast. (Passifloraceae) 


artius var. a 
tenuata Meissn. (Lauraceae) 

Gynobalanus fendleri Meison. (Lauraceae) 
Piper ovale iperaceae 

Peperomia charaiana C. (Piperaceae) 
Acalypha villosa Jacq. var. intermedia Muell. 


Arg. 
i penninervia Muell. Arg. (Euphorbi- 


Pilea urticella Wedd. (Urticaceae) 


Stelis corüfolia Lindley (Orchidaceae) 
Tillandsia tovarensis D rt 
Tillandsia longibracteata B Ardens di 


Bilbergia filicaulis Griseb. ADA 


Aechmea cymoso-paniculata Baker ER 


aceae 
Aechmea fendleri André ex Mez (Bromeliaceae) 
Sloanea fendleriana Benth. (Elaeocarpaceae) 
Blakea grisebachii Cogn. ^H ee 


Malanea fendleri Standley (Rubiace 


Piper venezuelense E: 


Evodianthus funifer (Poit.) Ws es Bett fen- 
dlerianum Harling (Cyclanthace 
Peperomia venezueliana C. DC. (Pieria 


AUGUSTUS FENDLER'S 
VENEZUELAN COLLECTIONS 
OF FERNS AND FERN ALLIES! 


Alan R. Smith? and Carol A. Tod zia? 


ABSTRACT 


Augustus Fendler collected over 500 collection 2 of pteridophytes in Venezuela between 1854 and 


These collections are noteworthy because of the lar 
numbers thus far identified) among them. This Ios 


1858. 


types (44 collection 


collections and a historical account of the collections, including a portrayal of Fendler's fascination with fe 


Augustus Fendler collected in northern Vene- 
zuela between 1854 and 1858, where he amassed 
important collections of angiosperms, ferns, fern 
allies, lichens, mosses, liverworts, and fungi. Al- 
though a general account of his Venezuelan col- 
lecting is given in the previous paper (Todzia, 1989), 
this article deals specifically with his fern collec- 
tions. Fendler’s pteridophyte collections are note- 
worthy because they represent the first intensive 
gathering of ferns and fern allies from the Coastal 
Cordillera of Venezuela, and because they are widely 
distributed. Consequently, they have become type 
specimens of a great many fern species. The fol- 
lowing account has been assembled with the aid of 
Fendler’s correspondence with Daniel C. Eaton and 
Asa Gray, at the YU and GH archives, respec- 
tively. 

Fendler’s Venezuelan fern collections as well as 
his sets of flowering plants were distributed by Asa 
Gray. As with the angiosperms, Fendler initially 
retained the first set for himself. In 1859 he re- 
ceived a request from Thomas Moore of the Chel- 
sea Botanic Garden for a set, but since Moore had 
access to William Hooker’s herbarium containing 
set 2, Fendler decided to sell his first set to Daniel 
C. Eaton, who was then a student of Gray. He did 
this because Eaton agreed to publish a list of these 
collections, and because the collections would then 
remain in the United States, where Fendler would 


have easy access to them. In a letter to Eaton, 
Fendler stated, “The only set remaining with me 
is set No. 1 which I used for comparison and in 
which I felt a pride and a pleasure to lay up not 
only the most complete but also the finest speci- 
mens selected from the rest. There was a time 
when I thought I could never part with these plants. 
In bringing them up from their hidden places I 
have neither shun danger nor travelling expenses, 
and it was chiefly the search after Ferns, whic 

induced me to cross the principal chain of the 
cordillera of the coast upon four different tracks. 
I therefore need not say how dear they are to me, 
and I never would have sold them to anyone except 
to a Botanist, who is able to make the proper use 
of them" [Fendler to Eaton, 24 January 1859, 
St. Louis]. Fendler went on to say, *For my part 
I have neither the necessary books, time, nor her- 
barium to do it. Great resources undoubtedly are 
necessary to determine, which of the species are 
new and which are not; for I have reason to believe 
that in works on Ferns, with many species, a critical 
and patient overhauling is very desirable, in order 
to disperse the existing confusion. | remember to 
have seen in one herbarium the same species under 
four different names, one of these specimens was 
even disposed of under a new genus” [Fendler to 
Eaton, 24 January 1859, St. Louis]. In total, 


Fendler sent Eaton close to 500 collection numbers 


' We thank the archivists and librarians at the Missouri Botanical Garden, Gray Herbarium, and Yale University 


for aiding in finding some of the s 
M 


Steyermark, and H. van der 


ources used in this pape 
Werff concerning Fendler in Venezuela. R. C. Mor 
, G. K. Rogers, and J. Ewan provided helpful comments on various drafts of this manuscript. The 


with J. Ewan, R. C. 
oran, C. Blaney, N. 


r. We benefited from discussions 


junior author was supported by a National Museum Act Fellowship at the Missouri Botanical Garden during the 


course of the project. 


a of Botany, Herbarium, University of California, Berkeley, California 94720, U.S.A. 


3 Missouri Botanical Garden, P.O. Box 


Center, Dime of Texas, Austin, Texas 78713, 


299, St. Lo 


ANN. 


ouis, Missouri 63166, U. 
U.S.A. 


S.A. Current address: Plant Resources 


Missouni Bor. GARD. 76: 330-349. 1989. 


Volume 76, Number 1 
1989 


Smith & Todzia 
Fendler's Venezuelan 
Fern Collections 


with about 564 sheets. » ^ paid 10 cents a sheet 
for a total cost of $56.4 

In 1860, Eaton was Ea a Bachelor of 
Science degree from Harvard University for his 
dissertation enumerating Fendler's Venezuelan fern 
collections along with those collected by Wright in 
Cuba, and a few collections of Schott and Hayes 
from Panama and Colombia. A year later, Eaton's 
dissertation was published (Eaton, 1861). Fendler, 
however, was not pleased with this work. In the 
following long passage, we can see what a keen 
observer Fendler was. “In regard to Mr. Eaton’s 
Filices I am sorry to say that I was very much 
disappointed. Being in possession of the names of 
Dr. Engelmann's set of ferns as determined by 
Prof. Mettenius,* I was able to compare them with 
those given by Eaton and find that the names of 
146 sp. agree with those determined by Mettenius, 
while the names of 101 spec. disagree. I am not 
able to say who is in the right, but should be inclined 
to depend more upon the one that has the longest 
experience in the matter [Mettenius J. Such differ- 
ences of different authors are most discouraging 
to the student of botany and a severe drawback in 
the acquisition of knowledge. But what concerns 
me more immediately and of which I think I have 
reason to be solicitous, is the readiness with which 
Mr. Eaton throws different, quite distinct, species 
under one and the same name. I regret this, the 
more it may be inferred as if I purposely manu- 
factured species for the sake of selling one and the 
same species over again under different numbers. 
Having pointed out in Mr. Eaton's set some of the 
more obscure characters of many species very 
much resembling each other, I thought I had done 
something to prevent mistakes. It always seemed 
to me, that authors who only see 1 or 2 dried 
specimens of each species ought to pay at least 
some slight regard to the opinion of a collector who 
had the best opportunities, and for 4 years availed 
himself of those opportunities to study the habits 
of the different ferns, and to observe them in their 
growing state in the different phases of their growth, 
not on single specimens or in single localities only, 
but in hundreds of different places and individuals." 
Fendler continued: “There may be a dozen species 
about which I was rather doubtful, but all the rest 
I consider genuine species; and as soon as I shall 
have time to examine Dr. Engelmann's set, I shall 
endeavor to point out the differences in species 
considered the same by Mr. Eaton. That even the 


g H. Mettenius (1823-1866), Ne pteridol- 
t B. 


* Geo 
ogist, ie fern herbarium is now mostly 


most experient [sic] authors on ferns may some- 
times take 2 different species to be one and the 
same is proved by Prof. Mettenius calling No. 161 
'a young specimen of 162.' Now the former (No. 
161) is not only different in form but has also 2 
to 3 oval tubers, all the fronds die at a certain 
season of the year, and new fronds make their 
appearance rather suddenly but not before the 
commencement of the rainy season; the latter (No. 
162) is a perennial plant to be seen at all times of 
the year and never possessing tubers” [Fendler to 
Gray, 24 March 1868, St. Louis]. In fact, these 
two numbers do represent two species (Appendix 
I), and Fendler was correct. 

Nevertheless, Fendler's friendship with Eaton 
grew, as evidenced by Fendler's 15 letters to Eaton 
written from Trinidad between 1878 and Fendler's 
death in 1883. In Trinidad, as in Venezuela, he 
seems to have been preoccupied with collecting 
ferns and seems to have particularly relished dis- 
cussing his Trinidad fern collections with Eaton. 
Fendler, however, much preferred Venezuela over 
Trinidad and in his letters to Eaton, Engelmann, 
and Gray from Trinidad, he often mentioned how 
much he would like to return to Colonia Tovar. In 
his last letter to Eaton before his death, Fendler 
stated, “I long for the cool alpine air (damp though 
it be) for the deep shades and the eternal silence 
of those primeval forests, where a quarter of cen- 
tury ago my footsteps were wandering in search 
of the treasures of botany—especially those of 
ferns—again to go and see them, and to draw them 
forth from out their hiding places in the deep dells, 
or to bring them down from the mountain's high 
summit” [Fendler to Eaton, 15 June 1882, Port- 
of-Spain, Trinidad]. At 69, forever the fervent 
collector, Fendler asked Eaton if the professor could 
find subscribers to about 40 sets of Venezuelan 
ferns [Fendler to Eaton, 15 June 1882, Port-of- 
Spain, Trinidad]. Fendler often wrote to his cor- 
respondents that he longed to return to the “land 
of perpetual peace ... the happy region of the 
ferns." 


FENDLER'S FERN NOTEBOOK AND COLLECTIONS 


Fendler's field notebook in the GH archives is 
not chronological and consecutive, but rather a 
catalog of species in taxonomic order. Thus, the 
number sequence begins with 1 (Marattiaceae), 
proceeds through the more primitive fern families 
(Osmundaceae, Schizaeaceae, Hymenophyllaceae, 
Gleicheniaceae, Cyatheaceae), and then continues 
with the more advanced ferns, genus by genus, up 
to number 324. Numbers after 324 were collected 


Annals of the 
Missouri Botanical Garden 


after a trip to the United States and after he had 
distributed some of his material. It is obvious that 
the notebook was compiled after many of the col- 
lections had already been made and put into se- 
quence. Fendler combined collections made on dif- 
ferent dates and at different places under a single 
number. For example, Fendler 6, Anemia villosa, 
was collected on nine different dates (according to 
data on the sheet at YU) and most likely at several 
different localities during the years 1854, 1855, 
and 1856. 

Fendler probably revisited many collection sites 
in an attempt to amass additional material of a 
given number, because he intended to collect in 
large sets for sale to as many individuals or herbaria 
as possible. It appears that he tried to collect in 
sets of 20, although he gathered 40 sheets of 
Fendler 23, Trichomanes reptans. 'The number 
of collections appears in the upper right-hand cor- 
ner of the labels on sheets at YU but does not 
appear on the labels of Fendler collections in other 
herbaria. In some cases, Fendler kept an account 
of material of varying stages, e.g., he collected 26 
young and 12 old specimens of Marattia laevis 
(Fendler 3). In other instances, he recorded the 
number of specimens in different categories, e.g., 
he collected 20 specimens of no. 94, Paesia glan- 
dulosa, and 5 more with the rhizome. For many 
numbers through about 324, specimens were col- 
lected in rather large sets. Thereafter, the sets were 
generally much smaller, often only a few sheets. 
Most of the numbers after 324 were gathered in 
the years 1855 to 1858, whereas most collections 
of the numbers before 325 were made in 1854, 
with additions to the set made in 1855 and 1856. 

Fendler's keen eye is evident by the fact that, 
even though he often collected the same species 
(or number) at different times and/or localities, 
there are very few mixed gatherings. He must have 
had a very discriminating eye for differences. When 
in doubt, he often gave aberrant collections Greek 
letter designations a and 8 under the same number, 
or labeled the additions “ad 56”; sometimes these 
represent noteworthy taxonomic variations (occa- 
sionally different species), sometimes only forms of 
little taxonomic significance. The following are tax- 
onomically mixed collections: 16, 31, 64, 72, 73, 
83, 98, 117, 143, 179, 181, 185, 187, 221, 
224, 232, 2587, 259, 293, 294, 322, 324, 364, 
378, 380, and 473. Some of these probably rep- 
resent inadvertent mixing or misnumbering, where- 
as others are in large genera of great taxonomic 
difficulty, e.g., Elaphoglossum, Thelypteris, and 
Selaginella. 

Fendler's collection numbers range from 1 to 


499. Of these, 44 are thus far identified as types. 
We have seen specimens or notebook information 
for all numbers except 2 and 339. In addition, 
neither we nor Eaton (1861) saw specimens num- 
bered 337, 345, 346, 384, 392, 401, 427, 442, 
458, 462, 478, 484, or 490. The notebook at 
Harvard contains entries for numbers 58 to 499; 
inexplicably, the early numbers are missing. 

Because Fendler gave the same collection num- 
ber to plants collected at different times and lo- 
calities, it often is impossible to determine the lo- 
cality and date of a particular specimen. In a few 
cases there is only a single date on the specimen 
and in the notebook. In the list that follows (Ap- 
pendix I) dates are given only where there is 
reasonable certainty that the entire gathering was 
made on a single date at a single place. For those 
numbers that were collected on more than one 
date, we give the total number of dates (as indicated 
from the notebook and on the specimens at YU). 
For all numbers, the size of the set, when known, 
is given. 

Fendler's Venezuelan pteridophytes were widely 
distributed, and specimens have been located in 
the following 16 herbaria: B, BM, BR, CU, F, G, 
GH, GOET, K, MO, NY, P, PH, S, US, and YU. 


LITERATURE CITED 


ALSTON, A. H. G., A. C. JerMY & J. M. RANKIN. 1981. 
The genus Selaginella in tropical South America. 
Bull. Brit. Mus. (Nat. Hist.), Bot. 9: 233-330. 

BARRINGTON, D. S. 1978. A revision of the genus Tri- 
chipteris. Contr. Gray Herb. 208: 1-93. 

CHRISTENSEN, C. 1920. A monograph of the genus 

ryopteris, Part I. The tropical American pinnatifid- 
gcn species. Kongl. Danske Vidensk. Selsk. 
Naturvidensk. Afd., Ser. 8. 6: 1-132. 


CoPE Du. E.B. 19 Ctenopteris in America. Philipp. 
J. Sci. 84: 381-475. 
Eaton, D. C. 1861. Filices Wrightianae et Fendler- 


ianae, — nempe in Insula Cuba a Carolo Wright et 
n DER ab Aug. Pode ann. 1854-60, (non- 
lis ib inter rjec tis), E t 


Men . Amer. Acad. Art 

Evans, A. M. 1969. loierporibo relationships in the 
Polypodium pectinatum-plumula complex. Ann. 
Missouri Bot. Gard. 55: 193-293. 

HENNIPMAN, E. 1977. A monograph of the fern = 
Bolbitis i O Leiden Bot. Serie 
xu + 3 

"a K. iT 1957. A revision of the genus Lindsaea 

n the New World with notes on allied genera. Acta 
Bot. Neerl. 6: 97-290. 

LELLINGER, D. B. 1984. e Aor sce 
Pp. 9-46 in B. Maguire et al., The of the 
Guayana Highland— Part XII. Mem. New. Yor k Bot. 
Gard. 38: 1-84. 

Maxon, W. R. & C. V. Morton. 1938. The American 
species of Dryopteris, subgenus Meniscium. Bull. 
Torrey Bot. Club 65: 347-376. 


Volume 76, Number 1 
1989 


Smith & Todzia 333 
Fendler's Venezuelan 


Fern Collections 


MoRTON, C. V. 1947. The American species of Hy- 
menophyllum section Sphaerocionium. Contr. U.S. 
Natl. Herb. 29: 139-201. 

. B. LELLINGER. 1966. The Polypodiaceae 
subfamily ei in Venezuela. Mem. Ne 
ork Bot. Gar -49. 

SMITH, A. R. 1980. "Taxonomy of Thelypteris subgenus 

Me cis including vane (Pteridophy- 

ta). POS . Publ. Bot. -38. 

Pteridophytes ee E an anno- 
ta ted E Published by the author 

STOLZE, R. G. A taxonomic revision of the genus 
Cnemidaria (Cyatheaceae). Fieldiana, Bot. 37: 


ToDzIA, C. A. 1989. Augustus Fendler's Venezuelan 
plant collections. Ann. Missouri Bot. Gard. 76: 310- 
329 


TRYON, A. F. A monograph of ES fern genus 
cie Contr. Gray Herb. 200: 54-174. 

TRYON, R. 76. A revision of the E. Cyathea. 
Contr. Ti Herb. 206: 19-98. 

WrssELs Borer, J. G. 1962. The New World es 
of Trichomanes sect. EU Rias aia and Micr 
gonium. Acta Bot. Neerl. 11: 277-330. 


APPENDIX I 


Determinations of Fendler's pteridophyte collections 


M. Tryon; AT = Alice F. Tryon; ME = A 
DL = David B. mais) ts RM = Robbin 


= Benjamin Vligaard. (s) = data from specimen a at YU; 
(n) = data from Fendler notebook at GH. 
1. ea moritziana C. Presl, vel aff.; MO, YU. 
"M fs 000-7,000 ft. (s). 
2. Not loca 
3. Maratia laevis E. Smith; MO, YU— 2. [3/38]; 
6,500 
4. UT m sp.; i cited by Eaton (1861) as B. 
decompositum Mart. & Gal. 
5. Osmunda regalis de MO, YU. [2/24]; 5,900 ft. 


Dm 


(s). 

. Anemia villosa Humb. € Bonpl. ex Willd.; MO, 
YU; det. JM. EE pegar dé 000 ft. (s). 

. Anemia ferrugine EAS: MO, 
YU; det. JM. (2/20); 6.300 fe. ( 

. Anemia hirsuta (L.) Sw.; MO, Tu det. JM. [6/ 
17]; 5,500 ft. (s). 

. Anemia oblongifolia = dade MO, YU; det. JM. 
[4/20]; 5,500-6,000 

p Anemia oblongifolia (Cav. ) Sw.; YU. [?/1?]. 

. Anemia oblongifolia v ) Sw.; MO— 2, YU; det. 
JM. [6/20]; 6,000 ft. 


-J 


co 


Nel 


o 


11. Anemia pastinacaria Mo ritz ex Prantl; MO— 2, 
YU; det. JM. [7/20]; 6,000 ft. (s). 
12. Anemia sp., undescribed?; YU. 11/9/54 [1/2]; 


3,500 ft. (s). 
128. Anemia sp., undescribed?; YU. 


13. Anemia phyllitidis (L.) Sw.; MO, YU; det. JM. 
UR $ 000-4,000 ft. (s). 

14. Lygo 

15. pénis hirsuta 
9]; 5,200 ft. (s). 

16. Trichomanes (Pachychaetum) rigidum Sw.; MO— 

2, YU— 2. [13/30]; 6,000-7,000 ft. (s). 

16. Tricheinalis (Lacosteopsis) diaphanum H.B.K.; 

MO 


MO, YU. [6/19] (s). 
; MO, YU; det. JM. [3/ 


m venustum Sw.; 


(L.) Sw. 


17. Trichomanes (Lacosteopsis) diaphanum H.B.K.; 
MO— 2, YU. [7/50]; 6,000-7,000 ft. (s). 

18. Trichomanes s peris dd diaphanum H.B.K.; 
MO— 2, US— 2, YU. [2/21] (s). 

19. dedans acoscopsis diaphanum H.B.K.; 
US— not se 

20. Tc (Lácosteopsis) radicans Sw.; MO— 

S, YU—3. [10/30]; 6,000-6,500 ft. (s). 


MO— 2, US, YU— 2. [6/20]; 5,000-6,000 ft. (s). 

22. dec (Lacosteopsis) capillaceum L.; MO— 
2, US, YU. ES 6,000- 7,000 ft. (s). 
)reptans Sw.; B, G 

H, MO 4, US, YU; cited by Wessels Boer (1962). 

naval 6 000- 7,000 ft. (s). 
24. deer ae) pusillum Sw.; B, 
G, GH, : p by Wessels 


Boer "es py 6, 500 ft. 
4. Trichomanes (Di ymaglosun) pusillum Sw.; 
YU. 18/3/56 [1/7] (s 
25. Trichomanes DE MEN e (Fourn.) W. 
idymoglossum ovale Four ziv P Bot. 
France 19: 240. 18 LOTYPE, p PES, B, 
— Cited by Wessels ho (1962) 
24/7/56 [1/2); 6,000 ft. (s). 
ad 25. Trichomanes pao ovale (Fourn.) 
W. Boer; YU. [2/2 
26. duas p. e bus robustum Fourn.; MO, 
US, YU. [4/21]; 5,500 ft. (s). 
2T Hymenophyllum (Ptychophyllum) fucoides (Sw.) 
— 2, US, YU. [14/20]; 7,000-8,000 ft. 
(s). 


278. J, 7] ( Di L L 1] 
Sw.; YU. [3/ 6] (s). 
28. Hymenophyllum Arana axillare Sw.; MO— 
U. [16/21]; 6,500 ft. 
29. Hymenophlum iode d M tunbrigense 
J. E. Smith; iu US, YU. 9/11/54 [1/5]; 
7, 000- 8, 000 ft. 


fucoides (Sw ) 


- MO—2, YU. id 6,000-7,000 ft. ja 
31. Hymenophyllum (Sphaerocionium) sp., unde- 
scribed? YU. [5/15] 6,500- 7,000 ft. (s). 
3l. 


in Martiu ; MO. 

31 . Hymenophyllum (Sphaerocionium) hirsutum (L.) 
Sw. ., ve 

29. L 2 í( M. y Met ex 
Kuhn, Linares 35: 391. 1868. ISOTYPES, MO, US. 
YU. 


ad 32. Hymenophyllum (Mecodium) i GN Mett. 
ex Kuhn; YU. 26/7/56 [1/?] (s 

33. Hymenophyllum PEN uda microcarpum 
Desv.; US, YU; cited by Morton (1947). [3/20]; 
6, 500- pe 200 ft. (s). 


x * ae > IQ 


MO, US, YU. 3a} 6,200 ft. 9. 


334 


Annals of the 
Missouri Botanical Garden 


in Martius, Fl. Bras. 1(2): 291. 1859. LECTOTYPE 
Wen by Lellinger, 1984), US; ISOLECTOTYPES, 


w 
an 


" Pod oU (Mecodium) asplenioides Sw.; 
MO, US. 


37. Hymenophyllum (Sphaerocionium) hirsutum (L.) 
Sw.; GH, MO, NY, YU; cited by Morton (1947). 
[4/20]; 6,000 ft. (s). 

378. Hymenophyilum lr rc hirsutum (L.) 
Sw.; YU. 29/1/57 [1/1 

38. Hymenophyllum rie ME fragile (Hed- 

wig) C. Morton; GH, MO— 2, NY, US, YU; cited 

b Morton (1947). 16/24 6,000- 7,000 ft. (s). 


Sturm, Bot. 
ISOSYNTYPES, GH, MO— 2, 
ton (1947). [4/21]; 6,200 ft. (s). 

40. Sticherus rubiginosus (Mett.) Nakai; MO, US, 
YU—2. [3/15]; 8,000 ft. (s). 

41. Sticherus nudus (Moritz ex Reichardt) Nakai; MO, 
YU. [4/17]; 6,000-7,000 ft. (s). 

42. Sticherus bifidus par ) Ching; MO, YU—2. [3/ 
11]; 6,000-7,000 ft. 

43. Sticherus penniger (Martius) Copel.; 
[5/19]; 6,000 ft. (s). 

44. Dicranopteris pne I Me Underw.; MO, 
YU. [4/20]; 6,200 

45. Diplopterygium inpr (Hook.) A. R. Smith; 
MO, US— 2, YU. [4/20]; 6,500 ft. E 

46. Cyat thea speciosa Willd.; ted 
by R. Tryon (1976). 17/9/55 10204 4 Pon Pn 


Zeitung (Berlin) 17: 298. 1859. 
2, US, YU. Cited by Mor- 


MO, YU. 


(s). 
468. Cyathea speciosa Willd.; YU. 12/12/56 [1/1] 
S}. 

47. Cyathea villosa Willd.; MO, YU; cited by B 
rington (1978) as Trichipteris villosa (Willd.) R. 
Tryon. 3/12/54 [1/21]; 6,500 

i ee. quadripinnata (J. Gmelin) C. Chr.; 
MO, YU—3. [2/21]; 7,800 ft. (s). 

49. "cwn vd panni m (Riba) Cir sa MO— 2, YU— 

4. [4/18]; 6,000-6,800 ft. 

496. Cyathea m (Riba) Tanase YU. 12/ 

ae [1/ ile 


IS 
co 


en , YU— 3; cited by 
R Tryon (910) e n o ft. (s). 
51. Cyathea fulva & Gal.) Fée; YU. [3/?]; 
6,500 ft. 

52. D fulva (Mart. & Gal.) Fée; F, GH, MO— 

, YU— 2; cited zx R. Tryon (1976). [3/21]; 
^ ,000- 7,000 ft. 

. R squamata ` (Klotzsch) Domin; F, GH, MO— 

se are ISOTYPES of Alsophila cara- 

casana Klotzsch var. fendleriana Domin, Pterid. 

Dominica 95. 1919. Cited by Ecran 1978) 

as Trichipteris eb CH (Klotzsch) R. Tryon. 
[2/20] 6,500-7,000 ft. 

. Cyathea gibbosa (Klotach) Domin; GH, MO, US, 
YU; cited by Barrington (1978) as Trichipteris 
gibbosa (Klotzsch) Barrington. 26/8/54 [1/14] 
6,000-6,500 ft. (s). 

. Cyathea pungens s (Willd.) Domin; GH, MO, NY, 
YU — 2; cited by Barrington (1978) as Mie IgE 
procera (Willd.) R. Tryon. [2/20]; 6,500 ft.; 
side (s) 

. Cyathea caracasana (Klotzsch) Domin var. cara 
casana; GH, MO, YU — 2; cited by R. Tryon (1976). 
[3/13]; 6,000-6,500 ft. (s). 


en 
w 


— 


on 
> 


en 
en 


on 
an 


ad 56. Cyathea caracasana (Klotzsch) Domin var. ca- 
racasana; YU. 28/6/56 [1 /?] (s). 

57. Dicksonia sellowiana Hook.; YU; 19/6/54 [1/ 
16]; 6,800-7,800 ft. (s). 

. Dennstaedtia cicutaria (Sw.) T. Moore; YU. 1/ 
6/54 [1/17]; 6,500 ft.; Colonia Tovar (n) 

. Dennstaedtia dissecta (Sw. Moore; MO. [3/ 
?]; 6,000 ft.; Colonia Tov 

, Denastaedtia arborescens (Willd, ) Ekman ex Max- 
on; MO. [4/?]; 6,500-7,500 ft.; Colonia Tovar 


on 
© 


[21] 
o 


[o 
© 


(n). 
. Saccoloma domingense (Sprengel) C. Chr.; MO— 
2, YU— 2. [6/20]; 6,000-7,000 ft.; Colonia Tovar 


a 
- 


(n). 

618. Saccoloma domingense (Sprengel) C. Chr.; YU. 
29/1/57 [1/1] (s). 

62. RIS, theciferum (H.B.K.) T. Moore; MO, 

U. [10/28]; 6,000 ft.; Colonia Tovar (n). 
63. Linda klotzschiana Moritz in Ettingshausen; 
U. [7/16]; 6,000-6,800 ft.; Colonia Tovar, also 

iun Maracai and hora (n). 

63a € B. Lindsaea klotzschiana Moritz in Ettingshau- 
sen; BR, G, GH, GOET, K, MO, PH, US; cited by 


638. Lindsaea klotzschiana Moritz in Ettingshausen; 
YU. 12/12/56 [1/8] (s). 
64. e viscosa Karsten; YU — 2. [4/20]; 6,500 
; Colonia Tovar (n). 

64. H polenit gne Karsten?; MO. 

s Ka iis a 17/6/56 
; highest mou ntis, AUN 

646. Hypolepis v iscosa Karsten; YU. 

65. Cheilanthes microphylla “(Sw .) Sw.; MO. [5/?]; 


.; La Victoria, also bebween Caracas 


e 
66. et ia parallelogramma (Kunze) C. Presl; MO, 
U—4. 20]; 6,500 ft.; Colonia Tovar (n). 

ad 66. Hypolepis paraltelogramma (Kunze) C. Presl; 
YU. 6/1/57 [1/? 

67. Adiantopsis se / ) dia YU. [2/?]; Maya a 

mi. S of Colonia Tova ] 

68. Adiantum braunii Mett. ex Kuhn; MO, YU. 25/ 
10/54 [1/20]; 3,500 ft.; a few mi. SW of Colonia 
[Tovar] (n). 

69. Adiantum tenerum Sw.; MO. [3/?]; 3,000-3,500 
ft.; lower valleys around Colonia [Tovar] (n). 

70. Adiantum tenerum Sw.; MO, YU. [2/7]; 3,200 


ft.; Biscaina, S of Colonia [Tovar], also near Caracas 


n). 

7l. Adiantum poiretii Wikstróm; MO, YU. [2/15]; 
3,000-4,000 ft.; a few mi. SW of Colonia Tovar, 
also near Caracas (n). 

72. Adiantum henslovianum Hook.; YU. ia 4,000 
t.; a few mi. SW of Colonia Torari 

72. Adiantum capillus. veneris L.; 

73. Adiantum concinnum Humb. & Bonpl. ex Willd.; 
YU. [3/14]; 3,000-4,000 ft.; Caracas, also a few 
mi. S of Colonia [Tovar ] (n). 

ad 73. Adiantum concinnum Humb. & Bonpl. ex Willd.; 
YU. 19/1/56 [1/?] (s). Same sheet as 73. 

73. Adiantum lunulatum Burm. f.; MO, YU 

73/76. Adiantum concinnum Humb. & Bonpl. ex 
Willd.; M 

74. Adiantum ES Re a Presl; YU. [4/4]; 3,000- 
3,500 ft.; 6-9 mi. S of Colonia [Tovar] (n). 
. Adiantum concinnum Humb. & Bonpl. ex Willd.; 


Volume 76, Number 1 
1989 


Smith & Todzia 335 
Fendler's Venezuelan 


Fern Collections 


MO—2, YU— 2. [5/20]; 3,000-4,000 ft.; 6-9 
] 


mi. S of Colonia [Tovar 


ad 75. Adiantum MES d EN & Bonpl. ex Willd.; 


YU. [4/7] (s 
. Adiantum concinnum Humb. & Bonpl. ex Willd.; 
YU. [3/17] (s). 

76. Adiantum concinnum Humb. & Bonpl. ex Willd.; 
YU. 28/6/55 pa 5,000-6,000 ft.; 9 mi. S of 
Colonia [Tovar] (n 

77. Adiantum concinnum Humb. & Bonpl. ex Willd.; 
YU. [3/2]; 2,000-3,000 ft.; La Victoria to Bis- 
caina (n). 

78. Adiantum henslovianum Hook.; MO. [2/?]; 3,000- 


4,000 ft.; around Caracas (n). 
. Adiantum patens Willd.; MO, YU. [4/16]; 3,000- 
4,000 ft.; a few mi. S and SW of Colonia [Tovar] 


~- 
kel 


(n). 
. Adiantum polyphyllum Willd.; MO, YU— 2. P 
20]; 3,000-4,000 ft; a few mi. S and SW of 


eo 
© 


œ 
han 


. Adiantum deflectens ih: LECTOTYPE, US (cho- 
sen by Lellinger, Proc. c. Washington 89: 
703. xdi of A. deco: var. ps dom eron., 

ot. . Syst. 34: 487 1904, as “tremula”; 
ISOLECTOTYPES, MO — 2, YU. [2/20]; 2,000-3,000 
ft.; between La Victoria and Biscaina, S of Colonia 
Tovar (n . 
J oe lunulatum Burm. f.; MO, YU. [2/12]; 
500 ft.; near La Victoria and south of Laguna 
2 Valencia 
. Adiantum pulverulentun L.; YU. [3/16]; 2,500- 
,000 ft.; a few mi SW of Colonia [Tovar], 
also Piedernales 

83. Adiantum odia sum Sprengel; MO. 

84. Adiantum serratodentatum Willd: MO, YU. [2/ 
18]; 2,500-3,500 ft.; Biscaina and Piedernales in 
savannahs at margin di woods (n). 

. Adiantum villosum L.; MO, YU — 2. [4/8]; 2,000- 
3,000 ft.; Men La Victoria, between Ca- 
racas and La Guayra (n). 


ad 85. Adiantum miens e Humb. & Bonpl. ex 
Willd.; YU. 


oo 
N 


eo 
w 


œ 
en 


86. Alanha pulverulentum L.; MO, YU. [8/20]; 
3,000-4,000 ft.; a few mi. S and SW of Colonia 
Tovar, also Piedennales (n). 

. Adiantum petiolatum Desv.; YU. 25/1/55 [1/ 
2]; 2,000 ft.; Piedernales, near Güigüe (n). 

. Adiantum macrophyllum Sw.; MO. [7/21]; 2,500- 
3,500 l valley of the Tuy River, Maya Pieder- 
nales (n). 

89. Pellaea ovata (Desv.) Weath.; MO— 2, YU; det 
AT. ecd 6,500 ft.; a few mi. S of Colonia j 
in a savann aid rocks (n). 

i Cheilanthes marginata .B.K.; 

000 ft.; a few mi. W = Colonia 

91. Haryopteris pedata (L. . 
Hicken; MO; YU— 2; He RT. [6/20]; 3,000- 
4,000 ft; lower valleys S and SW of Colonia Tovar 
(n). 

. Cheilanthes concolor (Langsd. € Fischer) R. & A. 
Tryon; MO, YU. [2 ore 2,000-3,000 ft.; npn 

i een Mens and Guay 
and between Maracai as Choroni (n). 

. Histiopteris incisa (Thunb.) J. Smith: MO, YU— 
3. [2/25]; 7,500 ft.; highest mountains east of 
Colonia Tov 


ar (n). 
. Paesia dnia (Sw.) Kuhn; MO, YU— 2. [2/ 


ec 
"d 


eo 
e 


O 
[e] 


O, YU. [3/20]; 


Tovar (n 


NO 
N 


o 
o 


o 
B 


25] 7,500 ft.; 
Tovar (n). 
95. Pteris gigantea Willd.; MO, US, YU— 2. [3/18]; 
3,500 ft.; lower valleys S of Colonia [Tovar] (n). 
. Pteris grandifolia L.; MO, YU — 2. [2/14]; 2,000- 
2,500 ft.; between Caracas and La Guayra, also 
above Hacienda Curisal (n). 
97. Pteris consanguinea Mett. ex Kuhn, Linnaea 36: 
89. 1869. ISOSYNTYPES, K — photo US, MO, YU— 
3. [2/16]; 6,700 ft.; mountains south of Colonia 
[Tovar] (n). 
98. Pteris podophylla Sw.; MO, US, YU— 2. [4/26]; 
5,500-7,000 ft.; valley of the Tuy and headwaters 
of San Carlos River (n). 
98. Pteris propinqua Agardh; YU. [?/3]; 7,000 ft. 
(s). 


highest mountains E of Colonia 


Ne) 
Dm 


99. Pteris propinqua Agardh; YU. 25/1/55 [1/3]; 


iigúe (n 

U. 5/6/55 Mk 4 

t.; small distance Bore Biscaina, 9 mi. 
of Colonia [Tovar] (n) 

101. Pteris deflexa Link; MO, YU. [5/10]; 6,000 ft.; 
Colonia Tovar (n). 

deflexa Link; MO, US, YU—3. [7/20]; 
7,000 ft.; a few mi. W of Colonia Tovar, high 
mountai ise 

103. Pteris quadriaurita Retz.; MO, YU. [3/1]; 4, T. 

t.; a few mi. S an of Colonia [Tovar] (n 


1038. Pteris quadriaurita Retz.; YU— 2. 26/1/57 d / 
1] (3). 


104. dei arachnoideum (Kaulf.) Maxon; MO, 
YU—3. [4/20]; 5,000-7,000 ft.; Colonia Tovar 


(n Er 
105. Pteris a E MO, US, YU. [7/20]; 6,000- 
6,5 ] 


. savannah a few mi. S of Colonia [Tovar 


(n). 
106. Blechnum iia RE L.; MO— 2. (3/?]; 4,000- 
it ns S of Colonia, also Caracas (n). 

107. Blechnige dam TED L.; YU; cited by Eaton 
(1861). [3/?]; 6,800 ft.; a few mi. W of Colonia 
Tovar (n). 

108. Blechnum occidentale L.; YU; 
(1861). 27/5/54 s 6,000 s 
of Colonia [Tovar 

109. Blechnum n ur B. occidentale vi x 

. polypodioides Raddi; MO— 2. [3/7]; 2,0 
3,500 ft.; near Caracas, also low valleys S of D 
(Tovar ], Piedernales (n). 
110. E polypodioides iur MO— 2. [4/?]; 


ited by Eaton 


a few mi. S 


000 ft.; near Caracas, La Victoria (n). 
1108. “Blechnum id e YU; cited un Tai (1861) as B. 
lypodioides 


111. piece sp.; YU; cited by Eaton (1861) a 
longifolium H.B.K. 25/1/55 [1/?); 2,300 4 
Piedernales (n). 

112. Blechnum x caudatum Cav. [B. fraxineum Willd. 
x B. occidentale L.]; MO. [3/?]; 3,000-4,000 
ft.; lower valleys a few mi. S and SW of Colonia 
Tovar (n). 

113. Blechnum x caudatum Cav. [B. fraxineum Willd. 

x B. occidentale L.]; MO. [3/?]; 5,500 ft.; Tuy 


t.; Macarao valley, 9 mi. above Las Aguntas 


n). 3 
115. Blechnum sp.; YU; cited by Eaton (1861) as B. 


336 


Annals of the 
Missouri Botanical Garden 


DNI H.B.K. [2/?]; 3,000 S Piedernales, 
also jd n Caracas ber La Gua 
. Blechn pr YU; cited by Eaton (1861) as B. 
fendleri ous 25/1/55 [1/?] 2,300 ft.; Pie- 
dernales (n). 
. Salpichlaena volubilis (Kaulf.) J. Smith; MO— 2. 
[3/?]; 6,000-7,00 
ch 


. Blechnum cordatum (Desv.) Horn: MO. [3/?]; 
ade 7,200 ft.; a few mi. W of Colonia [Tovar] 


y is cordatum (Desv.) Hieron.; MO. [5/?]; 
t.; a few mi. S and SW of Colonia Tovar 


. Blechnum sp.; YU; cited by Eaton (1861) as Lo- 
aria procera Sprengel [possibly B. cordatum 

(Desv.) Hieron.]. 16/8/55 ue 3,000 ft.; be- 
tween Caracas and La Guay 

121. Blechnum aff. ee ae ) Hie eron.; MO. 24/ 
6/55 [1/?]; 8,000 ft.; highest mountains 12 mi. 
E of Colonia Tovar (n). 

. Blechnum sp.; YU; cited by Eaton (1861) as Lo- 
maria procera pin [possibly B. cordatum 
(Desv.) Hieron.]. No data in notebook. 

. Asplenium rutaceum (Willd.) Mett.; MO, US, YU. 

[3/21]; 6,500 ft.; Colonia Tovar and seaside of 

mountains (n). 

Asplenium cristatum Lam.; n YU. [15/23]; 

6,000- 7,000 ft.; Colonia Tova 

ad 124. mM cristatum Lam.; YU. 25/1/55 [1/ 


124. 


1248. ER aff. cristatum Lam.; MO, US, YU. 
[5/11] rachis winged. 

Asplenium flabellulatum Kunze; MO, US, YU— 

3. [6/23]; 6,000-7,000 ft.; Colonia Tovar (n). 

Asplenium flabellulatum Kunze var. partitum 

Klotzsch; MO. 

Asplenium radicans L.; MO— 2, US, YU. [5/13]; 

6,000- 7,000 ft.; around Colonia Tovar (n). 

1268. Asplenium radicans L.; ?/4]. 

127. Asplenium radicans L.? ?; M0— 2, US, YU. [67 
21] 4,000- 2 ft; a few mi. S and SW of 
Colonia [Tovar] (n). 

1278. amago abeliul atum Kunze var. partitum 

tzsch; YU. 8/1/57 [1/1] (s). 

128. ers cristatum (Desr.) Alston; MO, US, YU. 
[6/18]; 6,500 ft.; Colonia Tovar (n). 

1288. Pe n cristatum (Desr.) Alston; YU. 9/2/ 

7 [1/1] (s 
129. Diplazium caracasanum (Willd.) Kunze 


O, US, YU. [1/165 6,500 ft.; 


125. 
125. 


126. 


a 


x T. 
Colonia 


Tova iy 
1298. Diplazium caracasanum (Willd.) Kunze ex T. 
e; MO, YU. 8/1/57 [1/9] (s). 
; Asplenium pumilum Sw.; MO, US, YU. [6/24]; 
2. ~- he 000 ft.; a few mi. SW of Colonia [Tovar], 
also r La Victor ia and between Caracas and La 


Gua yh ). 
131. pr ien obtusifolium L.; MO, US, YU. [3/22]; 
5,000-5,500 ft.; around Colonia Tovar, in wet 


. Asplenium sq um L.; , YU— 2; cited by 
Morton & Lellinger (1966) at GH, NY. 8/10/55 
[1/11]; 7,200 ft.; 5 mi. W of Colonia Tovar, upon 
rocks (n) 

. Asplenium formosum Willd. MO, US, YU. [5/ 
27); 4,000 ft.; a few mi. SW of Colonia Tovar, 
also several mi. above Choroni (n). 


. Asplenium monanthes L.; MO, US, YU. [2/23]; 
4,000 ft.; a few mi. SW of Colonia [Tovar] (n). 
One other sheet at MO is Asplenium cristatum 
Lam 


135. Asplenium harpeodes Kunze; MO— 3, YU. [13/ 
22); 6,500-7,500 ft.; around Colonia Tovar (n). 
. Asplenium laetum Sw., vel aff.; MO, US, YU. [5/ 
20]; 3,000-4,000 ft.; a few mi. S and SW of 
Colonia [Tovar] (n). 
137. Asplenium raddianum Gaud.; MO, US, YU—2. 
[14/20]; 6,000-7,000 ft.; Colonia Tovar (n). 


- 
w 
an 


ar (n). 
1388. ,Asplenium claussenii Hieron.; YU. 4/2/51 [17 


139. diplniun ruizianum Klotzsch; MO, YU. [3/20]; 
4,0 ; a few mi. S and SW of Colonia [Tovar] 


(n). 
1398. Asplenium ruizianum Klotzsch; US, YU. [2/18] 
S). 
139y. Asplenium claussenii Hieron.; YU. 25/1/55 [1/ 
1] (s). 
140. Asplenium cirrhatum Rich. ex Willd.; MO, YU. 
Mixed with 4. radicans L. at YU. [5/23]; 7,000 
ft.; seaside of mountains N of presas [Tovar] (n). 
141. Asp prs auritum Sw. var. usum Kunze ex 
Mart. O. [2/?]; 4,000 ft.; a yes mi. SW of 
Colonia d (n). 
Asplenium monodon Liebm.?; MO, YU. [6/20]; 
6,000 ft.; Colonia Tovar (n). 
ad 142. Asplenium monodon Liebm.?; YU. 4 (s). 
1428. Asplenium monodon Liebm.; YU. [2/1] (s). 
i ; MO, YU. [9/20]; 


142. 


143. Asplenium oligophyllum Kaulf.; MO 
nium abscissum Willd.; MO, YU. 4/10/ 


1438. Asplen 
54 [1/13] (s). 

143y. Asplenium melanopus Sodiro; YU. 19/12/54 
1/1] (s). 

144. Asplenium auriculatum Sw., vel aff.; MO— 2, YU. 


(1 1/22] 6,000-7,000 ft.; Colonia Tovar (n). 

. Asplenium alatum Humb. & Bonpl. ex Willd.; MO, 
US, YU. [6/20]; 6,500 ft.; Colonia Tovar fia. 

. Diplazium diplazioides (Klotzsch & Karsten ex 
Klotzsch) Alston; YU. 28/9/55 [1/2]; 6,500 ft.; 
seaside of mountains N of Colonia [Tovar] (n). 

147. Diplazium hians Kunze ex Klotzsch; US, YU— 
2. 28/6/54 [1/9]; 6,000 ft.; Colonia Tovar (n). 

. Diplazium hians Kunze ex Klotzsch ; MO, YU— 
2. [2/5]; 5,600 ft.; Tuy River valley below Colonia 
[Tovar], seaside of mountains (n). 

. Diplazium diplazioides (Klotzsch & Karsten ex 
Klotzsch) Alston; MO, US, YU. [5/9]; 7,000-8,000 
ft.; highest mountains around Colonia [Tovar] (n). 

. Diplazium aff. striatum (L.) C. Presl; MO, US, 
YU. [3/ 13]; 4,000 ft.; San Carlos River valley, a 
few mi. SW of Colonia [Tovar] (n). 

151. Diplazium mutilum Kunze; MO, US, YU. [3/11]; 

4,500 ft.; a few mi. S and SW of Colonia [Tovar] 


(n). 

. Diplazium centripetale (Baker) Maxon, vel aff.; 
MO, US, YU. 28/9/54 [1/10]; 6,000 ft.; seaside 
of mountains N of Colonia [Tovar 

. Diplazium celtidifolium Kunze; — 3, YU. 
aoe 6,000-7,000 ft.; around Colonia Tovar 


— 
en 
w 


Volume 76, Number 1 
1989 


Smith & Todzia 
Fendler's Venezuelan 
Fern Collections 


: m redi oral (L.) Urban; MO, US— 
, YU. 29/8/54 [1/26]; 5,000-6,000 ft.; Tuy 
em valley below Colonia [Tovar] (n) 

. Asplenium serra Langsd. & Fischer; MO, YU— 
3. [9/20]; 7,000 ft.; around Colonia Tovar (n). 

. Asplenium lividum Mett. ex Kuhn; IsoTYPES, MO, 
US. Cited by Morton & Lellinger (1966). 25/10/ 
54 [1/17]; 4,000 ft.; San Carlos River valley (n). 

. Asplenium praemorsum Sw.; MO, YU. [9/21]; 
E 000- 7,000 os around Colonia Tovar, also 15 

i. W of Caracas (n). 
1578. Asplenium | us Sw.; YU. 27/2/51 [17 
s). 


157?. 
158. 


pops praemorsum Sw.; YU. [2/?]. 
Thelypteris (Amauropelta) pilosohispida (Hook.) 
Alston; GH, MO, YU— 2. [9/20]; 6,000-7,000 
ft.; Colonia Toyar (n). 

159. Thelypteris( "P ) lip ] pope | (M M 
Mett.) Ching; B, CU, GH— , MO, US, U— 
[3/16]; 5,500-6,000 ft.; TR of mountains N 
of Colonia Tovar (n). 

. Didymochlaena truncatula (Sw.) J. Smith; MO, 
YU—3. [12/18]; 6,000- 7,000 ft.; Colonia Tovar, 
also seaside of mountains N of Colonia Tovar (n). 

. Nephrolepis occidentalis Kunze; MO— 2, YU. [2/ 

22]; 3,000-4,000 ft.; a few mi. S of Colonia Tovar, 


also La Victoria (n). “Plant annual from tubers' 


(s). 

d duris pendula (Raddi) J. Smith; MO— 2, 
YU— 2. [5/21]; 4,000-6,000 ft.; a few mi. S of 
Co bus [Tovar], also mountains N of Caracas (n). 


1628. Nephrolepis pendula (Raddi) J. Smith, YU. [?/ 


162. 


163. Athyrium skinneri (Baker) C. Chr.; YU. 25/1/55 
[1/1]; 3,000 ft.; ü 
164. Tectaria heracleifolia (Willd.) L. 
US, YU. ve au a 000 ft.; 

of Colonia 
1648. Tectaria DUE (Willd.) L. Underw.; YU. 


a few mi. S and 


165. Tectaria incisa Cav., p aff.; MO, YU—3. [6/ 

O ft.; a few mi. SW of Colonia Tovar (n). 

165 cional Tectaria notions (L.) Cav.; YU. 12/ 
2/56 [1/1] (s). 


1656. Tecra incisa Cav.; YU. 22/4/58 Le 4,000 
t.; seaside of mountains N of Petaqui 

166. Theoria incisa Cav.; YU. [3/1]; 2, 000- 4,000 ft.; 
a few mi. SW of Qulexia Tovar (n). 

167. Hemidictyum marginatum (L.) C. Presl; MO, YU— 
2. 7/3/57 [1/?]; 3,000 ft.; seaside of mountain 
range between Valencia and San Estevan (n). 

168. ARE Ed (Sw.) L. Underw.; MO— 2, US, 
Y / 34]; 7,000 ft.; a few mi. W of Colonia 


"i » 
169. [besos patula (Sw.) L. Underw.; YU. 21/10/ 
]; 3,200 ft.; Maya River valley, a few mi. 
S of Colonia Tovar fa ). 
170. C ate el denticulata (Sw.) Ching; MO. [4/?]; 
7,000-8,000 ft.; highest mountains around Colonia 


Tovar (n). 
171. Polystichum sp., aff. polyphyllum (C. Presl) C. 
esl; US; mixed with Polypodium sp. [3/?]; 8,200 
" highest mountains E of Colonia [Tovar], in an 
open glade (n). 
172. Pi ein muricatum (L.) Fée; US— 2. [2/?]; 
: ft.; Colonia igs 2 
173. stan sp-; d by Eaton (1861) as 


Aspidium PP Klotzsch f oeei P. muri- 


catum (L.) Fee]. [2/?]; 6,500 ft.; 


Colonia Tovar 
n). 
174. Poe muricatum (L.) Fée; US. [2/?]; 7,000 
; Colonia Tovar (n 
Dt platyphyllum (Willd.) C. Presl; YU; 
cited by Eaton (1861) as Phegopteris platyphylla 
(Willd.) Mett. [3/?]; 6,500 ft.; Colonia Tovar (n). 
. Thelypteris (Amauropelta) concinna (Willd.) Ching, 
vel aff.; CU, G, GH, MO, US. [2/?]; 2,500-3,000 
ft.; Biscaina and Hacienda Carisal S of Colonia 
Tovar (n). 
Thelypteris vat oboe opposita (Vahl) Ching; 
MO, US, YU. [2/7]; 3,200 ft.; Caracas (n 
Thelypteris Dh den pilosula (Met. ) R. 
Tryon; GH, MO, US, YU. [3/15]; 7,000 ft.; Co- 


lonia BOL (n). 
(A 


175. 


177. 


178. 


179. Thelypter (Will lld.) Ching; 
MO, XU! aoa rk 000 ft.; a few mi. W of 
Colonia Tova 


r (n 
; DE (Amauropeta) ees (Humb. & 
Bonpl. ex Willd.) Ching; 
E (Amauropelta) ‘pilosula (Mett.) R. 
Tryon; G , YU. [9/16]; 6,000-6,500 ft.; 


n: GH, MO 


n). 
ypteris (Amauropelta) oligocarpa (Humb. & 
Bonpl. ex Willd. ng; YU. 

‘ Fe pei má To Wem Navy COM 

vel aff.; , YU. [4/13]; 2 O ft.; 

zd E $ j^ E [Tera] pe puc e 
aracas and La Guayra (n) 

Thalyoter a Pu pilosula (Mett.) R. 

Tryon; YU. 24/6/55 [1/1]; 8,000 ft.; highest 

mountains E of Colonia Tovar (n). 

Thelypteris (Amauropelta) pilosula (Mett.) R. 

Tryon; YU. 13/10/54 VAN 6, T ft.; San Carlos 

River valley near Colonia [Tovar 

Thelypteris (Amauropelta) rudis Ko Proctor 

& T. concinna (Willd.) Ching; MO [T. rudis at US]. 

[4/13]; 5,800 ft.; a E mi. S of Colonia Tovar in 

wet places of savannah (n). 

ad 185. Thelypteris (Amaurapelta) rudis (Kunze) Proc- 

tor; YU. 1/6/54 [17 

Thelypteris Cnauropte proctorii A. R. Smith 

& Lellinger?; YU. 

: Thelypteris (Amauropelta) gs ig (C. Presl) R. 
Tryon; B, G, GH, MO—3, P, YU. [3/20]; 7,000 
[tia de mi. W of ro Tovar (n). 

y ON p ih e rudis (Kunze) Proctor; 
YU— 2. 27/8/55 [1/4]; 6,500 ft.; a few mi. 
of m Tova 


r (n). 
sp air (Amauropelta) cheilanthoides (Kunze) 


183. 


184. 


185. 


185. 


187. 
Proctor; G 

187. TM mop ide ipei (Kunze 
ex Mett.) C r. pachyrhac 

188. Thelypteris (Crain) grandis sed R. Smith var. 


A. R. Smith; MO, YU— 2. 
a few mi. S and SW of 
189. Thelypteris (CiBlosofus) patens (Sw.) Small var. 
patens; MO, YU. [3/10]; 3,000 ft.; river valleys 


near Caracas, also Biscaina S of Colon [Tovar] 


(n 
Thelypteris (Cyclosorus) patens (Sw.) Small E 
scabriuscula (C. Presl) A. R. Smith; MO, YU— 


190. 


Annals of the 
Missouri Botanical Garden 


o 


o 
d 


o 


a 


25/10/54 o 3,600 ft.; a few mi. SW of Co- 

lonia Tovar 

à o HN (Cyelosorus) hispidula (Decne.) Reed; 
U. [3/4]; 3 ,000 ft.; a few mi. W 

a Colonia enel 

. Thelypteris aa) patens (Sw.) Small var. 

patens; MO, YU. 20/2/54 [1/6]; 3,200 ne near 

Caracas 


(n 
s Thelypteris A ccn ios dg (Hieron.) 


ount 
193y. piero (Goniopteris) gemmulifera (Hieron.) 
A. R. Smith; YU. [3/6] (s). 

; sie nigrovenia (Christ) Copel.; MO, US, 
[3/16]; 4,000 ft.; a few mi. S and SW of ud 
Tovar (n). 

. Ctenitis nou (Langsd. & Fischer) Ching; 
3,200 ft.; Maya River, a few mi. 


(n). 
. Thelypteris (Goniopteris) tetragona (Sw.) Small; 
H, MO, YU. [3/9]; 3,000-4,000 ft.; a few mi. 
S and SW of Colonia Tovar (n). 
. Ctenitis oo enr sd. & Fischer) Ching; 
MO, US, YU— 2. [2/16]; 3, 200 i. valley of Maya, 
a few mi. E of Colonia [Tovar] (n 
Stigmatopteris nephrodioides lotus h) C. Chr.; 
MO, US, YU—2. 29/8/54 UA 5,600 ft.; 
River Tuy below Colonia. Tovar 


(R 


, or; 
GH, “MO, YU. [2/6]; 2: 000 ft.; 
between Caracas and La 
. Thelypteris (Goniopteris) uU (Poiret in Lam.) 
Morton; YU. 3,000-5,000 ft.; River Tuy below 
Colonia did also between Caracas and La Guayra 
(n); [2/2] (s 
; Tiehpiers Goniopers asterothrix (Fée) Proc- 
r; GH, M 2, YU. 19/12/54 [1/22]; 3,200 
íi. "Maya a a inr mi. S of Colonia Tovar (n). 
i Ctenitis subincisa (Willd.) Ching; MO— 3 = 
3. [8/20]; 6,000 ft.; Colonia Tovar (n). 
. Ctenitis sloanei (Sprengel) C. Morton; YU— 2. 5/ 
1/55 [1/1]; Piedernales near Güigüe (n). 
. Ctenitis ampla (Humb. & Bonpl. ex Willd.) Ching; 
, US— 2, YU. [4/6]; 3,000-4,000 ft.; a few 
mi. S and SW of Colonia Tovar (n). 
. Lastreopsis effusa (Sw.) Tindale subsp. divergens 
(Willd. ) Tindale; MO, YU— 2. [3/20]; 3,000-4,000 
ft.; a few mi. SW of Colonia Tovar (n 1). 
Thelypteris ( 4 ) 
Tryon; CU, GH, MO, Us YUS Ee. 7,500 
ft.; 9 mi. E of Colonia Tovar (n). 
. Grammitis } ith) P YU. [2/ 
4] 6/5/58; de 000- 8, 000 ft.; Colonia Tovar (n). 
. Grammitis Bari (J. D. Sm ith) C. Morton; MO, 
; these are ISOSYNTYPES of dar e oligo- 
um Mett. ex Kuhn, Linnaea 36: 132. 1869 (non 
Klotzsch. 1847). [4/21]; 7,000-8,000 i highest 


)Pr 
Piedernales, also 
n). 


LVR 


mountains E of Colonia Tovar, also between ara- 
cai and Choroni (n). Specimens at MO and YU 
mixed w enilis (Fée) C. Morto 


cited by Copeland (1955) as Ctenopteris wens 

ad Copel. [9/20]; 8,000 ft.; highest mountains 
ound Colonia Tov 

Grammitis cultrata (Willd. ) Proctor; US, YU. 

20]; 7,000-8,000 ft.; highest mountains ke di 


Colonia cds (n). 


21 Pues reper (Klotzsch) C. Morton; MO, 
US— 2, YU. [13/20] 7, 000- 8,000 ft.; riae 


mountains Me: Colonia Tovar and W to Valencia 


2118. ‘Canals sp.; YU; cited 2 Eaton (1861) as 
Polypodi ium truncicola Klotzsch. 
211, pt. Grammitis richomanoide: = ‘ AD US. 
212. ann concin .R.S ey 
de ISOTYPES of Po Was de. concinnum n Met 

nnaea 36: 132. 1869 (non Willd. 

1810). in [1/8]; 7,500 ft.; high mountains 

E of Colonia Tovar (n 

Grammitis chrysleri (Copel.) Mon As US, 

YU. [10/20]; 6,500 ft.; Colonia Tov 

2138. Grammitis asplenifolia ( (L.) ning YU. 6/5/ 

[1/8] (3). 


213. 


58 [1/8 
. Polypodium moritzianum Link; MO, US 
2. [9/30]; 6,000-7,000 ft.; Colonia Tovar (n). 


nue ‘Mee n; MO, 
US, YU. [3/20]; 6,000- 000 ft.; Colonia Tovar 
(n). 


. Grammitis firma (J. Smith) C. Morton; MO, YU. 
[6/20]; 6,000-7,000 ft.; Colonia Tovar (n). 

. Grammitis apiculata (Kunze ex Klotzsch) Sey- 
mour; MO, US, YU; cited by Copeland (1955) as 
Ctenopteris apiculata ose E eid [9/20]; 
6,000- 7,000 ft.; Colonia Tova 

. Grammitis xiphopteroides (Liebm A. R. Smith; 
MO, YU; cited by Copeland (1955) as Ctenopteris 
pilosissima (Mart. & Gal.) Copel. [14/20]; 6,000- 

8,000 ft.; Colonia Tovar (n). 

. Polypodium consimile Mett. var. consimile; 
GH, MO, US, YU; cited by Evans (1969). [6/20]; 
6,500 ft.; Colonia Tovar (n). 

2208. Polypodium consimile Mett. var. consimile; GH, 

S, YU; cited by Evans (1969). 

. Polypodium eurybasis C. Chr. var. eurybasis; MO, 
YU; det. ME at MO. At YU there are also spec- 
iq of P. plumula Humb. x Bonpl. ex Willd. 

P. dispersum A. M. Evans on the same sheet. 
[12/20]: 4 000. "8, 000 ft.; Colonia Tovar (n). 
i ; YU. [8/13] (s). 


ach- 
pm) (Hieron.) A. M. Evans; YU. 19/9/55 [1/ 
s). 


1] 

2216. Polypodium sp.; YU; cited by Eaton (1861) as 
. paradiseae Langsd. & Fischer 

222. Niphidium siet ss Cellier MO, YU; det. 


DL. [6/18]; 6,000-8,000 ft.; Colonia Tovar (n). 
. Niphidium crassifolium (L.) Lellinger; MO, YU; 
det. DL. [2/11]; 3,500-4,000 ft.; a few mi. SW 
of Colonia Tovar (n). 
mpyloneurum angustifolium (Sw.) Fée; YU; det. 
DL. 5/6/55 [1/5] 7,000-8,000 ft.; highest 
mountains E of Colonia Tovar, also between Mara- 
cal and Choroni (n). 
í ee chlorolepis Alston; YU. 29/1/ 
57 [1/1 
. Ca d um chlorolepis Alston; MO, YU. 3 
1/55 [1/6]; 5,000 ft.; a few mi. S of Colonia da 


(n). 
. Campyloneurum amphostenon (Kunze ex Klotzsch) 


Volume 76, Number 1 
1989 


Smith & Todzia 
Fendler's Venezuelan 
Fern Collections 


339 


ad 226. 


2288. Campyloneurum chrysop 
det 


e 


w 
on 


N 


ie MO, YU. ae 7,000-8,000 ft.; a few 
(n). 

oneurum E non (Kunze ex 

Klotzsch) Fée; YU; det. DL. 2/1] highest moun- 


tains (s). 


227. Campyloneurum phyllitidis (L.) C. Presl; MO, YU. 


[6/21]; 2,000-4,000 ft.; a few mi. SW of Colonia 
Tovar, also between Caracas and La Guayra, Pie- 
dernales (n). 


228. Campyloneurum chrysopodum (Klotzsch) Fée; det. 
M 


DL; — 2, YU; these are ISOTYPES of Campy- 
loneurum fendleri T. Moore, Ind. Fil. 224. 1861. 
5/10/54 [1/19]; 5,500 ft.; seaside of mountains 
north of Colonia Tovar (n). 

odum p Fee; 

L; MO, YU. 7/3/57 [1/6] (s 

. Ca amp S repens (Aubl.) C. m MO, YU; 
det. DL. [2/10]; 5,500 ft.; seaside of mountains 
N of Colonia Tovar (n). 
. Campyloneurum repens (Aubl.) C. Presl; MO, YU; 
det. DL. [5/20]; 6,000 ft.; seaside of mountains 
N of Colonia Tovar (n). 
. Campyloneurum decurrens (Raddi) C. Presl; YU; 
det. DL. [3/4]; 5,000 ft.; seaside of mountains N 


of Colonia Tovar (n ). 


1 C. Mor 
ton; CH, K, MO, ¥U— 2; cited de Maxon & Mor- 
ton (1938) as Dryopteris ensiformis C. Chr., in 


contains both species. 


2328. E eni p ardoren G ed & 
Wi 


onpl. ex 


U: 18/237 a 500 e 
Tuy B Rive. EE AR Tov 

. Polypodium pu Jacq.; YU. 18/10/54 
[1/1]; 7,000 ft.; a few mi. W of Colonia Tovar 


(n). 
. Polypodium fraxinifolium Jacq.; MO, YU. 24/6/ 
55 [1/20]; 7,000 ft.; high mountains around Co- 
lonia Tovar (n) 
. Polypodium fraxinifolium Jacq.; YU; cited by Ea- 
ton (1861). 28/9/55 (1/2) 7,000 ft.; seaside of 


n). 
. Polypodium aff. fraxinifolium Jacq.; YU. With 
eed spreading rhizome scales. 19/12/54 [1/ 
3]; 6,000 ft. a few mi. S of Colonia Tovar (n). 
d very doen from no. 236” (s). 
A Gina SORA b: YU. [2/3]; 4,000-5,000 
ft.; 


ange between Maracai and 


Choroni (n). 
2388. “Polypodium dissimile L.; YU. [?/1]. Mounted 


ame sheet as 238. 
. Pise um sp.; YU; cited by Eaton (1861) as P. 
WA Langsd. Fischer. Vy Ae 4 ‘000 


; a few mi. of Colonia Tova 
: Picadas sessilifolium Desv.; MO. "Yu. [6/10]; 
fi eee 000 ft.; high mountains around Colonia 
Tov 


à Polypodium tege pseudoaureum Cav.; 
MO, YU— 2. [10/18]; 5, aD 7,000 ft.; a few mi. 

S and SW of Colonia Tova 

. Polypodium posee ie oq R. Sm 
2, YU— 2. [2/23]; 3,000-4,000 ft.; 

and SW of Colonia Tovar (n). 


; MO— 


a p mi. S 


243. dar funckii Mett.; US— 2, YU. [2/14]; 
000 ft.; highest mountains E of Colonia Tovar 


D ). 

243?. dei sp.; YU; cited by Eaton (1861) as 

P. loric 

Polaca lasiopus Klotzsch; MO, US, YU. [10/ 

21]; 6,000-7,000 ft.; a few mi. S of Colonia Tovar 

(n). 

. Polypodium remotum Desv.; MO, US, YU. [6/ 
17]; 6,200 ft.; Colonia j| (n 

. Polypodium buchtienii Christ & Rosenstoc k; YU. 
[2/4]; 7, d 8,000 ft.; high mountains around 
Colonia Tovar (n). 

^ Pope steirolepis C. Chr.; MO, NY, US, YU. 
ISOTYPES of P. nigripes Hook., Sp. Fil. 5: 17. 1863 
(non Hassk., 1844). [5/17]; 3,500 ft.; a few mi. 
S and SW of Colonia Tovar (n). 

. Microgramma vacciniifolia (Langsd. & Fischer) 
Copel.; MO, YU. [2/6]; 1,000 ft.; seaside of moun- 
tains near San Estevan, also between Caracas and 
La Guayra (n). 

. Pleopeltis macrocarpa (Bory ex Willd.) Kaulf.; 
YU. 27/8/55 [1/4); 6,500 ft.; a few mi. of 
Colonia Tovar (n). 

. Pleopeltis macrocarpa (Bory ex Willd.) Kaulf.; 
MO, YU. [14/23]; 7,000 ft.; Colonia Tovar (n). 
251. xPleopodium leucosporum (Klotzsch) Mickel & 
Beitel pEr eltis macrocarpa USA i: bs 

Kaulf. x Polypodium thyssanolepis A 
Klotzsch]; MO, US, YU. 23/2/54 ee F| 000 
t.; mountains N of Caracas (n). 

. Polypodium thyssanolepis A. Braun ex Klotzsch; 
MO, YU. [4/20]; 3,000-4,000 ft.; valley of Ma- 
carao W of Caracas (n). 

. Polypodium polypodioides (L.) Watt var. bur- 
chellii (Baker) Weath.; MO, YU. [6/20]; 3,000- 

O ft.; a few mi. S and SW of Colonia Tovar 


244. 


(n). 

. Polypodium ursipes Moritz ex C. Chr.; MO, US; 
these are ISOSYNTYPES of P. ambiguum Mett. ex 
Kuhn, Linnaea 36: 134. 1869 (non Desv., 1827). 
[3/?]; 8,000 ft.; highest mountains E of Colonia 
Tovar (n). 

. Grammitis jamesonii (Hook.) C. Morton; US, YU 
7,000-8,000 ft.; Colonia Tovar (n); [5/20] (s). On 
same sheet as 35 YU. 

. Grammitis leptopoda 
US. [2/?]; 7,000 ft.; 


E Grammitis j 


(C. H. Wright) Copel.; MO, 
exe Tux r (n). 

h) Ching; YU; 
cited by Eaton | (1861) as ian. jungerm an- 
nioides Klotzsch. [2/?]; 6,800 ft.; Colonia Tovar 
(n). 

. Vittaria graminifolia Kaulf.; MO, YU. [9/20]; 
(n à 


6,700 ft.; Colonia Tovar 

: P pue pu Klotzsch; de oe by Al- 
ston et al. 81); erroneous citat 

. Vittaria moritzia a Kunze; 


na Mett. + V. stipita 
MO, YU. [6/19]; 6,000- 2.00 ft.; TUM Tovar 


n). 

B. Vittaria pom Fée; MO, YU. 6,000 ft. 
(n); [2/17] (s 

. Vittaria i Fée; MO, YU. [2/20]; 7,000 ft.; 
Colonia Tovar (n). 

. Polybotrya serratifolia (Fée) Klotzsch; MO— 2 
YU; det. RM. [5/17]; 6,000 ft.; Colonia Tovar TET 

. Polybotrya canaliculata Klotzsch; MO—2, 


340 


Annals of the 
Missouri Botanical Garden 


3; det. RM. [2/10 + 9] 5,000 ft.; seaside of 
mountain range o "Sus the € [Tovar] (n). 
m (Sw.) Urban forma fla- 
npl. ex Vid. ) Mickel; MO, 
River Tuy below Colony 


t2 
e 
m 


Bon 
U. [4/19]; 6,000 ft.; 


i Elaphoglossum erinaceum (Fée) T. Moore; MO, 
YU; det. JM. [6/5]; 5, tel 6,000 ft.; Tuy valley 
below Colonia [Tovar 

. Elaphoglossum eximium Mar ) Christ; GH, MO— 
2, PH, YU; these are ISOSYNTYPES of wis pia 
reichenbachii Moritz ex Kuhn, Linnaea 36: 
1869. [3/20]; 7,200 ft.; headwaters of San a 
River (n). 

. Elaphoglossum moritzianum (Klotzsch) T. Moore; 
GH, MO, NY, PH, US, YU; det. JM. [6/20]; 5,900 
ft.; valley of the Tuy River, below Colonia [Tovar] 
on rocks (n). 

i "e piloselloides (C. Presl) T. Moore; 

, YU; det. JM. 19/9/55 [1/17]; 
6, 600 ft., San Carlos valley in a cave in rocks in 
the seis (n). 

. Elaphoglossum hieracioides Mickel; HOLOTYPE, US; 
ISOTYPES, GH, MO, YU. [9/20]; 6,500 ft.; San 
Carlos valley (n). 

269. Elaphoglossum meridense (Klotzsch) T. Moo 
MO, NY, YU; det. JM. s 6,300 ft.; visiten 
range S of Colonia [Tovar] (n 

2698. Elaphoglossum deorsum T Vareschi; MO; 
det. JM. 


. Elaphoglossum bellermannianum (Klotzsch) T. 
Moore; MO, NY, US, YU; det. JM. [7/21]; 6,500- 
7,000 ft.; mountain range S of Colonia [Tovar]. 

: ies gets pe cuspidatum (Willd.) T. Moore; MO, 

NY, YU; det. JM. um 6,300 ft.; mountains S 
of Colonia [Tovar 

272. Elaphoglossum PAP (Fée) T. Moore; P; this 

is is HOLOTYPE of Acrostichum tenuiculum Fée, 
ém. Fam. Foug. 10: 6, t. 2, f. 2. 1865. ISOTYPE, 
YU. d 18}; 6,600 ft.; 7 mi. SW of Colonia [Tovar] 
w ocks in savannahs (n). 
273. pla tectum (Humb. & Bonpl. ex Willd.) 
oore; MO, YU; det. JM. [5/23]; 6,500 ft.; 
Colonia Tovar (n). 
ad 273. EE DIGNUS tectum (Humb. & Bonpl. ex 
illd.) T. Moore; YU. 3/7/55 [1/1] (s). 
274. TI tectum (Humb. & Bonpl. ex i ge ) 
T. Moore; YU. 27/8/55 (73h 6,600 ft.; 
W of Colonia [Tovar] (n 
275. Elaphoglossum succubus Mickel: HOLOTYPE, NY; 
ISOTYPES, MO, US. ivi 6,000 ft.; Maya valley 
near Colonia [Tovar] (n 
275a. Elaphoglossum affine es Gal.) T. Moore; 
not seen; cited by Mickel in Smith (1985). 
276. an huacssaro (Ruiz) Christ; MO, US; 
M. [3/?]; 6,000 ft.; mountain range S of 
Colonia [Tovar], in wet ground full of springs (n). 

277. Elaphoglossum sp.; YU; cited by Eaton (1861) as 

Acrostichum leptophyllum Fée. 23/8/55 [1/?]; 

7,500 ft.; Lagunassa, 12 mi. E of Colonia [Tovar] 


(n). 
278. A chrysopogon Mickel; MO, YU; det. 
JM. [2/14]; 6,900 ft.; 6 mi. SE of Colonia [Tovar ] 


(n). 
279. ae dd cg chrysopogon Mickel; HOLOTYPE, 
x TYPE, NY. [3/8]; 6,700 ft.; Lagunita, 6 


W of Colonia [Tovar], between een (n). 


. Elaphoglossum opens ed C. Chr.; MO, 
YU; det. JM. these are ISOSYNTYPES of en aphog- 
lossum pen anaes ‘Chr ist. [5/20] 6,300 ft.; 


. Elaphoglossum eximium (Mett. 
det. JM. irn > 800 ft.; aig sla of San Car- 
los River, 3 m olonia [Tov 

] Elaphoglossum irachoncuron (Fée) J. Smith; GH, 

, NY, US, YU JM. [2/20]; 6,500 ft.; 
ndi range S of Colonia [Tovar] (n). 

. Elaphoglossum ambiguum (Mett. ex Christ) Al- 
ston; MO, US, YU — 2; it are ISOTYPES of Acros- 
tichum ambiguum Pu ex sig Neue Denkschr. 
Allg. Schweiz. Ges. Ges n Naturwiss. 36: 60. 
1899. [5/20]; 6,400 n: Colonia [Tovar] (n). 

. Elaphoglossum UR (Mett. ex Christ) Al- 
ston; MO, det. JM. 28/9/54 [1/11]; 
6,000-7,000 ft.; cui of mountain range near 
Colonia [Tovar] (n 

. Elaphoglossum T (C. Presl) Brackenr.; MO, 
US, YU; det. JM. [2/6]; 6,300 ft.; mountain Tange 
S of Colonia [Tovar] (n). 

: igi pur aic (Hook.) T. Moore; MO, 

U; det. JM. [7/20]; Y 300 ft.; mountain 
aln S of eh Ton (à 

. Elaphoglossum macrophyllum (Klotzsch) Christ; 
MO, NY, YU; det. JM. [3/12], 6,300 ft.; mountain 
range S of Colonia [Tovar]. 

. Elaphoglossum sellowianum (Klotzsch ex Kuhn) 
C. Chr.; YU. [3/1]; 6,300 ft.; mountain range S 
of Colonia [Tovar] (n). 

. Not located, most likely an Elaphoglossum. 27/ 
9/55 [1/?]; 6,800 ft.; Colonia Tovar, plant easily 
drying, somewhat viscose, sticking to the paper, 
sterile 

, Elaphoglossum eie cds EHE ex Kuhn) Christ; 
BM, MO, NY, US, YU; these are ISOSYNTYPES of 
amas Lu attenuatum Kunze « ex Kuhn, Linnaea 
36: 56. 1869 (non Fée, 1853). [4/20]; 6,300 ft.; 
mountains S of Colonia [Tovar], on trunks of trees 
and upon the ground Va 

. Elaphoglossum sporadolepis (Kunze ex Kuhn) C 
Chr.; YU. 2/10/54 ins 6,300 ft.; mountains 


[ov] 
o 
— 


N 
No) 
N 


ISOSYNTYPES of Acrost ichum tova ritz ex 

Kuhn, Linnaea 36: 60. 1869. [4/2 0]; 6,300 ft.; 

mountain range S of Colonia [Tovar], upon the 

ground (n 

. Elaphoglossum sporadolepis (Kunze ex Kuhn) C. 

Chr.; MO— 2, YU; th ; 

rostichum corpulen e ex , Linnaea 

36: 61. 1869. n 6,300 ft.; mountain range 

S of Colonia [Tovar] (n). 

Elaphoglossum andicola (Fée) T. Moore; NY; cit- 

ed by Mickel in Smith (1985 

. Elaphoglossum sporadolepis (Kunze ex Kuhn) C. 

Chr.; MO, NY, US, YU; these are ISOLECTOTYPES 
of Acrostichum sporadolepis Kunze ex Kuhn, Li 
naea 36: 59. 1869. [5/21]; 6,300 ft.; mountain 
range S of Colonia [Tovar] (n) 

294, pt. Elaphoglossum sporadolepis ni ex Kuhn) 
C. ; B, YU; the former is the HoLoTYPE of 
Acrostichum compactum Mett. e aa Linnaea 

1869; cited by Mickel in Smith (1985). 

295. Elaphoglossum sellowianum (Klotzsch ex Kuhn) 


293. 


N 
No) 
E 


Volume 76, Number 1 
1989 


Smith & Todzia 341 
Fendler's Venezuelan 


Fern Collections 


C. Chr.; US, YU; these are ISOSYNTYPES of Acros- 
tichum sellowianum Klotzsch ex Kuhn, Linnaea 


36: 52. 1869. [10/22]; 6,400 ft.; Coloni Tovar 


(n). 
295a, pt. Elaphoglossum tenuiculum (Fée) T. Moore; 
MO, not seen; cited by Mickel in Smith (1985). 
Elaphoglossum iis cdm (Klotzsch ex Kuhn) 
hrist; YU; this is an ISOSYNTYPE of Acrostichum 
sellowianum Klotzsch ex Kuhn, Linnaea 36: 52 
1869. [4/16] (s 
Elaphoglossum i epis dn d ex Kuhn) C. 
Chr.; MO, YU; these are ISOSY pd Ep 
chum sporadolepis Kunze ex d “Lin 36: 
59. 1869; det. JM. [3/8]; 6,300 ft.; ba a S 
of Colonia [Tovar]. 
Pityrogramma trifoliata (L.) R. Tryon; MO, YU. 
[2/19]; 3,000-4,000 ft.; warmer parts of river 


2958. 


296. 


297. 


-J 


298. 


œ 
Y 
~ 
E 
o 
oR 
3 
3 
a 
O 
8 
o 
3 
E 
3 
BÉ. 
E 
E 
je] 
zi 
z 
o 
E 
gi 


zuelae 


AR A tartarea (Cav.) Maxon; MO, YU — 

3. [10/20]; 6,300 ft., Colonia Tovar (n). 

, Eriosorus flexuosus H. B. opel. var. — 
G, GH, K, MO, P, US, YU; cited by A. Tryon 
(1970). ee 7,400 ft.; 12 mi. E of Colonia 
[Tovar 

301. ieri hirtus (H. ue Copel. var. hirtus; B. 

: MO— 2, NY, YU— 2; cited by 
A. Tryon i. ao 3, 000 ft.; a few mi. W 
of Colon 

302. Hemionitis sni (L Yeh: YU; cited by Reim (1861) 

; Pieder- 


a (n). 

is palmata L.; YU. [2/18]; 2,500-3,000 
penes c a little S of the lake do Valencia, 
[also] between La Guayra and Caraca 


20); 6,000-7,000 ft.; 

. Antrophyum cajenense (Desv. ) Sprengel; MO, YU. 
Mixed on same sheet with Antrophyum brasilia- 
num (Desv.) C. Chr. at YU. [3/23]; 6,000- 7,000 
ft.; Colonia Tovar, near Tuy River (n). 

. Thelypteris (Amauropelta) aspidioides (Willd.) R. 
Tryon; G, MO, US, YU. [4/13]; 5,600 ft.; Tuy 
River below Colonia [Tovar]; San Carlos River (n). 

307. Thelypteris (Amauropelta) rupestris (Klotzsch) 

Reed; GH, MO, US. [2/7]; Colonia Tovar (n). 

. Blechnum fragile (Liebm.) C. Morton & Lellinger, 
vel aff.; MO. [3/?]; 7,800 ft.; 12 mi. E of Colonia 
[Tovar] (n). 

. Blechnum divergens (Kunze) Mett.; 
6,000- 7,000 ft.; Colonia Tovar (n 

. Blechnum lehmannii Hieron.; MO. [3/?E 6,000- 
7,000 ft.; Colonia Tovar (n). 

311. Bolbitis serratifolia (Mertens ex Peur. Schott; 

ted by Hennip- 
man (1977). [2/20]; 4, 000 ft; uibs of San Carlos, 
W of Colonia [Tovar] (n). 

: ds M a d reticulatum L.; MO. [3/?]; 6,500 
ft.; several m of Colonia [Tovar ]. 

: d rig andas Humb. & Bonpl. ex Willd.; 
MO, YU— 2. [4/22]; 6,500-7,500 ft.; a few mi. 
S of Colonia Tovar n). 


MO. [3/?]; 


w 
ES 
N 


w 
pea 
wo 


314. Lycopodium ee ran rs MO, YU. ees 7,000 
t.; a few mi olonia Tovar 

3148. Lycopodium dos Es YU: 19/1/56 [1725 
7,000 ft. (s). 

315. ap ee alopecuroides L.; YU. [2/17]; 6,000 
ft.; a few mi. S of Colonia Tovar, wet places in the 
wenn 

i Lycopodium reflexum Lam.; MO, YU. [7/20]; 

7,000 ft.; a few mi. W of Colonia Ed (n). 

2/7); 3,500 ft.; 


318. Lyco m cernuum L.; AA 500 ft.; 
highest mountains E of ¿ib Tovar (n 
319. Lycopodium taxifolium Sw.; 


U; dd . BO. 
[6/17] 6,500-8,000 ft.; und Colonia Fitar 


ycopodium myrsinites Lam.; MO, YU; det. BO. 

i 20]; 8,000 ft.; highest E, E of libi 
ar (n). 

821. Selana Ww yr (C. Presl) Spring in Martius; 
MO, NY, 22] 2,000- on ft.; valleys 
SW of Colonia da also La Victoria (n). 

3218. Ne o. pallescens (C. Presl) E YU. 23/ 
6 [1/1]; 2,000 ft.; [La] Victoria (s). 

Selaat viticulosa Klotzsch; MO, YU. 16/8/ 

55 [1/18]; 2,500 ft.; seaside of mountains between 

Caracas and La Cuayra a (n). 

. Selaginella cavifolia A. Braun; BM; this is an 
ISOSYNTYPE of S. Dod var. elongata A. Braun, 
Ann. Sci. Nat. Bot., 85. 1865; cited by 
Alston et al. (1981). 

. Selagin i Md Hg A. Braun; MO, YU; these 
are ISOSY of S. pu var. elongata 
Braun, RE Sci. Nat. , V. 3: 285. 1865. [4/ 
17]; 7,000 ft.; high Mund around Colonia To- 


322. 


var (n). 

. Selaginella flagellata Spring; BM, MO, E these 
are ISOSYNTYPES of S Am igua A. Braun, Ann 
Sci. Nat. Bot., t 1865; cited e ‘Alston 
et al. (1981). [2/22]; D 300 ft.; La Victoria and 
al (n). 

324, a Neg as qM A. Braun, Ann. Sci. 

o . 1865; SYNTYPES, B, BM, YU 
He by Alsion o et E (1981). At YU, this species 
and S. flag 

s Pa semicordata (C. P 
YU. 23/8/55 [1/18] 8,000 ft.; 


wo 
N 
¿ES 


q 
N 
en 


. Asplenium oligophyllum Kaulf.; MO, U. 
O ft.; a few mi. SW of Colonia Toter: river 
htiega (ny 17/5/55 [1/26] (s). 
ST. A er (Sphaerocionium) trichophyl- 
; GH; YU cited by Morton (1947). 25/ 
8/56 [1/20] 8,000 ft.; 10 mi. E of Colonia [Tovar], 
highest mountains (n); Lagunassa (s). 

. Hymenophyllum (Sphaerocionium) lanatum Fée; 
YU. 12/7/56 ne 7,500 ft.; a few mi. of 
Colonia [Tovar] (n 

ad 328. re HAB (Sphaerocionium) lan 
Fée; YU. 12/7/56 [1/10] high alum m Ta 
Meo dps s (s). 

329. Trichomanes (Acarpacrium L.; MO, 
US, NU 2. [2/20]; 6, "à 7,000 ft.; seaside of 
mountains north of Colonia Tovar (n). 

330. Lastreopsis exculta (Mett.) Tindale; MO. 28/6/ 

56, 8/1/57 [2/?]; 6,000 ft.; Colonia Tovar (n). 


342 Annals of the 
Missouri Botanical Garden 
331. Asplenium den igi L.; YU— 2. 12/7/56 [17 podium pendulum Sw. 12/12/56; 6,500 ft.; sea- 


?}; 6,000-7,000 ft.; Colonia Tovar (n). 
3318. oo auriculatum Sw.; YU. 22/4/58 [1/ 


332. depen serra Langsd. & Fischer; YU— 2. [?/ 

,000-8,000 ft.; Colonia Tovar (n). “Has been 

, but pinnae very different; 
also the meshes at E in the scales of rhizome 
smaller” (s). 

333. Asplenium serra Langsd. & Fischer; YU. 22/1/ 
56 [1/3]; 7,000 ft.; seaside of mountains N of 

) 


rts, n.s. 8: 195. 1861; svNTYPES, YU —3 sheets, 
collected on two different dates; ISOSYNTYPE, K. [2/ 
16]; 6,000 ft.; seaside of mountain range between 
Colonia [Tovar] and Chichiribichi (n). 
ad 335. nue fendleri D. Eaton; YU. 22/1/56 
[1/? 
336. Bolbitis portoricensis (Sprengel) Vd npa MO, 
YU — 2; cited by Hennipman (1977). One collec- 
tion at YU mixed with juveniles of Pteris altissima 
Poiret in Lam. 29/1/57 [1/13]; 3,500 ft.; veda 


of mountain range between Maracai and Choron 


.. Not located, Lycopodium (n). 17/7/56 [1/?] 
la cd ft.; high mountains E of Colonia Tovar, upon 
es (n). 


Gran mitis pilosissima (Mart. & Gal.) C. Morton; 
YU. 12/7/56 [1/1] 7,500 A Colonia Tovar (n). 
Not located, no data in notebo 

um ea (orah) C. Chr.; 


339. 
340. Blechnu 


341. Polypodiu at 

26/5/54 [1/1] 3,500 ft.; 

[Tovar] (n). 

342. Ophioglossum nudicaule L. f.; MO. 5,000 ft.; 

few mi. S of Colonia [Tovar], beneath low de 

at margin of woods (n). 

Polypodium bolivari Sota; YU. 3/9/56 [1/3]; 
,000 ft.; a few mi. S of Colonia Tovar (n); near 


. WA. Omit : 
Biscaina, S of Colonia 


343. 


Maya (s). 

. Cyathea villosa Willd.; B; this is the HOLOTYPE of 
Alsophila vernicosa Mett. ex K 
155. 1869; isorvPEs, GH, MO, YU; cited by Bar- 
rington (1978) as Trichipteris villosa (Willd.) R. 
Tryon. 21/8/56 [1/6]; 5,800 ft.; a few mi. S of 
Colonia [Tovar], in savannahs (n). 

345. Not located. 2/9/56; 6,000 ft.; Colonia Tovar (n). 

346. Not located. 5,000 ft.; beneath low eg) margin 

of woods, 


r^ VE Pra of. m ulinum i ing 


w 
D 


1 (n) 
ée) C. Ma MO. [2775 
O ft.; €— of mountain range between Ma- 
racai and Chor s » 
Grammitis aff s subtilis (Kunze ex Klotzsch) C. Mor 
ton; MO, YU. co [1/7]; 6,500 ft.; cesado 


of mountain range between Maracai and Choroni 


349. 


(n). 
350. Grammitis sp.; YU; cited by Eaton (1861) as Poly- 


~ of mountain range between Maracai and Cho- 
ni (n). 
351. eu mmitis asplenifolia (L.) Proctor?; YU. Juve- 
nile. 12/12/56 [1/1]; 6,000 ft.; seaside of moun- 
en Maracai and Choroni (n). 
i ; YU. ae ad ala 
delo near t Güigüe (n 
; Poly. odium loriceum L.; , YU— 2. [273]; 
,000 ft.; uide of r mountains "between Maracai 
and S (n). 
. Mi ramma reptans (Cav.) A. R. Smith; MO, 
YU. TA] 2,500 ft.; seaside of mountains be- 
tween Maracai and Choron ni (n) 


ok. is ex C. Chr.; 


355. Cochlidium rostratum (Ho 
MO. [2/?]; 6,000 ft.; in crossing the mountains 
from Maracai towards ni (n 


between Maracai and Choroni, and between Va- 

lencia and San Estevan (n ve 
Pityrogramma chrysoconia (Desv.) Maxon ex 

Domin; YU. [4/?]; 6, 000 ft: seaside of mountain 

range between arene and Chor oni (n). 

. Eriosorus hirtus (H opel. var. hirtus; 

YU; cited by A. dM [3/1]; 6,000- 7, 000 
ft.; a few mi. W of Colonia [Tovar] (n). 

; buon hispidulus (Kunze) Vareschi var. his- 
pidulus; K, YU; cited by A. Tryon (1970). 12/ 
12/56 [1/3]; 6,000 ft.; seaside of mountain range 
between Maracai and Choroni (n). 

. Thelypteris (Amauropelta) KEET (Willd.) R. 
Tryon; YU. 12/12/56 [1/2]; 2,500 ft.; seaside 


of mountain range between Maracai and Choroni 


357?. 


w 
e 
No) 


w 
On 
m= 


(n 

: Elaphoglossum peltatum (Sw.) Urban; MO, YU. 
[3/11]; 7,000 ft.; seaside of mountain range be- 
tween Maracai and Choroni (n). 

. Elaphoglossum pusillum (Mett. ex Kuhn) C. Chr.; 
GH, NY, PH; these are ISOSYNTYPES of Acrosti- 

hum fraseri Mett. ex Kuhn, Linnaea 36: 43. 1869. 
12/12/56 [1/?] 2,000 ft.; near Esmeralda above 
Choroni (n). 

; cia iis crinitum (L.) Christ; YU. [2/2]; 
2,500 ft.; mountain range between Ma- 
racai and: Chor oroni in 

Adiantum tetraphyllum Humb. & Bonpl. ex Willd.; 
YU. 12/12/56 064 2,000 ft.; seaside of moun- 
tain range between Maracai and Choroni (n). 

. Elaphoglossum pusillum IM ex Kuhn) C. Chr.; 
MO; det 

. Pteris aliisima Poiret in Lam.; YU. 12/12/56 
[1/1] 2,500 ft.; ~ of mountain iiis bed 
Maracai and Choroni 

: rii sede agita (Rad = MO, YU, 
[2/19]; 2 ntain range be 
tween ein = Chor d 

. Asplenium cirrhatum Rich. ex Willd.; YU. [2/1]; 
4,000 ft.; seaside of mountain range between 
racai and Choroni (n). 

. Asplenium tenerrimum Mett. ex Kuhn, Linnaea 
36: 97. 1869; isorYPES, MO, NY, US; cited by 
in & Lellinger (1966) as A. cuspidatum Lam. 

. tenerrimu ett. ex Kuhn) C. Morton & 
pe 21/11/56 [1/?]; 2,300 ft.; near La Vic- 
toria (n). 


Volume 76, Number 1 


Smith & Todzia 343 


1989 Fendler's Venezuelan 
Fern Collections 
369. Diplazium expansum Willd.; YU—2. 12/12/56 by Stolze (1974). [3/13]; 2,000-3,000 ft.; seaside 


w 
-l 
— 


[2v 
N 
w 


376. 


377. 


N 


378. 


[1/1]; 4,000 ft.; seaside of mountain range between 
Maracai and Charen ni (n). 


. Nephrolepis rivularis Mod Mett. La Krug; MO, 


YU. 29/1/57 [1/7]; 5,000 
range between Maracai and de oroni 


. rel (Steiropteris) praetervisa (Kuhn) A. 


R. Smith; K, YU; these are ISOTYPES of Aspidium 
praetervisum Kuhn, Linnaea 36: 111. 1869; e 
by Smith (1980). 12/12/56 [1/2]; 2,500 ft.; 


iim of mountains between Maracai and Choroni 


1 Thelypteris(Steiropteris ram (D. Eaton) Reed; 
YU— 


E of Aspidium fendleri D. Eaton, 

Mem. pes p» FON n.s. 8: 210. 1861; 

ISOTYPES, B, GH, K; cited by Smith (1980). [2/9]; 

500 ft.; seaside of mountains between Maracai 
(n 


2, 
and Choroni 


. Thelypteris (Steiropteris) decussata (L.) Proctor 


var. decussata; MO, YU — 2 cited by Smith (1980). 
[2/8]; 3,000 ft.; id etween Cho 
roni and Mariscal: also a p resi and San 
Estevan (n 


i Dnnstaedtid obtusifolia (Willd.) T. Moore; YU. 


12/12/56 [1/4] 6, 000 ft.; seaside of mountain 


. Saccoloma inaequale (Kunze) Mett.; MO, YU— 


2. [2/9]; 5,000 ft.; T of mountain range be- 
tween Maracai sid Choroni (n). 

Saccoloma elegans Kaulf.; YU. 12/12/56 [1/2]; 
3,000 ft.; seaside of mountain range between Ma- 
racai and Choroni (n). 

Lindsaea stricta (Sw.) Dryander var. stricta; B, 
BR, C, F, G, GH, GOET, K, MO, PH, US, YU; 
cited by Kramer (1957). [2/22]; 4,000 ft.; qe 


of mountain range between Maracai and Choro 


(n). 
Lindsaea lancea (L. ) Beddome var. lancea; MO, 


mountain range between Maracai and Choroni (n). 


other species o Lin saea. 


. = 638 (n s), which is Lindsaea klotzschiana 


Moritz in Ettingshausen. Without locality data in 


00 
. Li dsaea stricta (Sw.) Dryander var. stricta; cited 


by Kramer (1957). 12/12/56 [1/6]; 5,000 ft.; 
seaside of mountain range between Maracai and 


Choro 


P il V PAA T PEN Moritz in ld pouen 


YU. 12/12/56 [1/?]; 5,000 ft. 


. Grammitis taxifolia (L.) usd YU. 30/1/58 


[1/1]; 6,500 ft.; seaside of mountains between Agua 
Blanca and E bote (n). 


. Cyathea dm (Klotzsch) Domin; YU. 12/12/ 


56 [1/1] 5,000 ft; seaside of mountain range 
between Mirscai and Choroni (n). 


. Cyathea senilis (Klotzsch) Domin; YU— 2. [2/3]; 


6,500 ft.; seaside of mountain range between Ma- 
cai and Choroni and further west (n). 


ra 
. Not located. 29/1/57 [1/?]; 6,000 ft.; seaside of 


mountains between Maracai and Choroni (n). 


. Cnemidaria horrida (L.) C. Presl; MO. YU: cited 


ange between Maracai and Choroni, 


386. Cnemidaria karsteniana (Klotzsch) R. Tryon; MO; 
cited by Stolze (1974). [3/8]; 5,000 ft.; seaside of 


mountain range between Maracai and Choroni and 


387. Hymenophyllum (Sphaerocionium) lanatum Fee; 

, MO, YU; cited by Morton (1947). 12/12/ 

56 [1/11]; 6,000 ft.; seaside of mountains between 
Maracai and Choroni (n). 

. Trichomanes (Achomanes) crispum L.; YU. [2/ 
21]; 4,000-5,000 ft.; rd of mountains between 
Maracai and Choroni 

3886. m! (Achoman Sas) sp., * adde E 

89); YU. 29/1/57 [1/2] (s 
389. pha (Achomanes) sp., A MO, 
YU. [3/10]; 5,000 ft.; p of mountains be- 
tween Maracai and Choroni 

390. Trichomanes TIo: D pinnat um Hedwig; 

MO, YU. [2/ Ls 4,000 d 4 se aside of mountains 
between Maracai and nd ni (n). 

391. Elaphoglossum label um b Smith; YU. [2/4]; 

gn ft.; in crossing mountains from Maracai to 


(e) 
co 
eo 


Choroni (n). 
392. dd Wei 12/12/54 [1/?]; 6,500 ft.; Colonia 
ar (n). 


393. LI. J] (M. 

Kuhn; YU. 12/12/56 tr 6, 000 ft.; ds of 

mountain range between Maracai and Choro ni (n). 
Sw. 


l YU. 12/12/56 [1/3]. 6,500 ft.; seaside of moun- 


,500 ft.; Colonia Tovar, on rocks near a Ee 
(n); 4,000 ft.; (s). 
396. Schizaea elegans (Vahl) Sw.; MO. 12/12/56 [1/ 
?]; 4,500 ft.; seaside of mountain range between 
Maracai and Choroni (n). 

397. Selaginella diffusa (C. Presl) Spring; MO, YU. 12/ 
12/56 [1/12]; 4,000 ft.; p of mountain range 
between Maracai and Chor ni (n). 

398. Li d (n), not estad. 12/12/56 [1/?); 

,000 ft.; seaside of mountain range between Ma- 
racai and Choroni o 

399. Lycopodium cernuum L.; YU. 12/12/56 [1/3]; 
5,500 ft.; seaside p mountains between Maracai 
and Choroni (n) 

3998. Lycopodium cernuum L.; YU, mounted on same 
sheet as 399. 12/12/56 [1/1]; 6,000 ft.; moun- 
tains between Maracai and Choroni on dry south 
side (s 

400. Lycopodium thyoides Humb. & Bonpl. ex que 
s ud [1/1]; 6,500 jes 3 vag of mou 

aracai and 

401. Not Que "6/ hs 57 [17?]; biia be E 

of Colonia Tovar 

402. Selaginella ph ee spora A. Braun; YU. 12/ 
e be [1/3]; 5,000 ft.; deni of mountains be- 

en Maracai and Choroni n). 
403. Selaginella Sraten Song: YU. 12/12/56 [1/ 
4), 3, id t.; seaside of mountains between Maracai 
and Chor kp >. 
t A aih sp.; YU; cited by Eaton (1861) as 
decus decomposita Hook. [could be C. margina- 
a H.B.K.]. 7/3/57 [1/?] 3,500 ft.; seaside of 


EN 
co 
ER 


344 


Annals of the 
Missouri Botanical Garden 


ÉS 
© 
on 


408. 


ec 


409. 


ño) 


4 


— 
o 


424. 


NN 


mountain range between Valencia and San Estevan 


n). 
. Athyrium filix-femina (L.) Roth; MO, YU. 6/1/ 


57 [1/22]; 8,000 ft.; 12 mi. E of Colonia Tovar, 


highest mountains (n). 


. Grammitis moniliformis (Lagasca ex Sw.) Kau 


YU; cited by Eaton (1861). 29/1/57 [1/?]; 7, Dod 
ft.; seaside of mountains between Maracai and Cho- 
roni (n). 


. Polypodium hygrometricum Splitg.; YU. 28/1/ 


57 [1/1]; 3,000 ft.; seaside of mountains between 
Maracai and Choroni (n). 

Bolbitis hemiotis (Maxon) Ching; YU. 29/1/57 
[1/5]; 4,000 ft.; seaside of mountains between Ma- 
racai and Choroni (n) 

Campyloneurum occultum (Christ) L. D. Gómez; 
YU. 29/1/57 [1/3]; 6,000 ft.; seaside of moun- 
tains between Maracai and Choroni (n). 


. Campyloneurum magnificum T. Moore; YU—4 


sheets, collected on two different dates; these are 


MO— 2. [2/13]; 4,000 ft.; ire of mountains 
between ipa ip ipe the sea (n). 

Asplenium sp.; YU; cited E "Eston (1861) as 
salicifolium L. 12/2/58 [1/?] 4,000 ft.; don 
of mountains between Petaquire and Cariaca (n). 


seaside of mountains between Petaquire and the sea 


(n). 
; T sidus haria Desv.; YU. 9/2/57 [1/ 


2]; 5,000 ft.; seaside of mountains between Peta- 
quire eu the sea (n). 


. Polypodium aff ` fraxinifolium Jacq.; YU. 29/1/ 


57 [1/1]; 6,500 js seaside of mountains between 
Maracai and Choroni (n). 
Polypodium arena Sw.; YU. 29/1/57 [1/1]; 


,000 de seaside of mcn between Maracai 


Bonpl. ex Willd.) Ching; YU. 12/12/56 [L1 
2,500 ft.; seasi e of te untain range between Ma- 
racai and Chor 


: Polypodium (Phlebodium) aureum L.; YU. [2/1]; 


3,000 ft.; Via. of mountain range between Ma- 
racai and Choron 


n). 
. Polypodium lano pus Klotzsch; YU. 29/1/57 [1/ 


3]; 6,500 ft.; seaside of mountains between Maracai 
and Choroni (n). 


. Microgramma tecta (Kaulf.) Alston; MO, YU. [2/ 


22); 2,000 ft.; — of mountains between Pe- 
taquire and the s 


. Microgramma b copa ATA ) Copel.; MO, YU. 


[2/15]; 2,500 ft.; seaside of mountains between 
Maracai and Choron i (n). 


. Pleopeltis percussa (Cav.) Hook. & Gre 


; MO, 
YU. [3/3]; 4,000 ft.; ; seaside of mountains i 


. Polypodium sororium Humb. & Bonpl. ex Willd.; 


MO, YU. 9/2/57 [1/6]; 8,000 ft.; highest moun- 
tains E of Colonia Tovar (n). 


. Dicranoglossum furcatum (L.) J. Smith; YU. 9/ 


2/57 [1/2]; 3,000 ft.; seaside of coast range of 
mountains (n). 

Vittaria costata Kunze; YU. 29/1/57 [1/1] 5,500 
ft.; seaside of mountain range between Maracai and 
Choroni (n) 


425. gps Ap aei E ) Splitg.; YU. 29/1/57 [1/ 
To 2" ountain range between 
and Chor 
426. Ofer cervina (L) RR MO, YU— 3. 9/2/ 
E Lad 4|] 4,000 ft.; seaside of m ia range 
n Petaquire and Chichiribichi (n). 

427. Not bed 29/1/57 [1/?] 6,000 ft.; ind of 
mountain range between ym and = ni (n). 

428. Elaphoglossum herminieri x Fée 
. Moore; YU. 29/1/57 MURS 3, 500 ft: E 

of mountain range between Maracai and ‘Choro: 


(n 
429. Elaphoglossum iini an Presl) Brackenr.; YU. 


O ft.; seaside of ain range between Ma- 
racai sak Choroni (n); 29/1/57 [1/2] (s). 
430. Elaphoglossum sp.; d by E 861 


ft; seaside of mountain range between Maracai and 
Charon i (n). 

431. palpa ne Hieron.; MO, YU. [3/4]; 3,000 
ft.; seaside of mountain = oes Petaquire 


432. Asplenium auriculatum Sw: YU. 29/1/57 m 
1]; 6,000 ft.; seaside of mountain sas betwe 
Maracai and Choroni (n). 

433. Asplenium pteropus Kaulf.; MO, YU. [2/8]; 5,000 
ft.; seaside of mountain range between Petaquire 
and Choroni (n). 

434. Asplenium cirrhatum Rich. ex Willd.; YU. 9/2/ 
57 [1/3]; 5,000 ft.; — of mountain range 
between Petaquire and the 


ea (n). 
435. Diplazium pedatum itech YU. [2/1]; 5,000 


4356. vore pedatum Klotzsch; YU. 12/12/56 
[1/1] (s). 

436. Diplazium pedatum Klotzsch; MO, YU. 29/1/57 
[1/7]; 6,000 ft.; seaside of mountain range between 
Maracai = Choroni 

437. Th ola pte | PAS y J L| > y (NA 
Mett. rt YU. 26/1/57 yii 100 ft.; near 
pes N of Choroni (n 

438. Thelypteris oa pelta) balbisii (Sprengel) 
Ching: AU. 26/1/57 (LAE 200 ft.; near Colombia 

on the coast (n). 


439. lado aid bir uu sp., aff. gracilis (He- 


ward) Proctor; YU. 29/1/57 [1/1]; 3,500 ft.; 
aside of mountain range between Maracai and 
nia 1 (n). 


440. Thelypteris pop pelta) pilosula (Mett.) R. 
Tryon; YU. 6/1/57 we 4 id O ft.; highest 
mountains E of LR onia 

441. Thelypteris odes hispidula e ) Reed; 
YU. [2/1]; 4,000 ft.; a few mi. SW of Colonia 


Tovar (n). 
442. Not Ta 25/1/57 [1/?]; 300 ft.; seaside of 
mountains between Maracai and Choroni (n). 
443. Cyclopeltis semicordata (Sw.) J. Smith; MO. 25/ 
1/57 [1/?]; 200 ft.; between Choroni and Colombia 


(n). 

444. “Ctenitis microchlaena” (Fée) A. R. Smith, comb. 
ined.; YU. 9/2/57 [1/1]; 5,000 ft.; seaside of 
foU SUA between Petaquire and the sea (n). 

445. Thelypteris (Goniopteris) nephrodioides (Klotzsch) 
Proctor; YU. 9/2/57 [1/1] 3,000 ft.; seaside of 
mountains between Petaquire and the sea (n). 

446. Thelypteris (Goniopteris) paucijuga (Klotzsch) A. 
R. Smith, vel aff.; YU. 9/2/57 [1/3]; 4,000 ft.; 


Volume 76, Number 1 
1989 


Smith & Todzia 


345 


Fendler's Venezuelan 
Fern Collections 


> 
> 
- 


> 
> 
e 


449. 
450. 


A Ctenitis pulverulenta (Poiret) br ow 
y C 


s up of mountains between Petaquire and the sea 


K, YU— 
cite istensen (1920) as Dryopteris kar- 
D pe bun Hieron. 9/2/57 [1/3], 3,000 


e of mountains between Petaquire and the 


E 
. Dennstaedtia bipinnata (Cav.) Maxon; MO, YU — 


2. [3/5]; 4,000 ft.; seaside of mountains north of 
Colonia Tia (n). 
= 4506 (n). 


Dicranopteris seminuda (Klotzsch) Maxon; MO, 
— 2. [3/20]; E 500 ft.; high mountains NE of 


Colonia [Tovar 


4508. Dicranopteris prea E L. Underw.; 


451. 


— 


452. 


N 


4 


4 


on 
> 


4 


456 


` YU. 29/1/57 [1/2]; 3,000 ft; 


465. 


en 
e 


en 
"d 


. Trichomanes (Lacosteopsis) debile Bo 
. Danaea elliptica J. E. Smith; MO, YU. 3 


* Sela gt inella h 


(n). 
. Pleopeltis percussa (Cav.) Ho 


MO, YU. 9/2/57 [1/6] (s). 

Trichomanes (Acarpacrium) trigonum Desv.; YU. 

28/1/57 [1/1]; 4,500 ft.; seaside of mountain 

range between Maracai and Choroni (n) 

E iint nn (Didymoglossum) punctatum Poiret 
sp. sphenoides (Kunze) W. Boer; B, G, GH, 

MO, YU; cited by Wessels Boer (1962). 29/1/57 

[1/19]; 4,500 ft.; seaside of mountains between 

Maracai and Coran i (n). 

Trichomanes (Lecanium) membranaceum L.; MO, 

n 10/2/57 [1/20]; 3,000 ft.; Eee of moun- 

range between Petaquire and the sea (n). 


: Trichomanes EE) Didi (Bosch) 
H, M 


O, YU; cited by Wessels Boer 
(196). 10/2/51 [1/16]; 3.500 ft.; seaside of 


anii 

O, US; cited by Wessels Boer (1962). 
13/12/56 [1/?]; 5,500 ft.; seaside of mountains 
between Maracai and Choroni (n). 


Doddi n 1 


seaside of moun- 
tains between Maracai and Choroni (n) 


. Hymenophyllum (Sphaerocionium) elegans 


Sprengel forma minor C. Morton; GH, MO, YU; 
cited by Morton (1947). 29/1/57 [1/19]; 6,200 
ft.; seaside of mountains between Maracai and Cho- 
roni (n). 


. Hymenophyllum (Sphaerocionium) hirsutum (L.) 


v of mountain 


Sw. 29/1/57 [1/?] 6,000 ft.; se 
ridge between Maracai and Choron 


YU—2. 10/3/57 [1/14]; 5,000 ft.; 


mountain: 


seaside en e and 


of mountain range betwe 
Chichiribichi (n); 9/2/51 [1/10] (s). 


Martius 
MO, “YU. 9/2/57 [1/16); 2, 500 ft.; seaside of 
mountain range between Petaquire and Chichiri- 


qom ornatu tt.) Christ; K, YU; 
kel in Smith 6) [2/4]; 3,000 ft.; 
seaside of na range between Agua Blanca 


ok. & Grev.; YU. 
[?/1]; 6,500 ft.; Colonia Tovar (n). On same sheet 
as 421. 
Grammitis sp.; YU; cited by Eaton ven as Poly- 
podium truncicola Klotzs ch. [2/?]; 6,000 ft.; sea- 
side of mountains between Valencia d Sen Es- 
tevan and between Agua Blanca and Cumbote (n). 


469. 


. Elaphoglossum sp.; YU; cited by Eaton (1861) as 


Acrostichum perelegans Fée. 7/3/57 [1/?] 4,000 
ft.; seaside of mountain range between Valencia 
and Campanero (n). 


. Grammitis papal e (Copel.) Proctor; YU. 30/1/ 


58 [1/3]; 6,500 ft.; seaside of mountains between 


Agua Blanca and Cumbote 


(n). 
. Adiantum latifolium Lam.; YU. 7/3/57 [1/2]; 


1,000 ft.; 
side of mountain range (n). 

Adiantum petiolatum Desv.; YU. [2/4]; 500-1,000 
ft.; near San Estevan a few mi. N of Puerto Cabello 


between Valencia and San Estevan, sea- 


(n ). 
4698. Adiantum sp.; YU; cited by Eaton (1861) as A. 


470. 
471. 


4 


N 


473. 


476. 


Dm 


2. 


lfussii Kunze [may be A. petiolatum Desv.]. 
— 4699 (n), not located. 

Polypodium triseriale Sw.; YU. 7/5/57 [1/1]; 
7,000 ft.; high mountains E of Colonia [Tovar] (n). 
Thelypteris (Amauropelta) pachyrhachis (Kunze 
hachis; ve MO, YU 
[2/6]; 4,000 ft.; seaside of mountains between Agua 
te, and between Valencia and 


. Thelypteris (Cyclosorus) conspersa (Schrader) A. 


Smith; YU. [2/3]; 4,000 ft.; Macarao River 
valey W of En gr also re x mountains 
etween Valencia and San 
Thebipiena (Cyelosorus) hispidula; YU. (2/31. 
) Reed; 
YU; "this is an ISOTYPE of Polypodium stramineum 
Baker in Hook. & Baker, Syn. Fil. 316. 1867. [1/ 
4]; 2,300 ft.; Piedernales near Güigüe (n). 


. Thelypteris i) ariel vu d (Deane: ) Roed; 


YU. 7/3/57 [1/1] 2,5 
between Valencia and e ida evan 


Thelypteris ki cde pridiri (Book JR 
Tryon var. costalis (Baker) A. R. Smith; K; this is 
an ISOTYPE o T. costale Mett. ex Kuhn, 


Linnaea 36: 111. 1869. 7/3/57 [1/?]; 1,500 ft.; 
seaside of mountains between Valencia and San 
Estevan (n). 


. Thelypteris Mit cad yas pm m R. 
A. R. 


Tryon var. costalis (Bak h; YU. 7/ 
3/57 [D1ZIE 1, So ft.; o of rd range 


between Valencia and San Estevan (n) 


. Not located. 7/3/57 [1/?] 1,000 ft.; seaside of 


mountain range between Valencia and San Estevan 
). 


. Lindsaea stricta (Sw.) tng M var. stricta; G, 


GH, GOET, K, MO, PH, YU; by Kramer 

(1957). 7/3/57 [1/12]; 4, 000 pr . seaside of 

mountain range between Valencia and San Estevan 
n). 


. Cnemidaria spectabilis (Kunze) R. Tryon 


spectabilis, GH, K, MO, YU— 2; cited by Stolze 
(1974). 7/3/57 [1/9]; 4,500 ft.; seaside of moun- 
tain range between Valencia and San Estevan (n). 


P Hymenophyllum (Sphaerocionium) dependens C. 


US, MO, YU; cited by Morton (1947). 
29/1/58 [1/?]; 5,000 ft.; seaside of mountain 
range between Agua Blanca and Cumbote (n) 


. Dicranopteris pectinata (Willd.) L. - Underw.s ; MO, 


YU. 7/3/57 [1/6]; 6,000 ft.; 
between Valencia and San Evan yA ). 


. Dicranopteris pectinata (Willd.) L.  Underw.; MO, 


YU. 7/3/57 [1/5]; 5,500 ft.; 
between Valencia and San Estevan y 


346 Annals of the 
Missouri Botanical Garden 
484. Not located. 7/3/57 [1/?] 5,500 ft.; seaside of — 4. capillus- veneris L. p 


ÉS 
No] 
Oo 


mountains betw ween Valencia and San Este evan (n). 


. Schizaea poeppigiana Sturm in Martius; MO. 27/ 


2/57 [1/?]; 3.800 ft.; a few mi. S of Colonia Tovar 


(n). 
. = 322, Selaginella (n), not located. 7/3/57 [1/ 


?]; 4,000 ft.; seaside of mountains between Valencia 
and San Estevan n ). 


. Selaginella cruciformi is Alston ex Crabbe & Jerm 


BM; cited by us et al. (1981). 7/3/57 [1/7] 
3,000 ft.; seaside of mountains between Valencia 
and San Estevan 


. Lophosoria quadripinnata (J. Gmelin) C. Chr.; 
YU. [2/7]. 


. Polypodium fraxinifolium Jacq.; YU. 57 


E. 1]; 8,000 ft.; high mountains E of la Tovar 


(n). 
. Asplenium serratum L.; YU. 7/3/57 [1/3]; 3,500 


ft.; seaside of mountain range between Valencia 


and San n (n) 
. Not Heated. 1/3/51 [1/7]; 4,000 ft.; seaside of 


mountain range between Valencia and San Estevan 


" Cyathea barringtonii A. R. Smith ex Lellinger; 


YU; cited by Barrington (1978) as SII dada 

cordata (Klotzsch) R. Tryon. [2/2]; 6,300 ft.; sea 

-— of mountain range between Maracai and Cho: 
ni (n). 


. a villosa Willd.; YU. [2/4]; 5,800 ft.; sea- 


side of mountain range between Agua Blanca and 
Cumbote, also savannahs S of Colonia [Tovar] (n). 
Cy athea villosa Willd. 


2?. 
i Selaginella flabellata A. ) Spring var. latifrons A. 


a n. Sci. Nat. Bot., V. 3: 278. 1865; 
ISOT YPES , B YU; cited by Alston et al. 
(1981). 30/1/58 iiA] 3,000 ft.; seaside of 


ains between Agua Blanca and Cumbote (n). 


. Selaginella i a y" Braun; YU. [2/4]; 6,000 


ft.; Colonia Tov. 


ar (n). 
A Elaphog lossum variolatum Mickel; YU. 4,000 


ft.; seaside of mountain range between Agua Blanca 
and dia (n) 30/1/58 [1/1] (s). 


. Adiantum sp.; YU; cited by Eaton (1861) a 


penne veneris L. 20/2/58 [1/?]; 3,000 ft.; "Ha. 


cienda Curisal (n 


à Diplazium celtidifolium Kunze; YU. 22/4/58 [1/ 


1]; 4,500 ft.; €— of mountain range between 
Du and the sea (n). 


. Dipl azium favescen dise ) Christ; MO, YU. 22/ 


4/58 [1/7], 4,000 ft.; seaside of mountain range 


' between Petaquire and e e sea (n). 


. Rumohra adiantiformis (G. Forster) Ching; YU; 


cited by Eaton (1861) as Aspidium coriaceum Sw. 
22/4/58 [1/?]; 5,500 ft.; seaside of mountains 
een Petaquire and the sea (n). 


W 
s.n. Lindsaea klotzschiana Moritz in Ettingshausen; YU; 


originally det. L. quadrangularis Raddi. 


INDEX 


Below is an alphabetical list of pteridophyte species 
collected in Venezuela by Fendler. The numbers refer to 
the Fendler collection numbers in the list of determina- 


tions 


Adiantopsis radiata (L.) Fée—67 
Adiantum braunii Mett. ex Kuhn— 68 


A. concinnum Humb. & Bonpl. ex me — 13, pt; ad 
73; 73/76; 75; ad 75; 758; 76; 
A. UE Martius—81 


A. lunulatum Burm. f.— 73, pt; 82 

A. macrophyllum Sw.— 88 

A. patens Willd.— 79 

A. petiolatum Desv. P 469 

A. poiretii Wikstr. — 

A. polyphyllum Willd —80 

A. pulverulentum L.—83, pt; 86 

A. raddianum C. Presl — 74 

A. Ae gad Mira — 84 

A. tenerum Sw.— 69; 

A. Dinky e Hunb. & Bonpl. ex Willd.—ad 85; 
364 


A. les L.—85 

A. sp. — 4698; 

Anemia ferruginea de B. P var. ferruginea — 1 
A. hirsuta (L.) Sw.— 

A. oblongifolia (Cav.) rw —9; 98; 10 

A. pastinacaria Moritz ex Prantl— 11 

A. phyllitidis (L.) Sw.—13 

A. villosa Humb. & eye ex Willd.—6 

A. sp., undescribed? — 

Anat citrifolium (L. ) S litg. — 425 


M 


Aras ode dina M (Sw.) Ching — l 17, pt; 170 
Asplenium abscissum Willd.— 143, pt; 1438 

4. alatum Humb. & Bonpl. ex Willd.— 145 

A. auriculatum Sw.— 3318; 432 

A. auriculatum Sw., vel aff.— 144 

A. auritum Sw. var. obtusum Kunze ex Mett.— 141 


A. cristatum Lam.— 124; ad 124; 134, pt 

A. aff. cristatum Lam.—124 46 

A. flabellulatum Kunze— 125, 

A. flabellulatum Kunze var. um Klotzsch — 125, 
pt 


A. formosum Willd. — 133 


428 
A. n Liebm. 2142; ad 142 
A. s L.—131 
A. oligophyllum dide md pt; 326 
A. praemorsum Sw 1578 
A. pteropus Kaulf. ie 
A. pumilum d meu 
A. raddianum Gaud. — 
A. radicans L.—126; on ud 140, pt 
A. ruizianum "ir ia 13968 
A. rutaceum (Willd.) ~ai 23 
A. salicifolium L.— 33 
A. serra Langsd. " Fischer 15 332; 333 
L.— 


zm 
A. Gun Mett. ex Kuhn— 368 


Volume 76, Number 1 
1989 


Smith & Todzia 
Fendler's Venezuelan 
Fern Collections 


347 


É fl fina S ed Roth—405 
A. skinneri (Baker) C —163 


Blechnum X caudatu ely dor 113 


B. en 


— 340 
i^. 116; s 122 
hnum hito d —10 
Bolbitis hemiotis peel Ching — 408 

. portoricensis (Sprengel) Burn an— 336 
B. serratifolia AS unen, ex Kaulf.) Schott —311 
Botrychium sp.—4 
| sé ca amphostenon (Kunze ex Klotzsch) Fée — 
226; 
4 A (Sw.) oer pt 
chlorolepis Alston — 224, 225 
chrysopodum eg Fée 22 2288 
. decurrens (Raddi) C —231 

magnificum T. Hotes T 
occultum (Christ ) L. D. Gomez— 409 
. phyllitidis us ) C. Presl— 227 
repens (Aubl.) C. Presl— 229; 230 
Cheilanthes concolor (Langsd. & Fischer) R. & A. 


—— 


QA 
C. margin —90 
a micron seen ) Sw.—65 
a horrida (L.) C. Presl — 385 
C. karsteniana (Klotzsch) R. Tryon— 386 
C. spectabilis (Kunze) R. Tryon. var. spectabilis — 480 
Cochlidium rostratum (Hook.) Maxon ex C. Chr. — 355 
E L. E. Bishop — 356 
. & Bonpl. ex Willd.) Ching — 204 
ed.— 


C. nodes pal (Fée) A. R. Smith, comb. ine 
444 


C. nigrovenia (Christ) Copel. — 194 
G pulverulenta (Poiret) Copel. — 447 


g—2 
C. submarginalis (Langsd. & Fiche) Ching— 195; 197 
Cyathea barringtonii A. ex Lellinger — 491 
C. caracasana (Klotzsch) Doniin. var. caracasana — 56; 

ad 56 
C. ebenina Karsten 
C. fulva (Mart. & Cal. 5 m e—51; 52 
C. gibbosa (Klotzsch) vee cea 382 
C. pungens (Willd.) Domin—55 
C. senilis (Klotet) Domin— 383 

46; 4 


ct 
um 
Y 
hi 
3 
~, 
© 
an 
E] 
C 


D. moritziana C. Presl — 
Dennstaedtia arborescens (Willd. Ekman ex Maxon 


D. T" (Cav. a Maxon — 448 
D. cicutaria (Sw.) T. Moore —58 
D. dissecta (Sw. ) T. Mons «59 


D. obtusifolia (Willd.) T. Moore— 374 

Dicksonia sellowiana Hook.— 

peii 2 furcatum (L. = J. Leo a 423 

Dicranopteri L. Underw.— 44; 449; 
450 


D. pectinata (Willd.) L. Underw. — 482; 483 

D. seminuda (Klotzsch) Maxon — 450 

Didymochlaena truncatula (Sw.) J. Smith — 160 

Diplazium caracasanum (Willd.) Kunze ex T. Moore— 
129; 129 

D. celtidifolium Kunze — 153; 497 

D. centripetale (Baker) Maxon, vel aff. — 152 

D. cristatum (Desr.) Alston—128; 1 

D. diplazioides (Klotzsch & Karsten ex Klotzsch) Al- 
ston— 146; 1 

D. expansum Willd.— 

D. flavescens (Mett.) Christ — 498 

D. hians Kunze ex Klotzsch— 147; 148 

D. mac ope Desv. — 

D. iie Kunz 


151 
m Klo tesch— 435; a 436 
Ur 


Diplopterygium bancroftii (Hoo . R. Smith—45 
UM pedata (L.) Fée var. jb (Willd.) Hick- 


D. punit (Raddi) J. Smith — 366 
Dryopteris patula (Sw.) L. Underw. — 168; 169 
[in affine (Mart. & Cal.) T. Moore — 275a 
. ambiguum (Mett. ex Christ) eps 284 
. andicola (Fee) T. Moore— 293, p 
ellermannianum (Klotzsch) T. Moore —270 
. brachyneuron (Fée) J. Tg e 
. burchellii (Baker) C. Chr.— 


E 

E 

E 

E 

E 

E 

E 

E. cuspidatum (Willd.) T. Moore — 271 

E. deorsum (Karsten) Vareschi — 2698 

E. erinaceum (Fée) T. Moore— 264 

E. eximium (Mett.) Christ — 265; 281 

E. glabellum J. Smith — 391 

E. herminieri (Bory & Fée ex Fée) T. Moore — 428 
E. hieracioides Mickel— 268 

E. hoffmannii (Mett. ex Kuhn) Christ — 290 

E. huacssaro (Ruiz) Christ — 276 

E. lingua (C. Presl) E A Ea its 429 

E. lloense (Hook.) T. Moo 

E. macrophyllum (Klotzsch) ‘Christ —287 

E. meridense (Klotzsch) T. Moore — 269 

E iun 

E 
E 


. ornatum aey Christ— 


i 


E. pusillum (Mett. ex Kuhn) C. Chr.—362; 364, pt 
E. sellowianum (Klotzsch ex Kahn) C. Chr. — 288; 295; 
2956 


E. sporadolepis (Kunze ex Kuhn) C. Chr. — 291; 293, 
pt; 294; 296 

E. succubus Mickel— 2 

E. tectum Hum b. & E ex Willd.) T. Moore— 273; 


E. Jp e (Fée) T. Moore— 272; 295a 

E. tovarense (Moritz ex Kuhn) T. Moore— 292 
E. variolatum Mickel— 

E. sp.— 277; 289; 430; 466 


348 


Annals of the 
Missouri Botanical Garden 


Eriosorus flexuosus (H.B.K.) Copel. var. flexuosus — 300 


. . hirtus — 301; 

E. hispidulus (Kunze) Vareschi var. hispidulus 359 
Grammitis alfarii (J. D. Smith) 08, p 
G. apiculata (Kunze ex Klotz ub. iK 218 

G. asplenifolia (L.) Proctor — 2138; 351 

G. chrysleri (Copel.) Proctor — 213; 467 

G. concinna A. R. Smith — 212 

G. cultrata (Willd.) Proctor — 210 

G. fendleri (Copel.) Lellinger — 215 


longisetulosa (Copel.) C. Morton — 216 
moniliformis (Lagasca ex Sw.) Kaulf. —406 
phlegmaria (J. Smith) Proctor — 207 
pilosissima (Mart. & Gal.) C. Morton — 338 
senilis (Fée) C. Morton — 208, pt; 209 
cf. senilis (Fée) C. Morton — 348 
aff. subtilis (Kunze ex em C. Morton — 349 
. taxifolia (L.) Proctor — 
. trichomanoides (Sw.) Ga. 211, pt 
truncicola (Klotzsch) C. Morton— 211, pt 
xiphopteroides (Liebm.) A. R. Smith—219 
. sp.— 2118; 350; 465 
Hemidictyum px i ) C. Presl— 167 
Hemionitis palma — 
H. rufa (L.) Sw. 309 
Histiopteris incisa (Thunb.) J. Smith—93 
Hymenophyllum (Mecodium) apiculatum Mett. ex 
Kuhn— 32; ad 32; 
H. is dium) asplenioides Sw.— 36 


AAAHNAHAAAAHAHAAHA 
= 


ton— 
H. (Mecodium) fendlerianum Sturm in Martius—31, 
t; 
H. (Sphaerocionium) fragile (Hedwig) C. Morton— 38 
H. D As ic did Ki oponen (Sw.) Sw.— —-2f 27 B 
irs 


—30; Do 
Sphaerocionium) trichophyllum 'H. B.K.— 
Hymenophyllum) tunbrigense (L.) E ^s "cn 29 
Mecodium) undulatum (Sw.) 


(Sphaeroc 


Mu NR) un (L. Sw. i bs aff cL pt 
(Sphaerocioniu ios Sturm — 39 
(Sphaerocionium) lanatum ~ 328; ad Im 387 
(Sphaerocionium) micro iiid 

(Mecodium) polyanthos (Sw.) Sw 

( 

( 

( 


Sw. — 

) sp., a pt 

Hypolepis bogotensis Karsten —64, pt; 648, pt 

H. parallelogramma (Kunze) C. Presl—66; ad 66 

H. — Hieron. — 431 

osa Karsten— 64, pt; 648, pt 

Lastreopsis e (Sw.) Tindale subsp. divergens (Willd.) 
Tindale — 

L. exculta ar ) Tindale— 330 

Lindsaea klotzschiana Moritz in Ettingshausen — 63a; 
638; 379; 380, p 

L. lancea (L.) Beddome var. lancea — 378, pt 

L. stricta t ) Dryander var. stricta —377; 378, pt; 


RS EUER E balas 


» Pt; 
ee fendleri D. Eaton — 335 
Lonchitis hirsuta L.— 100 


Lophosoria quadripinnata (J. Gmelin) C. Chr. — 48; 487? 
Loxoscaphe theciferum (H.B.K.) T. Moore—62 


Lycopodium acerosum Sw.—317 

. alopecuroides L.— 315 

. cernuum L.— 318; ae 3998 
48 


PEPER 


. thyoides Humb. | 3 Bonpl. ex Willd. —313; 400 
L. sp.— 398 

Lygodium venustu um Sw. 

Marattia laevis J. E. Smith— 

Microgramma lycopodioides L ) MS —420 

M. reptans (Cav.) A. mith — 

M. tecta (Kaulf.) Alston— 419 

M. vacciniifolia (Langsd. & Fischor) Copel. — 248 
Nephrolepis occidentalis K 


Niphidium crassifolium (L.) ) Lelnger—223 
N. mortonianum Lellinger 
Olfersia cervina (L.) Md 

34 


hioglossum nudicaule L. f. —342 


Paesia glandulosa (Sw. ) Kuhn— 94 

Pellaea ovata (Desv eath. — 89 

Phanerophlebia juglandifolia (Humb. & Bonpl. ex Willd.) 
J. Smith — 233 

Aroa calomelanos (L.) Link — 298 

P. chrysoconia (Desv.) Maxon ex Domin — 357? 

P. tartarea (Cav.) Maxon— 29 

P. trifoliata (L.) R. Tryon — 297 

Plagiogyria semicordata (C. Presl) Christ — 325 

Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. — 249; 

0 


25 
P. nouus n ) Hook. & Grev. — 421; 464 
x Pleopod fickel & Beitel — 


51 
Polybotrya canaliculata Klotzsch — 262 
P. serratifolia (Fée) Klotzsch — 261 
Polypodium (Phlebodium) aureum L.—417 
bolivari Sota — 34 
P. buchtienii Christ & Rosenstock — 
P. camptophyllarium Fée var. demam (Hieron.) A. 
ans— 221 
P. consimile Mett. var. consimile — 220; 2208 
A. M. Evans— 221, pt 
B 


P. eurybasis C. pue t; 2218 
is fraxinifolium Jacq. — 234; 235; 236; 488 
P. aff. ea olium T — 237; 
P. funckii Mett. — 
o fuscopetiolatum A. R. Smith—242; 341 
P. hygrometricum Splitg. —4 
P. lasiopus Klotzsch— 244; 418 


P. moritzianum Link — 214 
P. plumula Humb. & Bonpl. ex Willd.— 
P. DOS pocite (L.) Watt var. bor P Baker) 
Weath. 
P. (Phlebodium) pseudoaureum Cav.— 241 
remotum Desv. — 245 


P. sessilifolium Desv. — 240; 413 

P. sororium Humb. & Bonpl. ex Willd. — 422 
P. steirolepis C. Chr. —247 

P. thyssanolepis A. Braun ex Klotzsch — 252 


Volume 76, Number 1 
1989 


Smith & Todzia 
Fendler's Venezuelan 
Fern Collections 


349 


HB triseriale Sw. — 352; 415; 471 
— 254 


Polystichum muricatum (L. ) Fée— 172; 174 
P. platyphyllum (Willd.) C. Presl—175 

P. sp., Es polyphyllum (C. Presl) C. Presl— 171 
E. sp.—173 

Pter 2d. arachnoideum (Kaulf.) Maxon — 104 
Pteris altissima Poiret in Lam.— 


Rumohra adiantiformis (G. Forster) Ching — 499 
Saccoloma domingense (Sprengel) C. Chr. —61; 618 
376 


unze) Mett 
Salpichlaena a (Kaulf.) 1. pun 117, pt 
Schizaea elegans (Vahl) Sw. — 


poeppigi 
Selaginella cavifolia A. Bra m saa 2 323; 494 
E cladorrhizans A. Braun—3 


S. cruciformis Alston ex Crabbe & er. 487 
S. diffusa (C. Presl) Spring — 397 
S. flabellata (L.) Spring var. latifrons A. Braun— 493 
S. flagellata Spring— 324, pt; 40 
S. haematodes (Kunze) Spring in Martius — 461 
S. pallescens (C. Presl) Spring — 321; 3218 
S. porphyrospora A. Braun— 402 
S. viticulosa Klotzsch — 258, pt; 322, pt 

sp.— 486 

Sticherus bifidus (Willd.) Ching — 42 
S. oritz ex Reichardt) > aiibi 


Stigmatopteris daa Pernah) C. Chr.— 198 
Tectaria ibus rv i. .) L. Underw.— 164; 1648 
Cav 6 


T. incisa av : 


a Cav 
T. Aronet eq ) n n var. plantaginea —412 
x: Mae ai Cav.— 165 additional 
lypteris ncm) arborescens (Humb. & Bonpl. 


illd.) C. M 
T. Aioria) e dioides (Willd.) R. Tryon— 306; 


T. (Goniopteris) asterothrix (Fée) Proctor — 201 

T. (Amauropelta) balbisii (Sprengel) Ching —438 

T. (Amauropelta) cheilanthoides (Kunze) Proctor — 187, pt 
T. Foro concinna (Willd.) Ching— 179, pt; 


9, pt 
T. Lou malta) concinna (Willd.) Ching, vel aff. — 176 
Cyclosorus) conspersa (Schrader) A. R. Smith — 473, pt 
T. epic ad decussata (L.) Proctor var. decussata — 
373 


T. (Amauropelta) deflexa (C. Presl) R. Tryon—186 
T. (Amauropelta) diplazioides (Moritz ex Mett.) Ching — 


T. (Meniscium) ensiformis (C. Chr.) R. Tryon— 232, p 
, (Steiropteris) fendleri (D. Eaton) Reed — 372 
T. (Goniopteris) gemmulifera (Hieron.) A. R. Smith— 
193; 193y 


T. (Cyclosorus) grandis A. R. Smith var. aequatorialis 
. Chr. . Smith— 1 
T. (Cyelosorus) hispidula (Decne.) Reed— 191; 441; 
t; 

T. sb leprieurii E ) R. Tryon var. costalis 
(Baker) A. R. Smith— 7 

T. (Goniopteris) nephrodioides (Klotzsch) Proctor — 445 

T. (Amauropelta) corpo ames & Bonpl. ex Willd.) 
Ching — 179, pt; 181, 

T. (Amauropelta) o Bu e Vahi) Ching— 177 

T. (Amauropelta) opposita (Vahl) Ching, vel aff. — 182 

T. (Amauropelta) aii Ae (Kunze ex Mett.) Ching 
var. pachyrhachis — 187, 472 

T. E patens (Sw.) Small var. patens — 189; 


T. tor did patens ye ) Small var. scabriuscula (C. 
Presl) A. R. Smith— 


T. Oone ST je (Klotzsch) A. R. Smith — 446 
e adc pennata (Poiret in Lam.) C. Morton— 
200 


T. 

T. (Amauropelta) pilosohispida (Hook.) Alston— 158 
T. (Amauropelta) ria d (Mett.) R. Tryon— 178; 180; 
T. 
: 


(Goniopteris) ken n (Bory) Proctor — 199 
(Steiropteris) praetervisa (Kuhn) A. R. Smith —371 
a eek opelta) proctorii A. R. Smith & Lellinger — 


» pt 
T. Vue. pteroidea (Klotzsch) R. Tryon— 206 
T. MEE ii rudis (Kunze) Proctor —185, pt; ad 


T. qum. rupestris (Klotzsch) Reed— 30 

T. (Meniscium) salzmannii (Fee) C. De, pu pt 
T. (Amauropelta) scalaris (Christ) a e pt 

T. (Goniopteris) straminea (Baker) R 

T. (Goniopteris) tetragona (Sw.) em TM 

T. sp., aff. gracilis (Heward) Proctor — 439 
Trichomanes (Lacosteopsis) capillaceum L.— 22; 395 


T. (Achomanes) crispum L.— 

T. (Lacosteopsis) debile Bosch — 459 

T. (Lacosteopsis) eat vos H.B. T p pt 17; 18 

T. (Microgonium) ekmanii W. Boe 

T. (Didymoglossum) lineolatum (Bosch) Eu —454 

T. (Lecanium) membranaceum 

T. (Didymoglossum) odia (Fourn.) W. Boer — 25; ad 25 

T. (Neurophyllum) pinnatum Hedwig — 390 

T. (Acarpacrium) polypodioides L. — 329 

T. (Didymoglossum) punctatum Poiret subsp. sphe- 
noides (Ku Boer — 

T. (Didymoglossum) ME E — 24; ad 24 

T. (Lacosteopsis) radicans — 

T. (Didymoglossum) reptans ey 23 

T. (Pachychaetum) rigidum Sw.— 16, pt 

T. (Achomanes) robustum Fourn.— 26 

T. (Lacosteopsis) rupestre (Raddi) Bosch — 21; 456 

T. (Acarpacrium) trigonum Desv. 

T. (Achomanes) sp., undescribed? — 3888; 389 


Vittaria costata Kunze —424 

V. gardneriana Fée— 2598 

V. graminifolia Kaulf. —258, pt 
V. moritziana Mett. — 259, pt 
V. remota Fee— 260 


V. stipitata Kunze — 259, pt 


NOTES 


A NEW ECHEANDIA 
(LILIACEAE) FROM 
VENEZUELAN GUAYANA 


Echeandia bolivarensis Cruden, sp. nov. TYPE: 
Venezuela. Bolivar: Igneous forested slopes, 
Serrania de Pijiguao, 160 km SW of Caicara 
del Orinoco, 6%35'N, 66%45'W, 100-125 m, 
12 Sep. 1985, Steyermark, Holst & Manara 
131761 (holotype, MO, on two sheets, 
3472626, 3472627; isotype, UC). 


Regio penaria radicum ab cormo 4-5 cm, 4-5 cm 

longa. Fotis basalia 6-10, anguste i margi- 

rulatis, 45-60 cm longa, 11-18 mm 

lata. iin cabe 4 4-5 ramosus. Folia caulina 4-5 

longiattenuata. Tepala flava. Filamenta squamata, linear- 
ia. Antherae librae. Capsula late oblonga 


Storage region of the roots 4-5 cm from the 
corm, 4—5 cm long. Basal leaves 6-10, ascending, 
narrowly lanceolate above the base, bases persis- 
tent as a fibrous collar, minutely denticulate-ser- 
rulate, usually 45-60 cm long and 11-18 mm 
wide. Scape glabrous, 98-118 cm high, 5-branched. 


ANN. Missouni Bor. Garb. 76: 350. 1989. 


Cauline leaves 4-5, long-attenuate. Tepals yellow, 
ca. 10 mm long. Filaments minutely to noticeably 
scaled, linear, 6.5 mm long. Anthers free, 2.4-3 
mm long, appearing to dehisce laterally. Ovary 2 
mm long. capsule broadly oblong, 10-11 mm long, 
6.5-7.5 mm wide. Seeds black, colliculose, 2.5- 


3 mm across. 


Known only from the type locality, this species 
is easily distinguished from other yellow-flowered 
South American Echeandia species by the com- 
bination of scaled filaments, many branches, and 
the storage organs developing at some distance 
from the corm. It is unusual for yellow-flowered 
species to occur at such low elevations. 


— Robert William Cruden, Department of Bota- 
ny, University of lowa, lowa City, lowa 52242, 
U.S.A 


SOLANUM TOLIAR AEA, 
A NEW SPECIES FROM 
MADAGASCAR 


Solanum toliaraea D'Arcy & Rakotozafy, sp. 
nov. TYPE: Madagascar: D'Arcy & Rakoto- 
zafy 15460 (holotype, MO; isotypes, K, P, 
TAN) 


rutex armatus spinis brevibus validis rectis, foliis ro- 
tundis parvis stellato-tomentosis; flores solitari v-geminati, 
calyce inermi sed in statu fructu accrescenti, echiniformi, 
acinum tegenti 


Slender, branched shrub or tree to 3 m tall; 
twigs slender, at first appressed tomentose with 
reddish hairs, soon puberulent with reduced buff- 
colored trichomes and deeply furrowed, armed with 
short, stout-based, puberulent 
spines. Leaves mostly paired in fasciculate short- 
shoots, entire, rotund, apically rounded, ed 
rounded, truncate or cordate, mostly 8-20 m 
long, thick, felty with dense short-stalked, elate 
hairs, the costa evident, the minor venation o 


nearly straight, 


scure; peduncles canaliculate adaxially but alten 
appearing terete because of the dense tomentum, 
mostly 5-10 mm long, with occasional small straight 
spines. Inflorescences solitary flowers or paired fas- 
cicles from the leafy short-shoots; pedicels resem- 
bling the petioles but somewhat longer. Flowers 
with calyx ca. 4 mm long, lobed about 44 way 
down, the lobes convex, obtuse, tomentose, un- 
armed; corolla pale purple, mauve, or white, ca. 
10 mm long, lobed 44 way down, tomentose outside 
except on the membranaceous inter-lobe fringing 
areas, glabrous within, the nervature drying con- 
spicuous within; stamens 5, equal, the anthers yel- 
low, slender, 5 mm long, opening by minute ter- 

minal pores; ovary glabrous, drying somewhat lobed, 
the style slender, exserted slightly beyond the sta- 
mens, with a few short, gland-tipped simple hairs 
near the base. Berry green, juicy, 4-lobed, ca. 5 
mm diam., entirely but loosely enveloped in the 
calyx, the spiny, accrescent calyx echinoid, subglo- 


ANN. 


seeds (Descoings 2375) 
compressed, dark, 2.5 mm long 


bose, ca. 2 cm diam.; 


At time of flowering the calyx of this species is 
devoid of spines, in contrast to its copiously spiny, 
echinoid condition when in fruit. 

Collections of this species were first confused 
with Solanum dubium Fresen of Ethiopia, which 
has similar spiny accrescent calyces, but that species 
is herbaceous and has much larger, lobed leaves. 
The species described here differs from the other 
two arborescent species of southwestern Madagas- 
car in its accrescent, spiny calyx and in its rotund, 
concolorous leaves. Solanum toliaraea is sympat- 
ric, at least in part, with other arborescent sola- 
nums, S. bummeliifolium Dunal and S. heineanum 
D'Arcy & Keating, which differ in usually spineless 
calyces and porrect, scutellate hairs. 

Solanum toliaraea was seen as scattered indi- 
viduals in didieriad woodlands of southwestern 
Madagascar. Most collections were made from 
plants growing on sand near the coast just north 
of the major city of Tulear and hence just north 
of the Tropic of Capricorn. 

The name toliaraea is derived from the Mala- 
gasy spelling for the city of Tulear. 


e seen. ll aa TULEAR (Toliary): bush 

a Didie sur dunes, embouchure de Fiherenana, Be- 
landa, Nox 1956, Bokir 10472 (MO, P); scrub forest 
on pure ei sand, 8 km N of Tulear on road to 
Morombe, 5 1975, I 30778 (MO); road to 5 
km N of Belalanda, 17 May 1983, D'Arcy & Rakotozafy 
15460 (MO, K, P, TAN); Sable, Manombo, région de 
Tuléar, Decary 16214, 18714 (P); route de Manombo 
sur sable, 7 Jan. 1957, Descoings 2375 (MO); bush á 
Didierea sur dunes, embouchure de Fiherenana, Bela- 
landa, May 1960, Keraudren 690 (P); bush à Euphorbiés, 
route de Monombo, prés de la cóte, environs de Tuléar, 
3 Nov. 1960, Léandri & Jean de Dieu 3627, 3635, 
3804 (P); thorn scrub of Euphorbia and Didierea on 


Missouni Bor. GARD. 76: 351-352. 1989. 


352 Annals of the 
Missouri Botanical Garden 


t 
o 
o 


FIGURE 1. Solanum toliaraea. — A. Habit. — B. Flower. —C. Fruit. — D. Stamen. — E. Seed. — F. Indument. — 
G. Trichome from leaf. (E after Descoings 2375, other fe after Diy & Hakot 15460 [MO].) 


sand dunes behind littoral, Route Nationale no. 9, 15km — — William G. D'Arcy, Missouri Botanical Gar- 
N of Tulear, 12 Nov. 1978, Lorence 1940 (MO). den, P.O. Box 299, St. Louis, Missouri 63166, 
U.S.A., and Armand Rakotozafy, Centre National 
de Recherches de Tsimbazaza, B.P. 4096, An- 
tananarivo 101, Democratic Republic of Mad- 


This study was supported by a grant from the 
National Geographic Society. Logistical assistance 
in the field was provided by the Missouri Botanical 
Garden's Madagascar Research Program with sup-  “845Car. 
port from the W. Alton Jones Foundation. 


NOTE CONCERNING 
FESTUCA HENRIQUESII 
(GRAMINEAE) IN 
NORTHERN PORTUGAL 


The species Festuca henriquesii Hackel (Gra- 
mineae), first described in 1883, is endemic to 
Portugal. In various floras and subsequent publi- 
cations, namely Coutinho (1939), Sampaio (1946), 
Litardiére (1952), and Tutin et al. (1980), the 
location of this species has been given as restricted 
solely to the Serra da Estrela, a mountain range 
in northeastern Portugal. Little information is avail- 
able concerning its ecology, except that specimens 
have generally been collected from high places in 
this mountain range. 

Recently, however, I recorded this species from 
a grassland area in the Serra do Gerés, in northern 
Portugal (Hale, in press). I had believed that this 
was the first record for the species outside of the 
Serra da Estrela range. The location, in grazed 
grassland within an area generally of ericaceous 
shrubland, is known as ““Lagoas” because of two 
small nearby lakes. The plants were growing on 
an acid soil over granitic parent material at an 
altitude of 1,150 m (grid reference X326 071; 
Carta Militar 31). 

Further investigation revealed, however, that 
this record for the species in the Serra do Gerés 
was not unique. Besides two records from the Serra 
da Estrela, three specimens of F. henriquesii exist 
in the Herbarium of the Institute of Botany “Dr. 
Gonçalo Sampaio” in Porto, collected from the 
Serra do Gerés and nearby areas (Table 1). Despite 
the existence of these specimens, however, this 
species appears not to have been previously re- 
ported from these mountain ranges. 

It is clear from these records that Festuca hen- 
riquesii is not restricted to the Serra da Estrela; 
the range includes the serras of Gerés, Peneda, 
and Barroso. The species might be a glacial relict 
rined to a few mountain ranges and not noticed 

ide of the Serra da Estrela; alternatively, 
its dictation may be expanding. In view of the 


apparently discontinuous distribution, these other 
mountain ranges being nearly 200 km from that 


TABLE 1. F. henriquesii records outside Estrela. 
Date of 
Location Habitat Collection 
Lamas do Homem, 
Gerés Dry ground 6/7/1948 
Barca do Homem, 
Gavieira, Peneda Grassland -/1/1961 
S. Fins, Montalegre, 
arroso Roadside 11/8/1978 
Lagoas, Gerés Grassland 6/7/1984 


of Estrela, it is quite possible that the species might 
turn up in other intervening mountain ranges in 
the north of Portugal. 

I thank the staff of the Herbarium of the In- 
stitute of Botany “Dr. Gonçalo Sampaio,” Porto, 
for confirming the identification of the “Lagoas” 
specimen and for access to the previous records 
of the species. 


LITERATURE CITED 


COUTINHO, A. X. P. 1939. Flora de Portugal (Plantas 
asculares), 2nd edition. Bertrand (Irmaos) Lda., 


isbon. 

Hare, W. H. G. Variation in pue les composition within 
and between some mountain grasslands in the Parque 
Nacional da Peneda- tdi Agronomia lusitana 43 
(in press). 

pore. R. 1952. Contribution à l'étude des 
Festuca du odia Agronomia lusitana 14: 31 
51. 


SAMPAIO, C. 
prensa Moderna Lda., Por 

Turin, T. G., V. H. Heywoop, E A. Burces, D. H 
VALENTINE, S. M. WALTERS & D. A. WEBB Won 
1980. Flora ad oe: Volume V. Cambridge Univ. 
Press, London 


1946. Flora REA 2nd edition. Im- 


—W. H. C. Hale, Instituto de Botanica, Rua do 
Campo Alegre 1191, 4100 Porto, Portugal. Pres- 
ent address: School of Environmental Science, 


University of Bradford, Bradford BD? 1DP, U.K. 


ANN. MISSOURI Bor. GARD. 76: 353. 1989. 


PASSIFLORA CITRINA, 
A NEW SPECIES IN 
SECTION XEROGONA 
(PASSIFLORACEAE), 
FROM MESOAMERICA 


The center of diversity for Passiflora L. subg. 
Plectostemma Masters sect. Xerogona (Raf.) Killip 
is in northern Central America, where at least six 
species occur in Guatemala alone. The section is 
notable in the genus Passiflora for its unusual 
capsular fruit. Killip (1922, 1924, 1927) described 
five species of this section from Central America 
and southern Mexico that had been confused with 
P. rubra L. or P. capsularis L. Newly described 
here is the yellow-flowered P. citrina, yet another 
member of this section that has been hidden under 
misapplication of the name P. capsularis. 


Passiflora citrina MacDougal, sp. nov. TYPE: 
Honduras. Comayagua: El Carrizal, 14 km 
NW of Siguatepec, 900 m, 27 June 1971, 
Molina & Molina 26057 (holotype, F; iso- 
types, EAP, NY, US). Figure 1. 


Passiflora pubescens scandens; caulis sub5-angulatus; 
petioli eglandulosi; folia bilobata vel trilobata, lobis laterali- 
bus acuminatis vel rotundis, lobo centrali obtuso vel ob- 
soleto, marginibus integris, angula inter lobos laterales 
(30- )35- 55(-70); bracteae nullae vel raro floribus brac- 
ra, 2-4 mm 


et ¿uo adnata; androgynophorum 21-28 mm lon- 
ium dense p fructus fusiformis vel 
bovoideus, conspicue 6- ina 6- 7(8) sulcata. 


9s 
B 


carinathus: sem 


Small vine, pubescent to hirsutellous throughout 
with 0.10-1.0(-1.2) mm + erect, slightly an- 
trorsely inclined or bent trichomes. Stems subpen- 
tangular, striate, densely pubescent with 0.2-0.8- 
mm bent trichomes. Stipules 4-7(-11) x 0.5-0.9 
mm, linear-triangular, falcate, the apices not ne- 
crescent. Petioles 0.3-1.2(-2.0) cm long, eglan- 
dular. Laminas (2-)3-6.5(-9) x (1.5-)3-7(-9.5) 
cm at fertile nodes, widely obovate (obtriangular 
to widely oblong) in general outline, rounded to 
truncate (very shallowly cordate) at the base, en- 
tire, adaxially sometimes variegated along the main 
veins of the lateral lobes, evenly pubescent or often 
the trichomes in 2 distinct size classes and then 


the leaf scabrous and lightly hirsutellous, abaxially 
densely pubescent, 2(-3)-lobed 0.07—0.4(—0.6) the 
distance to the base, the lateral lobes acuminate 
to rounded, the central lobe (acute) obtuse to ob- 
solete, the angle between the lateral lobes (30-) 
35-55(-70P, the ratio of lateral to central lobe 
lengths (1.0-)1.2-2.0(-2.5), the ratio of laminar 
width to length (0.50-)0.6-0.9(-1.1); juvenile 
leaves variegated, deeply 2-lobed, the lobes acu- 
minate, the angle between the lobes 80-100? or 
more; laminar nectaries absent. Posture of shoot 
apex + straight. Prophyll of vegetative ramifying 
ud 1, narrowly lanceolate-triangular. Peduncles 
(solitary) geminate at the nodes, (0.3-)0.7- 
3.2(-4.3) cm long, uniflorous, ebracteate or rarely 
with one bract near the apex, 2-4 x 0.2-0.4 mm, 
linear to linear-triangular. Flowers yellow or bright 
yellow (to greenish yellow), sometimes drying with 
reddish brown speckles, inodorous, cylindrical in 
bud, the ventricose hypanthium 5-9 mm diam., 
sometimes with some of the trichomes borne on 
slightly raised bases; floral stipe 1.5-4 mm long; 
sepals 20-28(-34) x 2.5-5 mm, narrowly oblong- 
lanceolate or slightly oblanceolate, basally connate 
mm, ecorniculate, strongly 3-veined, the veins 
raised and hirsutellous; petals 19-28 x 1.7-2.6 
mm, narrowly oblong-lanceolate, basally adnate to 
the sepals; coronal filaments in 1 series, 9-14 mm 
long, erect, variably connate much of their length 
and adnate to the perianth, free distally 3-6 mm, 
pale yellow with yellow tips, not banded; operculum 
1.0-2.1 mm long, membranous, only barely plicate 
near the erose apex; nectary 5(10)-ventricose, an 
additional annulus absent; limen (disk) ca. 2 mm 
diam., closely surrounding the base of androgyn- 
ophore; staminal filaments connate 21-28 mm along 
the androgynophore, the free portions 4.5-6.0 mm 
n anthers 4.4-6.0 mm long; ovary 2.8- 
4.2(-5.0) x 1.5-2.3 mm, obovoid-e 
ly pubescent, the trichomes mostly 0. 2-0.5 mm; 
styles 5.0-6.8 mm total length; stigmas ca. 1.3 
mm diam. Fruit (2.5-)3-4.3 x 1.2-2.3 cm, fu- 
siform-ellipsoid or slightly obovoid, basally conical 


ANN. MISSOURI Bor. GARD. 76: 354-356. 1989. 


Volume 76, Number 1 
1989 


Notes 


or slightly attenuate, apically conical, acutely 
6-carinate with thin 1-2-mm-high ridges, pubes- 
cent, reddish, dehiscent?, the stipe 1-8 mm long; 
arils unknown; nearly mature seeds 3.5-4 x 2.2- 
2.4 mm, slightly obcampylotropous with the cha- 
lazal beak strongly inclined toward the raphe, trans- 
versely sulcate with 6—7(-8) sulcae, the ridges 
continuous, smooth, and rather flat. 


Habitat and distribution. 
is endemic to the pinehills country of central to 


Passiflora citrina 


western Honduras and adjacent eastern Guate- 
mala. There it has been collected in moist (often 
open) pinewoods, Pinus-Liquidambar forest, or 
edges of moist mixed forest; it usually is climbing 
over grasses, tall herbs, and small shrubs in thick- 
ets. 

Vernacular names. The name “moco” is re- 
corded on an herbarium sheet from Chiquimula, 
Guatemala. ““Calzoncillo,”” 
for several of the bilobed passifloras in Mesoameri- 
ca, is recorded from Intibucá, Honduras. 


a vernacular name used 


Misapplied names. Passiflora capsularis 
auct., non L.: Killip (1936, 1938), in part; Stand- 
ley & Williams (1961), in part. 


The specific epithet refers to the color of the 
flowers. 


Additional specimens exam GUATEMALA 

( P IMULA: La Cumbre, old road to Cet Ran l, 500 

28 Sep. 1971 (fl), Molina * Tr m 26821 (F); 
Volcán Quezaltepeque, 3-4 m of Quezaltepeque, 
1,500-2,000 m, 8 Nov. 1939 a Ted 31508 
(F). HoNDURAS. COMAYAGUA: Temagua, 2,000 ft., 5 Aug. 
1933 (fl), Edwards P-640 (F), Trincheras, 20 km N of 
Siguatepeque, 4,500 ft., 29 July-10 Aug. 1951 (fl, fr), 
Howard et al. 637 (MICH, NY); Trincheras on = road, 
ca. 14?39'N, 87%55'W, 1,050-1,100 m, 1988, 
MacDougal et al. 3048 (BM, MEXU, MO TEFH) near 
Cerro rp: km post 127.5 on Ed between Sig- 
uatepeque ago Yojoa, 9 Nov. 1988, 
MacDougal » al. 3469 (BM, MEXU, MO, US, TEFH); 
Barranco de Trincheras, 1,300 m, 29 Aug. 1955 (fl), 
Molina 5826 Aa, Baran x Tuned s, 3 km a Mon- 
tanuela, 1, m, 28 Mar. 1964 (fl), Molina 13637 (F); 
8 mi. W of N igu E e, hio m, 7 Sep. 1975 (fl, fr), 
Molina & Molina 31167 (F, MO); 24 a S of Lake 
Yojoa along road to Siguatepeque, 1,400 m, 26 Mar. 
1976 (fl), Pilz & Pilz 1572 (MO); Barranco de Trinch- 
eras, 18 k of Siguatepeque, 1,200 m, 15 Apr. 1951 
(f), Williams & Molina 17991 (F, US). COPÁN: 5 km al 
S 


ar. 1973 (fl, fr), J. R. 


anquilla, cerca de Yashse, 1,500 m, 1 
Molina 6510 (F, US); El Duraznillo, Cordillera, Opalaca, 


ug 


C D 


FIGURE 1. Variation in pare of leaves of Passiflora 
a —A. Molina 26. Molina & Molina 

6. —C. Molina & Molina 26057.— D. Molina & 
[ien 31167. 


2,000 m, 24 May 1964 (fl), Molina & Molina 14086 
(F). ocorEPEQUE: 17 km NE of Nueva Ocotepeque, 13 
Aug. 1970 (fl), Harmon & Dwyer 3785 (MO); 41 km 
al NE de Nueva Ocotepeque, camino al San Pedro Sula, 
1,490 m, 12 June 1985 (fl), E. Martínez S. & O. Téllez 
12932 (DUKE, MEXU); El Moral on Cordillera Mer- 
endón, 1,600 m, 27 Aug. 1968 (fl), Molina 22263 (F, 
NY). 


Passiflora citrina may be referred to subg. 
Plectostemma sect. Xerogona because of the sub- 
pentangular stem, absence of petiolar or laminar 
nectaries, absence or near absence of floral bracts, 
elongate fruits with six "keels, and seeds with a 
transversely grooved sclerotesta having a chalazal 

eak that is sharply angled towards the raphe. 
Mature fruits are unknown; however, a fruit that 
is probably of full size with nearly mature seeds is 
found on the isotype at NY. On a different collec- 
tion, Howard et al. 637, the collector observed 
the fruits to be “red,” but no fruits are included 
with the specimens. It is unknown whether the fruit 


356 


Annals of the 
Missouri Botanical Garden 


of this species is six-valvately dehiscent, as is usual 
in the other members of sect. Xerogona. 

Passiflora citrina is remarkable for its elongate 
yellow flowers that are tubular in the basal half. 
The androgynophore is distinctively long and sets 
the species apart from all other members of sect. 
Xerogona except P. sanguinolenta Masters from 
southern Ecuador. Though very similar to P. ci- 
trina, P. sanguinolenta has nearly triangular stems, 
peduncles solitary at the nodes, pink to purplish 
red flowers that retain a diminutive second series 
of coronal filaments, and seeds with slightly ru- 
gulose ridges. Both species have slightly elongate 
flowers of similar morphology, flowers that have 
departed from the usual bowl-shaped hymenopter- 
an-adapted groundplan in sect. Xerogona (Lind- 
man, 1906) by reduction and fusion of the coronal 
filaments and partial connation and adnation of the 
perianth. In P. citrina the sepals are free from 
each other to within 2-6 mm of their bases, but 
they are submarginally adnate to the edges of the 
petals; the petals, in turn, are adnate to the connate 
portion of the coronal filaments. The net effect is 
a cylindrical structure formed by the adnation of 
the basal half of the perianth to the tubular and 
basally connate coronal filaments. The coronal fil- 
aments are free only in the distal 3-6 mm and are 
usually not visible on herbarium sheets without 
dissection. In dissected and living specimens, I could 
detect no trace of a second series of coronal fila- 
ments near the operculum. 

The derived floral morphology of P. sanguin- 
olenta is related to a shift in pollinators: the species 
is primarily hummingbird pollinated in its native 
habitat (C. Hofmann, pers. comm.). It is probable 
that P. citrina has also shifted away from polli- 
nation by hymenopterans. 

The butterfly Heliconius charitonius was ob- 


served using MacDougal et al. 3048 as a host 
lant. 

Floral bracts are fundamentally absent in P. 
citrina, but in Molina 6510 the peduncles occa- 
sionally bear a single bract at the apex. 

This species was first identified as undescribed 
while the author was a National Science Foundation 
Fellow at Duke University, and I gratefully ac- 
knowledge the financial support of the NSF, the 
assistance of the curators from the herbaria cited 
above, and Donald E. Stone for inspiration and 
guidance during my doctoral studies. The Jessie 
Smith Noyes Foundation generously provided post- 
doctoral support for research on Passifloraceae at 
the Missouri Botanical Garden. 


LITERATURE CITED 


KiLLiP, E. P. 1922. New passifloras from Mexico and 
entral America. J. Wash. Acad. Sci. 12: 255-265. 
1924. New species of deer from tropical 
America. J. Wash. Acad. Sci. 14: -116. 
= | New cesar il Ped South Amer- 
ica and Mexico. J. Wash. Acad. Sci. 17: 423-427. 
———, 1936. Passifloraceae of : Mayan region. 
XIII. Pp. 299-328 in Botany of the Maya Area 
Misc. papers I-XIII, Publ. inen Inst. Wash. 
461. 


The American Te of Passiflora- 


1938. 
ceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 1- 


13 
LINDMAN, C. A. M. 1906. Zur Kenntnis der Coro 
der einiger Passifloren. Pp. 55-79 in F. R. Kjellman 
ae rear Studier Wert 
STANDLEY, P. C. & L. O. WILLIAMS. 19 Passiflorace- 
ae. id 118- 146 in Flora of Pediat Fieldiana: 
Bot. 24(7). 


—John M. MacDougal, Missouri Botanical Gar- 
den, P.O. Box 299, St. Louis, Missouri 63166- 
0299, U.S.A 


A NEW SPECIES OF 
STRYPHNODENDRON 
(FABACEAE: MIMOSOIDEAE) 
FROM AMAZONIAN ECUADOR 


Stryphnodendron Mart. (Fabaceae: Mimoso- 

ideae) is a genus of 25 known species, primarily 
Brazilian in distribution. It is well represented in 
Amazonia by forest trees and in the Brazilian cer- 
rado region by shrubby species (Occhioni Martins, 
1974, 1981; Occhioni, in prep.). Recent collec- 
tions by the senior author and associates in eastern 
Ecuador, in the course of fieldwork for a floristic 
study of the trees of the region, have revealed a 
new species of Stryphnodendron at the western 
edge of the Amazon basin and in the adjacent 
foothills of the Andes. 
Stryphnodendron porcatum Neill & Occhioni 
Tropical Moist Forest, alluvial soil, 
floodplain near mouth of Rio Coca, secondary 
forest and pasture, 250 m, 00%25'S; 77°00'W, 
15 Sep. 1986 (fl, frt), David Neill & Walter 
Palacios 7359 (holotype, QCNE; isotypes, 
MO, RFA, QCA, QAME, AAU, NY, K, GB, 
US, about 10 additional isotypes to be dis- 
tributed). Figure 1. 


Str lend i (Aubl.) Benth. subsp. glan 
duloso Forero foliis | simile, e calyce et corollae bbs 
aureopubescentibus recedit; a speciebus ceteris Tr 
ornamento, hase natione in rule cristarum in fruct 


valvis differ 


Trees attaining 40 m with trunk to 60 cm diam. 
Outer bark grayish brown, lenticellate. Young 
branchlets, inflorescence axis, and petioles with 
minute dense brown scurfy tomentum. Leaves 15- 
16(-22) cm long; petiole terete, puberulent 3.5- 
5.0 cm, with a 1-mm verruciform gland about 2.5 
cm from its base; rachis villous 11-12(-18) cm, 
with rounded glands between the distal 5-8 pairs 
of pinnae. Pinnae 11-13 pairs, opposite or subop- 
posite, 5—6(-9) cm long, with small rounded glands 
between the pairs of leaflets, excepting the 2-3 
basal pairs. Leaflets 10-13-jugate (6- 7-jugate in 
the proximal 1-2 pinnae), alternate, elliptic, 6— 
10(-14) x 3-4(-5) mm, the apex retuse or slightly 
emarginate, the base unequal; venation inconspic- 


uous; upper surface glabrous, lustrous; lower sur- 
face sparsely and evenly villous, the midrib and 
margins more densely so, with a dense tuft of villi 
near the base of the blade; margins slightly revolute 
and thickened; petiolules 0.5 mm, villous. Inflo- 
rescence spicate, cylindric, axillary, solitary or 2 
together; peduncle terete, brown-scurfy, 1-1.5 cm 
long; rachis densely pubescent, 5-8 cm long. Flow- 
ers hermaphroditic, or sometimes all staminate, 
3.5-4 mm long, bracts conchiform, deciduous. Ca- 
lyx cupular, 1 mm with 5 deltoid lobes, densely 
golden pubescent. Corolla narrowly campanulate, 
2-2.5 mm; lobes acute, 1 mm; basal half of corolla 
glabrous or with a few scattered golden hairs, the 
lobes densely golden pubescent externally. Stamens 
10, free; filaments 3-4 mm; anthers elongate, 0.5 
mm, with a stipitate deciduous apical gland. Ovary 
1.5 mm, short-stipitate, densely white-pilose; style 
2 mn, filiform, glabrous or with a few hairs. Le- 
gume subligneous, 10-12(-14) x 2-2.5(-3) cm, 
slightly curved, dark brown with scurfy epidermis, 
the epicarp ornamented with a pattern of ridges 
aligned at an angle from the center of each valve 
trending proximally toward the edges, forming a 
repeated, irregular chevron-like pattern. Seeds 
transverse, ellipsoid-oblong, 8-10 x 5 mm, dark 
brown, surrounded by sweet gelatinous pulp. 


Additional specimens examined. ECUADOR. NAPO: 
Canón de Los Monos, m al N de Coca, 250 m 
00°20" S, 77°01'W, 5 Apr. 1985 pam Neill, Baker, Pa. 
FA) Estación 


1987 (flr), Neill, giu Cerón & Manning 7837 (MO, 
QA ME); Reserva Biológica Jatun Sacha, Rio Napo, 8 km 
rio abajo de Misahualli, 450 m, 01%08'S, 77*30'W, 2 
Oct. 1986 (fir.), Walter Palacios 1369 (MO, QAME, 
RFA). PASTAZA: 2 km al NE de Mera, 1,100 m, 01?27'S; 
78°06'W (sterile), Neill, Baker, Palacios & Zaruma 
6161 (MO, QAME). MORONA-SANTIAGO: Taisha, 500 m, 
02%23'S; 77%30'"W, Oct. 1975 (fl, frt), E. L. Little, Jr. 
et al. 690 (QAME, US); Sucúa, 1,200-1,500 m, 02°34’S, 
78°10'W, 7-9 Oct. 1975 (fir, frt), E. L. Little, Jr. et 
al. 670 (QAME, US). 


ANN. MISSOURI Bor. GARD. 76: 357-359. 1989. 


358 Annals of the 
Missouri Botanical Garden 


Ady & 


TRANS 
AVS 
CANO , 
CANS y 
CANCINO Uy 
VO ER NAM) 
AO eee n GY 
CONES cO CO 
DA A] £ V as 


x 
DI 
q 
$C 


"p, 


% 


sss 
hh 


í WA ME 


NA SE 


eS 
BT 
MARS 

Cp e 

SIS 

CL 


S 


jx 
GAZA 
“J 


= YN 
ia POS 
SOS ec 
0227 
22) 

>> 

NA 


Lae) 

AN 
CURG, 
LA» 

- N hy - 
AZ, 
AL 


j^ 


J^ 


Gh, 
SR 
ME 
222 25 
ANN 

inf 

202 


f 
E 
-- 


A 
*-3 
b 
LA y 
AS 


Y 
/ » 
amar 
x 


a. - 
À 


ss 

SA 

=z 

Net 
> 
Ni 
y 

Wo 

\ TN 


TN Y . fs y LA ys WZ", Lp 
OR e 
b PY. APR YY EN 
E OVV DO A ez MES 
DEW e tyre © Ss ee VW De AY 

; SAN 
S O 


ver 
i Nef, 

UY: 
Ñ Yn ñ 
Y 


^N 


FIGURE 1. Stryphnodendron porcatum. — A. Habit. — B. Pair of leaflets, showing upper (right) and lower (left) 
surfaces, and small glands between each pair of leaflets. — C. Pistil. — D. Flower bud. — E. Open flower. — F. Mature 
fruits, showing pattern of ridges on epicarp. 


Volume 76, Number 1 
1989 


Notes 359 


Stryphnodendron porcatum is probably most 
closely allied with S. guianense (Aubl.) Benth. subsp. 
glandulosum Forero, known from Peru, Bolivia, 
and Amazonia in western Brazil, which has similar 
pubescence of foliage, including the tuft of hairs 
near the base of the blade, and densely pilose ovary. 
In S. guianense subsp. glandulosum, however, 
the calyx and corolla are short-puberulent exter- 
nally, whereas in 5. porcatum the calyx and corolla 
lobes are densely pubescent with longer, golden 
hairs, while the corolla tube is nearly glabrous with 
scattered golden hairs. The typical subspecies of 
S. guianense, which ranges widely in Amazonia 
and the Guianas, lacks the tuft of hairs at the base 
of the leaflet blades, and all floral parts are glabrous. 

e most distinctive feature of 5. porcatum, 
which readily separates it from all other species in 
the genus, is the striking ornamentation of the 
legume, an irregular chevron-like pattern of ridges 
on the surface of the valves. The specific epithet 
refers to this sculptured surface of the fruits (Latin 

orca — *'ridge between two furrows made in plow- 
ing”; Stearn, 1966). 

Some individuals of S. porcatum are evidently 
female-sterile; the flowers of Palacios 1369 com- 
pletely lack pistils. 

To our knowledge, this is the first published 
report of the genus Stryphnodendron for Ecuador. 
No other species are known to occur sympatrically 
with S. porcatum in the Amazon region of the 
country. 

Stryphnodendron porcatum occurs on rich silty 
alluvial soils of river floodplains, and in Amazonian 
Ecuador, on upland Oxisol hills up to 1,500 m on 
the lower slopes of the Andes. In primary forest it 
attains a height of 40 m with a diameter at breast 
height of 70 cm. In the region around Coca, Napo 
Province, Ecuador, 5. porcatum is also frequent 


in secondary forests and is often left in pastures 
after the surrounding forest is cut. 

The sweet pulp surrounding the seeds is similar 
to that of Prosopis and Gleditsia. Several trees at 
the INIAP-Payamino agricultural experiment sta- 
tion near Coca stand in experimental pastures es- 
tablished for raising tropical hair sheep. The sheep 
eat with great relish the Stryphnodendron pods 
that fall to the ground, apparently attracted by the 
sweet pulp. The new species therefore shows some 
promise for inclusion in agronomic trials as part of 
the agroforestry and silvo-pastoral production sys- 
tems involving sheep, pastures, and forage legumes 
now being developed for and implemented by set- 
tlers in the Napo region of Ecuador. 

The support of the U.S. Agency for International 
Development (USAID) and the Forestry Director- 
ate of the Ecuadorian Ministry of Agriculture, for 
the research in eastern Ecuador, is gratefully ac- 
knowledged. The authors thank Enrique Forero for 
bringing us together through correspondence, and 
Alejandro Suárez for preparing the illustration. 


LITERATURE CITED 
MIR E. M. Stryphnodenron (Fabaceae: Mimoso- 
l. Neotrop. Monogr. (in preparation). 
Occo t E. M. 19 Stryphnodendron 
m. Ás especies de naroste; sudeste e 
id de Brasil IL Leandra 4-5: 
1981. Stryphnodendron Mert. (Legumino- 
sae-Mimos oideae) com especial jm aos taxa 
amazonicos. Leandra 10-11: 3-100. 
SrEARN, W. T. 1966. Botanical TA Hafner, New 
York 


— David A. Neill, Missouri Botanical Garden, 
P.O. Box 299, St. Louis, Missouri 63166, U.S.A., 
and Elêna Maria de Lamare Occhioni, Universi- 
dade Federal do Rio de Janeiro, Departamento 
de Botânica, Rio de Janeiro, Brazil. 


A NEW SPECIES OF 
HYBANTHUS (VIOLACEAE) 
FROM PANAMA 


Hybanthus is a very poorly known genus of 
150 species of herbs, trees, and shrubs from the 
Old and New World. The last synopsis of the genus 
throughout its range enumerated 32 species in the 
New World (Melchior, 1925); Morton (1944) pro- 
vided the most recent synopsis of the Central Amer- 
ican species. The new species proposed here, Hy- 
banthus jefensis, is clearly distinct from all species 
in Central America and is not placed within any 
species presently known from Colombia (Smith & 
Fernández-Pérez, 1954). 


Hybanthus jefensis Todzia, sp. nov. TYPE: Pan- 
ama. Panamá: Cerro Jefe, near Rio Indio, 
2,100-2,200 ft., 17 Feb. 1968, Dwyer 1969 
(holotype, MO). Figure 1. 


Frutices caulibus juvenibus strigulosis; stipulae anguste 
triangulares in siccitate iioii folia alterna abel 
labra in siccitate rubropunctata in pagina inferna; in 
rescentiae axillares de ec eir in brachyblastis, foribus 
1-4 in pedicellis 1-2 cm longis; petalum panduratum 
0.7-1 cm longum rotundatum ad apicem; capsulae 7-8 
mm vieni 


Shrubs 1-2.5 m tall; older stems terete, reddish 
brown, glabrous, sparsely branched; younger stems 
terete to quadrangular, dark to light brown, strig- 
ulose with hairs curving upwards, with leaves erect 
and + adpressed to stems; stipules narrowly tri- 
angular, 1.5-2 mm long, ca. 0.5 road at 
base, drying chartaceous, brown. Leaves alternate, 
elliptic to narrowly obovate, 3-12 cm long, 1-3.8 
cm broad; petioles terete, 0.5-1 cm long, glabrous; 
the blade acute at apex, attenuate at base, smooth, 
glabrous, drying chartaceous with red dots and 
reticulations beneath, each side with 5-7 usually 
obscure, gradually arcuate lateral veins; the margin 
crenulate with 8-17 glandular teeth on each side. 
Inflorescences 1-4-flowered, axillary, fasciculate 
on short shoots ca. 2 mm long; bracts inserted at 
base of pedicels, lanceolate, 1-2 mm long, acute, 
drying chartaceous, brown; bracteoles on lower half 
of pedicels 0.5-1 mm long, lanceolate; pedicels 1— 
2 cm long, slender, glabrous, articulate above the 
middle. Flowers white, zygomorphic; sepals 5, lan 
ceolate, 2-2.5 mm long, acute, glabrous; lateral 
petals 4, 4-5 mm long, falcate, with irregularly 


crenate margins; anterior petal pandurate, 0.7-1 
cm long, with claw 4-5 mm long, sparsely pilose, 
lip 3-5 mm long, 4-5 mm broad at widest point, 
rounded at apex, glabrous; stamens 5, sessile, free 
or connate; filaments very short, the anterior 2 
stamens with a subglobose gland dorsally; anthers 
1-1.5 mm long, sparsely pilose dorsally and at 
base; connective appendages ca. 2 mm long, 0.5- 
l mm broad, rounded at apex, membranaceous; 
ovary globose, ca. 1 mm long, glabrous, with 3 
locules containing 1-2 ovules each; style 2-3 mm 
long, curved at apex. Capsules loculicidally and 
elastically 3-valvate, 7-8 mm long, glabrous, with 
3—6 seeds, often with persistent style and sepals; 
seeds globose to ovoid, 2-3 mm diam., smooth, 
black. 


Distribution, habitat, and phenology. Endemic 
to the Cerro Jefe region of the Prov. Panamá, 
Panama, at 650-1,000 m elevation in disturbed 
and undisturbed premontane wet forest. Flowering 
and fruiting October, November, December, Jan- 
uary, and February 


Representative specimens examined. PANAMA. 
PANAMÁ: region of Cerro Jefe, ca. 1,000 m, 3 Oct. 1969, 
Correa et al. 1598 (MO); between Cerro Azul and Cerro 
Jefe, 24 Dec. 1967, Dressler 3270 (MO); between Cerro 
Jefe and La Eneida, 17 Jan. 1968, Dwyer et al. 8195 
(MO); 6.5 km by road N of Lago Cerro Azul, 650-730 
m, 13 Jan. 1974, Nee 9292 (MO). 


This species is distinctive by virtue of its small 
crenulate leaves with red dots and punctations when 
dried; axillary inflorescences fasciculate on short 
shoots ca. 2 mm long; stipules that are chartaceous 
when dry; and pedicels 1-2 mm long. The anterior 
petal, one of the most diagnostic characters of the 
genus, is pandurate, 0.7-1 cm long, and rounded 
apically. In southern Central America H. jefensis 
appears to be most similar to H. guanacastensis 
Standley from which the new species differs by its 
chartaceous (vs. indurate) stipules, crenulate (vs. 
doubly serrate) leaf margins, shorter anterior petals 
(0.7-1 cm long vs. ca. 1.2 cm long), and smaller 
capsules (0.7-0.8 cm long vs. 1.5-2 cm long). 
Hybanthus jefensis is probably most closely related 
to H. sylvicola Standley & Steyerm. of southern 


ANN. MISSOURI Bor. GARD. 76: 360-362. 1989. 


Volume 76, Number 1 Notes 361 


es So le 

FicurE 1. Hybanthus pr — A. Habit with close-ups of leaf margin and leaf axil showing bracts at base of 
fascicles. — B. Flower. — C. Close-up of anthers and style with sepals and petals removed. — D. Close-up of an anterior 
anther. (A-D from Do rs )—E. Fruit (from Dwyer 8195). 


Mexico and Guatemala but differs by having strig-. the region of Cerro Jefe in central Panama (Lewis, 

ulose (vs. glabrous) young branches, leaves with 1971). In addition to the three Hybanthus species 

(vs. without) red punctations beneath, and lip 0.7- reported in the Flora of Panama (Robyns, 1967), 

1 cm vs. 1.1-1.3 cm long. two other species are presently known from 
Hybanthus jefensis is yet another endemic to ama (D'Arcy, 1987; pers. obs). 


362 Annals of the 
Missouri Botanical Garden 


KEv TO HYBANTHUS IN PANAMA 


la. Shrubs or small trees, woody; flowers in fascicles borne on short shoots. 
. Shrubs or small trees 1-7 m tall; capsules 0.7-2 c 
3a. uli gu drying white, indurate; leaves aps serrate, » drying uniform in color beneath, without 
ots; anterior petal 1.2-4 cm long; capsules 1-2 c 
ri Anterior petal up to 4 cm long; capsules 1.5-2 cm s seeds 4-5 mm ae as H. prunifolius 
4b. Anterior petal ca. 1.2 cm long; capsules 1-1.3 cm long; seeds 2-3 mm long „u 
anacastensis 


m gua 
3b. Stipules drying brown, chartaceous; leaves crenulate, drying with red dots and reticulations iow 
nterior petal 0.7-1 cm long; capsules 0.7-0.8 cm long 


a H. jefensis 
2b. Shrubs less than 1 m tall; capsules 0.3-0.6 c cm p 
ni ot w 


H. galeottii 


attenuatus 
thiemii 


Sb. Leaves alternate, ovate to oblong-ovate, acute at apex; plants perennial 


oo M. 1925. Violaceae. /n: A. Engler & K. 
antl, Die Natürlichen Pflanzenfamilien, ed. 2. 21: 
77. 


I thank Guy Nesom for providing the Latin 
description, Doris Lee Tischler for the handsome 
illustration, and James W. Grimes, George Rogers, Morr on G. V. 1044. “The genas bas incon: 
and Henk van der Werff for reviewing the manu- tinental North America. Contr. U.S. Natl. Herb. 29: 
script. This work was supported by a Noyes Foun- 2 
dation grant to the Missouri Botanical Garden. 


74-82. 

RoBvNs, A. 1967. Violaceae. In: R. E. Woodson, Jr. 
& R. W. Schery (editors), Flora of Panama. Ann. 
Missouri Bot. Gard. 54: 65-84. 

LITERATURE CITED SMITH, L. B. € A. FERNÁNDEZ-PÉREZ. 1954. Revisio 

D’Arcy, W. G. 1987. Flora of Panama. Checklist and Violacearum Colombie: Caldasia 6(28): 83-181. 


1 
Index. Part 1: The Introducti d Checklist. M . f 
-n Sep a eae d 17. E A, Todzia, Department of Botany, Uni- 


Lewis, W. H. 1971. High floristic endemism in low versity of Texas, Austin, Texas 78712, U.S.A. 
cloud forests of Panama. Biotropica 3: 78-80. 


BOOK REVIEW 


Mora-Osejo, Luis Eduardo. 1987. Estudios Mor- 
fológicos, Autoecológicos y Sistemáticos en 
Angiospermas. Academia Colombiana de 
Ciencias Exactas, Físicas y Naturales, Serie 
Jorge Alvarez Lleras 1: 1-195. Retail price: 
$15.00. (Available from Dr. L. E. Mora-Ose- 
jo, Academia Colombiana de Ciencias Exactas, 
Fisicas y Naturales, Apartado Aéreo 44763, 
Bogotá D.E. 1, Colombia.) 


Morphological, Autecological and Systematic 
Studies in Angiosperms is an important addition 
to the increasing botanical literature of high sci- 
entific quality being produced in Latin America. It 
has been published by the Colombian Academy of 
Sciences as the first volume of a new series called 
“Serie Jorge Alvarez Lleras.” The actual text is 
188 pages long, with 75 figures, including several 
color photographs, and a bibliography with over 
90 citations. The text is divided into two parts. 
The first part deals with the morphological and 
autecological studies and includes five chapters (1— 
5), while the second deals with the systematic stud- 
ies and is made up of one chapter (6). 

The author indicates that one of his objectives 
is to present to a broader audience the “typolog- 
ical method, which has been developed in Ger- 
many but is not well known among Spanish- and 
English-speaking phytomorphologists. In the first 
chapter he discusses the general principles of the 
typological method, including such concepts as ho- 
mology, position, continuity, similarity, etc.; the 
procedures employed in the study of specific prob- 
lems; the contribution of typology to our under- 
standing of the organization and architecture of 
the angiosperms; and the relationship of typology 
with taxonomy and phylogeny. In Chapter 2, the 
typological theory and its relation to the evolution 
of the inflorescence is discussed. Chapter 3 deals 
with new concepts developed by the author on the 
diversity and evolution of organizational patterns 
found in the floral meristems in angiosperms, based 
on his observations of a number of families, in- 
cluding Melastomataceae, Araliaceae, Bignoni- 
aceae, Asteraceae, Cyperaceae, Juncaceae, Po- 


aceae, Bromeliaceae, and Scheuchzeriaceae. 
Chapters 4 and 5 deal with the author's original 
typological studies in Lilaea, Triglochin, and 
Scheuchzeria, which include the study of the ar- 
chitecture of the meristem and its correlation with 
the reproductive strategies of these plant groups. 
In Chapter 6, Mora-Osejo gives an example of the 
applicability of the typological method to the taxo- 
nomic study of Oreobolus R. Br., a genus of Cy- 
peraceae. 

There are several aspects of this publication that 
I find outstanding, the most important being the 
thorough discussion of the theory and practice of 
the typological method. The author explains how 
this method allows for a comparative analysis of 
architectural patterns in plants as opposed to a 
purely descriptive analysis. The method also allows 
for interpretation of information derived from the 
detailed analysis of plant structures through the 
understanding of the interconnections among them. 
At the same time, it provides valuable tools for 
understanding adaptations to the environment as 
well as reproductive strategies in angiosperms. 

Aside from its obvious scientific value, the reader 
is also very favorably impressed by the good quality 
of the printed product and by the quantity and 
quality of the illustrations, which include numerous 
schematic representations, color and black and white 
photographs, details of plant parts, and more. 

The volume is a substantial contribution to our 
understanding of this important subject, as are most 
of Professor Mora-Osejo's publications. His study 
of the biology of the Haloragaceae, published as 
part of his treatment of the family for the Flora 
de Colombia, is another example of his scientific 
excellence. It is to be expected that this volume 
will be given proper consideration by phytomor- 
phologists in the United States and in Latin Amer- 
ica. Its contents are well worth careful study. This 
work is the result of many years of careful research 
by one of Latin America's most prominent scien- 
tists.— Enrique Forero, Missouri Botanical Gar- 
den, P.O. Box 299, St. Louis, Missouri 63166, 
U.S.A. 


ANN. MISSOURI Bor. GARD. 76: 363. 1989. 


NOTICES 


THE 1988 Jesse M. GREENMAN AWARD 


The 1988 Jesse M. Greenman Award has been 
won by John H. Wiersema for his publication “A 
Monograph of Nymphaea Subgenus Hydrocallis 
(Nymphaeaceae)," which appeared in Systematic 
Botany Monographs, volume 16. This monograph 
is derived from a Ph.D. dissertation submitted to 
the University of Alabama, under the direction of 
Dr. Robert R. Haynes. Dr. Wiersema based his 
classification on numerical taxonomy, scanning 
electron microscopy of seeds and pollen, chro- 
mosome numbers, floral biology, artificial hybrid- 
ization, flavonoid chemistry, and general mor- 
phology. Fourteen species are recognized in the 
subgenus, which is restricted to the American trop- 
ics and subtropics, and one new subspecies is pro- 
posed. 

This Award is named for Jesse More Greenman 
(1867-1951), who was curator of the Missouri 
Botanical Garden Herbarium from 1919 until 1943. 
A cash prize of $250 is presented each year by 


the Garden, recognizing the paper judged best in 
vascular plant or bryophyte systematics based on 
a doctoral dissertation that was published during 
the previous year. 

Nominations for papers published during 1988 
are now being accepted for the 21st annual award, 
which will be presented in the summer of 1989. 
Reprints of such papers should be sent to: Green- 
man Award Committee, Research Division, Mis- 
souri Botanical Garden, P.O. Box 299, St. Louis, 
Missouri 63166-0299, U.S.A. In order to be con- 
sidered for the 1989 award, reprints must be re- 
ceived by June 1, 1989. 


REPRINTS 


Copies of “The Genus Xyris (Xyridaceae) in 
Venezuela and Contiguous Northern South Amer- 
ica" (Ann. Missouri Bot. Gard. 75: 522-722. 1988) 
can be obtained by sending $15.00 to: Dr. Robert 
Kral, Herbarium, Box 1705, Station B, Vanderbilt 
University, Nashville, Tennessee 37235, U.S.A. 


Volume 75, Number 4, pp. 1169-1670 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN was published on 
8 March 1989. 


Tropical florulas 


Recognizing the utility of floras of restricted tropical areas to workers in a variety of biological 
disciplines, Missouri Botanical Garden staff members have produced several during the past few 
years. Available titles are: i 


La Flora de Jauneche 

C. H. Dodson, A. H. Gentry & F. M. Valverde 

This is the first in a series of florulas of the life zones of Ecuador. It treats the plants of the 
Jauneche field station, located in tropical moist forest between Quito and Guayaquil, at an altitude 
of 50-70 meters. The station occupies 130 hectares and has a known flora of 728 species. All 
are treated, and each is illustrated with a line drawing of its habit and details of critical features. 
The book is bilingual: all information is in both English and Spanish. 

xxxix + 512 pages, 184 plates, 14 figures; soft bound. Published by Banco Central de Ecuador. 
1986. $25.00 


" i | 

Flora of the Río Palenque Science Center 

C. H. Dodson & A. H. Gentry 

This flora is similar in plan to the Jauneche work, but it treats more species, about 1100. The Rio 

Palenque station occupies 167 hectares at an altitude of 150-220 meters in the tropical rainforest 
e zone. 


xxx + 628 pages, 278 plates, 22 figures; soft bound. Published as Selbyana, vol. 4, 1978. $35.00 


Flora of Barro Colorado Island 

Thomas B. Croat 

About 1350 species of vascular plants are described and many illustrated from Barro Colorado 
Island, the Smithsonian Institution's 15.6 sq. km reserve in the Panama Canal. “Croat's well- 
executed effort sets the precedent for the kind of work perquisite to fuller understanding." David 
Barrington, Plant Science Bulletin. 


added to orders requiring invoices. No shipments are made until payment is received. Mail form with your 
check or money order, payable to Missouri Botanical Garden, to: 
Department Eleven 
PO Beg 0 5 a 
St, Louis, MO 63166-0299, USA. 
Please send the publications circled above to: — 


n Payment enclosed. 


Papua ee en AB, PESE PMI 
decur a D e el aid (FA DO Ton wil hi: 
Address m ; 35 ; UP E ^ added to total). 


Prices Are Suasect To CHANGE Wirnout Nonig ——-— * 


CONTENTS 


Tessmannianthus, an Arborescent Genus of Melastomataceae New to Panama Frank 
Almeda 


The Genus Argostemma (Rubiaceae-Argostemmateae) in Borneo Birgitta Bremer .... 
Studies in Neotropical Paleobotany. VII. The Lower Miocene Communities of Panama— 
The La Boca Formation Alan Graham 


A Revision of Mesoamerican Psychotria Subgenus Psychotria (Rubiaceae), Part I: Intro- 


duction and Species 1-16 Clement W. Hamilton 67 
Dianthoveus: A New Genus of Cyclanthaceae Barry E. Hammel & George J. 
ilder 112 
Anatomia Sisi eth de Lacandonia schismatica (Lacandoniaceae) J. Márquez > 
| n, M. Engleman, A. Martínez-Mena, E. Martinez & C. Ramos cc ic Bi 
Buen (Triuridales): Una Nueva Familia de Mexico Esteban Martínez & Clara deii 
— Hilda Ramos Td 128 | 
Systematics of the Amazon Lilies, Eucharis and Caliphruria (Amaryllidaceae) p» 
WF Me o od st i aai caa e Roa AE 136 
The Systematics and Evolution of Ludwigia Section Microcarpium NAE | 
Ching-I Peng A : 
Two New Species of the Lupinus lanatus Complex Ana Maria Planchuelo & David T 
| Riu UII Quam EUIS Etras qu quest bRucd be DuC x wwe A uri 303 
Augustus Fendler's Venezuelan Collections of Ferns and Fern Allies Alan R. Smith E 
CUA A A ee x bs 
NOTES 2 


- A New Echeandia (Liliaceae) from Venezuelan Guayana Robert William Cruden ...... 350 — 
Solanum toliaraea, a New Species from suis William G. D'Arcy & Armani 
Rakotozafy = 


351 
Note Concerning Pues henriquesii (Gramineae) i in Northern Portugal W. E e y. 3 


| Passiflora. citrina, a New Species in Section Xerogona (Passifloraceae), "s Mero 

John M. MacDougal H 

A New luus of Stryphnodendron (Fabaceae: Mimosoideae) from Amazonian Ecua- ats 
dor David A. Neill & Eléna Maria de Lamare Occhioni „a fares 

_ A New Species of Hybanthus re from Penama. Carol A. Todzia A ts Tart 
BOOK REVIEW... 7 A 
SC NOBGES 25s . | RUDI E Es 


— 


Cover illustration. Solanum toliaraea D'Arcy & Rakotozafy, by John Myers. 


bdsm 76 
‘umber 2 


qe 


ES 
——— 


Volume 76, Number 2 
Summer 1989 


Annals of the 
Missouri Botanical Garden 


The Annals, published quarterly, contains papers, primarily in systematic botany, con- 
tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the 
Garden will also be accepted. Authors should write the Editor for information concerning 
arrangements for publishing in the ANNALS. Instructions to Authors are printed in the back 


of the last issue of each volume. 


Editorial Committee 


. George K. Rogers 
Editor, Missouri Botanical Garden 


Amy Scheuler 
Editorial Assistant, 
Missouri Botanical Garden 


Glenda Nau 
. Magdalen Lampe 
. Publications Staff 


Marshall R. Crosby 


Missouri Botanical Garden 


Gerrit Davidse 
Missouri Botanical Garden : 


John D. Dwyer 
Missouri Botanical Garden & 


Saint Louis University 


Peter Goldblatt 


Missouri Botanical Garden 


Dale E. Johnson 


Missouri Botanical ore 


| Henk van der We 


Missouri Botanical Garden E ? 


For, subscription. information contact Department 
= — Eleven, P.O. Box 299, Ste Laue, MO 63166. Sub- __ 
scription price is $75 p 1e U.S., $80 Canada — 


and Mexico, $100 all other countries. Four i issues — 


ES A volume. Ss 


OER AN Mee EL S S : Eleven, P.O. Box 299, St. joue, me 63166. 
© Missouri Botanical Garden 1989 | 


. THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. — 


The ANNALS OF THE MISSOURI BOTANICAL a 


(ISSN 0026-6493) is published quarterly 


Missouri Botanical Garden, 2345 Tower Grove Av 
enue, St. Louis, MO 63110. Second class. 


paid at St. Louis, MO and sonal aie e 
n 


PosrMASTER: Send address changes to A^ 


THE MISSOURI BOTANICAL GARDEN, 


Volume 76 
Number 2 
1989 


Missouri 
Botanical 


Garden 


NEW SPECIES AND 
COMBINATIONS IN TROPICAL 
AMERICAN POLYSTICHUM 
(DRYOPTERIDACEAE) 


David S. Barrington? 


ABSTRACT 


A taxonomic history for the fern genus Polystichum in Latin America and an analysis of the vi dein characters 
at the species level in the genus provide the context for the recognition of new and overlooked taxa. Three new 
species are described from Latin America, and a new combination is provided for a Polystichum species from the 


southern Andes 


In the course of preparing a treatment of the 
genus Polystichum for the Flora Mesoamericana, 
I have discerned a set of new and neglected species 
from Central America and the northern Andes, 
which I report here. The species now included in 
the genus Polystichum have only once been in- 
cluded in a monograph (Mettenius, 1858); there 
has been no modern monographic work on the 
genus in the New World. An inquiry into species 
and speciation phenomena in Costa Rica (Barring- 
ton, 1985a, b; Barrington, et al., 1986) has led 
in turn to an inquiry into the significance of hybrids 
with respect to species concepts in the ferns (Bar- 


rington et al., in press). I report here on the mor- 
phological characters important in distinguishing 
species of Latin American Polystichum. This re- 
view of characters provides the context for the 
descriptions of species, which make up the central 
contribution of this work. 


TAXONOMIC HISTORY 


The genus Polystichum Roth, terrestrial ferns 
characterized by fastigiate leaves with inequilateral 
leaf segments and peltate indusia (when present), 
is prominent and diverse in montane regions 


'T thank Luis Diego Gómez P. for his energy and support of my fieldwork in the Sierra Talamanca, Steven R 
Hill for his critical reading of the Latin descriptions, and Lynn Bohs for her thorough review of the manuscript. 
Several nomenclatural and taxonomic problems were resolved with Rolla Tryon's help. The curators of the following 


herbaria supported this project through loan of materials: A, AA 


U, CR, ENCB, F, GH, MO, NY, UC, US, USNA, 


. The reped of Vermont provided support for laboratory and fieldwork through its Institutional Grants BRSG- 


79 ad BSCI85- 


? Pringle ANN Department of Botany, University of Vermont, Burlington, Vermont 05405-0086, U.S.A. 
ANN. MISSOURI Bor. GARD. 76: 365-373. 1989. 


366 


Annals of the 
Missouri Botanical Garden 


throughout the world. The generic name originates 
with Roth (1799), but it was K. B. Presl (1836) 
who first circumscribed the genus as it is now 
recognized. The most important influence on the 
taxonomic history of Polystichum was the treat- 
ment of W. J. Hooker (1862), in which most of 
the twice-pinnate taxa in the world were combined 
into one species. As a result, the morphological 
and taxonomic diversity of species was left un- 
studied by the otherwise productive taxonomists of 
the late nineteenth century. Christensen (1906) 
maintained Hooker's counterproductive species 
concept in Polystichum, which has perpetuated 
the tendency for taxonomists to ignore the diversity 
of twice-pinnate species. Christensen did recognize 
4l forms, many of which were associated with 

names whose types represent distinct species. 
atin America and eastern Asia are the two 
centers of diversity for the genus Polystichum. 
Although the diversity and biology of species are 
well known in Asia (Tagawa, 1940; Daigobo, 1972), 
we know little about the genus in the New World 
tropics. Maxon (1909) was the first to recognize 
natural species in the American tropics, but his 
work was confined to the West Indies. Only re- 
cently (A. R. Smith, 1981; Stolze, 1981) has sim- 
ilar progress been made in defining the continental 
species. In the past few years, the extent of the 
taxonomic problem has become clear with the dem- 
onstration of extensive hybridization and allopoly- 
ploidy in the American tropics (Barrington, 1985a, 
" popa et al., 1986). I reviewed the present 
mic status of the genus Polystichum in Latin 


America (Barrington, 1985c). 


MORPHOLOGY 


The analysis leading to the descriptions of the 
new species provided here and the work on hy- 
bridization and polyploidy have revealed the most 
useful characters for delineating Polystichum 
species in Latin America. Since there has been no 
previous morphological review for the genus in the 
Neotropics, I provide an analysis of the important 
characters for Polystichum. I hav 
mented on ontogenetic and environmentally in- 
duced variation, which is unusually confusing in 


the genus 


also com- 


INDUMENT 


Indument in Polystichum consists entirely of 
scales and scale homologs (squamules and micro- 
scales); true hairs, bristles, and spines are absent, 
at least in Latin American species. The general 
form, texture, and frequency of scales are impor- 


tant in delineating taxa of Polystichum. Particu- 
larly important are histological features of the scales: 
they may be conform (sensu Tryon, 1970) or they 
may have a margin of cells different in shape and 
orientation from the scale body. Scale color is im- 
portant too, although variable with position on the 
plant and age of the plant. Tagawa (1940) con- 
tended that the smallest scales on the abaxial lam- 
ina surface (the microscales) are important in de- 
fining subgeneric groups in Polystichum. In Latin 
America, variation in microscales is extremely con- 
servative, consistent with Tagawa's contention. Most 
species have the acicular microscales typical of 
sect. Metapolystichum 

I have followed Smith (1981) in describing the 
processes that arise from the edges of scales and 
squamules as cilia, although this is an oversimpli- 
fication. [n fact, scales and squamules bear a con- 
tinuum of processes from cilia (long and delicate) 
to setae (short and stiff). I have chosen to call all 
of these processes cilia and use adjectives to de- 
scribe their length and stiffness. The kinds and 
proportions of these cilia on the different scales 
and squamules are important taxonomic characters 
in Polystichum. 


LAMINA 


The dissection and texture of the lamina provide 
critical characters for distinguishing species of Po- 
lystichum, although they are phenotypically and 
ontogenetically variable. Particularly confusing is 
the tendency for older plants to have larger, more 
dissected leaves. Nevertheless, species can be cir- 
cumscribed within the context of the ontogenetic 
variation encountered. In addition, the shape, angle 
of attachment, and revoluteness of leaf segments 
(pinnae and pinnules) are all important, although 
phenotypically variable (especially revoluteness). 
Development of the spinules (extensions of vein 
tips) varies among species. The complex interaction 
of ontogeny, environment, and genotype that shapes 
Polystichum leaves perhaps explains the recog- 
nition of large, poorly resolved taxa by both Hooker 
and Christensen. 


SORI AND SPORANGIA 


The placement of the sorus relative to the vein 
tips is important in Polystichum, even though it 
is variable across a segment. Indusium presence 
or absence is an important character, although the 
indusium is sometimes pal in ee Po- 
lystichum species. In addition, , Shape, 
and histology of the inden vary among, but 
usually not within, species. Color is sometimes use- 


Volume 76, Number 2 
1989 


Barrington 
Polystichum 


ful but is variable with age. The design of the 
perispore is usually constant within species. 

Color of the indument is a useful character in 
Polystichum, as in many ferns. The color names 
used in the descriptions here are the National Bu- 
reau of Standards color names as defined in Na- 
tional Bureau of Standards Special Publication 
440 (and its predecessor, Circular us iir 
by comparison of indument with the BS 
Color-Name Chart (a supplement to D Special 
Publication 440), which is a partial set of color 
chips representing the color centroid. I standardize 
color terminology in this way, rather than using 
the Ridgway-Dade terminology and standardization 
(Stearn, 1983), because those color names are 
much less familiar, and the color chips on which 
they are based are much harder to locate. Indu- 
ment is variable and systematically useful in all 
three dimensions of the color centroid (hue, value, 
and chroma); it is also variable in these dimensions 
with position on the plant and age of the individual. 


HYBRIDIZATION AND SPECIES 
DEFINITION 


Hybrids between the new Costa Rican species 
described here and several other species were en- 
countered during fieldwork. Several of these hy- 
brids have been described elsewhere (Barrington, 
1985a, b). All of them meet two criteria of hy- 
bridity: intermediacy and spore irregularity. Since 
all of the hybrids have irregular spores, they are 
probably sterile and hence provide evidence of 
species-level divergence between their progenitors 
(Barrington et al., in press). This evidence provides 
a first working hypothesis for species definition in 
tropical American Polystichum and the basis for 
recognizing the new species described here. 


DESCRIPTIONS OF NEW SPECIES 


Polystichum concinnum Lellinger ex Barring- 
ton, sp. nov. TYPE: Costa Rica. San Jose: Villa 
Mills, S of Pan American Highway about one 
km E of Hotel La Georgina, ca. 3,100 m, 3 
Jan. 1980, D. S. Barrington 694 (holotype, 
GH; isotype, VT). Figure 1 


de cal terrestris. Caudex erectus base ascen- 
dente. Squamae petiolorum anguste lanceolatae, coria- 
ceae, a oran. cilia proximalia squa- 
marum  petiolorum longa enuiaque, distalia brevia, 
rief: saepe . Lam- 
ina 0.4-2.0+ m longa, bipinnata, indument o pagina e 
rhachidis abaxialis constans e squamis lanceolatis um- 
brinis. Pinnae patentes attenuatae, m is medianis circa 
7-18 cm longis. Pinnulae crenatae patentes fere planae 


vix spinulosae circa 0.8 cm longae 0.4 cm latae, squamis 
minimis acicularibus, pallidibus. Sorus terminalis in ramo 
acroscopico venae locatus, indusio peltato circa 0.5-0.7 
m ; 


m diametro 


Plant terrestrial, the spreading-ascendent leaves 
to 2 m long. Stem erect from an ascendent base, 
sometimes with weak lateral branches. Petiole with 
8 vascular bundles, 2 large adaxial and 6 small 
abaxial; petiole scales narrow-lanceolate, 
ceous, conform, blackish red to dark reddish brown 
with sharply delimited narrow lighter borders, the 
color transition to border abrupt; the proximal cilia 
frequent, long and delicate, the distal cilia scat- 
tered, short, indurate, often with recurved hooklike 
processes. Petiole squamules unguiculate and del- 
tate, wrinkled, very dark red with deep orange 
borders, with cilia scattered on the unguiculate 


coria- 


squamules to dense on the deltate squamules. Lam- 
ina 0.4 to more than 2 m long, chartaceous, acu- 
2-pinnate, abaxial rachis scales strong 
brown, ciliate and lanceolate to sparsely ciliate and 
on each side of the 


minate, 


unguiculate. Pinnae ca. 20- 
rachis, attached at right angles to the rachis, at- 
tenuate, the median pinnae ca. 7-18 cm long, 
basal pinnae about %, as long; pinna-rachis squa- 
mules strong brown, sparsely ciliate, lanceolate or 
0-25 on each 
side of the pinna rachis, attached at nearly right 
angles to the pinna rachis, flat, truncate at the 
base and obtuse to rounded, crenulate (the basal 
acroscopic pinnules of each pinna often pinnatisect 


rarely unguiculate. Pinnules ca. 1 


above the auricle), weakly spinulose, about 0.8 cm 
long and 0.4 cm wide, the basal pinnules of each 
pinna abruptly shorter than the next distal pinnules; 
microscales of costules and veins acicular, pale, 
without glandular cells. Sorus terminal on an ac- 
roscopic vein branch, covered by a thin, peltate, 
casually deciduous indusium ca. 0.5-0.7 mm diam., 
the indusium flange of cubical to radially elongate, 
slightly sinuate cells organized into radial files, the 
indusium-flange margin a single row of narrow, 
radially elongate cells. Spores with a well-developed 
cristate perispore. 


Distribution. Endemic to high, wet forests of 
the Sierra Talamanca, Costa Rica and Panamá, 
2,000-3,000 m, occasionally to 3,400 m in dis- 
turbed areas above treeline. 


The affinity of this large, graceful species is with 
the large group of indusiate and two-pinnate species 
most diverse in southern Mexico. Polystichum con- 
cinnum is most similar to two Mexican species, P. 
rachichlaena Fée and P. erythrosorum A. R. 
Smith: all three are large indusiate, flat-leaved, two- 
pinnate species with crenate pinnules, and they are 


368 Annals of the 
Missouri Botanical Garden 


FIGURE 1. Polystichum concinnum. — A. Leaf. — B. Two pinnules, the left one with sori removed. — C. Indusium, 
from above (upper) and side. — D. Rachis scale. — E, F. Rachis squamules. —G. Pinna-rachis squamule. — H. Medial 
edge of petiole scale (below), petiole scale apex.— I. Petiole scale. 


Volume 76, Number 2 
1989 


Barrington 369 


Polystichum 


all confined to high-altitude forests. Polystichum 
rachichlaena is notable for the large size (over 1 
mm diam.) and persistence of its indusia; it also 
has petiole scales virtually devoid of cilia. The 
lamina and sorus of P. concinnum are similar to 
those of P. erythrosorum, but the petiole scales of 
P. erythrosorum are er, marginate, and 
strongly bicolorous (basally lustrous and black with 
brownish orange borders). In addition, the long, 
delicate cilia of P. concinnum are lacking in P. 
erythrosorum, and the small distal cilia are not 
dark and indurated as they are in P. concinnum. 
Polystichum concinnum is well represented in her- 
baria; it has been determined most often as either 
Polystichum muricatum (L.) Fée (which differs in 
having short petiole-scale cilia) or Polystichum 
ordinatum (Kunze) Liebm. (which differs in having 
narrow leaf bases and strongly bicolorous petiole 
scales with thick bases). 

David B. Lellinger, of the United States National 
Herbarium, first distinguished P. concinnum and 
coined its species epithet. 

Several chromosome counts of plants from the 
Talamanca range yielded n — 41 pairs at meiosis I. 


Additional specimens examined. COSTA RICA. CAR- 
TAGO: Villa Mills, 1 km N of Restaurant La Georgina, 30 
May 1978, Barrington 576 (GH, VT). sAN JOSÉ: Cerro 
Chirripó, SW slopes, along ridge trail from Canaan to 
summit, 8,500-10,000 ft., 22 & 27 Aug. 1967, Evans, 
Lellinger & Bowers 73 (US). PANAMA. CHIRIQUÍ: Cuesta 
de Las Palmas, southern slope of Cerro de la Horqueta, 
1,700-2,100 m, 18 Mar. 1911, Maxon 5527 (US 


Hybrids. | Hybrids between this species and the 
distinctive P. speciosissimum have been collected 
four times from roadcuts on the Pan-American 
Highway in the Sierra Talamanca area of Costa 
Rica (Barrington 705, 1274, 1516, VT; W. D. 
Stevens 14317, MO). Hybrids with the following 


species are discussed under that taxon. 


Polystichum talamancanum Barrington, sp. 
nov. TYPE: Costa Rica. Cartago: Cerro de la 
Muerte, roadcut 0.4 mi. W of highest point 
on Pan-American Highway, SE aspect, 3,330 
m, 4 Jan. 1980, D. S. Barrington 708 (ho- 
lotype, GH; isotypes, CR, VT). Figure 2 

Filix terrestris. Rhizoma repens frondibus fasciculatis. 
luteofuscae, ciliis longis tenuibus 


cundorum 12-70 cm longa, rigide erecta in locis apricis, 
indumento paginae rhachidis abaxialis constanti e squamis 


longae. Pinnulae patentes fere (70-80? e rhachidi), re- 
volutissimae in locis apricis crescente, vix spinulosae, circa 


cm longae, 0.4-0.5 cm latae, squamis minimis vid 
cularibus pallidis. Sorus terminalis insidens ramo acrosc 
venae, indusio l1 nullo vel spathulato, raro 


pico 
peltato, 0.05-0.07 mm lat 


Plant terrestrial, the rigidly erect leaves to 60 
cm long. Rhizome short-creeping, with small lateral 
branches. Petiole with 2 large adaxial and 6 small 
abaxial vascular bundles; petiole scales lanceolate, 
herbaceous or rarely chartaceous, conform except 
at cilia, medially dark yellow-brown, the border 
similar or distally moderate olive-brown, the color 
the cilia proximally 
common, long, arising from broad bases of several 


transition to border gradual; 


cells + perpendicular to the scale axis, thin and 
delicate; petiole squamules unguiculate, strong 
brown, + flat, with broad and short cilia common 
at base. Lamina of fertile leaves 12 to 70+ cm 
long, rigidly erect (spreading-erect in shade), 
lanceolate, 2-pinnate; abaxial rachis scales narrow- 
lanceolate, uniformly medium to deep orange-yel- 
low, with short cilia throughout and long cilia dis- 
tally; abaxial rachis squamules like those of the 
petiole, deep orange, crumpled, the cilia well de- 
veloped at squamule base. Pinnae about 40 on 
each side of the pinna rachis, attached at about a 
40? angle to the rachis, acuminate to short-atten- 
uate, the median pinnae ca. 5-8 cm long, basal 
pinnae /-%, as long; abaxial pinna-rachis squa- 
mules deep orange with long cilia at the base and 
scattered short cilia distally, the larger long.lan- 
ceolate, the smaller unguiculate. Pinnules ca. 11- 
14 on each side of the pinna rachis, attached at 
about a 70-80? angle to the pinna rachis, strongly 
revolute in sun (only the sides of the pinnules 
revolute, not the tips), barely revolute in shade, 
cuneate at base and acute-tipped, crenulate (the 
basal acroscopic pinnules of each pinna often pin- 
natisect above the auricle), with spinules only at 
auricle and pinnule apex, ca. 0.7-0.9 cm long, 
0.4-0.5 cm wide (wider at the well-developed au- 
ricle), the basal pinnules of a pinna about the same 
length as the more distal pinnules; microscales acic- 
ular, pale, without glandular cells. Sorus terminal 
(rarely subterminal) on an acroscopic vein branch, 
without true indusia or less commonly with a spat- 
ulate or rarely reniform to peltate indusium 0.05- 
0.07 mm diam., the spatulate indusia composed of 
a narrow proximal portion with cells elongate par- 
allel to the axis and a broader distal portion with 
cells irregularly placed and shaped. Spores with a 
well-developed long-cristate perispore. 


Distribution. Endemic to the subalpine rain- 
páramo areas and disturbed sites in the tropical 


370 Annals of the 


Missouri Botanical Garden 


(— 


Ss 


ist SE a S SS 
SAL 
FIGURE 2. Polystichum talamancanum. — A. Le 


Pinnule of plant from sunny site.—D. Petiole squa 
—H. Medial edge of petiole scale. —I. Petiole scale. 


af. —B. Pinnule of plant from shade site, sori removed. —C. 
mule. —E, F. Rachis squamules. —G. Pinna-rachis squamule. 


montane rainforest of Costa Rica and western Pan- 


and an unknown diploid species that is also a pro- 
amá, 3,000-3,500 m. 


genitor of Costa Rican plants of the widespread 
tetraploid P. orbiculatum (Desvaux) Gay (Bar- 
rington, 1985b; Barrington et al., 1986, and un- 
published data). (Polystichum orbiculatum is the 


Polystichum talamancanum is the allotetra- 
ploid derivative of P. concinnum (described above) 


Volume 76, Number 2 
1989 


Barrington 
Polystichum 


correct name of the widespread Latin American 
alpine Polystichum commonly known as Polysti- 
chum polyphyllum (Presl) Presl.) Polystichum 
talamancanum is prominent in rock clefts and on 
roadbanks along the Pan-American Highway above 
treeline in Costa Rica. It tends to have more patent 
pinnae and flat pinnules in shade. The indusia are 
variously developed: some sporophytes never have 
indusia; others regularly produce spatulate indusia. 
This variation complicates discrimination of P. con- 
cinnum, P. talamancanum, and their backcross 
hybrid. 
Polystichum talamancanum, endemic to the 
region above treeline in the Sierra Talamanca, may 
be confused with Polystichum orbiculatum and 
allied taxa. Polystichum orbiculatum in Costa Rica 
has flaccid petiole scales that are paler and more 
orange than those of P. talamancanum and has 
long, delicate cilia throughout. In addition, P. or- 
biculatum has apically as well as laterally revolute 
pinnules, and it is never indusiate, whereas 
talamancanum has pinnules that are revolute only 
laterally and sometimes has indusia. 

n the Andes from Venezuela to Peru, an array 
of plants has been collected from those like P. 
orbiculatum in Costa Rica to plants much like P. 
talamancanum. The Andean plants most like P. 
talamancanum differ from it in lacking indusia 
and having broader, larger petiole scales with nar- 
row white borders bearing short, stiff cilia; if they 
are hybrids, their progenitors differ from those of 
P. talamancanum and are best determined as P. 
pycnolepis (Kze. ex Kl.) Mett. The taxonomy and 
evolution of P. orbiculatum and its allies in the 
Andes is a complex and intriguing problem: the 
hybrid origin of P. talamancanum on the Sierra 
Talamanca is likely to be one part of a much 
broader, Andean set of evolutionary events, in- 
cluding the origin of P. orbiculatum and other 
high-altitude Polystichum species. 


Additional specimens examined. COSTA RICA. CAR- 
TAGO: Cerro de la Muerte, Pan-American Highway, 5 km 
above Millsville (about 8 km above Nivel), Cordillera de 
Talamanca, 3,400-3,500 m, 25 July 1949, Holm & 
lltis 590 (MO). SAN JOSÉ: Cerro Asunción del Macizo de 
Buena Vista, 3,325 m, 31 de 1964, Jiménez M. 2235 
(CR). PANAMA. CHIRIQUÍ: mit of Chiriquí Volcano, 
3,374 m, 27 Feb. 1918, Killip 5459 (US). 


Hybrids. Polystichum talamancanum hybri- 
dizes with three species on the Sierra Talamanca 
in Costa Rica, making recognition of the taxa in- 
volved difficult. It hybridizes with its diploid pro- 
genitor P. concinnum to yield a highly variable 
triploid backcross, reported by Barrington (1985b) 
as P. muricatum X P. polyphyllum. Represen- 
tative specimens of this hybrid are Barrington 


596, 719, and 1278 (all CR, GH, VT), all from 
Cerro de la Muerte, Costa Rica. In addition, sterile 
putative hybrids of P. talamancanum with P. or- 
biculatum (Barrington 1283, CR, GH, VT) and 
P. speciosissimum (Barrington 1500, CR, GH, 
VT) have been collected from disturbed sites above 
treeline in the Sierra de Talamanca. 


Polystichum bulbiferum Barrington, sp. nov. 
TYPE: Ecuador. Prov. Zamora-Chinchipe: road 
Loja-Zamora, km 39, 1,600-1,650 m 
(79%04'W, 3°59’S), 21 April 1973, L. Holm- 
Nielsen, S. Jeppesen, B. Løjtnant & B. Øll- 
gaard 4095 (holotype, AAU). Figure 3. 


Filix terrestris. Rhizoma parvum, ascendens. Squamae 
petiolarum lanceolatae, papyraceae, emarginatae (con- 
formes), umbrinae, ciliis longis tenuibusque. Lamina 0.3- 
0.4 m longa chartacea, acuminata, pinnata, abaxialiter 
ha Pe : up 


1 r r I I 
LE . E E “reg 


bus fuscis. Pinnae patentes, ena Aisa: falcatae, 
attenuatae, vix spinu ulosae; pinnae mediana e circa 7 cm 
longae 1.5 cm latae, i 


Sorus abaxialis, in venis acroscopicis fasciculi venarum. 


Plant terrestrial. Stem ca. 0.5 cm long, 1.0 cm 
diam., erect from an ascendent base. Petiole scales 
lanceolate, papyraceous, conform, light yellow- 
brown, petiole-scale cilia long and delicate; petiole 
squamules narrow-lanceolate, crumpled, pale or- 
ange-yellow, with abundant short cilia. Lamina 0.3- 
0.4 m long, chartaceous, acuminate, 1-pinnate, 
with a scale-covered bulbil abaxially near the distal 
end of the rachis; abaxial rachis scales filiform, 
deep brown, with very sparse short cilia. Pinnae 
ca. 15 on each side of the rachis and attached to 
it at right angles, flat, auriculate, falcate-attenuate, 
crenate (crenulate to entire above), weakly spi- 
nulose at auricle and pinna apex; the median pinnae 
ca. 7 cm long and 1.5 cm wide, the basal pinnae 
ca. ^4, as long; pinna-rachis squamules abundant, 
dark brown, filiform, without cilia; microscales of 
costules and veins acicular, pale, without CA 

cells. Sorus abaxial on an acroscopic vein bran 
indusia absent. Spores with a well-developed folded 
perispore. 


In its leaf dissection and pinna shape, Polysti- 
chum bulbiferum resembles the Madagascar species 
P. kalambatitrense Tard. and P. maevaranense 
Tard. Among New World polystichums, it is the 
only one-pinnate species with bulbils attached below 
the apex of the leaf. In contrast to the Madagascar 
species, it is exindusiate, as are all of the polysti- 
chums endemic to the northern Andes. The detec- 
tion of a morphologically distinctive species such 


372 Annals of the 
Missouri Botanical Garden 


Qs 


as Sa M 
FIGURE 3. Polystichum bulbiferum. — A. Leaf with bulbil on rachis (at arrow). — B. Pinna outline. —C. Portion 


of pinna with sori removed. — D. Petiole squamule.— E, F. Rachis squamules.— G. Medial edge of petiole scale. 
—H. Petiole scale. 


Volume 76, Number 2 
1989 


Barrington 373 


Polystichum 


as P. bulbiferum only within the last 15 years 
suggests that major morphological trends in the 
diversity of the genus in Latin America may still 
be undocumented by collections, in spite of recent 
intensive work. This strongly suggests that main- 
taining substantial collecting programs in tropical 
forests between 2,000 and 4,000 m is a priority 
if we are to represent the morphological diversity 
in this and other genera adapted to the wet montane 
forests. 


Additional specimen examined. | ECUADOR. ZAMO- 


1979, Luteyn, Lebrón-Luteyn & McAlpin 6634 (AAU). 
New CoMBINATION 


P ly ick bint i (Hooker & Ar- 
nott) Barrington, auod. nov. Aspidium sub- 
integerrimum Hooker & Arnott, The Botany 
of Captain Beechey's Voyage 52. 1832. TYPE: 
Chile. Biobio: **Conception" 
collector unknown (holotype, K). 


[Concepción ], 


This distinctive species has black, coriaceous 
petiole scales, and its two-pinnate leaves have widely 
spaced pinnae and widely spaced, coriaceous pin- 
nules with auricles pinnatifid only in the basal pin- 
nules of a pinna. It may be well represented in 
other herbaria besides GH. Some specimens from 
northern Peru reminiscent of P. subintegerrimum 
have browner petiole scales, more dissected and 
less coriaceous pinnules, and no indusia. These may 
be hybrids or a hybrid species including a Peruvian 
progenitor and P. subintegerrimum. 


i-i si prias CHILE. I Lonco (Con- 
cepción), 22 Sept. 1928, Espinosa 1, 2, 5 (GH); Con- 
cepción, Nov. 1937. i Jaffuel Rd LA ARAUCANIA 
(CAUTÍN): Depto. Victoria, along the road to Laguna Mal- 
na ye de la Culebra and Laguna Verde, sandy places, 

1,050 m, 15 Mar. 1939, J. L. Morrison & R. 
^ae dad 17522 (GH). Los LAGOs: Chiloe, Fuerte de 
Ahui (Ancúd), Jan. 1929, M. R. Espinosa 7 (GH); Chiloe, 
Ancúd, n 1868, R. O. Cunningham s.n. (GH); Corrál, 
oth H. Gunckel 2050 (GH); same locality, 16 

H. Gunckel 2082 (GH); Niebla, 13 Dec. 
1931, H. poe 2865 (GH). 


Exindusiate specimens: PERU. ANCASH: Depto. Yun- 
gay, N side of first lake, Llanganuco, 11,500 ft., 30 July 
1960, S. G. E. Saunders 517 (F); 10 km NE of Yungáy, 
near Laguna Llanganuco, 3,450 m, 22 Oct. 1965, Tryon 
& Tryon 6557 (GH). 


LITERATURE CITED 
BARRINGTON, D. S. 1985a. Hybridisation in Costa Rican 


jr tichum. Proc. Royal Soc. Edinburgh 86B: 335- 


19 The morphology and origin of a new 
Polystichum hybrid from Costa Rica. Syst. Bot. 10: 


i Dm The present evolutionary and taxo- 
mic status of the fern genus Polystichum: the 
1984 Botanical Society of America Preridophyte Sec- 


. WERTH. "Ls 

reticulátion; and species concepts in ferns. Amer. 

Fern J. 78 (in press). 

. PARIS & T. A. RANKER. 1986. System- 

die alerce from EEA and stomate size in the 
ferns. Amer. Fern J. 76: -159. 

"a C. 1906. P Filicum. Hagerup, Co- 


Dicono * "1912. erp be studies on the fern 
nus Polystichum in Japan, Ryukyu, and Taiwan: 
i. Rep. Tokyo Bunrika Daigaku, Sect. B 1 


HOOKER, W.J. 1862. Species Filicum, Volume 4. W. 
Pamplin, London. 

MAXON, hu R. 1909. Studies in HUP a de 
fer . 2. Contr. U.S. Natl. Herb. 13: 1-43. 
Mos. n 1858. Ueber einige Farngattu ur. IV. 
Phegopteris und Aspidium. L. Broenner, Frankfurt 

a.M. 


PnEsL, C. B. 1836. Tentamen Pteridographiae. T. Haase, 


rague. 
RorH, A. W. 1799. Tentamen Florae Germanicae, 
Volume 3. j. G. Mueller, Leipzig. 
SMITH, Ar B 1981. Part 2. Pteridophytes. In: D. E. 
(editor), Flora of dai California Acad- 
y of Sciences, Gan Fra 
Speirs ^w. T. 1983. ey Iun: David and Charles, 
Newton Abbot, England. 
STOLZE, R. G. 1981. Ferns and fern allies in Guatemala. 
part 2. Polypodiaceae. Fieldiana, Bot. n 522. 
Tacawa, M. 40. Polystichum in Non. Korea and 
Formosa I. Acta ae Geobot. 9: 119-138. 
Tryon, R. M., JR. 1970. The classification of " Cy- 
aibescoas. Contr. Gray Herb. 200: 3-53. 


NEW OR NOTEWORTHY 
ORCHIDS FOR THE 
VENEZUELAN FLORA. 
VII. ADDITIONS IN 
MAXILLARIA FROM THE 
VENEZUELAN GUAYANA' 


Germán Carnevali? and Ivón Ramírez? 


ABSTRACT 


T w Maxillaria species from the Venezuelan Guayana, M. foldatsiana and M. santanae, are described and 
illustrated. One highly variable species, M. auyantepuiensis, is discussed, and a new subspecific taxon is proposed. 
Maxillaria tenuis is noted as a new record for the Venezuelan flora. 


Maxillaria Ruiz & Pavón is the largest genus 
of the subtribe Maxillarinae, with about 400-4 
species ranging from Mexico and the West Indies 
to southern Brazil. The genus reaches its highest 
development in the Andes, but le America, 
southern Brazil, the Amazonian a and the 
Guayanas are well represented. The species of 
Maxillaria usually grow in places of heavy rainfall 
but often can be found in places with a marked 
dry season. They grow in almost every vegetation 
association and at any elevation from sea level to 
near the snow line but are especially common in 
rainforest or cloud forests at 500-2,000 m. Most 
members of the genus are epiphytic, but it is not 
unusual to find lithophytic or even terrestrial species, 
particularly in open, humid places. The vegetative 
morphology is very variable, ranging from tiny 
plants less than 1 cm high to plants 2 m long. 
They are usually pseudobulbous, sympodial plants, 
but pseudobulbless, almost monopodial species are 
not unknown. Rhizomes may be short, making 
caespitose plants, but many species have long 
creeping primary stems; these may be appressed 
to substrate for their entire lengths or only attached 
at their bases to the host, making erect, ascendent, 
or pendulous plants 

Maxillaria is characterized generically by sol- 


itary flowers originating from the pseudobulb base 
or from the internodes of the rhizome (the Cama- 
ridium alliance). The flowers may be minute to 
very large and showy and are variable in many 
respects, although the column is always basally 
produced into a more or less well-developed col- 
umn-foot to which the labellum is hinged. Orni- 
thidium Salisb. has a rigidly attached, unhinged 
labellum, and thus we feel it should be recognized, 
even though many authors do not do so. As a 
member of the subtribe Maxillarinae, all Maxil- 
laria species have four dorsiventrally flattened pol- 
linia in two unequal pairs, more or less well-de- 
veloped, usually semilunate viscidium, and variously 
sized and shaped stipites or no stipe at a 

In Venezuela there have been recorded about 
75 species of Maxillaria, of which 47 are known 
in the Venezuelan Guayana. All major groups with- 
in the genus are represented in southern Venezuela. 
In terms of numbers of species, Maxillaria is only 
surpassed in the Venezuelan Guayana by Pleu- 
rothallis R. Br. (ca. 57 species) and Epidendrum 
L. (ca. 51 species). The next-largest genus is Oc- 
tomeria R. Br. (ca. 29 species). 

A survey of Maxillaria species, associated with 
the Orchidaceae treatment for Steyermark's Flora 
of the Venezuelan Guayana (Carnevali et al., in 


' We are grateful to Lic. Bruno Manara for the drawings and criticisms of the Latin Wr es and to Dr. Julian 


S 
suggestions and a reading of the manuscr 
santanae. For cr 


of the Venezuelan 
o Foldats, Gilberto Morillo, and jai Carnevali for their 


ipt. We a are indebted to Mr. Darko Elakovich for his drawings of M. 
cial access to the orchid collection of the Herbario Nacional de Venezuela (VEN) we thank the 


is x a iai ee ew its curators Dr. Gilberto Morillo and Lic. Libia Laskowski. 


otanico de Car , Herbario Nacion 


al de Venezuela, INPARQUES, Apartado 2156, Caracas 1010-A, 


Venezuela. Current ace Mison Bonae Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. 
ANN. MISSOURI Bor. GARD. 76: 374-380. 1989. 


Volume 76, Number 2 Carnevali and Ramírez 375 
9 Venezuelan Maxillaria 


FiGURE 1. A, B. Maxillaria foldatsiana. — A. Flowering habit. — B. Labellum, flattened. — C, D. Maxillaria 
tenuis. — C. Flowering habit. —D. Labellum, flattened. 


prep.) has revealed: 1) a new subspecies of M. THE MAXILLARIA AUYANTEPUIENSIS COMPLEX 
auyantepuiensis Fold. and a new species closely 
related to it, 2) an undescribed taxon in the Di- 
crypta. group of the genus, and 3) a new record 
for the Venezuelan flora, M. tenuis C. Schweinf. 


Our studies of the complex of taxa centered 
around M. auyantepuiensis Fold., which includes 
M. desvauxiana Reichb. f., M. neophylla Reichb. 


376 


Annals of the 
Missouri Botanical Garden 


f., and a few others, have revealed a previously 
undescribed species and a new infraspecific taxon 
referable to M. auyantepuiensis. Here we provide 
the descriptions of both new taxa. 


Maxillaria foldatsiana Carnevali et I. Ramirez, 
sp. nov. TYPE: Venezuela. Bolivar: Uaipante- 
pui, 1,200 m, T. Koyama & G. Agostini 
7378 (holotype, VEN). Figure 1A, B. 


Species haec M. antepuiensi affinis, sed differt: 
labello triplo longiore « q latiore (duplo longiore quam 
latiore in M. a ntepuiensi) et lobulo medio conspicue 
unguiculato (subsessili i in M. auyantepuiensi). 


Apparently epiphytic, erect herbs, 27-30 cm 
high. Rhizome abbreviated with aggregated pseu- 
dobulbs. Pseudobulbs narrowly ovoid, 4.7-5 c 
long, 1.3-1.6 cm wide at their broadest, mono- 
phyllous, concealed by appressed, scarious sheaths. 
Leaves coriaceous, erect, narrowly elliptic, acute, 
basally attenuate into a subcylindric pseudopetiole 
4-4.5 cm long, leaf total length 23-24.5 cm, the 
blade 18.5-20 cm long, 2.3-2.6 cm wide. Inflo- 
rescences originating from the pseudobulb base, 
flowers solitary; peduncle subterete, entirely con- 
cealed by 4-5 tubulose, scarious sheaths, 3-5 cm 
long. Flowers medium-sized for the genus, appar- 
ently vertically erect. Perianth segments scarcely 
open. Floral bracts elliptic, acute, 1.4-1.5 cm long, 
0.4-0.5 cm wide. Pedicellate ovary apparently 
subcylindric, 15-17 mm long. Dorsal sepal nar- 
rowly oblong-elliptic, acute, 20-20.5 mm long, 4- 
4.5 mm wide. Lateral sepals narrowly triangular, 
acute, 20-21 mm long, basally 3.5-4 
Petals narrowly elliptic, slightly obovate, acute or 
narrowly obtuse, 19 mm long, 4-4.5 mm wide. 
Labellum parallel to the column, in general outline 
narrowly oblong, 17-18 mm long, 4.5 mm wide 
at its broadest, slightly pandurate-trilobate above 
the middle; central lobe elliptic, obtuse-rounded, 
slightly concave, 7 mm long, 2.5 mm wide, basally 
attenuate in a short unguiculus about 2 mm lon 
and wide, the narrowly oblong longitudinal callus 


mm wide 


apically rounded and centrally concave, 7 mm long, 
0.8 mm wide. Column straight or slightly recur- 
vate, hemicylindric, ventrally concave, 8 mm long 
and 2 mm wide at the thickened apical zone. Cli- 
nandrium apiculate. 


This species is very similar to M. auyantepuien- 
sis, from which it differs in its inflorescences longer 
than the pseudobulb (vs. subequal or shorter), nar- 
rowly elliptic floral bracts (vs. widely elliptic), petals 
to 4.5 mm wide (vs. at least 6 mm in M. auyan- 
tepuiensis), and conspicuously narrower labellum 


(in M. auyantepuiensis it is approximately twice 
as long as wide vs. more than three times longer 
than wide in M. foldatsiana). Further, the labellum 
of M. foldatsiana is less trilobulate. As M. fol- 
datsiana is only known from the type collection, 
and the differences between the two species are 
not deep, we feel that more material may show M. 
foldatsiana to be only a variation of M. auyan- 
tepuiensis. However, we have studied many spec- 
imens of M. auyantepuiensis (alive and dried) and 
have never noticed intergradation in the differential 
characters, so we think that they are best treated 
as closely related but distinct species. 

This apparently scarce species has been col- 
lected only once despite repeated collecting trips 
to the summits of the tepuis in the last ten years. 
Perhaps it is endemic to Uaipantepui. 

We take great pleasure in naming this species 
after our friend and colleague Dr. Ernesto Foldats, 
orchidologist. He was the first to notice the new 
species as undescribed. 

Maxillaria auyantepuiensis is very common in 
the Gran Sabana, in southeastern Bolivar, Vene- 
zuela. Surprisingly, it is seldom collected. This 
species comprises two infraspecific taxa, one of 
which has remained undescribed. 

Maxillaria auyantepuiensis subsp. auyante- 
puiensis is terrestrial or lithophytic, always grow- 
ing on sandy soils or on acid sandstone, forming 
large colonies in more or less open places as scrub 
or savannalike tepui formations. It is usually ex- 
posed to full sun or resides under the light shade 
of shrubs. Occasionally it reaches be p iin 
margin or even its interior, provide 
light and is always over sandstone DUREE or 
sandy soil. Morphologically, M. auyantepuiensis 
subsp. auyantepuiensis is highly variable with fu- 
siform to (rarely) spherical pseudobulbs and sessile 
to long-petiolate leaves. The sheaths that conceal 
the pseudobulb do not defibrate but remain intact. 
The inflorescences vary from shorter than to longer 
than the pseudobulb and are always conspicuously 
peduncled. The flowers are held in an upright or 
suberect posture. The perianth segments are rel- 
atively long and narrow with petals 18-19 mm 
long, 4.3-4.5 mm wide (width/length = 0.38- 
0.42) the dorsal sepal is 20-21 mm long and 3.5 
mm wide (width /length = 0.16-0.17) 


Maxillaria auyantepuiensis Fold. subsp. epi- 
phytica Carnevali & I. Ramirez, subsp. nov. 
TYPE: Venezuela. Bolivar: bosque hümedo en 
las vacinidades del km 119 al sur de El Do- 
rado, 130 m, 12 Jan 1964, J. A. Steyermark, 


Volume 76, Number 2 


Carnevali and Ramírez 377 


1989 Venezuelan Maxillaria 
TABLE 1. Characters distinguishing subspecies of M. auyantepuiensis. 
Subspecies 
Characters auyantepuiensis epiphytica 
Substrates Sandy soil or sandstone outcrops Epiphyte on trees 
Habitats Various tepuilike associations Rain and cloud forest 
Pseudobulb sheaths Intact Defibrating 
Pseudobulb shapes Spherical to ellipsoid Spherical 
Leaves Sessile to long-petiolate Always long-petiolate 
owers Erect to suberect Spreading horizontally 
Petals length x width 18-19 5-16 x 5 mm 
Width/length 42-0.45 0.30-0.38 
Dorsal sepal length x width 20-21 x 3.5 mm 16-17 x 4.5-5 mm 
Width/length 0.16-0.17 0.27-0.31 


G. C.K. & E. Dunsterville 92993 (holotype, 
VEN) 


Subsp. haec subsp. auyantepuiensi vaginis dissolutis, 
tepalis proportione latioribus differt itemque habitu epi- 
phytico et in silvis pluvialibus incola. 

The subspecies are distinguished in Table 1. 
Both subspecies have been illustrated in Dunster- 
ville & Garay (1965: 181, 498). Plants A, C, and 
D there depict the variation to be expected in M. 
auyantepuiensis subsp. auyantepuiensis. Plant B 
represents our new M. auyantepuiensis subsp. epi- 
phytica. 

The new subspecies comprises populations of 
epiphytic plants from rain or cloud forest at vari- 
able altitudes from 600 to 1,400 m, with spherical 
or broadly ellipsoid pseudobulbs that are small in 
proportion to leaf length and are concealed by 
defibrating sheaths. These pseudobulbs occur on a 
short to long, obviously creeping rhizome, which 
adheres to the bark of the host tree. The leaves 
are always long-petiolate and very elongate with 
respect to pseudobulb length. The almost-epedun- 
culate inflorescenses are always shorter than the 
pseudobulbs, and the flowers are almost held hor- 
izontally. The flowers are of proportionately smaller 
dimensions than in subsp. auyantepuiensis, with 
relatively broader perianth segments (dorsal sepal 
16-17 mm long, 4.5-5 mm wide (width/length 
0.27-0.31). The labellum is almost identical in the 
two subspecies. 

Maxillaria auyantepuiensis subsp. epiphytica 
is superficially similar to the Ecuadorian species 

. neophylla Reichb. f., but this has an acute 
central labellum lobe. The new subspecies also sug- 
gests a vegetatively reduced M. desvauxiana 
Reichb. f. from the Amazonian Basin, but their 
labella are very different, and M. desvauxiana does 
not have defibrating sheaths concealing the pseu- 


dobulbs. 


The new subspecies has a wider distribution than 
the typical subspecies and extends beyond the 
Guayana boundaries. We know of a population in 
Guatopo, Estado Miranda, of the Coastal Range, 
represented by Patrszeck s.n., 13 Feb. 1963 
(VEN), also a population from Barinitas, Estado 
Barinas, in the Andean foothills, from which we 
have living material but no voucher. 

The typical subspecies appears to be restricted 
to the acid sandy soils of the Roraima Formation, 
and this may explain a limited range in geographic 
terms even though it appears to be locally abundant 
in its natural habit. 


Paratypes. VENEZUELA. BOLÍVAR: Selva hümeda con 
árboles de un promedio de 25-30 m de altura en el 
drenaje del Rio Cuyuni, km 140-141.5 al sur de El 
Dorado, 1,300-1,380 m, 22-28 Dec. 1970, J. A Stey- 
ermark, & E. Dunsterville 104324 (VEN); 
lir Guaiquinima, Río Szczrbanari (Río Carapo), 750 

, 20-25 Jan. 1977, J. A. Steyermark, G. C. K. & E. 
Boc dh 113219 (VEN). MIRANDA: Guatopo, 13 Feb. 

63, Patcrzck s.n. (VEN) (Colección en líquido Duns- 
terville 252). 

Study of the Dicrypta group of the Venezuelan 
Guayana has revealed an apparently undescribed 
taxon: 


Maxillaria santanae Carnevali & I. Ramirez, 
sp. nov. TYPE: Venezuela. Territorio Federal 
Amazonas: Departamento Atabapo, Cerro 

Marahuaca, porción central. Bosque con ár- 

boles hasta 20 m, aprox. 1,000 m. Floración 

en cultivo en el orquideario del colector, 11 


Nov. 1986, G. Santana 1 (holotype, VEN). 
Figure 2 


Species haec M. Mobi des et M. superfluae ~~ 
sed ani nostra habito conspicue parviore, petalis e 
labello pro portione latioribus et labello unicalloso cinn 
barino vel cinnabarino-roseo lobo centrali late triangulari 
discrepat. 


378 


Annals of the 
Missouri Botanical Garden 


FIGURE 2. Maxillaria santanae. — A. Flowering habit. — B. Foliar apex. — C. Lateral view of flower at anthesis. — 


D. Perianth segments, flattened. — E. Labellum and colum 


Small to medium-sized, epiphytic, caespitose, 
erect herbs, 11-23 cm high. Rhizome short and 
proportionately thick, entirely concealed by scar- 
ious sheaths. Roots proportionately thick and long, 
pale brown. Pseudobulbs aggregate, ovoid-ellipsoid, 


n. 


ancipital, 2.1-3 cm long, 0.6-1 cm wide, apically 
unifoliate, clothed with 4—5 sheaths, the innermost 
2-4 of these with foliar blades, the blades falling 
after 2-3 years of growth, leaving the sheaths 
persisting and becoming grayish brown in fresh or 


Volume 76, Number 2 
1989 


Carnevali and Ramírez 379 


Venezuelan Maxillaria 


dry condition. Leaves fleshy-coriaceous, erect or 
patent-erect, the outer leaves 10-15 cm long from 
the articulation, the apical leaf 14-21 cm long, 
7-10 mm wide when fresh, 5-9 mm wide when 
dry, basally attenuate and forming a pseudopetiole 
1-2.3 cm long, the blade apex acute and deeply 
oblicuous-bilobulate (one of the halves of the blades 

mm longer than the other). Inflorescences one- 
flowered, originating from the leaf-sheath axils one 
at a time. Peduncle subterete, fleshy, 17 mm long, 
concealed by 3-5 subimbricate, distichous sheaths. 
Floral bract similar to the peduncle bracts, nar- 
rowly triangular, acute, membranose, 6 mm long, 
2.5-3 mm wide. Pedicellate ovary subterete, 9- 
10 mm long, 2.5 mm thick. Flowers medium-sized, 
campanulate, erect or suberect; perianth segments 
fleshy, subparallel to the column, greenish yellow, 
the petals with or without a few small brown-orange 
dots, the labellum pale orange or rose-lilac with 
orange-brown basal patches and a salmon-colored, 
tomentose callus. Sepals elliptic-oblong, acute; the 
laterals slightly oblicuous, 11-13 mm long, 4-5 
mm wide, with a dorsally thickened apex. Petals 
narrowly obovate-elliptic to narrowly elliptic, 
subfalcate, acute, 10- 10.5 mm long, 2.7-3.2 mm 
wide. Labellum basally articulate with the column- 
foot, in overall shape elliptic, subtrilobate about the 
apical fourth, 9-9.5 mm long, 5-5.5 mm wide 
between the margins of the lateral lobes; lateral 
lobes suberect and making a concave labellar disc; 
central lobe broadly ovate or broadly triangular 
obtuse, 3.8-4 mm long, 2.8-3.2 mm wide, fleshier 
than the rest of the labellum and with a central 
concavity originating where the callus ends, the 
disc with a thickened, waxy, tomentous longitudinal 
callus, this 6-7 mm long, 1 mm wide. Column 
semiterete, slightly thickened, apically arcuate, 6.5— 

m long; column foot 2-3 mm long. Anther 
subconic, 2 mm long, obtuse, the basal margins 
erose. 


Paratypes. VENEZUELA. BOLÍVAR: confluencia del Cano 
Makarupai con el Río Akaruai, unos 10-15 km al SO de 
la Misión de Wonken, ca. 850 m, en bosque macrotermico 
medio/ medio, caracterizado por una gran abundancia de 
epifitas debido a los rápidos de la confluencia. Floración 


en cultivo, Maracay, 2 Jan. 1984, G. Carnevali 1445 
(VEN). 


This relatively scarce species has a wide range 
within the Venezuelan Guayana. One of the authors 
collected living plants of it in 1982 in the sur- 
roundings of Peray-tepui (W of Sta. Elena de Uai- 
ren, Gran Sabana, Estado Bolivar), but no voucher 
was made. The affinities of M. santanae lie with 
all the species of the Dicrypta group: M. crassi- 
folia Reichb. f.; M. superflua Reichb. f., and per- 


haps M. bicallosa (Reichb. f.) Garay, M. char- 
tacifolia Ames, and others. This is a difficult group 
in need of revision—the characters distinguishing 
the component members are not well defined and 
seem unreliable. Among the Dicrypta group, M. 
santanae can be easily recognized by its small 
vegetative stature; very narrow leaves; broadly 
ovate or broadly triangular, obtuse central labellum 
lobe; and orange-yellow or rose-lilac labellum, which 
has a callus in the disc and lacks another in the 
central lobe as is usual in other species of the group. 
We are pleased to name this species after Dr. 
Gustavo Santana, an enthusiastic orchid collector 
and knowledgeable amateur orchid grower. He col- 
lected the living plant from which the holotype 
specimen was prepared and kindly provided fresh 
vegetative and floral material for study. 


Maxillaria tenuis C. Schweinf., Bot. Mus. Leafl. 
11: 289, T. 18, fig. 1, c-d. 1945. TYPE: Perú. 
Loreto: 
forest, Klug 1045 (holotype, 

C, D. 


Mishuyaco, near Iquitos, 100 m, in 


US). Figure 


Small to medium-sized epiphytic herbs. Rhizome 
abbreviated. Pseudobulbs cylindric, ellipsoid or ob- 
longoid, apically unifoliate, 1.5-2.3 cm long when 
mature, 5-7 mm wide, concealed by oblong-ovate, 
acute, and acuminate sheaths, these verruculous- 
pustulose without, the innermost bearing leaf blades. 
Leaves coriaceous, 11-14 cm long, 6-8 mm wide, 
the blades linear, acute, the base attenuate into a 
short pseudopetiole 3-15 mm long. Inflorescences 
erect, 1-flowered, solitary, originating from the base 
of the young pseudobulbs. Peduncle subterete, 1.5— 
3.2 cm long, concealed by 2 elliptic, acute, acu- 
minate sheaths, these verruculose without, 8-11 
mm long, slightly longer than internodes. Floral 
bracts almost identical to the last peduncle bract, 
9-11 mm long, about 6 mm wide. Ped y 
subterete, longitudinally sulcate, verruculose, 11- 

2 mm long. Flowers medium-sized but large for 
the plant, with white tepals and a yellow and white 
labellum having purple nerves. Perianth segments 
subcoriaceous, parallel to the column. Dorsal sepal 
very narrowly elliptic, acute and acuminate, con- 
cave basally, 18-27 mm long, 3-4 mm wide. Lat- 
eral sepals narrowly oblong to linear-lanceolate, 


acuminate, slightly oblicuous, 18-27 mm long, 4 
mm wide basally, decurrent on the column-foot 
and forming with it a short mentum. Petals linear- 
lanceolate, acuminate, 16-23 mm long, 2-2.3 mm 
wide. Labellum fleshier than the other perianth 
segments, broadly elliptic to elliptic-ovate overall, 
trilobate near the apical third, 9.5-10 mm long 


380 


Annals of the 
Missouri Botanical Garden 


between the apices of the extended lateral lobes, 
5-6 mm wide; lateral lobes erect and narrow, end- 
ing in a free ovate-rounded portion about 1 mm 
long with the apical margins minutely erose; central 
lobe very fleshy, 3.5-4 mm long, 2 mm wide, 
triangular ovate, acute or obtusate, minutely apic- 
ulate subapically, the ventral face rugulous-papil- 
lose and longitudinally sulcate. Disc along its entire 
length with an oblongoid callus, 5-6 mm long, with 
an apically rounded surface. Column proportion- 
ately short, about 4.5 mm long, semiterete, basally 
produced into foot about 2 mm long. 


Additional CRM examined. VENEZUELA. TERRI- 
TORIO FEDERAL AMAZONAS: wp Pe 2 San pid de 
Rio Negro, on he road to Solano, 120 m, Dec. 
1986, H. L. Clark 8098 (VEN). 


Distribution. Known from the rain forests be- 
tween 80 and 200 m in the Amazonian Basin of 
Peru, Brazil, and Venezuela. 


This is the first record of this species for Ven- 
ezuela. Since the Venezuelan material shows a few 
differences with the other known populations, here 
we offer a complete description with the full range 
of variation. Maxillaria tenuis seems to be rather 
variable. The type material from Iquitos, Amazo- 
nian Peru (Klug 1045, US), has sepals about three 
times longer than labellum (27:9.5 mm) and a 
triangular-ovate central labellar lobe. The second 
collection known, collected in Amazonian Brazil (G. 
A. Black 48-2676, IAN; G. A. Black 48-2747, 
IAN), has a very narrow central lobe (half the 
width of the type material) and an apparently bi- 
lobated labellar callus. Based on this Brazilian ma- 
terial, a var. amazonica Pabst was described (Pabst, 
1955). All persons experienced with Maxillaria 


——— & F. = 


herbarium material know how difficult it is to re- 
hydrate and reconstruct labella in these species, 
especially the central lobe and calluses, so we won- 

er if the narrower central lobe and bilobated callus 
are not artifacts. Provided they are not, we wou 
think that var. amazonica could stand as a good 
variety or subspecies, even though in Pabst & 
Dungs (1975) they seem to have rejected Pabst’s 
variety without explanation. The third population 
of the species to be sampled is the one we report 
here, which seems to depart from the type and 
from Brazilian material in having shorter sepals, 
less than two times longer than labellum (18: 10 
mm). The labellum is identical with that of the type 
material. By orchid standards this seems to be a 
good difference for the recognition of at least a 
subspecies, but we feel that the evidence is too 
scanty, as is the known material, and further ma- 
terial could bridge the gap between the three known 
populations of this extremely variable species. 

Maxillaria tenuis is a well-marked species, es- 

pecially its vegetative morphology: oblongoid, elon- 
gate pseudobulbs, linear leaves, and verrucose-pus- 
tulose sheaths. San Carlos de Rio Negro is only a 

undred meters away from Colombia, across the 
Río Negro, so we would expect to find M. tenuis 
also in Colombia. 


LITERATURE CITED 


DUNSTERVILLE, G. C. K. & L. GaRaY. 1965. ring 
Orchids Illustrated, IIT. Andre Deutsche, 
PABST, G. 


3: 132-133, T. 34.B. 
1975. Orchidaceae Brasiliensis 
2: 183-18 


CONTRIBUCION AL GENERO 
MIMOSA (MIMOSACEAE)! 


Renée H. Fortunato? 


ABSTRACT 


A taxonomic revision of four species of the genus Mimosa L. from A 
as a species, and its affinity and taxonomic position are discu 
iven. Mimosa petraea Chodat 


(Burk.) Fortun., stat. nov. is proposed 
of | ans ich. and M. niederleinii Burk. are 


is presented. Mimosa dar ae 
ssed. Synony 


et Hassl. is restricted to s 


Paraguayan territory. All these taxa are briefly described and their geographic distributions are given 


Como parte de la revisión que se esta realizando 
del genero Mimosa L. para la Argentina, Bolivia, 
Paraguay, y Uruguay, se da a conocer el estudio 
taxonómico de cuatro especies presentes en el área. 
Estas especies pertenecen a la sección Mimosa 
(Bentham, 1842, 1875) por presentar flores ha- 
plostémonas. 

Dentro de dicha sección, segün ciertos carac- 
teres exomorfológicos (hábito, presencia de acú- 
leos, nümero de pinnas, nümero de foliolos) se 
agruparia a M. pauperioides y M. balansae en 
la serie Pectinatae Benth. y a M. niederleinii y 
M. petraea en la serie Pedunculosae Benth. 

En el sistema de clasificacion de Bentham, la 
ubicación taxonómica de muchas de las especies 
de Mimosa en cada serie es dudosa. A través del 
estudio biosistemático que se esta efectuando en el 
género para Austroamerica se espera poder se- 
leccionar los caracteres de valor taxonómico. 


mi 


. Mimosa pauperiodes (Burk.) Fortun., stat. 
nov. M. balansae M. Mich. var. pauperioides 
Burk., Darwiniana 8: 146, fig. 22. 1948. TIPO: 
Argentina. Corrientes: 4 Dec. 1941, Birabén 
5027 ns LP). Iconografia: A. Burkart 

n N. N. Bacigalupo, Col. Ci. 

INTA 53): 503, fig. 224, e-1. 


Sufrütice prostrado; tallos estrigosos, delgados, 
aculeados, a veces inermes; acüleos 1-2 de 0.5- 
3 mm long., infraestipulares o internodales (ge- 
neralmente ramas aculeadas e inermes en la misma 
planta). Pinna uniyugada de 1.5-5 cm long.; fo- 


liolos 4-18-yugados de 2-9 mm long. x 0.6-2 
mm lat., angostamente elipticos, asimetricos en la 
base, pubescencia combinada de pelos estrigosos y 
sedosos en ambas caras, a veces solo estrigosos en 
la cara abaxial. Inflorescencia con pedúnculos 1l- 
2 por axila de hasta 11 cm long., mayores que las 
hojas. Flores rosadas; cáliz hialino; corola de 2.5- 
3 mm long., glabra, 4-lobulada, lóbulos pubérulos 
en el ápice. Craspedio subtoruloso, hispido y pu- 
bérulo en replo y artejos. 


Distribución. Común en los suelos arenosos 
del estado de Rio Grande do Sul en Brasil y en las 
provincias argentinas de Entre Rios y Corrientes. 


Material selecto examinado. ARGENTINA. CORRIEN- 
TES: Dep. Curuzú Cuatiá: 12 km del acceso N a Curuzú 
Cuatiá, Cristóbal et al. 1961 (CTES). Dep. Empedrado: 
ruta 12 y Rio Empedrado, Schinini 12099 (CTES). Dep. 
Mercedes: L. R. Parodi 6232 (BA, SI); Millán 347 
(BA); Mercedes, Krapovickas et al. 20333 ( 

Justino Solari, Ibarrola 2587 (SI); ruta 14, km 37, 

SW de Carlos Pellegrini, Schinini e SI); Ayo. 
P mino a Ita Cora, Tressens et al. 
2269 (CTES); camino a la feria, orillas del Ayo. Las 
Garzas, P. Rodrigo 608 (LP). Dep. San Martin: Arrocera 
Drews, 8 km N de Carlos Pellegrini, Krapovickas et al. 
29215 (CTES). ENTRE RIOS: Dep. Federación: ruta 14, 

m 513, Burkart et al. 29331 (SI); ruta 14 al N de 
Federación, Troncoso et al. 2493 (SI); Santa Ana, Bur- 
kart 26234 (SI); A. L. Cabrera et al. 19353 (LP). Dep. 
Concordi R. Parodi 3962 (SI); ruta 14, km 15 al 
N ] : 


SI). BRASIL. RIO GRANDE DO S 
Porto Alegre, sur Xavier 17 (SI). 
Mimosa balansae var. pauperioides fue dife- 


1948) de M. balansae var. 


renciada (Burkart, 


' Se agradece a la Lic. Maria Mercedes Arbo por la lectura critica del manuscrito y a la Fundación Robert Bass 
por la beca otorgada para consultar la importante colección depositada en el Field Museum of Natural , History, 
Asi 


mismo, se hace extensivo el agradecimiento a los curadores de los herbarios aqui citados, 


a, INTA, Castelar 1712, Prov. de Buenos Aires, Argentina. Investigadora del 
T). 


Consejo Nacional de pia Científicas y Técnicas (CONICE 


ANN. 


Missouni Bor. GARD. 76: 381-385. 1989. 


382 


Annals of the 
Missouri Botanical Garden 


balansae por presentar foliolos más angostos y el 
nervio medio prominente. En el material estudiado 
se observó que ademas de los caracteres citados 
por Burkart, también se podia diferenciar a M. 
balansae var. balansae por poseer inflorescencias 
mayores que las hojas, y tallos inermes o con l- 
2 acüleos internodales y/o infraestipulares. Todas 
estas diferencias se mantienen constantes, e inclu- 
sive son más importantes que las existentes entre 
M. balansae y otras especies afines. Por lo aqui 
descrito, se propone considerar a M. pauperioides 
como especie independiente de M. balansae. 

En los ejemplares inermes es facil confundir a 
M. pauperioides con M. petraea. Ambas presen- 
tan afinidad en el tipo de hábito y nümero y forma 
de los foliolos; no obstante se diferencian por el 
largo del pedünculo, pubescencia, y tamano de los 
foliolos. 

Segün Bentham (1842, 1875), la serie Pecti- 
natae Benth. se caracteriza por agrupar sufrütices 
pequenos o raro arbustos, generalmente con acü- 
leos infraestipulares o internodales, pinna uniyu- 
gada, foliolos multiyugados (a veces hasta 6-8- 
yugados) pedúnculos 1(2) axilares o formando 
racimos compuestos apicales (desarrollo posterior 
de las hojas), racimos globosos, flores 4-meras, y 
craspedio setoso a hispido. A su vez, la serie Pe- 
dunculosae Benth. se diferencia de Pectinatae casi 
ünicamente por incluir especies inermes. En e 
estudio de ambos grupos supraespecificos, se ha 
observado que la presencia de acüleos es un ca- 
racter variable en muchas especies del género, 
como ocurre con M. pauperioides. Este taxon solo 
presenta algunas ramas con uno o dos acüleos 
pequenos, y a veces es una planta totalmente iner- 
me. Si se siguiera el criterio de Bentham, erró- 


Pedunculosae. Por otro lado, esta especie es más 
afin a M. petraea de la serie Pedunculosae. Como 
se ha senalado, en su mayoria, los caracteres di- 
ferenciales descriptos por Bentham para las series 
del género Mimosa no responden a un patrón 
estable. 


2. Mimosa balansae M. Mich., Mém. Soc. Phys. 
Nat. Genève 28(7): 52, fig. 18. 1883. TIPO: 
Paraguay: Trinidad, Mar. 1875, Balansa 
1479 (lectotipo, G, selecionado aqui; isolec- 
totipo, F); Balansa 1479a (sintipo, G, frag- 
mento F; fotografia n° serie F (ex Bt) — 1397, 
F, G, SI); Villa Concepción, in pratis, Balansa 
1835 (sintipo n.v.). Iconografia: A. Burkart 

in N. Troncoso & N. Bacgaupo Col. Ci. INTA 
6(3): 501, fig. 224, a-d. 1987 


Mimosa balansae M. Mich. var. robusta Chodat et Hassl., 
Bu a Herb. Boissier ser. x 4: 488. 1904. TIPO: 
doi guay: in campis San Bernardino, Nov. 1898/ 

9, Hassler 3507 (holotipo, G; isotipo, 
EU balansae M. Mich. 


ran aco: ad ripam occident 
$23°20', 23°30’, Oct.? 1903, Hassler 2355 (ho- 
lotipo, G). 


Sufrütice de + 15-80 cm alt., 
trado, robusto a + grácil; tallos con pubescencia 
pubérula y estrigosa, + patente, esparcida a densa, 
raro glabros; acúleos 2-5 mm 


erecto a pros- 


long., rectos a re- 
curvos, infraestipulares, presentes en ternos o va- 
riando su número de 1-6 (a veces 1 o 2(3) in- 
ternodales), raro sin acúleos. Pinnas uniyugadas; 
foliolos (3-)8-25-yugados, 5-15 mm long. x 2- 
elipticos a angostamente elipticos, 
estrigosos y pubérulos en ambás caras, hasta com- 


5 mm lat., 


pletamente glabros en el epifilo, nervio medio sub- 
central, poco visible en la cara abaxial (solo a veces 
en la base y/o en el ápice), borde estrigoso-pu- 
bérulo, no formando un margen nerviforme. Inflo- 
rescencia racemosa; pedúnculos 1-2 por axila, me- 
nores que las hojas. Cáliz 0.25-0.5 mm long., 
hialino, truncado a brevemente ciliado en el borde; 
corola 2-3.5 mm long., membranácea, glabra, ló- 
bulos pubérulos en el ápice. Craspedio 1-2 cm 
long. X 0.4-0.5 cm lat., subtoruloso, pubescente. 
Distribución. Crece preferentemente en es- 
partillares. Su distribución se extiende desde Bo- 
livia, Paraguay hasta Uruguay, S de Brasil y NE 
Argentino. 


Material pd examinado. GENTINA. CORRIEN- 
TES: Dep. Berón de Astrada: ars 3969 (LIL); 46 
km W de Ita Ibaté, Valencia, Schinini sae (CTES). 
Dep. Ca n 15 km E de Corrientes, camino a San Luis 
del Palmar, ruta i d 3247 (CTES): "16 km ENE 

e Cor maed 
(CTES): Molina Panta, PEE thas $e Cri 
(CTES, SI); Schinini 14134 (CTES, F); Perichón, Schi- 
nini & Martínez Crovetto 12252 (CTES). Dep. Empe- 
drado: Ea. Las Tres Marias, Pedersen 3063 ind Dep. 
General Paz: Navarro, ruta 5, : 
Corrientes, Anzótegui 336 (CTES). Dep. Merce ede s: Ea. 
Ita Caabo, Pedersen 5355 (LP); Jofré, Ibarrola 2698 
(LIL). Dep. Monte Caseros: Monte Caseros y empalme 
con ruta 14 a Curuzú Cuatia, Nicora 4630 (SI); en pajonal 
inundable, prox. 


Schinini et al. 17246 (CTES). Dep. 
Saladas: Fa. Rinesi, Pedersen 6627 (LP). Dep. S 
me: Vivero Forestal, paso de la Patria, Carnevali 2 
SI); San Cosme, Clos 5929 (BAB); ruta 12, entre Co- 
rrientes y Paso de la Patria, Nicora & Cámara Hernán- 
dez (BAA 272, CTES). ENTRE Rios: Dep. Concordia, 
Estación Agronómica, rig 967 (SI). Dep. Federa- 
cion: Barra del Mocor Troncoso et al. 1311 (SI). 
FORMOSA: Dep. Patirio, Estanislao del Campo, por ruta 81 


~ 


Volume 76, Number 2 
1989 


Fortunato 383 


mosa 


17 km hacia el N, Cabral, Molina & Fortunato 1194 
(BAB); Insfrán 1175 (BAB). Dep. Pilcomayo: Clorinda a 
Formosa, Morel 1685 (LIL); km 5, ruta 11, Laguna 
Blanca, Morel 2263 (CTES, LIL); al SE de Pto. Porteño, 
a 2 km, Morel 2350 (CTES, LIL); Clorinda, Morel 1150 
(CTES). Sin dep.: común en el campo Guaycalec, Jor- 
gensen 3212 (MO, SI). Sin loc., Jorgensen 3218 (MO). 
MISIONES: Dep. Candelaria, Meyer 18956 (LIL); Ya- 
bebyry, Montes 654 (CTES 


al. 28318 (SI). Sin dep.: Cerro Cora, Martínez Crovetto 
9496 (BAB). BRASIL. RIO GRANDE DO : Uruguaiana, 
Barra do Quarai, Rambo 4247 (SI). MATO GROSSO: Campo 
Grande, Nienstedt 401 (RB). PARAGUAY. ALTO PARAGUAY: 
Palmas Chicas, Chaco paraguayo, Rojas 7783 (SI); Pues- 
to Casado, Rojas 2139, 3042 (SI); Olimpo, Arenas 339 
(SI). AMAMBAY: 20 km S de Bella Vista, Krapovickas & 
Schinini 3267 1 (CTES). per bre Botánico, pra- 
dera entre gramineas, Roja 6 (SI); campos bajos 
entre Luque y Trinidad, pou P (SD); Yaguarón, 
orillas del m Yaguaron, Krapovickas et al. 12291 
(CTES, SI); Trinidad to Asunción, Osten 9168 (G); Ipa- 
caraí, Rojas 14278 (BAF, SI). CONCEPCIÓN: San Salvador, 
Alto Paraguay, Ea. Arrecife, Rojas 3036 (SI). CORDI- 
LLERA: Cordillera de Altos, Fiebrig 1005 (F, G); San 
Bernardino, Osten 9169 (G). GUAIRA: Azucarera de Te- 
bicuary, Ayo. Ihaca, Schinini 5864 (G, SI). PARAGUARI: 
Paraguari, Jardin he la Iglesia, M. Sanchez 19 (BAB 
SAN PEDRO: m S de San Estanislao, 24°40'S, 
56°21'W, See et eek 807 (BAB, PY); Distrito Lima, 
Pedersen 9440 (SI). Sin loc.: Pto. Yatayba, Daguerre 
(BA 28/924); Hiaty, Jorgensen 3624 (BAB, F, LP, SI); 
Central Paraguay, Moro URUGUAY. 


(SI); orilla del Rio Uruguay, Santa Sofia, Rosengurtt 
B-3659 ¥% (SI). SALTO: prox. Salto Grande, Osten 5373 
(SI) 


Se ha observado en el campo y en el material 
de herbario que M. balansae forma un complejo 
donde los caracteres descriptos (hábito, pubescen- 
cia, número de acúleos, largo de la pinna, y número 
y largo de foliolos) aparecen combinados en los 
diferentes ejemplares. Esta variabilidad imposibilita 
realizar una división infraespecifica natural de la 
especie, por ello se considera que los epítetos va- 
rietales robusta y rojasiana son solo probables 
formas ecológicas que presenta el complejo citado. 

Mimosa balansae var. rojasiana fue descripta 
por Hassler con foliolos glabrescentes en la cara 
adaxial. Al estudiar el material típico se pudo ob- 
servar que los foliolos son pubescentes en ambas 
caras. 

De los tres sintipos citados por Micheli en la 
descripción original de M. balansae, se han podido 
estudiar Balansa 1479 y 1479a (G, F). En esta 
nota se selecciona y se propone como lectotipo a 
Balansa 1479 depositado en el herbario G por ser 
un ejemplar más completo Es fr, y tres hojas de 
herbario) que Balansa 147 


3. Mimosa niederleinii Burk., Darwiniana 7: 
523. 1947. TIPO: Argentina. Misiones: Va- 


rana, 23 Ago. 1887, Niederlein 1107 (ho- 
lotipo, SI). Iconografia: A. Burkart, Darwinia- 
na 8: 162, fig. 26. 1948. 


Mimosa niederleinii Burk. var. riograndensis Burk., 
Darwiniana 7: 526, fig. 4. TIPO: Brasil. Rio Grande 
do Sul: Estrela, Mar. 1899, Dutra 382 (holotipo, 
HB; isotipo, SI). 

Arbusto o sufrütice de + 0.5-1.5 m alt., as- 
cendente; tallos ramificados, estrigosos y pubérulos, 
inermes. Estipulas 3-5 mm long. x 0.5 mm lat., 
subuladas a lanceoladas; pecíolo 0.4-1.5 cm long., 
pubescente; estipelas 2-3 mm long. x 0.25-0.5 
mm lat., subuladas a lineares; pinna uniyugada 3- 
5 cm long.; foliolos 18-55-yugados, 4-8 mm long. 
X 0.5-2 mm lat., oblongos, asimétricos en la base, 
papiráceos a subcoriáceos, discoloros, cara adaxial 
glabra, brillante, cara abaxial pubérula, levemente 
estrigosa o hirsuta, raro glabra a subglabra, con 
] -2(-3) nervios, nervio medio central a subcentral, 
prominente, borde ciliado, formando un margen 
nerviforme (o no marginado). Inflorescencia en ra- 
cimos espiciformes 1.5-3.2 cm long., axilares. Flo- 
res rosadas; bráctea 0.5-1 mm long. x 0.25 mm 
lat., l-nervia; cáliz + 1 mm long., 2-partido a + 
4- dentado, escarioso, blanquecino, lacerado en el 
ápice, pubérulo en la base; corola 1.5-2 mm long., 
4-lobulada, 
Craspedio 1-2 cm long. x 0.3- 


con pubescencia retrorso-estrigosa. 
0.5 cm lat., 
a subtoruloso, 3-4-articulado, artéjos 5-7 mm 
long., subcuadrados, estrigoso-pubérulos. 


recto 


Distribución. 
pos húmedos y modificados del planalto brasileño. 
Es dudosa su presencia en el territorio argentino, 
siendo citada para la provincia de Misiones, uni- 


Vive preferentemente en cam- 


camente sobre la base del ejemplar tipo. 


Material selecto examinado. BRASIL. PARANÁ: Bo 


28704 (SI); Osorio, Fazenda do Ayo. prox. Osorio, Rambo 
46755 (SI); Caracol, prox. Canela, Rambo 52807 (SI). 

En 1947, Burkart describió para Rio Grande 
do Sul a M. niederleinii var. riograndensis; pos- 
teriormente, en 1979 la citó para el Estado de 
Santa Catarina. Segün él, esta variedad se dife- 
renciaba por presentar foliolos pubérulos en la cara 
abaxial y pinnas 24—54-yugadas, mientras que la 
var. niederleinii se caracterizaba por poseer folio- 
los con la cara abaxial pubérula y estrigosa y pinnas 


384 


Annals of the 
Missouri Botanical Garden 


19-32-yugadas. El material tipo consultado de M. 
niederleinii var. riograndensis posee la mayoria 
de los foliolos con pubescencia pubérula, pero, tam- 
bién, se pueden observar algunos pelos estrigosos 
mezclados, como ocurre en la variedad tipica. Con 
respecto al nümero de foliolos por pinna, es con- 
veniente senalar, que el material tipo de M. nie- 
derleinii es un ejemplar pobre y las pinnas no 
poseen completos los pares do foliolos. Por otro 
lado, en el material brasileno, se han observado 
varios ejemplares con pinnas 30-50-yugadas pero 
con pubescencia pubérula y estrigosa en la cara 
abaxial de los foliolos. Ante la variabilidad obser- 
vada en los ünicos caracteres diferenciales, se pro- 
pone en esta nota considerar en la sinonimia de 
M. niederleinii a la var. riograndensis. 
ta de M. niederleinii para la Argentina fue 
realizada sobre la base del ejemplar tipo coleccio- 
nado en Varana (?), Misiones, por Niederlein en 
la expedición de la Comision Nacional de Limites. 
En 1948, Burkart senala que dicha localidad per- 
tenece a la parte centro-oriental de la provincia de 
Misiones, en la region de Fracrán. A pesar de la 
especificación posterior, desde 1887 hasta el mo- 
mento, no se ha encontrado nuevamente material 
de esta especie en la Argentina, en cambio si es 
común hallar ejemplares recientemente coleccio- 
nados en el SE brasileno; existe la posibilidad que 
el material citado fuera coleccionado en el estado 
de Rio Grande do Sul y no en el territorio argentino. 
Esta especie posee una variedad en el Edo. de 
Santa Catarina, Brasil: M. niederleinii var. pseu- 
dolepidota Burk. Esta variedad presenta tallos, 
peciolos, pedünculos, y craspedio con pubescencia 
densa, estrigosa o setulosa, y ramificada. 


4. Mimosa petraea Chodat et Hassl., Bull. 
Herb. Boissier ser. II 4: 548. 1904. TIPO: 
Paraguay: in glareoris prox. Chololó in valle 
fluminis, Y-acá, Dec. 1900, Hassler 6619 
(holotipo, G; isotipo, F; fotografia n? serie 
F(G)— 28277, MO, F, SI) 


Sufrútice pequeño 10-30 cm alt., inerme; tallos 
con pubescencia estrigosa o estrigosa y pubérula, 
a veces con pequeñas glándulas caedizas. Estípulas 
2.5-5 mm long. X 0.5 mm lat., 
linear-triangulares, 3-nervias, glabras, ciliadas en 


lanceoladas a 


el margen. Peciolo 1-3 mm long.; pinnas uniyu- 
gadas 15-30 mm long.; estipelas 0.25-0.5 mm 
long., lineares, pilosas; foliolos 12-28-yugados 2- 
3.5(-4.5) mm long. x 0.75-1 mm lat., oblongos 
a triangulares, con 1 -(2) nervios visibles en la cara 
abaxial, nervio medio subcentral, cara adaxial gla- 
bra, cara abaxial + pubérula, a veces con algunos 


pelos estrigosos (pubescencia estrigosa, más abun- 
dante en los foliolos inferiores de la pinna). Inflo- 
rescencia axilar; pedúnculo 1(2) en la axila de las 
hojas superiores, 5-10 mm long., menores o igua- 
les que las hojas; racimos globosos a subovoides de 
+ 4 mm long. X 4 mm lat. (sin filamentos); bráctea 
2-3 mm long. x 0.5-0.75 mm lat., lanceolada, 
aquillada, mayores que las flores, glabra a pubérula 
en el ápice, 1-nervia, margen con glándulas cae- 
dizas y ciliado en el Y, apical, caediza; botón floral 
ovoide; cáliz de + 0.5 mm long., membranáceo, 
glabro, ciliado en el borde; corola de 1.5-2 mm 
long., 4-lobulada, turbinada, lóbulos de 0.75-1 mm 
long., l-nervios, glabros a levemente pubérulos; 
filamentos estaminales libres hasta la base. Cras- 
pedio 12-17 mm long. x 4-5 mm lat., levemente 
toruloso, comprimido, estrigoso y pubérulo, 3-4- 
seminado, artejos de 4-5 mm long.; replo 0.5 
mm lat. Semilla + 2.5 mm long. x 2.5 mm lat. 
(inmadura). 


Distribución. Vive en lomadas con rocas aflo- 


rantes en las serranias de Paraguay central. 
Material adicional examinado. PARAGUAY. CORDI- 
LLERA: Itacurubi, Schinini 3207 (CTES, F). PARAGUARI: 
Piraretá-Piribeby, Schinini 2594 (CTES, F, SI); links vom 
Piribebuy und sublich von ati: Serrania nordlich von 


Fal des Y-agui-guazu, Fiebrig 840 (F, LIL) 


En 1948 y 1952, Burkart citó a esta especie 
para la Argentina sobre la base del ejemplar: ““Mi- 
siones, Bonpland, P. Jorgensen 325, IX-1909 (BAB 
34712)". En el material estudiado solo se pudo 
encontrar un ejemplar de herbario que si bien posee 
los mismos datos citados en la exsiccata por Bur- 
kart, no coincide en la fecha de colección. En la 
etiqueta del material se lee XI- 1909 y no IX-1909. 
Esta diferencia probablemente se deba a un error 
de impresión. Por otro lado, el ejemplar Jorgensen 
325 (BAB 34712) no pertenece a M. petraea, sino 
a una forma más pequeña y pubescente de 
oligophylla M. Mich. Al revisar todo el material 
depositado en el herbario del Depto. de Botánica 
Agricola, INTA, Castelar (BAB), se pudo compro- 
bar que existen más ejemplares de P. Jorgensen 
325, los cuales corresponden a distintos nümeros 
de BAB y a diferentes especies de Mimosa, por 
ej.. BAB 30213, 30214, 30224 = M. oligophyl- 
la; BAB 32116, 32117, 32118 = M. paupera 
Benth., etc. A pesar de la mezcla existente entre 
el material y el número de BAB, ninguno de los 


s 
cita de Burkart (1948), hasta el momento 
no se ha encontrado esta especie en la Argentina, 
por lo que se cirscunscribe su distribución geográ- 
fica al territorio paraguayo. No se descarta que en 


Volume 76, Number 2 


Fortunato 
Mimosa 


el futuro pueda ser coleccionada en las provincias 
l 


En el material examinado de M. petraea (Bur- 
kart, 1948), fue citado otro ejemplar: “Paraguay, 
P. Jorgensen 4450,” el cual tampoco corresponde 
a la variedad tipica, sino a M. petraea var. hirtula 
Burk. Dicho ejemplar, al igual que el tipo y demás 
especimenes estudiados de var. hirtula se diferen- 
cian de var. petraea por ser un sufrütice erecto a 
procumbente en la base con pinnas 1-6.5 cm long., 
y generalmente inflorescencias mayores a iguales 
que las hojas. 


LITERATURA CITADA 


BENTHAM, G. 1842. Mimosa. In: W. J. Hooker, J. Bot. 
: 358-416. 
. 1875. Revision of the suborder Mimoseae. 
Trans. Linn. Soc. London 30: 335-665. 
BURKART, A. 48. Las especies de Mimosa de la flora 
Argentina. ae 8: 9-231, figs. 1-26 
9 as Leguminosas Argentinas mese 
y Cultivadas: 2nd edition. Acme, Buenos 
1979. Mimosoideae. In: P. Reitz, FL j m 
Cutan, 


A REVISION OF 
MESOAMERICAN 
PSYCHOTRIA SUBGENUS 
PSYCHOTRIA (RUBIACEAE), 
PART II: SPECIES 17-477" 


Clement W. Hamilton? 


GROUP 3. THE GRACILIFLORA GROUP 


Shrub or sometimes small tree; young stems 
glabrous or sometimes ferrugineous-pubescent; 
stipules sheathing or not, triangular or ovate or 
lanceolate, the apex often bifid or biaristate (Fig. 
2f), uniform in color, glabrous or less commonly 


fringed or ferrugineous-puberulent, caducous. Leaf 


blades usually elliptic, sometimes narrowly so, or 
oblanceolate or slightly obovate, drying reddish 
or greenish gray, brown, or black; secondary 
veins 3-14(-16) pairs, diverging (45?-)55?- 

90(-100%), eucamptodromous or brochidodro- 
mous, with collector vein only in P. jimenezii, the 
axils with domatia (Fig. 4b) or not (with tufts of 
hair only occasionally in P. bakeri); tertiary veins 
orthogonal reticulate (sometimes percurrent in P. 
marginata). Inflorescences panicles of cymes or 
rarely (P. bakeri and sometimes P. parvifolia) of 
glomerules, usually pedunculate; secondary axes 
usually in 1 pair per rank, if more than 1 pair 
then size-differentiated, often delicate or diffuse in 
structure and diverging at right angles (Fig. 7h); 
bracts rarely conspicuously enlarged (to 6 mm long 
in P. jimenezii). Corolla tubes (1.5-)2-4(-5) mm 
long (Fig. 8), the lobes usually without apical ex- 
tensions (sometimes with blunt protuberances in P. 
graciliflora). Fruit when dry usually ellipsoid to 
spherical (obloid in P. philacra); persistent calyx 
a beak or not evident; seed dorsal surface with 
3-5 (6-8 in P. parvifolia) usually deep longi- 
tudinal furrows (Fig. 10d), the ventral surface with 
2 deep or broad longitudinal furrows (plus several 
irregular furrows in both surfaces in P. philacra; 
ventral surface often concave in P. graciliflora 


[ Fig. 10c 


The most commonly encountered species in the 
group, Psychotria marginata (distribution, Fig. 
20), does not obviously closely resemble any other 
species. Psychotria graciliflora (Fig. 21), the oth- 
er widespread species, is the centerpiece of the 


morphologically cohesive P. graciliflora complex, 
which includes also P. bakeri (Los Santos, Pana- 
ma), P. laselvensis (Costa Rica), P. liesneri (east- 
ern Panamá Province), P. orosiana (Costa Rica 
and Panama), P. parvifolia (Costa Rica and west- 
ern Panama), and P. philacra (eastern Darién 
Province; see Fig. 22). Psychotria jimenezii, a 
Costa Rican species (Fig. 23), is unique in the group 
in having a collector vein, inflorescence bracts as 
long as 4-6 mm, and seed dorsal surface with 
usually only three longitudinal furrows, but it too 
may belong to the P. graciliflora complex. 

Of the nine species in the group, six appear 
distylous. All six show between-morph asymmetry 
in floral part lengths: Psychotria graciliflora, P. 
jimenezii, P. laselvensis, P. marginata, and P. 
parvifolia have longer-exserted parts in pins than 
in thrums, and in P. orosiana pin reproductive 
organs are longer at both levels. The only positively 
derived state is seen in P. liesneri, which is long- 
homostylous. Available flowering material of P. 
bakeri and P. philacra is insufficient to determine 
their breeding systems. 


17. Psychotria bakeri Dwyer, Ann. Missouri 
Bot. Gard. 67: 347. 1980. TYPE: Panama. 
Los Santos: Loma Prieta, Cerro Grande, 720- 
840 m, 8 June 1967 (fl), Lewis et al. 2249 
(holotype, MO; isotypes, F, GH). Figure 22. 


Shrub to 1.5 m; young stems glabrous, the bark 
smooth; stipules sheathing, the sheath 2.5 mm 
long, biaristate with extensions 2-3 mm long, gla- 
brous, caducous, leaving a red-brown ridge. Leaves 
petiolate; petioles 3-5 mm long, glabrous, flat above; 
blades membranous, narrow-elliptic, the apex acute, 
the base attenuate, (3-)4.5-7 x 1-1.6 cm, gla- 
brous above and below, drying pale red-brown; 
secondary veins 6-7 pairs, diverging 45°-55°, 
brochidodromous, arcuate near margin, not ele- 
vated, glabrous, the axils occasionally with minute 


' Part I: Introduction and Species 1-16 appears in the preceding issue of the Annals of the Missouri Botanical 
l 


dcin hire 76: 67- 
2 Cen 


r for Urban Hortlealuie: GF-15, University of Washington, Seattle, Washington 98195, U.S.A. 
ANN. MISSOURI Bor. GARD. 76: 386-429. 1989. 


Volume 76, Number 2 
1989 


Hamilton 
Mesoamerican Psychotria, Part II 


300 km 


105 100 95 


90 85 80 


L L 


FicuRE 20. Distribution of Psychotria marginata in Mesoamerica. 


tufts of white hairs; tertiary veins inconspicuous. 
Inflorescences terminal or pseudoaxillary, panicles 
of glomerules; panicle branched to 2-3 degrees; 
main axis 1.5-3.5 cm long, the peduncle 1-3 cm 
long; secondary axes in 1 rank, the first-rank axes 


2, 0.4-0.8 cm long; cymes branched to 2 degrees; 
bracts and bracteoles linear to triangular, often 
cleft, 1-2 mm long, puberulent within, the margins 
ciliate. Flowers sessile; calyx cup-shaped, the tube 
1 mm long, the lobes 5, triangular, 0.5-0.7 x 


300 km 


105 100 95 


L 


"T T ' T 


-F 


T.C.4 


90 85 80 


L L 


FIGURE 21. Distribution of Psychotria graciliflora in Mesoamerica. 


388 Annals of th 
Missouri ieee Garden 
124 
^» 

114 
- x 10 4 

y 

vo 
2? : A ni 
rae 
r^g só 
100 km 
e) 
S E 
85 84 83 81 80 79 78 7 
FIGURE 22. 


and P. pr (square 


0.6 mm, glabrous; corolla white, the tube cylin- 
drical, 1.5-3 x 1-1.5 i 
throat, the lobes 5, narrow triangular, 1-2 x 1 
mm; stamens 5, the filaments not seen in pins, 1.5 
mm long in thrums, the anthers 1 mm long; style 
not seen in pins, ] mm long in thrums, the branches 


mm, white pubescent in 


linear. Fruit not seen. 


Distribution (Fig. 22). Known only from the 
type collection from Los Santos, Panama at around 

O m elevation in premontane wet forest with 
tropical-equatorial climate. It was collected in flow- 
er on e8 


Psychotria bakeri may be recognized by its 
small narrow-elliptic (usually 4.5-7 x 1-1.6 cm) 
leaves with acute apex and small (1.5-3.5 cm long) 
panicles of glomerules. Only the thrum morph has 
been seen, but one collection does not suffice to 
suggest thrum-monomorphy. 


18. Psychotria graciliflora Bentham in Oer- 
sted, Vidensk. Meddel. Dansk Naturhist. For- 
en. Kjebenhavn 1852: 35. 1853. Mapouria 
graciliflora (Benth.) Oerst., Amér. Centr. p. 
17, t. 14., fig. 6. 1863. Uragoga graciliflora 
(Benth.) Kuntze, Revis. Gen. Pl. 2: 960. 1891. 
TYPE: Costa Rica. Alajuela: nr. Naranjo, ca. 
1,350 m, (fl), Oersted s.n. (holotype, C, n.v., 
photo, F neg. no. 22832; isotypes, K, US). 
Figures 10c, 21. 


Distributions of Psychotria bakeri q circle), P. liesneri (triangle), P. parvifolia (solid circles), 
e). 


cic oaxacana Standley, Contr. U.S. Nan. Herb. 
1926. TYPE: Mexico. Oaxaca: nr. Sto 
ATEN 480 m, 18 June 1895 (fl), E. W. Ne bon 
2688 (holotype, US) 
Psychotria vallensis Dwyer, Ann. Missouri Bot. Gard. 
: 438. 1980. TYPE: Panama. Coclé: La Mesa nr. 
chicken farm, 3 Jan. 1974 (fl, fr), Dwyer 11867 
(lectotype designated herein, ro 630054; iso- 
lectotypes, F, GH, MO— 2 shee 


Shrub 1-3 m tall; young stems glabrous, the 
bark gray, smooth to shallowly furrowed; stipules 
ovate, biaristate with often puberulent extensions 
3-4(-6) x 1-1.5 mm, glabrous, caducous, leaving 
a pale ridge with red-brown fringe. Leaves sub- 
sessile to petiolate; petioles 1-7 mm long, glabrous, 
flat above; blades membranous, elliptic, the apex 
acuminate or sometimes acute, the base attenuate 
to narrowly cuneate, (2-)3-8(-11.5) x (0.9-)1.3- 
3(-3.5) cm, glabrous above and below, drying gray- 
green to deep red-brown; secondary veins (4)5- 
8(9) pairs, diverging (67?-)70?-80*(-959?), eu- 
camptodromous to brochidodromous, constantly 
arcuate, barely elevated below, glabrous, the axils 
lacking domatia or hairs; tertiary veins inconspic- 
uous to evident, the intersecondaries often evident, 
orthogonal reticulate. /nflorescences terminal or 
pseudoaxillary, delicate spreading panicles of cymes; 
panicle branched to 3 degrees; main axis 3-5 cm 
long, the peduncle 2.5-3.5 cm long; secondary 
axes in (2-)3 ranks, the first-rank axes 2, 0.8- 
1.5 cm long, the second-rank axes 2, 0.2-0.6 cm 


Volume 76, Number 2 
1989 


Hamilton 389 


Mesoamerican Psychotria, Part II 


long, the third-rank axes 2, 0.1-0.2 cm long; cymes 
branched to 1-2 degrees; bracts triangular to lin- 
ear, conspicuous, 1-3 mm long, glabrous, some- 
times fringed; bracteoles irregular, 0.5-1 mm long, 
fringed. Flowers sessile to pedicellate, the pedicels 
to 1.5 mm long; calyx cup-shaped, the tube 0.3- 
0.5 mm long, the lobes 5, barely evident to tri- 
angular to linear, to 1 mm long, puberulent outside; 
corolla white, the tube cylindrical, 2-2.5 x 
white pubescent in throat, the lobes 5, linear often 
with blunt protuberance near apex, 1-2 x 0.5 

mm; stamens 5, the filaments 2-2.5 mm long in 
pins, 3-3.5 mm long in thrums, the anthers 1 mm 
long; style 4-5 mm long in pins, 2.5 mm long in 
thrums, the branches linear. Fruit when dry ellip- 
soid to spherical, 4-5 mm long, 3-4 mm diam., 
maturing red, drying red-brown to black; persistent 
calyx a beak to 1 mm long; seed dorsal surface 
with 4 longitudinal furrows, the ventral surface 
road shallow longitudinal fur- 


l mm, 


concave or with 


rows (Fig. 10c). 


Distribution (Fig. 21). Widespread from Mex- 
ico through Panama, missing only in El Salvador. 
It is noteworthy that such a common and wide- 
spread species in Central America is rare at best 
in South America and apparently absent from the 
Antilles. Psychotria graciliflora has the greatest 
elevation range of any species of the subgenus, 
ranging 0—2,500 m, and occurs in tropical moist 
to lower montane rainforest with equatorial and 
tropical climate. It has been collected in flower 
January- August, primarily March-July, and in fruit 
throughout the year, primarily August- February. 


Selected specimens examined. MEXICO. CHIAPAS: 
Mpio. Cintalapa, 16 km NW of Rizo de Oro along a 
Colonia Figueroa, 1,600 m, 27 Apr. 1972 
(fl), Brsedious 24913 (MEXU); 8 Jan. 1973 (fr), Breed- 
~ & Smith 31291 (MEXU); Mpio. = fel ap 

o Laguna Ocotal Grande, 800 m, 6 Feb. 1973 (fr), 
Breedlove 33059 (MEXU, MO); SE side of c Tres 
Picos and ridges nr. summit, 2,100-2,500 m, 28 Mar 
1973 (fl), Breedlove 34380 (MEXU); ix Corzo, E bias 
of Cerro Tres Picos nr. Cerro Bola along a logging rd. 
SW of Colonia Agronomos Mexicanos, 1,500 m, 10 Dec. 
1972 (fr) Breedlove Thorne 30180 (MEXU). 
GUERRERO: Mpio. Atoyac, 19 km NE of Atoyac, nr. San- 
tiago La Unión, 770 m, 3 Nov. 1979 (fr), Koch et al. 
79293 (MEXU). HIDALGO: Mpio. Xochicoatlán, Malila, 
1,450 m, 16 May 1969 (fl), Puig 4604 (ENCB, MEXU). 
OAXACA: Dto. Putla, 10 km al NE de Putla, carr. a 
Huajuapán, 990 m, 20 June 1982 (fl), Cedillo & Torres 
1481 (ENCB, MEXU); Pluma Hidalgo, N of Pochutla, 
July 1943 (fr), Leyva s.n. (US); Dto. Villa Alta, forest 
between Los Llanos de Ozumazui and Rio Chiquito, 
17*35'N, 95°55'W, 500 m, 1 May 1939 (fl), Schultes 
& Reko 732 (F); Tuxtepec, Chiltepec, 14 Sep. 1961 (fr), 
Sousa 936 (MEXU — 2 sheets); Dto. Miahuatlán, 16 km 
al SE de piedra larga, camino al Progreso, 23 Sep. 1982 
(fr), Torres & Cedillo 1397 (ENCB. MEXU); Dto. Teo- 


titlán, “Río Uluapan” 8 km al E de San Bartolomé Ay- 
antla, carr. Huajuapán-Jalapa de Díaz, 880 m, 20 July 
1982 (fl, fr), Torres et al. 845 (ENCB, MEXU). QUERE- 
TARO: Mpio. de Tancoyol, La Parada, 1,300 m, 16 Apr. 
1969 (fl), Puig 4500 (ENCB). SAN LUIS POTOSÍ: Las Can- 
oas, hills, 21 Aug. 1891 (fr), Pringle 5041 (GH); Mpio. 
de Xilitla, ejido de Xilitlilla, 1,000 m, 5 May 1959 (fl), 
Rzedowski 10577 (ENCB, MEXU). rABASCO: Mpio. Her 
oica Cárdenas, km 21 de la carret. Cardenas—Coatza- 
coalcos, 19 May 1981 (fl), Magaña & Curiel 201 (ENCB, 
MEXU). TAMAULIPAS: nr. Gómez Farías, Rancho del Cielo, 
,260 m, 1 May 1967 (fl), Gómez- Pompa 2019 (MEXU); 
Jaumave, 1930 (fr), Viereck 680 (US). vERACRUZ: Mpio. 


Catemaco, cerro al NE de Coyame, lado 


(ENCB, F, MEXU); Atoyac, E May 1937 (fl), 
1414 (A, F, K, MEXU, MO, NY); Río Vista, Río Coatza 
coalcos, nr. Jesús iO 20 m, 9 Aug. 1971 (fr), 
Nevling & Gómez-Pompa 2564 (ENCB — 2 sheets, GH, 

MEXU); Mpio. A "p Soloxuchil, 17?16'N, 
94°37'W, 152 m, 2 Apr. 4 (fl), M. Vázquez et al. 
312 niei F, MEXU); Mpio. Minatitlán, orilla de Río 
Uxpanapa, un poco arriba del poblado de Uxpanapa, 120 
m, 26 May 1981 (fl), Wendt et al. d MEXU). 


GUATEMALA. ALTA VERAPAZ: Sebol, m E of San Luis 
Road, between Achiote and Sebol, 23 Apr. 1964 (fl), 
Contreras “4485 (MO); à m 


HU un ANGO: between m and Rio Ixcan, Sierra de los 
Cuc anes, 150-2 , 23 July 1942 (early b 
d 49271 "m n Puerto Barrios, 28 
ari o os D (GH, vic. garis, 75- 225 
15-3 922 (fl), epit 24291 (GH, NY, 
US). PETEN: pn De ego, Rio de la Pasión, 16 Apr. 1935 
(A), M. Aguilar H. 499 (F, GH, K, MO, NY); La Cumbre, 
on km 138/139 on San Luis-Cadenas r 


forest between Finca Yalpemech and 
m, 28 Mar. 1942 (fl), uia 45431 (F); along n 
Sant label, TY mouth of Rio Sebol and El Porvenir, 
00 m, 21 Apr. 1942 (fl), -— Pss iud ae 
a EZALTENANGO: ce Pireneos, below a de 
Jesús, 1,350-1,380 m, 11 Mar. 1939 pss de 
68196 (A). DM. TOLEDO: Columbia Forest Station, 24 
June 1972 (fl), rr 9903 (DUKE, NY); 16 km from 
Punta Gorda, rd. to San Antonio, 31 July 1980 (early 
fr), Dwyer 15141 (MO 3 sheets); Golden Stream, upper 
reach, 4 May 1944 p Gentle 4581 (DUKE—2 a 
MO). HONDURAS. ATLANTIDA: nr. Tela, Triunfo, 0 m, 
Dec. 1927 (fr), sí 53755 (A, US); foothills dien 
of La Ceiba, 25 June 1938 (fr), Funcker et al. 8019 
(GH, K, MO, NY, US). coMAYAGUA: Pito Solo, Lake 
Yojoa, 600 m, 24 Aug. 1932 (fr), Edwards 469 (A, US). 
CORTÉS: along Rio Lindo, nr. Lake Yojoa, 490 m, 8 Aug. 
1948 (early fr), L. O. Williams & A. Molina R. 14562 
(GH). SANTA BARBARA: Montaña Santa Barbara above 
Sauce nr. Lake Yojoa, 1,000 m, 7 Aug. 1948 (fl), L. O. 
Williams & A. Molina R. 14487 (GH). vono: Subirana, 
1,100 m, Oct. 1937 (fr), Hagen & Hagen 1114 (NY). 
NICARAGUA. BOACO: Cerro Alegre, San José de los Re- 
mates, 12%36'N, 85°44’W, 1,100-1,180 m, 11 Feb. 
1983 (fr), P. P. Moreno 20232 (MO). cHONTALES: Cerro 
Oluma, ca. 3 km SE de Cuapa, 12?18'N, 85?20'W, 700- 
740 m, 3 Jan. 1984 (fr), Grijalva et al. 3376 (MO). 
GRANADA: Volcán Mombacho, lado N, Plan de las Flores, 
1,100 m, 5 June 1975 (fl), Atwood 306 (MO). JINOTEGA: 
Volcán de Yalí, 1,500 m, 31 May 1975 (fl), Atwood 275 
(MO); “Filas el Portal" NE del Cerro Kilambé, 13?37'N, 


390 


Annals of th 
Missouri Botanical Garden 


85?40'W, 600-900 m, 26 Mar. 1981 (bud), P. P. Mor- 
eno & Sandino 7599 (MO); Macizos de Penas Blancas, 
El Gusaneras, 13?15'N, 85?41'W, 1,200- 
an. 197 


el C 

980 (fl), Araquistain & P. P. Moreno 2490 
(MO); La De N of Sta a de Ostuma, 1,600 m 

12 Mar. 1967 (fr), A. Molina. R. 20580 (F, NY, US); 
ridge along rd. between La Danta and La Luna, 12?40' 

85?43'W, 960-1,000 m, 30 July 1978 (fl), W. D. Ste- 
vens 9597 (MO). RÍO SAN JUAN: nr. Caño Chontaleño, 20 
km NE of El Castillo, 200 m, 18-21 Apr. 1978 (fl), Veill 
& Vincelli 3483 (MO). Rivas: Isla de Ometepe, N slope 
of Volcán Maderas on trail from Balgue to Laguna Mad- 
N, 85°32’W, 23 Jan. 1981 (fr), W. Hahn 
482 (MO). ZELAYA: rd. between Nueva Guinea and Ver- 
dún, 11?39'N, 84°26'W, 240 m, 17 Aug. 1983 (early 
fr), J. Miller & Sandino 1104 (MO); Rio Punta Gorda, 
Atlanta, desembocadura del Cano el Guineo, 11%33'N, 
, P. P. Moreno & 
Sandino 12858 (MO); Cano Costa Riquita, S of rd. be- 
tween Colonia Nueva León and Colonia Naciones Unidas, 
11?43'N, 84*18"W, 150-180 m, 6-7 Nov. 1977 (early 
fr), W. D. Stevens 4953 ee Us RICA. ALAJUELA: 
rd. from Canas to Upala, 13. 1 N of Bijagua, 100- 
150 m, 26 June 1976 (early fr), Co 36449 (CR, MO); 
vic. San Rafael de Guatuso, on Rio Frio, 10%4.3'N, 
84?48'W, 80-100 m, 4 Aug. 1949 (fl, early gi Holm 
& Iltis 861 (NY); 12 km NNW of San Ramón on rd. to 
San Lorenzo, 10?10'N, 84°29'W, 1,100 m, 25 ren 1983 


550 m, 29 Apr. 1983 (fl), Liesner et al. 15230 (CR, 
MO); San Carlos, Villa Quesada, 850 m, 12 Mar. 1940 
(fl), A. Smith 2612 (F, GH, MO); Alfaro Ruiz, San Luis, 
1,500 m, 20 Apr. 1940 (fl), A. Smith 2647 (F, NY); 
vic. Fraijanes, 1,500-1,700 m, 12-13 Feb. 1926 (fr), 
Standley & Torres 47600 (US). CARTAGO: along Camino 
Raiz de Hule, SE of Platanillo n 1,200-1,400 m, 
1 July 1976 (fl), Croat 36785 (CR, MO); 10 km S of 
Cartago, 1 km S of Palo Verde, 946 N, 83°57'W, 1,450 
m, 21 Apr. 1983 (fl), Liesner & Judziewicz 14537 (CR, 
MO); N of Muñeco, Rio Sombrero, ca. 1,400 m, 25 June 
1972 (fl), Primack et al. 196 (DUKE); Cerro de La 
Carpintera, 1,500-1,850 m, Feb. 1924 (fr), Standley 
35521 (US). GUANACASTE: Parque Nacional Rincón de la 
bora SE slopes of Volcán Sta. Maria, 10?47'N, 85*18'W, 
900-1,200 m, 27-28 Jan. 1983 (fl, early fr), Davidse 
et al. 23455 (CR, MO); 3 km N of Rio Naranjo, W of 
Tenorio Volcano, 550 m, 3 June 1972 (fl), Lent 2589 
(DUKE, MO). HEREDIA: Finca La Selva, OTS field station, 
Rio Puerto Viejo, 100 m, 5 Feb. 1981 (fr), Folsom 8756 
(DUKE); 1 May 1981 (fl), Folsom 9957 (DUKE); Vara 
Blanca de Sarapiquí, 1,500-1,750 m, July-Sep. 1937 
(fl), Skutch 3137 (GH, K, MO, ae US). LIMON: llanuras 
e Sta. Clara, 300 m, Apr. 1896 (fl), Donnell- dis 
6602 (GH, K, NY, US—2 | drenajes de los Rio 
Parismina y oo - = 3 Oct. 1951 (fr), Shank 
& Molina 4283 (GH - Guapiles, 300-500 m, 
i 2-13 Mar. S (8), ‘Standley 37279 (US). deer 
NAS: San Vito ava, Finca Las Cruces, 1,300 m 
May 1971 (fl), Burch 4587 “MO, NY); Cañas E 
Finca Loma Linda, 8?44'N, 8255 W. E m, 25 Feb. 
1973 (fl), Busey 627 (F, GH, MO); Rio Negro between 
La Union and Cortu, 8 Aug. 1974 (early fr), Croat 26543 
(GH, MO); Rio Burt, upper slopes of Cerro Burt, 9°01'N, 
82*52'W, 1,800 m, 19 Aug. 1983 (early fr), Davidse 


et al. 23772 (CR, MO). SAN JOSÉ: La Hondura, 1,300- 
1,700 m, 16 Mar. 1924 a Fred 37831 (US); La 
Palma, 1,600 m, 17 Mar. 4 (fl), Aun oed 38200 
(US); between Volcán Barba pir Iraz i. 
San Vicente, ca. 1,450 m, 15 Jan. 1968 m Wilbur & 
Stone 9665 (CR, DUKE, MO). PANAMA. CANAL AREA: 
Coco Solo, 11 Oct. 1972 (fr), Gentry 6475 (MO); vic. 
Fort Sherman, 15 Jan. 1924 (fr), Standley 31070 (US); 
per 14 July 1966 (fl), Tyson et al. 4562 (DUKE, 
H, MO). cuirigui: Palo Santo, 3 mi. N of Volcán, 19 
ÓN 1971 (fr), Croat 13594 (F, MO); along Rio Colorado, 
8%50'N, 82°43'W, 1,200-1,400 m, 17 Mar. 1983 (fl, 
fr), Hamilton & Stockwell 3424 (CR, MO); 11 July 1983 
(fl, early fr), Hamilton & Krager 3784 (CR, MO); 7.5 
mi. from bridge over Rio Chiriquí Viejo on rd. to Río 
Sereno, 1,350 m, 7 Apr. 1979 (fl), Hammel et al. 6863 
(MO); Finca Ojo de Aqua, 8°51'N, 82%46'W, 1,300 m, 
14 Oct. 1981 (early fr), Knapp 1580 (DUKE, MO); 
ridge 7 mi. N of Los Planes de Hornito, IRHE Fortuna 
Hydroelectric Project, 8?45'N, 82*12'W, 1,000 m, 13 


Pilón, 900-1,173 m 
(GH, MO, NY); Cro Pilón, 5 bm 
1,045 m, 13 June 1975 (early fr), Mori 6551 (MO, US). 
COLÓN: along Rio Boquerón above manganese mine, 
9°20'N, 79°35'W, 100-200 m, 13 Dec. 1981 (fr), Knapp 
& Sytsma 2444 (MO); between France Field and Cativa, 
9 Jan. 1924 (st), Standley 30178 s DARIÉN: El Real, 
Apr. 1966 (fr), Bristan 83 (GH, MO); Rio Morti, 250 
m, 18 Sep. 1967 (early fr), Duke 14184 (MO, NY); SE 
of Rio Chico, ca. 10 km upstream from Nazareht, 8°15'N, 
77°35'W, 500 m, 21 Dec. 1980 (fr), W. Hahn 163 
(MO); E slope of Cerro Sapo, 450 m, 2 Feb. 1978 (fr), 


Hammel 1256 (MO); trail SE of Manené to Rio Coasi, 
24 Dec. 1980 a Hartman van iin vic. Cam- 
pamento Buena Vista, Rio Chucu onfluence 


OV 
with Rio Tuquesa, p p 1959 (i "Sein et a “921 (GH, 
MO, US); 10 km NE of Jaqué, headwaters of Rio Pa- 
varando, 420 m, 31 Jan. 1981 (fr), Sytsma & D'Arcy 
3390 (ENCB, MO); 3 mi. N of Santa Fe, 15 e di 
(fl), Tyson et al. 4634 (DUKE, MO). Los SANTOS: 
km SW of El Cortezo, 450-600 m, 28 Oct. 1979 pes 
fr), iui 5428 (MO). PANAMÁ: Rio La Maestra, 0- 
: , 4 Dec. 1936 (fr), Allen 25 (A, F, GH, MO); 5 

ÓN of Cerro Azul on rd. to Cerro Jefe, 720 m, 13 
Nov. e (fr), Blum et al. 1699 (SCZ); Maje, trail ca. 
5 mi. Rio Majé, 200 m, 10 Nov. 1970 (fr), Foster 
& C 2010 (DUKE, F, MO); Las Cumbres, 26 
Nov. 1974 (fr), J. Gómez ? (MO); Rio Ipeti, S of Pan- 
Am. Hwy., 9%03'N, 78°25'W, ca. 100 m, 17 Sep. 1982 
(early fr), Hamilton & D'Arcy 1335 (MO); Nuevo Ar- 
raiján, 1 km de la carret. Panamericana, 22 Sep. 1974 
fr), J. Rodríquez 2 (MO). san BLas: Rio Kwadi, ca. 6 
air miles SW of Mulatupu, ca. 100 m, 20 Sep. 1967 
(early fr), Duke 14214 (MO, NY). VERAGUAS: “Cerro 
Tute” ridge up from former Escuela Agricola, Santa Fe, 
8*35'N, 81?05"W, 800-1,100 m, 15 July 1983 (early 
fr), Hamilton & Krager 3949 (MO); Cerro Tute, ca. 10 

NW of Santa Fe, 750-1,000 m, 18 May 1975 (fl), 
Mori 6237 (MO —2 sheets). 


— 


Types of P. oaxacana and P. vallensis have 
morphologies well within the normal range of P. 
graciliflora and are therefore included. 

Psychotria graciliflora may be recognized by 
its fairly small (usually 3-8 x 1.3-3 cm) glabrous 


Volume 76, Number 2 
1989 


Hamilton 391 
Mesoamerican Psychotria, Part II 


leaves drying grayish or red-brown and with few 
ege 5-8) secondary veins diverging usually 

*-80", its small (main axis 3-5 cm long) i 
i ce and its small (4-5 x 3-4 mm) el- 
lipsoid to spherical fruits often drying black. Leaves 
in Honduras and Nicaragua are at the upper end 
of the spectrum in terms of size and number of 
secondary veins, whereas Costa Rican specimens 
generally have the smallest leaves with the fewest 
secondaries. Calyx lobes are generally barely evi- 
dent to triangular, but in Darién and eastern Pana- 
má provinces the lobes are linear. 

The exserted style in the pin morph is noticeably 
longer than the exserted stamens in the thrum (4— 
5 vs. 2.5-3 mm). 

19. Psychotria jimenezii Standley, J. Wash. 
Acad. Sci. 15: 288. 1925. TYPE: Costa Rica. 
San Jose: La Hondura, ca. 1,400 m, 16 Mar. 
1924 (A), Standley 37892 (holotype, US). 
Figure 23 

i dione wendlandiana Oersted ex Mg RR. J. Wash. 

cad. Sci. 18: 9. 1928. TYPE: Costa Rica. San 
Migue, 13 May 1857 (fl), Wendland 781 were 
, fragment, F, photo, F. neg. no 
a US) 

Treelet or shrub, 2-5(-7) m tall; young stems 
ferrugineous-pubescent, the bark smooth; stipules 
lanceolate to ovate, shortly bifid, 6-9 x 3-5 mm 
glabrous, often ciliate, caducous, leaving a pale 
ridge with red-brown fringe. Leaves petiolate; pet- 
ioles 0.3-2(-3.2) cm long, puberulent, terete; blades 
membranous, elliptic, the apex acuminate, the base 
attenuate to cuneate to , (7-)8-15 x 
(2-)2.5-5(-5.5) cm, glabrous above, puberulent 
on midvein below, drying red-brown, paler below; 
secondary veins (1 1—)12—14(—16) pairs, diverging 
(60°-)75°-100°, brochidodromous with collector 


vein undulating near margin, constantly arcuate, 


» 


cordate 


prominulous below, glabrous, the axils lacking do- 
matia or hairs; tertiary veins inconspicuous to ev- 
ident, orthogonal reticulate, the intersecondaries 
often evident. 
doaxillary, panicles of cymes; panicle branched to 
4 degrees; main axis 6-7.5 cm long, the peduncle 
rarely lacking or 2.5-5 cm long; secondary axes 
in 3(-4) ranks, the first-rank axes (2)4(6), the 
longest pair 1.5-2.5(-4), the medium pair when 
present 0.5-1.6 cm long, the shortest pair when 
present reduced to 0.3 cm long, the second-rank 
axes 2 or 4, the longer pair 0.6-2.2 cm long, the 
shorter pair when present 0.2-1 cm long, the third- 
rank axes 2 or 4, the longer pair 0.4—0.8 cm long, 
the shorter pair when present 0.2-0.6 cm long; 
cymes branched to 2 degrees; bracts lanceolate, 


Inflorescences terminal or pseu- 


4-6 mm long, ciliate; bracteoles triangular, often 
split, to 1 mm long. Flowers subsessile, the pedicels 
o 0.5 mm long; calyx cup-shaped, the tube 0.5 
mm long, the lobes 5, triangular, barely evident to 
0.2 mm long, glabrous, often ciliate; corolla cream, 
the tube cylindrical, 2-2.5 x 1.2 
bescent in throat, the lobes 5, lanceolate to ovate, 
1-1.2 x 0.7 mm; stamens 5, the filaments 2 mm 
long in pins, 3 mm long in thrums, the anthers 0.7 


mm, white pu- 


mm long; style 3.5-4 mm long in pins, 2 mm long 
in thrums, the branches linear. Fruit when dry 
ellipsoid to spherical, 3.5-5(-6) mm long, 3.5-4.5 
mm diam., maturing red, drying red-brown, some- 
times sparsely puberulent; persistent calyx a beak 
0 mm long; seed dorsal surface with 3(-4) 
deep longitudinal furrows, the ventral surface with 
2 longitudinal furrows. 


Distribution (Fig. 23). 
band across northern Costa Rica from eastern 
Guanacaste to Cartago and western Limón, at el- 
evations of 200—1,600 m, mostly above 1,000 m, 
in tropical wet to premontane rainforest with equa- 
torial-mountainous climate. It has been collected 
in flower February-May and in fruit August- March. 


Found in an east-west 


Selected specimens examined. COSTA RICA. ALAJUE- 
LA: nr. Qda. Guillermina, N side Volcán Arenal, 10?29'N, 
84°42'W, 500 m, 21 Apr. 1973 (fl), Lent et al. 3412 
(DUKE, F, NY); Guadalupe de Zarcero, 1,625 m, 30 
May 1938 (fl), 4. Smith 699 (F—2 sheets, GH, MO, 
NY); Villa Quesada, 600 m, 15 May 1939 (fl, fr), 4. 
Smith 1793 (MO, NY, US); 2 km N of Bijagua, rd. to 
San Miguel, 2 Feb. 1976 (fr), J. Utley & K. Utley 2923 
(CR, DUKE). GUANACASTE: 30 km E of Pan-American 
Hwy. on rd. to Upala, 180 m, 25 Aug. 1980 x Kress 
et al. 9594 (DUKE); Los Ayotes, Tilarán, 600-700 m, 
2] Jan. 1926 (fr), Standley & Valerio 45411 (F, US). 
LIMÓN: below La io 
to Guapiles, 10°03'N, 83%58'W, 1,100-1,200 m, 16 
Dec. 1966 (fr), Burger 3909 PT ep m 2 sheets, 
NY). PUNTARENAS: Monteverde, 1,500 m Apr. 1981 
(fl), Haber 447 (MO). SAN xut: above Rio dura at 

aja La Hondura, E m, 31 Dec. 1974 (fr), J. des 
17883 (NY, US tS n Rafael nr. Heredia, 1,118 m 
Feb. 1890 (8), adia 1974 (US). 

Psychotria jimenezii may be recognized by its 
ferrugineous-pubescent aspect, elliptic leaves often 
with cordate bases, secondary veins usually 12- 
14 diverging 75°-100° with undulating collector 
vein, and inflorescences with secondary axes usu- 
ally in two unequal pairs per rank in usually three 
ranks. 

The exserted pistil in the pin morph is longer 
than the exserted stamens in the thrum morph 


(3.5-4 vs. ca. 3.3 mm). 


20. Psychotria laselvensis C. Hamilton, Phy- 
tologia 64: 228. 1988. TYPE: Costa Rica. He- 


392 Annals of the 
Missouri Botanical Garden 
12 4 
UJ 
11 7 
E 
e 
^ e 107 
A 

Q 

UNA ^ A 
Lo 9 Q 
100 km £ 
a 
85 84 83 82 ef. 80 79 78 7 


FicunE 23. 


redia: Finca La Selva, OTS field station, Rio 
Puerto Viejo just E of its junction with Río 
Sarapiqui, ca. 100 m, 19 May 1980 (fl), Ham- 
mel 8706 (holotype, DUKE; isotype, CR). 
Figure 23. 


Shrub 1.5-4 m tall; young stems glabrous, the 
bark smooth; stipules triangular, 3-4 x 
glabrous, caducous, leaving a pale ridge with red- 
brown fringe. Leaves subsessile to petiolate; peti- 
oles 1-7 mm long, glabrous, grooved above; blades 
membranous to chartaceous, elliptic, the apex acu- 
minate, the base caudate to subcordate, (5-)7.5— 
13(-16) x (1.5-)2.5-5 cm, glabrous above and 
below, drying red-black; secondary veins (8-)10- 
13 pairs, diverging (65?-)80?-85?, brochidodro- 
mous, constantly arcuate, prominulous below, gla- 
brous, the axils often with domatia below; tertiary 
veins evident, reticulate. /nflorescences terminal, 


3 mm, 


spreading panicles of cymes; panicle branched to 
4 degrees; main axis 6.5-15 cm long, the peduncle 
4-9 cm long; secondary axes in 4-5 ranks, the 
first-rank axes 2 or 4, the longer pair 1.8-4.2 cm 
long, the shorter pair when present 0.5-0.7 cm 
long, the second-rank axes 2 or 4, the longer pair 
0.6-1.8 cm long, the shorter pair when present 
0.2 cm long, the third-rank axes 2, 0.3-0.9 cm 
long, the fourth-rank axes 2, 0.1-0.4 cm long, the 
fifth-rank axes 2, 0.2 cm long; cymes branched to 
l degree; bracts lanceolate to triangular, 2-3 mm 


Distributions of Psychotria jimenezii (circles), P. laselvensis (square), and P. orosiana (triangles). 


long, glabrous; bracteoles lanceolate, 0.5-1 mm 
long, glabrous. Flowers pedicellate, the pedicels 
ca. 0.5 mm long; calyx cup-shaped, the tube 0.3 
mm long, the lobes 5, triangular, barely evident, 
minutely ciliate; corolla white, the tube cylindrical, 
2.5-3 x 1.2 mm, white pubescent in throat, the 
lobes 5, lanceolate, 2 x 1 mm; stamens 5, the 
filaments 2 mm long in pins, 3-3.5 mm long in 
thrums, the anthers 1-1.2 mm long; style 5-5.5 
mm long in pins, 2.5-3 mm long in thrums, the 
branches linear, recurved. Fruit not seen. 


Distribution (Fig. 23). 
type locality, the Organization for Tropical Studies 
field station at Finca La Selva, Río Puerto Viejo, 
Heredia, Costa Rica, at ca. 100 m elevation in 
tropical wet forest with equatorial climate. Psy- 
chotria laselvensis has been collected in flower 


Known only from the 


February-May and with immature fruit in July. 


Additional specimens examined. COSTA RICA. HERE- 
DIA: Finca La Selva, OTS field station, Río Puerto Viejo, 
100 m, 1 May 1981 (fl), Folsom 9958 (CR, DUKE); 4 
May 1981 (fl), Folsom 10004 (DUKE); 18 Feb. 1980 
(f), Hammel 7778 (DUKE); 1 July 1981 (early fr), 
Hammel 10938 (DUKE); 27 Mar. 1982 (fl), Hammel 
11491 (DUKE). 


Psychotria laselvensis may be recognized by 
its resemblance to P. graciliflora and P. orosiana 
and its generally larger inflorescences and leaves. 
It differs from P. orosiana in having leaf blades 


Volume 76, Number 2 
1989 


Hamilton 393 


Mesoamerican Psychotria, Part II 


to 13(-16) cm (vs. 10.5 cm) long, more secondary 
veins (usually 10-13 vs. 7-9), inflorescence sec- 
ondary axes commonly 4 (vs. 2) in the first rank, 
and usually shorter corolla tubes (2.5-3 vs. 2.5- 
5 mm). These three closely related species— P. 
graciliflora, P. orosiana, and P. laselvensis— 
form a continuum with regard to many quantitative 
characters, but their recognition as species is 
straightforward. 

The exserted pistil in the pin morph is conspic- 
uously longer than the exserted stamens in the 


thrum (5-5.5 vs. 3.5-4 mm). 


21. Psychotria liesneri Dwyer, Ann. Missouri 
Bot. Gard. 67: 387. 1980. TYPE: PANAMA. 
PANAMÁ: NE of town of Cerro Azul, 20 km by 
road from Interamerican Hwy., 10 Dec. 1974 
(fr), Mori & Kallunki 3658 (holotype, MO). 
Figure 22. 


Shrub 2-3 m tall; young stems minutely pu- 
berulent, the bark pale, irregularly fissured longi- 
tudinally; stipules sheathing, triangular-ovate, 2- 

5 X 1.5 mm, puberulent, red-brown ciliate, ca- 
ducous, leaving a pale ridge with red-brown fringe. 
Leaves petiolate; petioles 2-4 mm long, minutely 
puberulent, flat and furrowed above; blades mem- 
branous to subcoriaceous, elliptic, the apex long- 
acuminate, the base attenuate, the margin inrolled, 
(4-)4.5-6(-6.7) x (1-)1.5-2.2 cm, glabrous 
above, puberulent on midvein below, drying dark 
green-brown to red-brown; secondary veins 8-10 
pairs, diverging 70°-80°, brochidodromous, straight 
then arcuate near margin, prominulous below, gla- 
brous, the axils usually with often minutely ciliate 
domatia; tertiary veins inconspicuous, orthogonal 
reticulate. Inflorescences terminal, 
cymes; panicle branched to 2-3 degrees; main axis 
1.5-3 cm long, the peduncle 1-1.5 cm long; sec- 
ondary axes in (1—)2(—3) ranks, the first-rank axes 
2, 0.3-1 cm long, the second-rank axes 2, 0.2- 
0.5 cm long, the third-rank axes 2, 0.2 cm long; 
bracts triangular, ca. 0.7 mm long, puberulent, 
ciliate; bracteoles triangular, to 0.5 mm long, pu- 
berulent, ciliate. Flowers sessile to subpedicellate, 
the pedicels to 0.5 mm long; calyx cup-shaped, 
the tube 0.3 mm long, the lobes 5, triangular, 0.3 
X 0.7 mm, puberulent; corolla green-white, the 
tube cylindrical, 1.5 x 1 mm, white pubescent in 
throat, the lobes 5, lanceolate, 0.8 x 0.7 mm; 
stamens 5, the filaments 1.3-1.5 mm long in pins, 
not seen in thrums, the anthers 0.5 mm long; style 
2.2 mm long in pins, not seen in thrums, the 
branches lanceolate. Fruit spherical when dry, 
(4-)5 mm long, (4-)5 mm diam., maturing red, 


panicles of 


drying black; persistent calyx inconspicuous; seed 
dorsal surface with 5-6 longitudinal furrows, the 
ventral surface with 2 longitudinal furrows. 


Distribution (Fig. 22). 
type region, Cerro Jefe, Cerro Azul, and Goofy 
Lake (Lago Cerro Azul), eastern Panamá province, 
Panama, at 600-700 m elevation in premontane 
wet forest with equatorial-mountainous to tropical- 
equatorial climate. It has been collected in bud in 

ovember, in flower in February, with immature 
fruit in September, and in fruit in November- 
December. 


Known only from the 


Additional specimens examined. PANAMA. PANAMÁ: 


Cerro Jefe, nr. 
(fl), Duke 15237 ( 
toward Cerro Jef 

(GH, MO); 26 Nov. 1966 (fr), Dos 7098 (GH, 
US); Cerro Jefe, ca. 1 mi. upstream from Frizzel's Finca 
Indio, 9 Sep. 1970 (early fr), Foster & Eor 1851 
(DUKE, F, MO, PMA); Cerro Azul, Feb. 1968 (fl), Gó- 
mez-Pompa et al. 3436 (MEXU). 


Psychotria liesneri may be recognized by its 
small elliptic (usually 4.5-6 x 1.5-2.2 cm) often 
subcoriaceous leaves with inrolled margins, 
brochidodromous secondary venation, and small 
(1.5-3 cm long) inflorescences with secondary veins 
usually in 2 ranks. 

In both flowering collections, the negligible sep- 
aration between style and stamens suggests that 
the species is long-homostylous instead of distylous. 


22. Psychotria marginata Swartz, Prodr., 43. 
1788. Uragoga marginata (Sw.) Kuntze, Re- 
vis. Gen. Pl. 2: 961. 1891. Myrstiphyllum 
marginatum (Sw.) Hitchc., Annual Rep. Mis- 
souri Bot. Gard. 4: 95. 1893. TYPE: Jamaicae 
australis: (fl), Swartz s.n. (holotype, S, n.v., 
photo, A; isotype, B- Willdenow 4068). Cf. 
also Swartz, Fl. Ind. Occid. 400. 1797. Fig- 
ures 4b, 7h, 8, 20 


Psychotria nicaraguensis Bentham in Oersted, Vidensk. 
M Dansk Naturhist. Foren. Kjebenhavn 1852: 
34. 1853. Uragoga nicaraguensis (Benth.) Kuntze, 
Revis. Gen. Pl. 2: 957. 1891. TYPE: Nicaragua. San 
Juan, (fr), Oersted 11634 (holotype, C, n.v., photos, 
GH, NY, US; isotype, K). 


Shrub 1-3(-4) m tall; young stems glabrous, 
the bark smooth; stipules sheathing, lanceolate, 9— 
13 x 2.5-4 mm, glabrous, caducous, leaving a 
pale ridge with red-brown fringe. Leaves petiolate; 
petioles 0.5-2(-2.5) cm long, glabrous, flat or 
E I blades membranous, oblanceolate, 

inate, the base attenuate, 


rd y3- 106 pee x : (3- )4—-6.5(- 1.5) cm, glabrous 


394 


Annals of the 
Missouri Botanical Garden 


above and below, the margin often ciliate, drying 
red-gray to sometimes dull red-brown; secondary 
veins (9-)12-14(-16) pairs, diverging (55?-)65?- 
75%, eucamptodromous, constantly arcuate, ele- 
vated below, glabrous, the axils usually with do- 
matia along midvein below 
evident to inconspicuous, orthogonal reticulate to 
percurrent. /nflorescences terminal or pseudoax- 
illary, diffuse panicles of cymes (Fig. 7h); panicle 
branched to 5 degrees; main axis (7-)10-15 cm 
long, the peduncle 4-7 cm long; secondary axes 
in (4-)5-6 ranks, the first-rank axes 2(4), the 
longer pair 2-8 cm long, the shorter pair when 
present 0.5 cm long, the second-rank axes 2, 
(0.5-)1.5-5.5 cm long, the third-rank axes 2, 
(0.4—)0.7-2.5 cm long, the fourth-rank axes 2, 
(0.2-)0.4-1 cm long, the fifth-rank axes 2, 0.2- 
0.7 cm long, the sixth-rank axes 2, 0.3 cm long; 
cymes branched to 1-2 degrees; bracts triangular, 
1.5 mm long, glabrous; bracteoles lanceolate, O 
mm long, glabrous. Flowers pedicellate, the pedi- 
cels 1-2.5 mm long; calyx cup-shaped, the tube 
0.3-0.5 mm long, the lobes 5, 
triangular, to 0.2 mm long, often puberulent with- 
out; corolla pale yellow, the tube cylindrical, 2- 

-1.2 mm, short white pubescent in throat, 
the lobes 5, lanceolate, 1-1.5 x 0.7 mm, stamens 
5, the filaments 1.5-1.8 mm long in pins, 3 mm 
long in thrums, the anthers 0.8-1 mm long; style 
4—4.5 mm long in pins, 2-2.5 mm long in thrums, 
the branches linear, recurved (Fig. 8). Fruit when 
dry spherical to slightly ellipsoid, 3.5-4 mm long, 
3.5-4 mm diam., maturing red, drying black; per- 
sistent calyx inconspicuous or sometimes a beak 
to 0.7 mm long; seed dorsal surface with 4-5 deep 
longitudinal furrows, the ventral surface with 2 
deep longitudinal furrows. 


ig. 4b); tertiary veins 


not evident to 


Distribution (Fig. 20). Widespread from 
Mexico through Panama, at elevations of 0- 1,000 
m, usually below 500 m, in tropical moist to pre- 
montane rainforest with equatorial to tropical cli- 
mate. Psychotria marginata occurs also in Cuba, 
Jamaica, Colombia, Venezuela, Ecuador, and Peru. 
It has been collected in flower throughout the year, 
primarily September—April, and in fruit throughout 
the year. 


Selected specimens examined. MEXICO. CHIAPAS: 
Mpio. Las Margaritas, low ridges at conten 2: Rio 
Ixcán ith Rio Lacantum e dat té), 300 m a Mat 


tro Arqueológico nn k, Mpio. Ocosingo, 350 n 1, 23 

1 (fr), Meave et al. B-8 (MO). OAXACA: km 5 
carret. Tuxtepec- Pueblo Viejo a 500 m a la derecha, 15 
June 1964 (fl), Chavelas PS Pérez 4 (MEXU, SLPM). 
TABASCO: Teapa, (fl), Linden s.n. (F ex P). VERACRUZ: 


Mpio. Hidalgotitlan, 6 km E del campamento Hermanos 
Cedillo, 17?00'N, 94°35'W, 110 m, 13 Mar. 1974 (fl), 
B. Dorantes 2537 (MEXU — 2 sheets); 4 mi. S of Tam- 
pico Alto, MEX 180, 18 July 1971 (early fr), Dwyer et 
al. 97 (MO). GUATEMALA. ALTA VERAPAZ: Chahal, 200 m 
from airfield on village rd., in corozal, 2 Oct. 1968 (fl), 
Contreras 7793 (F); vic. Laguna Sapalá (Chajvovuch), 2 
km SW of Sibicté, 280 m, 11 Mar. 1942 (fl, fr), Stey- 
ermark 44914 (A, F); pd 350 m, Mar. 1904 4 (Â), 
Tuerckheim 8533 E , US— 2 sheets). BAJA VERA- 
Paz: Unión Barrios, E K bn 154, 7 June 1975 (fr), 
Lundell & Feind 19383 (LL). HUEHUETENANGO: be- 
tween Ixcan and Rio Ixcan, Sierra de los Cuchumatanes, 
150-200 m, 23 July 1942 qn, Steyermark 49228 (F, 
GH). IZABAL: Puerto Méndez, on To , 8 Sep. 
1980 (fr), Contreras ui (MO) ; vic. Duirgus. 75- 225 
m, 15-31 May 2 (fl, fr), Stendiey 24534 (F, GH, 
MO, NY, US); nr. Puerto Barrios, 0 m, pr.-6 May 
1939 (fl), Standley 72549 (F, NY). PETÉN: San Diego, 
Rio de la Pasión, 11 Apr. 1935 (fl), M. Aguilar H. 509 
< NY— 2 sheets); La Cumbre, km 135 of 
Cadenas Road, bordering Chacte River, 2 Oct. 1966 (fr), 
Contreras 6312 (DUKE, F); San Luis, km 51-52 of rd., 
10 July 1959 (fr), Lundell 16270 (DUKE); forest be- 
tween Finca Yalpemech along Rio San 
Diego on Rio Cancuen, 50- 
Mal 45376 (F, NY). Bu BELIZE: Burrel Boom, 
vic. 2nd Ferry, 2 Aug. 1979 (fr), edid 14943 (MO). 
CAYO: oe 16 Mar. 1931 (fl), Bartlett 12043 
(F, K, NY), Humming Bir wy., 35 mi. section, high 
ridge, 18 Oct. 1955 (fr), Gentle 8908 (F, GH, US). STANN 
CREEK: S s E Agricultural pue nr. Stann Creek, 
23 Mar. 1967 (fl), Dwyer et al. 5 MO — 2 sheets); 
Stann Creek Railway, back of sy l Ras 1939 (fl), 


Gentle 2685 (A, K —2 sheets, NY, US); Middlesex, 
60 m, 15 daly 1929 (early fr), Schipp 245 (A, F, K, 
MO, US); 2 June 1939 (fr), Gentle 2818 (A, F, K, 


NY, US); along un Branch SE 
75 m, 18 Nov. 


vic. San José 


of Humming Bird Gap, 
76 (fl), Proctor 36591 (MO). TOLEDO: 
Ns Indian village, 10 km N of Columbia 
Forest Station, 13 June 1973 (fr), Croat 24386 (DUKE, 
F, MO— 2 sheets); Columbia Forest Station entrance, 11 
dune 1973 (fr), Dwyer 11095 (MO—4 sheets); Temash 
River, high icm 28 Feb. 1945 (fl), Gentle 5241 (DUKE, 
F, NY). HONDURAS. ATLANTIDA: Lancetilla Valley, 16 km 
SE of Tela, forest preserve, 10-150 m, 3 Aug. 1977 
(fr), Croat PY (MO, US); Montana Lancetlla: 10 
m, 17 Mar. 1962 (fl), 4. Molina ja 10309 (F); mountain 
Nombre de Dios between Saladito and San Francisco, 200 
m, 26 Apr. 1967 (early fr), A. Molina R. 2086 64 (F); 
vic. La Ceiba, slopes o of Mt. Cangrejal, 16 July 1938 (fr), 
Yuncker et al. 8457 (F, GH, K, ge NY, US). CORTÉS: 
nr. Agua Azul, iuis Yojoa, 650 14 Apr. 1951 (fl, 
fr), L. O. Williams & Molina 17929(F. GH, US). cRACIAS 
^ DIOS: Qda. Tiro, Rio Platano, 15?43'N, 84°50'W, 4 
Apr. 1981 (fr), ade 1215 (F, MO). OLANCHO: 30 
km NE Culmi, Mata de Maiz, O de Montaña Punta de 
Piedra, 700 m, 1-4 May dl bi C. Nelson & Vargas 
2738 (MO). NICARAGUA. CHONTALES: nr. Sto. Domingo, 
9 Apr. 1961 (fl, fr), Bunting £ ea 1145 (DUKE, F, 
NY, US). JINOTEGA: Comarca . Cruz, el Calvario, al 
SW del Cerro Kilambe, 13°34’ M "B5*40' "Ww, 900- 1 ,000 
m, 27 Mar. 1981 (fr), P. P. wines 
Kayaska, Rio Bocay, 13%51'N, 
7 Mar. 1980 (fr), W. D. s et ai. 1653 30 no 
MATAGALPA: 4 km al E de Rio Blanco, “Wana Wana,” 
carret. Tuma, 12%55'N, 85?11'W, 280-300 m, 5 May 


Volume 76, Number 2 
1989 


Hamilton 395 
Mesoamerican Psychotria, Part II 


1984 (fr), P. P. Moreno 24101 (MO). RÍO SAN JUAN: Rio 
Sabalos, 11%02'N, 84°28’W, 50 m, 19 Feb. 1984 (early 
fr), P. Moreno 23117 (MO). Rivas: Isla Ometepe, 
faldas del lado N del Volcán Maderas, 11?27'N, 85°30'W, 
00-1,200 m, 19 Jan. 1983 (early fr), P. P. Moreno 
19687 (MO); Isla Ometepe, Volcán Concepción, 11°33'N, 
85?38'W, 600-900 m, 13 Feb. 1984 (fr), Robleto 218 
ELAYA: region of Braggman's Bluff (Puerto Ca- 
bdo 1928 (fl), Vice 185 (F, K, US); Cerro Way- 
lawás (Penas Blancas), ca. m al S de Wany, 13?30' 
84?45'W, 28 Oct. 1982 di Grijalva & Burgos 1651 
(MO); area de la Bahia de Bluefields, Rio Escondido, entre 
Sta. Martha y La Finca, Ginney Point, 0-30 m, 19 Mar. 
1949 (fr), 4. Molina R. 1885 (F, GH); Cano Montecristo, 
desembocadura del Caño El Consuelo, 11%35'N, 83%51' 
10 m, 7 Feb. 1982 (fr), P. P. Moreno 15072 (MO); El 
Guásimo, camino a El Dos, NE de Siuna, 13%48'N, 
84°39'W, 360-380 m, 25 Feb. 1983 (early fr), P. P. 
Moreno & Robleto 20743 (MO); Monkey Point, 1 km 
al S sobre la playa, 11%35'N, 83*39'W, 0-5 m, 22 Oct. 
1981 (A), P. P. Moreno & Sandino 12195 (MO); Colonia 
Kirai. 14°41'N, 84%04'W, 0-50 m, 3 Mar. 1979 ud 
T 4000 (MO); m e 2 km S of Wan 
13?42'N, 84°50'W, 0- m, 16 Mar. 1979 (fr), Pipoly 
4749 (MO); carret. NL Waslala, Rio Las Carpas 
y Rio Babasca, 13?15'N, 85*32'W, 540-580 m, 4 Mar. 
1982 (fl), Sandino 2427 (MO); 5 km N de colonia Jacinto 
Baca Jérez, 11%54'N, 84?24'W, 160 m, 20 Oct. 1984 
(fr), Sandino 4667 (MO); ca. 14.5 km W of Río Wawa 
ferry on rd. from Puerto Cabezas to Rosita, at Cario 
Kauhru Tingni, 14?06'N, 83?40' W, 10 m, 2 May 1978 
(early fr), W. D. Stevens 8638 (MO); rd. from Constancia 
to Laguna Siempreviva, 13%58'N, 84%40'W, 290-360 
m, 21 Feb. 1979 (fr), W. es Stevens 12409 (MO); vic. 
Neptune Mining Compan onanza, 14°01’ 
84°35'W, 200- Ps 25 Feb. 1979 (fr), W. D. Stevens 
12985 (MO); ca. 6 km upriver from Barra de Punta 
Gorda, S side, 11?30'N, 83?49'W, 8-10 m, 30 Sep. 
1981 (A), W. D. Stevens 20736 (MO). COSTA Rica. 
CARTAGO: Rio Turrialba, 500 m, Mar. 1894 (fl, early fr), 
Donnell- on 4832 (GH, K, US— 2 sheets); Atirro, 600 
m, Apr. 1896 (fl, fr), Bonnell Smith 6598 (GH, K, US). 
HEREDIA: Had La Selva, eld statio 
Viejo, 100 m, 17 May 1982 (fr), uie 12295 (DUKE); 
17 Nov. 1982 (8), McDowell 77 R, D Ü 
S 


: , 17 Mar. 1973 (fl, fr), Lent 3282 (CR, 

F, NY); Rio a below Cairo, Finca Montecristo, 
25 m, 18-19 Feb. 1926 (fr), Standley & Valerio 48555 
(US); Sipurio, Mar. 1894 (fl), Tonduz 8667 (US). PUNTA- 
mi. of Rincón de Osa, 8?*42'N, 83°31'W, 

30 m, 4-7 June 1968 (fr), Burger & Stolze 5441 (CR, 
F, MO, NY); Rio Barú, nr. Dominical, 9?1 7'N, 83?52'W, 
E 20 m, 20 Feb. 1977 (fr), Burger et al. 10654 (CR, 
F, MO); Corcovado National Park, 28 Nov. 1978 (fr), 
pem 11545 (MO); Río Volcán, 48 km SE of San Isidro 
El General, 300 m, 1 Mar. 1966 (fr), 4. Molina R. et 
al. 18192 (CR, F). san JOSÉ: Lado de Vargas, Tabarcia, 
cg 920 m, 22 Apr. 1963 (fr), A. Jiménez M. 661 
(CR, F, NY); entre Pedernal y Candelarita, 900 m, 29 
May 1966 (fr), 4. Jiménez M. 3965 (F, GH, NY); vic. 
El General, 850 m, Aug. 1936 (fl), Skutch 2873 (GH, 
K, MO, NY, US). PANAMA. BOCAS DEL TORO: above Qda. 


Huron on Cerro Bonyik, 150-360 m, 13 Apr. 1968 (fl), 
Kirkbride & Duke 591 (F, MO, NY); Almirante, just 
of Dos Milla, 20 Aug. 1964 (fi), McDaniel 5129 (MO); 
vic. Chiriqui Lagoon, Old Bank Island, Ns Feb. 1941 (fr), 
Wedel 2011 (GH, MO, NY, ). CANAL AREA: Barro 
Colorado Island, 3 May 1969 (A, fr), Foster HA (DUKE, 
F); 20 Dec. 1931 (fl), Wetmore & A , F, GH, 
MO); Cerro Galera, just W of Thatcher WERDE "8°55! N, 
79°35'W, 100 m, 15 Feb. ly = fr), i pint & 
Palmer 2963 (CR, E herman and nearby, 
9°20'N, 80°00'W, 0 dps ne 1983 (fr), Ham. 
ilton & Stockwell 3095 MO) Vignes Rd., 9 kn NW 
of Gamboa, 150 m, 11 Feb. 1974 (early fa. Nee 9598 
F, GH, MO); hills N of Frijoles, 19 Dec. 1923 (fl), 
Standley 27444 (GH, US). CHIRIQUÍ: vic. San Felix, 0- 
O m, Sep. 1911 (fl), Pittier 5742 (NY, US). COCLE: 


~ 


rs walk N of Alto Calvario, 
between La Junta and Limón, 800-1,000 m, 11 Oct 
d (fr), Folsom 5859 (MO). coLón: N of Rio Guanche. 
00-200 m v. 1975 (fl), Davidse & D'Arcy 
ie (MO, US); 6 mi. SW of aot 19 July 1970 
(fr), Luteyn ig (DUKE, F, MO); 4 km NW of Sala- 
manca, l E of Buenos Aires, E 410 m, 30 
Dec. 1973 (i. Nee 9091 (F, MO); between France Field 
and Catival, 9 Jan. 1924 (fl), inae 30189 (US). 
DARIÉN: Rio Chico across from 
1962 eec i pid 5223 (GH, MO): 
m, 18-20 S 7 (fr), Duke 14271 (MO, NY); E 
a of Cerro [^ climbing up from Cana, 7?55'N, 
77°40'W, 5 000 m, 23 Sep. 1982 (fl, 2 E 
& n M (MO); Cerro Sapo, 75 1 Feb. 
1978 (fl, early fr), UH 1195 (MO); 0. E Y a mi. E 
of Manené, 2 Dec. 1980 (fl), Hartman 12088 (ENCB); 
ridges on Pun Moe Grande, NW of Ensenada El 
a 7224'N, 78°07'W, 0-100 m, 22 Jan. 1982 
Sn Knapp & Mallet 2995 (DUKE, MO); Rio Tuquesa, 
250 m, 8 July 1975 (early fr), Mori 7032 (US); ‘vail 


a Loma Disk. Cono pra 720-840 
1967 (fr), Lewis et al. 2201 (MO). PANAMA: 
Maestra, 0-25 m, 4 Dec. 1936 (fl), Allen 11 (MO); Cerro 
Campana, 750 m, 17 Aug. 1982 (fr), D'Arcy & Hamilton 
14977 (MO); nr. Jeniné, Rio Canita, 24 Sep. 1961 (fr), 
Duke 3886 (GH, MO, US): Piriati, S of Pan-Am. Hwy., 
9°00'N, 78°30'W, 200-400 m, 8 Aug. 1982 (st), Ham- 
ilton 524 (MO); nr. Lago Cerro Azul, 9?10'N, 79?25'W, 
-600 m, 3 Mar. 1983 (early fr), Hamilton & Krager 
3194 (MO); nr. Arenosa, S shore of Gatun Lake N of 
Chorrera, 9%05'N, 79°55'W, 0-50 m, 27 Feb. 1983 (fl), 
Hamilton & Stockwell 3121 (CR, MO); San José Island, 
22 Jan. 1946 (fl), Johnston 1218 (GH, US); 8 km N on 
El Llano-Carti Rd., E of Rio Terable, 9°15’N, 78%00'W, 
450 m, 19 Aug. 1981 (fr), Knapp 955 (ENCB, MO); 
3.0 mi. E of Canazas checkpoint, 1-2 mi. S of Pan-Am 
Hwy., 8%52'N, 78?15'W, 0-50 m, 27 Feb. 1982 (fl, 
early fr), ps p gs Chimán, 12 Dec. 1967 (fl), 
Lewis et al. 8 (MO); nr. Tapia River, Juan Díaz 
region, 1-3 he 1923 (early fr), Maxon & Harvey 6694 
(GH, US); nr. Arraiján, 15 m, 22 June 1938 (fl), Woodson 
et al. 772 (A, F, M SAN BLAS: between Rio Diablo and 
e Acuati, nr. Nargana, 3 Nov. 1967 (fl), Duke 14869 
O, US); pe hae of Rio Cuadi, Camp Diablo, 18 

ae "1967 (A), Duke et al. 3644 (MO). a AS: Gu 


396 


Annals of the 
Missouri Botanical Garden 


Island, Playa Rosario, 26 Aug. 1970 (fl), Foster 1614 
DUKE). 


Psychotria marginata may be recognized by 
its oblanceolate long-acuminate leaves drying usu- 
ally red-gray, eucamptodromous secondary vena- 
tion with axillary domatia, large (usually 10-15 
cm long) diffuse panicles of cymes with delicate 
axes diverging at right angles, and small (3.5-4 
mm) spherical fruit drying black. Psychotria mar- 
ginata is remarkably uniform in qualitative and 
quantitative aspects throughout its broad range. 

The exserted pistils in the pin morph are longer 
than the exserted stamens in the thrum (4-4.5 vs. 
3.5 mm). 


23. Psychotria orosiana Standley, J. Wash 
Acad. Sci. 15: 288. 1925. TYPE: Costa Rica. 
Cartago: vic. Orosi, 30 Mar. 1924 (fl), Stand- 
ley 39803 (holotype, US). Figures 2f, 10d, 
23 


Shrub or small tree 1.5-4(-5) m tall; young 
stems glabrous, the bark smooth; stipules lanceo- 
late, 3-5 x 1-2 mm, biaristate, with fringed ex- 
tensions, caducous, leaving a pale ridge with red- 
brown fringe (Fig. 2f). Leaves sessile to subsessile; 
petioles to 3 mm long, glabrous; blades membra- 
nous, elliptic to oblanceolate, the apex acuminate, 
the base attenuate to subcordate, (4.5-)5.5-10.5 
x 1.7-3.8 cm, glabrous above and below, drying 
green-black; secondary veins 7-9 pairs, diverging 
80°-90°, brochidodromous, constantly arcuate, 
barely elevated below, glabrous, the axils with do- 
matia below; tertiary veins evident to inconspic- 
uous, orthogonal reticulate. Inflorescences termi- 
nal or pseudoaxillary, panicles of cymes; panicle 
branched to 3-4 degrees; main axis 3.5-10 cm 
long, the peduncle 2.5-7.5 cm long; secondary 
axes in (2-)3-4(-5) ranks, the first-rank axes 2(4), 
1-2.5 cm long, the shorter pair when present 0.7 
cm long, the second-rank axes 2, 0.3-1.5 cm long, 
the third-rank axes 2, 0.2-1 cm long, the fourth- 
rank axes 2, 0.2-0.4 cm long, the fifth-rank axes 
2, 0.2 cm long; cymes branched to 1 degree; bracts 
triangular, to 1.5 mm long, fringed; bracteoles 
lanceolate, 0.5 mm long, glabrous. Flowers sessile 
to pedicellate, the pedicels 0.5-2.5 mm long; calyx 
cylindrical, the tube 0.5 mm long, the lobes 5, 
triangular to barely evident, to 0.3 mm long, the 
lobes puberulent outside; corolla yellow-white, the 
tube cylindrical, 2.5- 9 mm, white pu- 
bescent in throat, the lobes 5, lanceolate, 1.5 x 
| mm; stamens 5, the filaments 2.5-3.5 mm long 
in pins, 3.5 mm long in thrums, the anthers 0.7— 
1 mm long; style (4-)5.5-6.5 mm long in pins, 2 


mm long in thrums, the branches short, linear. 
Fruit when dry ellipsoid, 4.5—5(—6) mm long, 3.5- 
4(-5) mm diam., maturing red, drying black; per- 
sistent calyx a minute beak; seed dorsal surface 
with 4-5 deep longitudinal furrows, the ventral 
surface with 2 shallow longitudinal furrows (Fig. 


Distribution (Fig. 23). Known from the cen- 
tral cordillera in Costa Rica and western Panama, 
at 400-1,400 m elevation, usually above 1,000 
m, in premontane to low montane wet to rainforest 
with equatorial-mountainous climate. It has been 
collected in flower January-June and September 
and in fruit in March, June, September, and Oc- 
tober. 


Selected specimens examined. — CosTA RICA. ALAJUE- 


A: La Palma de San Ramón, 1,275 m, 30 May 1927 
(fl), Brenes 5534 ( F, NY). CARTAGO: Qda. s, l 
km of Muñeco, 1,240 m, 15 Sep. 1969 (fi), Lent 


to Limón, 9%58'N, 83°34'W, 450-525 m, 10 Ma 
(A), Liesner et al. 15377 (CR, MO); vic. Orosí, 30 Mar. 
1924 (fl), Standley 39768 (US). PUNTARENAs: hills NE 
of Quaker settlement of Monte Verde, 8 May 1971 (fl), 
J. Utley 148 (DUKE). PANAMA. CHIRIQUÍ: Proyecto For- 
tuna, propriedad del IRHE, desde la finca Pitti hasta el 
filo del Cerro Fortuna, 8%45'N, 82°15'W, 1,000-1,200 
m, 25 Sep. 1976 (fl, fr), M. Correa A. et al. 2726 (MO); 
ortuna dam site, 1,400-1,500 m, 14 Sep. 1977 (fl), 
Folsom et al. 5454 (MO); La Fortuna project, 1,300- 
1,400 m, 23 Mar. 1978 (fr), Bud 222 1 (MO); behind 
m N of Los Planes del 
E Fortuna Hydroelectric Project, 8%43'N, 
82°14" W, 17 June 1982 (fl, fr), Knapp & Vodicka 5522 
ei: Fortuna, TRHE Fortuna Proj- 
e 1982 (fl), Knapp & Vodicka 
5595 UAS: sera of Rio Dos Bocas, 11-13 
km beyond rund School at Santa Fe, 350-500 m, 
25 July 1974 (early fr), Croat 25731 (GH, MO). 
Psychotria orosiana may be recognized by its 
resemblance to P. graciliflora—leaves drying 
blackish, secondary veins diverging ca. 80°, inflo- 
rescence axes delicate—and by its larger (3.5-10 
vs. 3-5 cm long) inflorescences, longer corolla tube 
(2.5-5 vs. 2-2.5 mm), and seed dorsal surfaces 
with deeper longitudinal furrows having acute lon- 
gitudinal ridges between. 
e reproductive parts at both levels are longer 
in pin morphs than in thrums. 


24. Psychotria parvifolia Bentham in Oer- 
sted, Vidensk. Meddel. Dansk Naturhist. For- 
en. Kjebenhavn 1852: 35. 1853. Mapouria 
parvifolia (Benth.) Oerst., Amér. Centr. p. 
17,t. 14, fig. 1. 1863. Uragoga levyi Kuntze, 
Revis. Gen. Pl. 2: 956. 1891. Not U. par- 
vifolia (Muell. Arg.) Kuntze (based on Rud- 


Volume 76, Number 2 
1989 


Hamilton 397 
Mesoamerican Psychotria, Part II 


gea parvifolia Muell. Arg.). TYPE of P. par- 
vifolia Benth.: Costa Rica. Alajuela: nr. 
Naranjo, ca. 1,350 m, Apr. 1847 (fl), Oersted 
11637 (holotype, C, n.v., photo, F neg. 22842; 
isotype, K). Figure 22. 


Shrub 1.5-3 m tall; young stems ferrugineous- 
pubescent, the bark irregularly furrowed; stipules 
sheathing, lanceolate, (2.5-)4-6 x (1-)1.2-1.7 
mm, ferrugineous-pubescent, caducous, leaving a 
pale ridge with red-brown fringe. Leaves petiolate; 
petioles 2-10 mm long, ferrugineous-pubescent, 
flat above; blades membranous, elliptic to slightly 
obovate, the apex acute, the base cuneate, 
(1-)1.5-5.2 x (0.6-)0.8-2 cm, glabrous above 
and below, the midvein often ferrugineous-puber- 
ulent, drying dark green-brown to red-brown, paler 
below; secondary veins 3-5 pairs, diverging 45°- 
50°, brochidodromous, constantly arcuate, incon- 
spicuous to not evident below, glabrous, the axils 
lacking domatia or hairs; tertiary veins not evident. 
Inflorescences terminal or pseudoaxillary, panicles 
of cymes or glomerules; panicle branched to 2-3 
degrees; main axis (2-)4-12 mm long, the pedun- 
cle lacking; secondary axes in 1-2 ranks, the first- 
rank axes (1)2, 1-4 mm long, the second-rank 
axes 2, 1-3 mm long; cymes branched to 1 degree; 
bracts lanceolate, ca. 2 mm long, glabrous; brac- 
teoles lanceolate, 1 mm long, glabrous. Flowers 
pedicellate, the pedicels 0.5-3 mm long; calyx cup- 
0.3-0.5 mm long, the lobes 4, 

mm long, puberulent; corolla 
white, the tube cylindrical, 2.5-4 x 1-1.5 mm 
white pubescent in throat, the lobes 4, lanceolate, 
1.5-2 x 0.8 mm; stamens 4, the filaments 2 mm 
long in pins, 3.5-5 mm long in thrums, the anthers 
0.6-0.7 mm long; style 5 mm long in pins, 2-3.5 
mm long in thrums, the branches linear. Fruit when 
dry spherical, 4-5 mm long, 4-5 mm diam., ma- 
turing red, drying dark red-brown; persistent calyx 
not evident or a minute beak; seed dorsal surface 
with 6-8 shallow longitudinal furrows, the ventral 
surface with 2 deep and sometimes 2-4 shallow 
longitudinal furrows. 

Distribution (Fig. 22). 
tral cordillera in Costa Rica and western Panama, 
at 1,300-2,000 m elevation in regions of low mon- 
tane rainforest with equatorial-mountainous cli- 
mate. Psychotria parvifolia has been collected in 
flower April-September and December and in fruit 
December-May. 


shaped, the tube 


» 


Known from the cen- 


Selected specimens examined. COSTA RICA. ALAJUE- 

A: La Palma de San Ramón, Socorro, 22 Aug. 1927 
(st), Brenes 5681 (CR, F, NY); 15 km NW of San Ramón 
by air, Cerro Azahar, Rio San Pedro, 10%09'N, 84°34 W, 


1,400-1, 500 m, 14 May 1983 bar Liesner - a 15585 
1,700 m, 12-13 Feb. 


des 47571 (US). caRTAGO: 15 km S of Tapan 
rande de Orosi, "AFN. 

17 Dec. 1969 (fl), Burger & Liesner 6847 (CR, F. MO, 
NY); La Congreja, carret. Panamericana al S de Cartago, 


km 47, 1,950 m, 28 s 1965 (fl), A. Jiménez M. 3365 


, NE 
b. 1926 (early fr), 
lerio 49194 (US); Cerro a are NE o 
2,000-2,400 m, 3 Mar 6 (early fr), Standle ey & 
Valerio 50271 (US); e & dagen: Verg Aii 
Cerro d 00- 


84*48'W, 1,300 m, 18 Aug. 1976 (fl), Solomon 537 
(CR, MO). san JOSÉ: vic. Sta. María de Dota, 1,500— 
1,800 m, 14-26 Dec. 1925 (fl, fr), cw 42857 (A, 


1925-8 Jan. 1926 (fl, fr), Standley & 
Valerio 44076 (US); La Hondura de San Jose, 1,300 m, 


0 
cy 6393 (CH. MO, NY); vic. El Boquete, 
1,000-1,300 m, 2-8 Mar. 1911 (fr), W. Maxon 4958 
). 


Psychotria parvifolia may be recognized by its 
small (usually 1.5-5.2 x 0.8-2 cm) membranous 
leaves with 3-5 inconspicuous secondary veins, 
minute (usually 0.4-1.2 cm long) inflorescences, 
and spherical fruits 4-5 mm in diameter. Collec- 
tions from Chiriqui, Panama, show more slender 
inflorescence axes, longer (3 mm) pedicels plus 
longer corolla tubes and filaments and styles than 
from Costa Rica. 

The exserted pistil in the pin morph is usually 
longer than the exserted stamens in the thrum (5 


vs. 3.8-5.3 mm) 


25. Psychotria philacra Dwyer, Ann. Missouri 
Bot. Gard. 67: 412. 1980. TYPE: Panama. 
Darién: Cerro Tacarcuna S slope, ridgetop well 
below summit, 1,250-1,450 m, 26 Jan. 1975 
(fr), Gentry & Mori 13916 (holotype, MO). 


Figure 22 


Shrub 3 m tall; young stems glabrous or red- 
brown puberulent, the bark mottled; stipules lan- 
ceolate, 4 x 1.2 mm, ferrugineous-pubescent, ca- 
ducous, leaving a pale ridge with red-brown fringe. 
Leaves petiolate; petioles 4-10 mm long, glabrous 
to sparsely puberulent, flat above; blades membra- 
nous, elliptic, the apex acuminate, the base atten- 

l. 


uate, 5-7 7-2.7 cm, glabrous above and 


398 


Annals of the 
Missouri Botanical Garden 


below, drying dull red-brown; secondary veins 6- 
7 pairs, diverging 60%-70%, eucamptodromous, 
constantly arcuate, prominulous below, glabrous, 
the axils lacking domatia or hairs; tertiary veins 
not evident. /nflorescences terminal, seen only with 
few fruits. Flowers not seen. Fruit when dry obloid, 
4.5-5.5 mm long, 5-6 mm diam., maturing red, 
drying dark red-brown; persistent calyx inconspic- 
uous or a minute beak; seed dorsal surface with 4 
deep irregular and several shallow irregular lon- 
gitudinal furrows, the ventral surface with 2 deep 
and several irregular longitudinal furrows. 


Distribution (Fig. 22). Known only from the 
type collection from Cerro Tacarcuna, eastern Da- 
rien, Panama, at 1,250- 1,450 m elevation in low 
montane rainforest with equatorial-mountainous 
climate. It was collected in fruit on January 26. 


Psychotria philacra may be recognized by its 
small elliptic (5-7 x 1.7-2.7 cm) leaves drying 
dull red-brown and with 6-7 secondary veins, ob- 
loid fruits, and seeds with shallow irregular furrows 
on both surfaces. 


GROUP 4. THE REMOT4 GROUP 


The description for the group in Mesoamerica 
is that of the lone species, Psychotria remota. Key 
recognition characters include leaves drying glossy 
red-brown and relatively few secondary veins for 
the large size of the blade. This group is larger in 
South America, including also P. anceps Kunth 
and P. cupularis (Muell. Arg.) Standl. 

Psychotria remota appears normally distylous, 
with no floral-part-length asymmetry between the 
morphs. 


26. Psychotria remota Bentham, J. Bot. 
(Hooker) 3: 225. 1841. Mapouria remota 
(Benth.) Muell. Arg., Flora 59: 459. 1876. 
Cf. also Martius, Fl. Bras. 6(5): 407. 1881. 
Uragoga remota (Benth.) Kuntze, Revis. Gen. 
Pl. 2: 962. 1891. TYPE: Guiana: on the Río 
Negro, (fl), Schomburgk 963 (holotype, K, 
n.v., photo, NY neg. no. 3421; isotypes, F, 
W, n.v.). Figures 4a, 7j, 10f, 24. 

Psychotria alboviridula K. pe jig: Koenigl. 
Bot Berlin 6: 208. 1914 PE: Brazil: Im 
Gahial des Alto Acre, Seringal, m Mannes Sep. 
1911 (fl), Ule 9846 (holotype, E Maur frag- 
ment, F, photo, F neg. no. 468; e, US). 


Tree 1.5-6 m tall; young stems id to 
sparsely minute-puberulent, the bark mottled, 
slightly ridged longitudinally; stipules sheathing, 
triangular, acuminate or biacuminate, 4-6 x 2.5- 


4 mm, glabrous, sometimes sparsely fringed, ca- 
ducous, leaving a pale ridge with red-brown fringe. 
Leaves petiolate; petioles 0.7-2.5 cm long, gla- 
brous, flat or grooved above; blades membranous 
to subcoriaceous, elliptic, the apex long-acuminate, 
the base cuneate, (8-)1 1-24 x (3-)3.5-8(-9) cm, 
glabrous above and below, drying glossy red-brown; 
secondary veins (7-)8-12 pairs, diverging 60°- 
70%, eucamptodromous to brochidodromous, con- 
stantly arcuate, prominent below, glabrous, the 
axils usually with prominent domatia below (Fig. 
4a); tertiary veins evident, orthogonal reticulate to 
percurrent, the intersecondaries often conspicuous. 
Inflorescences terminal, elongate panicles of cymes 
(Fig. 7j); panicle branched to 4 degrees, the axes 
winged; main axis 7-15 cm long, the peduncle 3- 
5 cm long; secondary axes in (4-)6-7 ranks, the 
), 1-5.5 em long, the shorter 
pair when present 1.2 cm long, the second-rank 
axes 2, (0.3-)0.5-1 cm long, the third-rank axes 
0.3-0.9 cm long, the fourth-rank axes 2, 0.2- 
0.6 cm long, the fifth-rank axes 2, 0.1-0.4 cm 
long, the sixth-rank axes 2, 0.1-0.3 cm long, the 
seventh-rank axes 2, 0.1-0.3 cm long; cymes 
branched to 1(-2) degrees; bracts and bracteoles 
triangular, 0.5-1.5 mm long, ciliate. Flowers ped- 
icellate, the pedicels 0.5-1 mm long; calyx cup- 
shaped, the tube 0.3-0.5 mm long, the lobes O or 
5, not evident to triangular, to 0.3 mm long, minute 
puberulent; corolla yellow-green, the tube cylin- 
drical, 1.5 x 1 mm, white pubescent in throat, 
the lobes 5, lanceolate, 1 x 0.6 mm; stamens 5, 
the filaments 1.5 mm long in pins, 2.5 mm long 
in thrums, the anthers 0.3-0.5 mm long; style 2.5 


first-rank axes 2(4), 


mm long in pins, 2 mm long in thrums, the branches 
club-shaped. Fruit ellipsoidal when dry, (6-)7-9 
mm long, (4-)5-6 mm diam., maturing dark red, 
drying dark red-brown; persistent calyx not evident 
or rarely a minute beak; seed dorsal surface with 
3 deep longitudinal furrows, the ventral surface 


with 2 longitudinal furrows (Fig. 10f). 


Distribution (Fig. 24). Scattered distribution 
in Caribbean Costa Rica and Panama, at 20- 1,000 
m elevation in tropical moist to premontane wet 
forest with equatorial to tropical-equatorial climate. 
It is more abundant in Colombia, Venezuela, the 
Guianas, Ecuador, In Central 
America, it has been collected in flower in March, 
April, September, and October and in fruit in 
March-May and September. 


Peru, and Brazil. 


Selected specimens examined. Costa RICA. HERE- 
DIA: Caño Negro, Río Sarapiquí, 23 km N of Puerto Viejo, 
20 m, 19 Apr. 1974 (early fr), Hartshorn 1444 (CR, 
MO). LIMÓN: 7 km SW of Bribri, 100-250 m, 4 May 


Volume 76, Number 2 
1989 


Hamilton 
Mesoamerican Psychotria, Part II 


399 


S 
v A. 
2 9 9d 
oO 
ve e 8 4 
100 km ° 
TN Oo 
U 
85 84 83 79 78 74 
FIGURE 24. Distribution of Psychotria remota in Costa Rica and Panama. 


1983 (fr), L. D. Gómez et se Aer (MO). PANAMA. 
COLÓN: Santa Rita lumber rd., 15 km E of Colón, 5 
Oct. 1969 (fl), Dressler & Lewis 5732 (MO, US). DARIEN: 
trail from Punta Guayabo Gra o Rio Jaque, 50-200 
m, 24 Apr. 1980 (fl), psi ^ Hahn 4407 (MO); 
between Rio Balsa and Rio Areti at their confluence, 100 
1966 (fl), Duke 8745 (MO); Puerta St. Do- 


—— Azul (Goofy Lake), 9°10'N, 
de Mar. 1983 (fl), Hamilton. & 


300-500 m, 30 Mar. 1 1973 (fr), Liesner 1280 (F, MO, 
NY); rd. past Cerro Azul, 28 Sep. 
Kallunki 2188 (GH, MO): El Llano-Carti rd., 16-18.5 
km N of Pan-Am. Hwy. at El Llano, 400- 450 m, 28 
Mar. 1974 (fr), Nee & Tyson 10937 (F, MO). VERAGUAS: 
0.6 mi. beyond Escuela Agricola Alto Piedra nr. Santa 
Fe, 730 m, 4 Apr. 1976 (fr), Croat & Folsom 34052 
(MO, NY). 

The inclusion of P. alboviridula within P. re- 
mota is discussed by Steyermark (1972; Mem. 
New York Bot. Gard. 23: 477) 

Psychotria remota may be recognized by its 
moderately large (usually 11-24 cm long) elliptic 
leaves drying glossy red-brown, few (usually 8-12) 
secondary veins generally with prominent domatia 
in the axils, reduced (except for the first rank) and 
winged secondary inflorescence axes, and large 
(usually 7-9 mm long) ellipsoidal fruit. 


GROUP 5. THE NERVOSA GROUP 


Shrub; young stems glabrous or red-brown to 
brown tomentose to puberulent; stipules usually 
sheathing, linear (Fig. 2c) or ovate, the apex some- 
times bilobed or paene uniform in color (except 
P. fosteri, with a darker red-brown midrib), gla- 
brous or fringed or red- brown pubescent, caducous. 
Leaf blades obovate or elliptic or oblanceolate, 
drying red-brown to green-brown (often chalky yel- 
low-green in P. fruticetorum), sometimes pale be- 
low; secondary veins (4-)5-17 pairs, diverging 
35°-65° (except 70?-80? in P. fosteri), eucamp- 
todromous or brochidodromous (with collector vein 
in P. chagrensis and P. fosteri), the axils usually 
with tufts of hairs (lacking in P. chagrensis, P. 
fosteri, and usually P. nervosa); tertiary veins re- 
ticulate or percurrent. Inflorescences panicles of 
cymes (often contracted in P. nervosa) or fascicles 
of flowers (in P. chagrensis and P. fosteri), not 
pedunculate (except P. erythrocarpa, P. fruti- 
cetorum, and often P. jinotegensis); secondary 
axes usually in 1 pair or less commonly 2 size- 
differentiated pairs per rank (4 equal axes per rank 
rarely in P. nervosa); bracts conspicuously en- 
larged in P. chagrensis, P. fosteri, and P. fruti- 
cetorum. Corolla tubes 2-4 mm long (6-7 mm in 
P. chagrensis), the lobes without apical extensions. 
Fruit ellipsoid when dry (narrow in P. nervosa); 
persistent calyx a tube to 5 mm long (Fig. 9d) 


400 


Annals o 
Missouri Botanical Garden 


or a beak or inconspicuous; seed dorsal surface 
with 4-5 deep longitudinal furrows (or 6-8- 
many irregular furrows in P. aguilarii and P. 
Jinotegensis), the ventral surface with 2 deep or 
shallow longitudinal furrows, sometimes incom 
pletely divided in P. nervosa or with additional 
irregular furrows in P. jinotegensis. 


Of the nine species, three fairly widespread ones 
do not closely resemble any others: Psychotria 
fruticetorum, with its biaristate stipules, leaves 
drying chalky yellow-green, pedunculate inflores- 
cences, and bracts as long as 8 mm; P. erythro- 
carpa, with its linear stipules and pedunculate 
inflorescences often of glomerules; and P. jinote- 
gensis, with its often pedunculate inflorescences 
and many irregular furrows on both seed surfaces. 
The P. chagrensis complex shares a collector vein, 
an absence of tufts of hairs in the axils of the 
secondary veins, and inflorescences of fascicles of 
flowers and with bracts conspicuously enlarged; the 
complex comprises the widespread P. chagrensis 
and P. fosteri, endemic to Coiba Island, Panama 
(Fig. 26). The P. nervosa complex includes that 
widespread species plus P. aguilarii (southern 
Guatemala), P. boquetensis (western Panama), and 
P. mirandae (southern Mexico); none of them de- 
viates morphologically from the predominant char- 
acter states of the group 

Of the nine species in the group, seven appear 
distylous. Of those, Psychotria fruticetorum has 
onger exserted parts in the pins than in thrums, 
and P. chagrensis has longer inserted parts in the 
pins. One species, P. mirandae, appears to be 
thrum-monomorphic. Only P. fosteri may not be 
evaluated for this character due to lack of adequate 
flowering material. 


27. Psychotria aguilarii npn & Steyer- 
, Bot. Ser. 
- ual. Chimalte- 
nango: rd. between alla and San 
Martin Jilotepeque, 1,500-1,700 m, 22 Dec. 
1940 (fr), Standley 80900 (holotype, F). Fig- 


ure 


Shrub 1-2(-6) m tall; young stems glabrous, 
the bark pale with shallow fissures; stipules sheath- 
ing, ovate, 7-10 x 2-4 mm, lacking fringe, gla- 
brous, caducous, leaving a pale ridge with irregular 
fringe. Leaves petiolate; petioles 2-4(-7) mm long, 
glabrous, flat above; blades membranous, narrow 
elliptic to obovate, the apex acuminate, the base 
attenuate, the margins crenulate, (8-)11-13 x 
(2.5-)3-4 cm, glabrous above, sparsely puberulent 
along midvein below, drying red-brown or some- 


times greenish; secondary veins 8-11 pairs, di- 
verging (45°—)50°-60°, eucamptodromous, slightly 
arcuate, prominent below, glabrous, the axils with 
minute tufts of hair; tertiary veins evident, retic- 
ulate to slightly percurrent. Inflorescences terminal 
or pseudoaxillary, sparse panicles of cymes; panicle 
branched to 2-3 degrees; main axis 3-4 cm long, 
the peduncle lacking; secondary axes in 2 ranks, 
the first-rank axes 2, 2-2.2 cm long, the second- 
rank axes 2, 0.5-1.2 cm long; cymes branched to 
1-2 degrees; bracts and bracteoles minute, linear, 
ca. 0.2 mm long, glabrous. Flowers sessile; calyx 
cup-shaped, the tube 0.5-1 mm long, the lobes 5, 
acute triangular, 0.2-0.5 x 0.3 mm, glabrous; 
corolla greenish white, the tube cylindrical, 3-3.5 
X ] mm, white pubescent in throat, the lobes 5, 
acute triangular, l- mm; stamens 5, 
the filaments 2 mm long in pins, 3.5 mm long in 
thrums, the anthers 1 mm long; style 3.5 mm long 
in pins, 2 mm long in thrums, the branches 
spathulate. Fruit when dry ellipsoid, 5-6 mm long, 
3-4 mm diam., maturing dark red, drying dark 
brown; persistent calyx not evident; seed dorsal 
surface with many irregular shallow longitudinal 
furrows, the ventral surface with 2 deep longitu- 
dinal furrows. 


Distribution (Fig. 25). Known only from the 
Guatemalan states of Chimaltenango, Escuintla, 
Guatemala, Huehuetenango, and Sacatepéquez, at 
elevations of 1,000-1,500 m, in tropical-moun- 
tainous climate. Psychotria aguilarii has been col- 
lected in flower in February and May and in fruit 
in August and December 


Additional specimens examined. GUATEMALA. 
ESCUINTLA: 6 km W of San Vicente Pacaya, 1,250 m, 
31 May 1970 bu Harmon 2431 (MO). GUATEMALA: 

Finca La Aurora, 1,500 m, 1938-1939 (fr), I. ps 
126 (F); 1939 cay fr), E Aguilar . 215 (F). HUEHUETE- 
NANGO: 


ermark 51133 (F). SACATEPÉQUEZ: nr. Barranco Hondo, 
SE of Alotenango, 1,000-1,260 m, 9 Feb. 1939 (fl), 
Standley 65024 (F). 


Psychotria aguilarii may be recognized first by 
its striking resemblance to P. nervosa. The former 
differs in being generally glabrous (as is P. nervosa 
sometimes) and in having leaves with crenulate leaf 
margins, inflorescence secondary axes in 2 (vs. 3- 
4) ranks, longer corolla tubes (3-3.5 vs. 2-3 mm), 
fruit less narrowly ellipsoidal (length/width — 1.5 
vs. ca. 2.2), and seed dorsal surface with many 
irregular shallow furrows (vs. 4—5 regular furrows). 


28. Psychotria boquetensis Dwyer, Ann. Mis- 
souri Bot. Gard. 67: 349. 1980. TYPE: Pan- 


Volume 76, Number 2 


Hamilton 401 


1989 Mesoamerican Psychotria, Part II 
"id ida - _ | - _ | E T.C.4 
, e 20 4 
T i: 
e 
e 
e a 
^. 
B + E le à + 15 4 
sp $a «? à : 
* 
ee 
a 
300 km + + ys 
eb 
105 100 95 90 85 , 80 


in Mesoamerica. 


ama. Chiriqui: Boquete, 1,080 m, 25 May 
1938 (fl), Davidson 709 (holotype, MO; iso- 
types, F, US). Figure 25. 


Shrub 0.5-2 m tall; young stems brown to- 
mentose, the bark gray, longitudinally grooved; 
stipules sheathing, ovate, bilobed with the lobes 2— 
6 mm long, (6-)8-11 x 2.5 mm, lacking fringe, 
glabrous or sometimes sparsely tomentose, ca- 
ducous, leaving a pale ridge with red-brown fringe. 


nous, elliptic to oblong, the apex long-acuminate, 
the base attenuate, (6-)9-15 x (1.5-)2-4 cm, 
glabrous to slightly puberulent along primary and 
secondary veins above, sparsely tomentose along 
veins below, the margins ciliate, drying red-brown, 
paler below; secondary veins (10-)12-15 pairs, 
diverging 40°-50°, eucamptodromous, slightly ar- 
cuate, prominent below, brown tomentose below, 
the axils with small tufts of hairs below; tertiary 
veins evident, orthogonal reticulate to slightly per- 
current. /nflorescences terminal or pseudoaxillary, 
sparse panicles of cymes; panicle branched to 3 
degrees; main axis 1.2-2.0 cm long, the peduncle 
lacking; secondary axes in (2)3 ranks, the first- 
rank axes 2, 10-15 mm long, the second-rank 
axes 2, 4-10 mm long, the third-rank axes 2, 2- 
3 mm long; cymes branched to (1)2 degrees; bracts 
appearing to consist of planes of long whitish hairs, 


FicURE 25. Distributions of Psychotria aguilarii (triangles), P. boquetensis (square), and P. fruticetorum (circles) 


l mm long, immediately subtending the flowers. 
Flowers sessile to subsessile, the pedicels to 1 mm 
long; calyx cup-shaped, the tube 0.5-1 mm long, 
the lobes 5, triangular to linear, (0.5-)1-1.5 x 
0.5 mm, glabrous; corolla white, the tube cylin- 
drical, 2-3 x 1 mm, white pubescent in throat, 
the lobes 5, linear-ovate, 1.5-2 x 0.8 mm; sta- 
mens 5, the filaments 2 mm long in pins, 3.5 mm 
long in thrums, the anthers 0.5-0.7 mm long; style 
3.5 mm long in pins, 2 mm long in thrums, the 
branches club-shaped in pins, linear in thrums, the 
branches puberulent. Fruit not seen. 


Distribution (Fig. 25). 
type locality near Boquete, Chiriqui, Panama, in 


Known only from the 


premontane wet forest with equatorial-mountainous 
climate at about 1,200 m. Psychotria boquetensis 
has been collected in flower in May and June. 


Additional specimens examined. PANAMA. CHIRIQUÍ: 
Boquete, 1,200 m, 30 June 1938 (fl), M. Davidson 849 
F). 


, 


~ 


Psychotria boquetensis may be recognized first 
by its striking resemblance to P. nervosa. Psy- 
chotria boquetensis differs in having often linear 
mm (vs. to 0.2 mm) 
long; in quantitative characters such as leaf and 
inflorescence size, P. boquetensis falls at the lower 
end of the P. nervosa spectrum. 


calyx lobes usually 1-1.5 


402 


Annals of the 
Missouri Botanical Garden 


29. Psychotria chagrensis Standley, J. Wash. 
Acad. Sci. 15: 105. 1925. TYPE: Panama. 
Canal Area: Barro Colorado Island, Gatun 
Lake, 17 Jan. 1924 (fr), Standley 31373 
(holotype, US). Figure 26 


Shrub 1-2.5 m tall; young stems glabrous, the 
bark smooth; stipules sheathing, the apex aristate 
and 2-4 mm long, the sheath 4-7 
glabrous, caducous, leaving a pale ridge. Leaves 
petiolate; petioles 2-5(-8) mm long, glabrous, flat 
or grooved above; blades membranous, elliptic to 
obovate, the apex acuminate, the base attenuate, 
(3-)4-7.5(-9.5) x (1-)1.5-2.5(-3.5) cm, gla- 
brous above and below, drying red-brown, often 
dull green below; secondary veins (6-)8- 10 pairs, 
diverging (45?-)55?-65?, brochidodromous with 
collector vein distinct, straight to slightly arcuate, 
scarcely elevated below, glabrous, the axils lacking 
domatia and hairs; tertiary veins usually incon- 
spicuous, the intersecondaries sometimes faint, the 
percurrent tertiaries rarely evident. /nflorescences 
terminal or pseudoaxillary, fascicles of several flow- 


3 mm 


» 


ers, 5-8 mm long, 5-8 mm across; peduncles 
lacking; bracts elongate, often irregularly divided, 
4-5 X 1.5-2 mm, glabrous, the margins often 
ciliate. Flowers sessile; calyx green, the tube cy- 
lindrical, 0.5-2 mm long, the lobes 5, linear, 2- 
2.5 x 0.5 mm, glabrous to puberulent, the margins 
ciliate; corolla white, the tube cylindrical, 6-7 x 
l mm, white pubescent in throat, the lobes 5, 
narrow-elliptic, 2-2. mm; stamens 5, the 
filaments 5-6 mm long in pins, 8 mm long in 
thrums, the anthers linear, 1 mm long; style 7-9 
mm long in pins, 2-5 mm long in thrums, the 
branches linear, sometimes recurved. Fruit when 
dry ellipsoid, 5-7 mm long, 3-3.5 mm diam., 
maturing red, drying deep red-brown; calyx per- 
sistent, the tube 0.5-2 mm long, the lobes 2-2.5 
mm long; seed dorsal surface with 4-5 deep lon- 
gitudinal furrows, the ventral surface with 2 shallow 
longitudinal furrows. 


Distribution (Fig. 26). 
on the Caribbean lowlands and slopes in Nicara- 
gua-Panama, with apparently disjunct populations 
in Veracruz, Mexico, and Izabal, Guatemala. 
range extends southward in Colombia and Peru. 
Psychotria chagrensis is found in Central America 
often near creek beds at elevations of O to 1,300 
m in tropical moist to premontane moist and wet 


Commonly collected 


forest with equatorial to tropical-equatorial climate. 
It has been collected in flower in December- 
September, primarily March-July, and in fruit 
throughout the year, primarily September- Decem- 
er. 


Selected specimens examined. MEXICO. VERACRUZ: 
San Andres Tuxtla, Estación de Biolo 
Tuxtlas, 150 m, 7 Nov. 196 
ENCB — 5 sheets, F, K, MEXU — 2 sheets 
MALA. IZABAL: Cerro San Gil, uppermost ridges and sum- 
mit, 1,200- 1,300 m, 26-27 Dec. 1941 (fr), Steyermark 
aged NICARAGUA. RÍO SAN JUAN: San Juan del Norte, 
O m, 27 Mar. 1971 (fr), Atwood 5258 (MO); El Castillo 
Viejo, (fl), Shimek & Smith 1893 (F). ZELAYA: Comarca 
del Cabo, swamp nr. Bilwaskarma, 0-100 m, 14 
1971 (fl), Atwood 46 78 (MO); Comarca del Cabo, ma- 
torrales del Río Leicus, region de Tronqueras, 60 m, 
Aug. 1965 (fr), 4. Molina R. 14925 (F, NY, US); "Ku- 
rinwacito," 13?08'N, 84?55'W, 90 m, 25 Mar. 1984 
, P. P. Moreno 23901 (MO); El Salto, along Rio Pis 
Pis, 14°04'N, 84°38'W, 100 m, 27 Feb. 1979 ide 
Pipoly 3572 (MO); 1.8 km N of base camp, 3 6 km S 
Cerro San Isidro, Rio Kama, Rio Escondido, 65 m, n 
Mar. 1966 (fl), Proctor et = n (F, NY); trail from 
Cerro Saslaya to S e Hormiguero, 13?44'N, 
84*57'W, 200-400 m, 13 Mar. ee a D. Stevens 
°57'N 


, MO). GUATE- 


PE 
== 


: 19 May 1968 (fl), Burger & Stolze 5032 (CR, DUKE, 
, MO); San Carlos, Quesada, 850 m, 12 Mar. 1940 

a A. Smith 2587 (A, F, MO). HEREDIA ~ La ape 

the OTS field station on Rio Puerto 100 m 

Mar. 1982 (fl), Folsom 9412 DS 6 Dec. 1982 i 

McDowell 1066 (DUKE); Tirimbina, 210 m 

1971 (fl), Proctor 32118 (LL). LIMON: Rio Toro Amarillo 

5.5 km al SO de Guapiles, 350 m, 10 Aug. 1 

Jiménez 1059 (CR, F, j 


~ 


Gamboa gate, 26 Sep. 197 
F, GH, MO, NY); Barro Colorado Island, 31 Ma ay 1969 
(fl), Foster 899 (DUKE— 2 sheets); 18 Aug. 


ilton & A] 3682 (MO). cocLÉ: hills N of El Valle 
de Antón, 1,000 m, 14 July 1940 (fl), Allen 2186 (F, 
GH); between Cano Blanco del Norte, Caño Sucio, and 
Chorro del Rio Tife, 8°43'N, 80°37'W, 200-400 m, 3 
Feb. 1983 (st), Davidse & Hamilton 23483 (MO); Alto 
Calvario, 7 km N of El Copé, ca. 900 m, 14 Jan. 1977 
is Folsom 1304 (MO); divide above El Cope, 8°40'N, 
?35'W, ca. 1,200 m, S 2 1983 (st), Hamilton s 
pelin 4167 (MO). coL i. W of Portobelo, ca. 
m, 9 Sep. 1979 (fr), Antonio 17 71 (MO); Santa ña 
idge lumber road, 3 Oct 8 (fr), M. Correa A. & 
End 1072 (DUKE, Moy hilado ca. 7 mi. on way 
ombre de Dios E of junction with road to Isla Grande, 
i Apr. 1980 (fl), D ee 13615 (MO); trail S of Rio 
BEA on ridge to Cerro Pan de Azucar, 200 m, 20 
4 (fr), — & Kallunki 2010 (MO). DARIÉN: 
amp el between Morti and Sasardi, ca. 400 m, 
14 Feb. 1967 (fr), "Duke 10027 (MO); ridges between 
Rio dis Valley and Pacific Ocean, 7°26'N, 78*05'W, 
300-500 m, 24 Jan. 1982 (A), Knapp & Mallet 3102 
(MO). de El Llano-Cartí Road, 13.8 km N of Pan- 


Volume 76, Number 2 Hamilton 403 
1989 Mesoamerican Psychotria, Part II 
A Vd. > T.C. 
e T S 
et 
zs dii. ü 
e 
e 
o 
+ SS 
* [ ] A Es 15 7 
x + , "E ki 
^ A^ 
es 
se 
L 101 


300 km 


100 95 


105 


85 > 80 


i L 


90 


E 26. 


Distributions of Psychotria chagrensis (solid triangles), P. erythrocarpa (solid circles), P. fosteri 


(olid itn P. jinotegensis var. jinotegensis (open triangles), and P. jinotegensis var. morazanensis (open circle) 


n Mesoamerica. 


American Hwy., 5 Oct. 1977 E disi et al. 5790 
(MO); Cerro Campana, 8?40'N, 5'W, 800 m, 17 
Aug. 1982 (fr), Hamilton & P794 po (MO); 23 July 
1983 (fl), cove et al. 4043 (MO). SAN BLAS: El Llano- 
Carti rd., m from Interamerican Hwy., 9?19'N, 
78*55'W, te 350 m, 19 Sep. 1984 (fr), Nee et al. 
3907 (MO); Cerro Habu, trail from Rio Sidro, 9°23'N, 
78°49'W, 420-600 m, 20 Dec. 1980 (fr), Sytsma et 
al. 2769 (ENCB, MO). 

Psychotria chagrensis may be recognized 
readily by its inflorescences consisting of fascicles 
of several flowers, its large (5-7 x 5 mm 
fruit with conspicuously persistent calyx usually 3- 
4 mm long, and small leaves (usually 4-7.5 X 
1.5-2.5 cm). Leaves appear at the lower end of 
the given size range in Costa Rica and at the higher 
extreme in Mexico and Guatemala. 

The inserted stamens of the pin morph are longer 
than the inserted pistil of the thrum (5.5-6.5 vs. 
2-5 mm) 


30. Psychotria erythrocarpa Schlechtendahl, 
Linnaea 9: 385. 1834 [1835]. Uragoga 
erythrocarpa (Schldl. Kuntze, Revis. Gen. 
Pl. 2: 960. 1891. TYPE: Mexico. Hda. de La 
Laguna (fr), Schiede 385 (holotype, B, de- 
stroyed, photo & fragment, F neg. no. 519; 
isotype, K). Figures 2c, 7c, 26 

Mapouria chamissoana Loesener, Verh. Bot. Vereins 
Prov. Brandenburg 65: 112. 1923. Psychotria 


chamissoana and St zm J. Wash. Acad. Sci 

17: 341. 1927. TYPE: Mexico. Tecolute, Jan. (fr), 
apor 1266 Da B, por photo, F neg. 
46). 


Shrub 1-3 m; young stems pale brown to red- 
brown tomentose or sometimes glabrous, the bark 
gray, furrowed longitudinally; stipules linear, 8- 
x 1.5-2 mm, glabrous to sparsely tomentose, 
caducous, leaving a pale ridge with long red-brown 

ringe to 1 mm long (Fig. 2c). Leaves subsessile 
to petiolate; petioles to 10 mm long, pale tomen- 
tose, flat and grooved above; blades membranous, 
obovate to sometimes elliptic, the apex acute to 
obtuse to rounded, the base attenuate, (3.5-)5- 
8(-10) x (1-)1.5-3.5(-5) cm, sparsely pale to- 
mentose to glabrous above, densely tomentose es- 
pecially near margins to glabrous below, drying 
green-brown above, paler green below; secondary 
veins 6-8(-9) pairs, diverging 35°-50°, eucamp- 
todromous to brochidodromous, straight to arcuate 
near margin, elevated below, dense pale tomentose 
to glabrous below, the axils sometimes with tufts 
of pale hair below; tertiary veins inconspicuous, 
orthogonal reticulate. Inflorescences terminal or 
pseudoaxillary, panicles of cymes or glomerules 

ig. 7c); panicle branched to 3-4 degrees; main 
axis (1.2-)2.5- cm long, the peduncle 
(0.5-)1.5-3 cm long; secondary axes in (2-)3 


pa 
N 


404 


Annals of the 
Missouri Botanical Garden 


ranks, the first-rank axes 2, (0.4—)0.8-2 cm long, 
the second-rank axes 2, (0.2-)0.3-1 cm long, the 
third-rank axes 2, 0.1-0.3 cm long; cymes 
branched to 1—2 degrees; bracts triangular, 1.5 x 
l mm. Flowers sessile to pedicellate, the pedicels 
to 1 mm long; calyx cup-shaped, the tube 0.5-1 
mm long, the lobes 5, barely evident, red-brown 
tomentose; corolla white, the tube cylindrical to 
slightly campanulate, 2-2.5 x mm, white 
pubescent in throat, the lobes 5, narrow-triangular, 
1 mm; stamens 5, the filaments 2-2.5 
mm long in pins, 3-3.5 mm long in thrums, the 
anthers 0.5-0.8 mm long; style 3.5-4 mm long 
in pins, 2-2.5 mm long in thrums, the branches 
linear in pins and club-shaped in thrums. Fruit 
ellipsoid when dry, 4.5-6 mm long, 3-4.5 mm 
diam., maturing red, drying brown, glabrous to pale 
puberulent when dry; calyx persistent as a beak to 

mm long; seed dorsal surface with 4 deep 
longitudinal furrows, the ventral surface with 2 
very deep longitudinal furrows. 


Distribution (Fig. 26). 
Mexico (primarily Veracruz and Chiapas) and up- 
land Guatemala, and extending to Cayo, Belize, in 
areas of subevergreen forest with usually tropical- 


Commonly collected in 


mountainous or sometimes equatorial to subtropi- 
cal-tropical climate. Psychotria erythrocarpa 
ranges in elevation 0-2,000 m, lower in Veracruz 
and higher in Chiapas and Guatemala. It has been 
collected in flower throughout the year, primarily 
April-July, and in fruit throughout the year, pri- 
marily August-January. 


Selected specimens examined. MEXICO. CHIAPAS: 

: mi. S of La Trinitaria, 
1965 (fl), Breedlove 10593 (ENCB, 
F, NY); 12 mi. E of La Trinitaria gre rd. to the Lagos 
de Monte Bello, 1,440 m, 6 Nov. 1965 (fr), Breedlove 


Breedlove 20102 (ENCB, MO, NY); Mpio. Villa Coro, 

ase o cs de Picos nr. Cerro Bola along logging 
rd. of a Agronomos Mexicanos, 1,500 m, 9 
Feb. 1972 WE (ries 23974 (MEXU, MO); Mpio. 
Cintalapa de Figueroa, 5-7 km NW of Rizo de Oro along 
a logging rd. to Cerro Baul and Colonia Figueroa, 100 
m, 19 Apr. 1972 (fl), Breedlove 24698 (DUKE, F, 
MEXU, MO i 


orre 
m, 24 aca 1972 (fr), Breedlove 27360 (DUKE, MO, 
NY); Mpio. Suchiapa, steep canyon 15 km SW of Su- 
chiapa along rd. to Villa Flores, 750 m, 3 Oct. 1972 (fr), 
Breedlove 28265 (DUKE, MEXU, MO, NY); Mpio. Cin- 


o a Cienega, 850 m, 5 
Oct. 1972 (fr), Breedlove 28450 (DUKE ENCB, MO, 
NY); Mpio. Soyaló, 5 km N of Soyaló along rd. to Bochil, 
1,300 m, 26 Oct. 1971 (fr), Breedlove & Thorne 21313 
(DUKE, F, MEXU, MO, NY); Mpio. Amatenango del 


Valle, creek “Chenek” Ha” " nr. Amatenango, 1,710 m 
cd e 1967 (fr), Ton 2678 (ENCB, F, MO); Monsecrata: 
3'N, 94°00'W, Apr. 1925 (fl), Purpus 14334 (A, 
F) Mpio. b des e Rancho Carmen along 
rd. from Acala to Venustiano Carranza, 780 m, 18 July 
1966 (fr), Laughlin 1329 (F, NY). GUERRERO: barr. 


1972 (fl), Gimate 658 (ENCB, MEXU, SLPM). 
JALISCO: Estación Biología Chamela, cerca de arroyo Pailles, 
8 Dec. 1970 (fr), Pérez 279 (MEXU). MORELOS: mon- 
tanas N de Cuernevaca, 2,000 m, 5 Nov. 1972 (fr), J. 
Vázquez 3981 (MEXU). oaxaca: 5 km S of Tuxtepec, 
limestone hill, less than 100 m, 30 Aug. 1974 (fr), J. 
'onrad & R. Conrad 3246 (F, GH— 3 sheets, MO, NY); 
Pochutla, Boquerón, camino de Forramoes, 150 m, 7 
May 1971 (fl), Conzatti et al 3265 (GH); Tehuantepec, 
Santa Lucia, 1,200 m, 5-6 July 1971 (fl), T. MacDougall 
H28(NY). PUEBLA: Mpio. de Jalpan, Ajenjibre, Nov. 1950 
(fr), Paray 322 (ENCB). san LUIS POTOSÍ: Mpio. de Cuidad 
el Maíz, El Salto, 400 m, 31 July 1960 (fl), L. González 
0. 138 (ENCB); Tamazunchale, 300 m, July 1937 (fr), 
Lundell & Lundell 7144 (F, MEXU, NY); El Abra, 10 
km E of Ciudad Valles, 1,500 m, 17 June 1956 (fl), 
Rzedowski 7772 (ENCB, MEXU, SLPM); Mpio. de Ta- 
masopo, San Nicolás de los Montes, 800 m, 28 May 
1959 (fl), Rzedowski 10664 (ENCB). SINALOA: Mazatlán. 
pio. Rosario, Chametla, Cuacoyolitos, 20 m, (fr), J. 
González Ortega 5875 (GH, MEXU, US). TAMAULIPAS: 
8 mi. N of Aldama, rd. to San Rafael, oak scrub, 330 
m, 10 Dec. 1959 (fr), M. Johnston 4945 (MEXU); vic. 
Tampico, 15 m, 1-22 May 1910 (fl), Palmer 478 (GH, 
K, MO, NY Y, US); Mpio. de Ocampo, 3 km NE of Flores 
Magon, 400 m, 7 Aug. 1969 (early fr), Puig 5135 
(ENCB); Mpio. de Gómez Farías, 2 km E of Gómez Farias, 
500 m, 11 June 1982 (fl), Valiente & Viveros 161 
ENCB, MEXU —2 sheets, MO). VERACRUZ: Mpio. Vega 
de Altorre, Canada de Mesillas, entrada por Santa Ana, 
360 m, 24 July 1981 (fr), Calzada 7740 (MEXU); Mpio. 
Coatepec, Cerro de la Palmas entre Jalcomulco y Tuza- 
mapan, 19%23'N, 96%48'W, 800 m, 1 Apr. 1979 (fl), 
Castillo & Tapia 501 (MEXU), Isla Lobos Om, 1 June 
1966 (f), E. Chávez s.n. (ENCB, MEXU); Mpio. de 
Naolinco, E de lava al S de La Concepción, 1,000 
m, Aug. 1973 (fr), Chazaro & Dorantes 255 (ENCB, 
MEXU); Mpio. Ozuluama, 5 km adelante de Ozuluama, 
hacia le ico, 


MEXU) c 


— 
T 


= 


pio. Apazapan, 2 km of Emiliano Zapata (= 

Carrizal) 19923! N, 96°38’ W, 400 m, 27 June 1980 (fl), 
Hansen & Nee 7522 (F, MO); Atoyac, 14 May 1937 
fl), Matuda 1484 (MEXU), Matuda S-4 (A, F, MO); 
Casitas-Gutiérrez Zamora, cerca ejido villa Sag c 
30 m, ms June 1970 (fl), Nevling & Gómez-Pompa 11 

MEXU); Coatzacoalcos, 21 Hos i. (fl), Tsai 
3106 (F) Cerro Monte de Oro, , 20 June 1972 
(fl), C. Vázquez Y. 614 (MEXU); Mao. Coetzala, as E 
con Omealca, 600 m, 16 de 1976 (fr), V. Váz 

458 (ENCB, MEXU — 2 sheets); Mpio. Pu ente National 
50 m, 16 Oct. 1970 (fr), Ventura — (ENCB, ee 
Mpio. de Totutla, Mata Oscura, 750 m, 25 Nov 

fr), enter 4575 (ENCB); Mpio e Dos Bins. he 
Gordo, 450 m, 15 Jan. 1975 (fr), aa 10809 (ENCB, 
MO). GUATEMALA. BAJA VERAPAZ: 13 km S of turnoff to 


~ 


aa 


Volume 76, Number 2 
1989 


Hamilton 405 
Mesoamerican Psychotria, Part II 


Salamá on hwy. to Cobán, 1,000 m, 14 July 1977 (early 
fr), Croat 41119 (MO); 5 km SE of Granados, 15 July 
1970 (early fr), Harmon & Dwyer 3105 (ENCB, F, GH, 
, US), 6 km from Rabinal, rd. to Guatemala City, 
thickets of Pachirax River, Sierra Chuacüs, 1,140 m 
Oct. 1972 (fr), 4. Molina R. & A. Molina 27784 (F. 
CHIQUIMULA: between Chiquimula and La Laguna, 500- 
1,000 m, 27 Oct. 1939 (fr), Steyermark 30684 (F). 
ESCUINTLA: sin. loc., 1942 (fr), J. I. Aguilar 1729 (F). 
GUATEMALA: Estancia Grande, 600 m, 8 Dec. 1938 (st), 
Standley 59208 (F). HUEHUETENANGO: between Nentón 
Las Palmas, via Yalisjao, Rincón Chiquite, aa ere 
Guaxacana, in Sierra de los Cuchumatanes, 800-1,20 


m, 30 Aug. 1942 (fr), Sod 51630 (F); otto 


6047a (GH, US). JALAPA: hills NE of Jalapa, rocky scrub- 
oak forest, 1,400-1,600 m, 10 Nov. 1940 (fl), Standley 
76783 (F); mountains along rd. between Jalapa and San 
Pedro Pinula, 1,4 nee wae m, 12 Nov. 1940 n Stand- 
ley 77070 (F). JUTIAPA: vic. Jutiapa, 850 24 Oct.- 
5 Nov. 1940 (fr), Standley 76317 (F, NY, US): quebrada 
etween ae (Dept. Jalapa) and 

00 m, 19 i 1940 (st), Standley 
77633 (F). QUICHÉ: sin. loc., 2 (A ), J. L Aguilar 1143 
(F); 1942 (fr), J. I. Aguilar p (F). za ACAPA: Loma El 
Picacho, above Santa Rosalía, dry SW -facing rocky s 
and bluffs of metamorphosed o Bis 1,200-1,600 
m, 15 Jan. rrt d diei 42705 (F, NY). BELIZE. 
CAYO: Cuev io 1 km al O de Augustine, 
400 m, 2 Sep. 1973 (fs), pose 4183 (MEXU). 


A minority of glabrous individuals appear in 
many populations of Psychotria erythrocarpa (D. 
Lorence, pers. comm.), so P. chamissoana, defined 
by its glabrous aspect, is included within P. ery- 
throcarpa. 

Psychotria erythrocarpa may be recognized by 
its usually pale brown to red-brown tomentose as- 
pect, obovate leaves with few (usually 6-8) sec- 
ondary veins diverging acutely (35?—50*), and small 
(usually 2.5-5.5 cm long) inflorescences with sec- 
ondary axes two per rank in three ranks. The fruits 
occupy the small end of the spectrum in Veracruz 
and the large end in Chiapas and southward. 


31. debo fosteri C. Hamilton, Phytologia 
64: 1988. TYPE: Veraguas: 
E men. northern tip of Coiba Island, 
26 Aug. 1970 (fl, fr), Foster 1605 (holotype, 
DUKE; isotypes, F, GH, MO). Figure 26. 


Panama. 


Shrub ca. 1 m high; young stems glabrous, the 
bark slightly furrowed longitudinally; stipules 
sheathing, narrow, with 2 triangular lobes, the 
sheath 8-12 x 1-3 mm, the lobes 2-3 mm long, 
fringed, with darker red-brown midrib leading to 
apex of each lobe, caducous, leaving a red-brown 
ridge with irregular fringe. Leaves petiolate; peti- 


oles 5-10 mm, glabrous, flat above; blades mem- 
branous, elliptic-obovate, the apex acuminate to 
mucronate, the base attenuate, (6-)8-12 x (2-)3- 

cm, glabrous above and below, drying green- 
brown above, deep red-brown below; secondary 
veins 11-14 pairs, diverging 70°-80°, brochido- 
dromous often with collector vein distinct, slightly 
arcuate, prominulous below, glabrous, drying deep- 
er red-brown than blade, the axils lacking domatia 
or hairs; tertiary veins inconspicuous, reticulate. 
Inflorescences terminal, fascicles of several flow- 
mm across; peduncles 
lacking; bracts broadly triangular, irregularly cleft, 


ers, ca. 8 mm long, 
ca. 5 X ca. 5 mm, fringed. Flowers sessile; calyx 
cup-shaped, the tube 1.5 mm long, the lobes 5, 
triangular, 0.5-1 x 0.5 mm, slightly fringed; co- 
rolla white, the tube cylindrical, 3.5 x 0.8 mm, 
white pubescent in throat, the lobes 5, triangular, 
mm; stamens 5, the filaments 3 mm long 
in pins, not seen in thrums, the anthers 0.5 mm 
long; style 4-4.5 mm long in pins, not seen in 
thrums, the branches linear. Fruit when dry ellip- 
soid, 5-6 mm long, 2.5-3 mm diam., maturing 
orange (probably to red), drying dark red-brown; 
calyx persistent, the tube 1—1.5 mm long, the lobes 
ca. 1 mm long; seed dorsal surface with 5 longi- 
tudinal furrows, the ventral surface with 2 longi- 
tudinal furrows. 
Distribution (Fig. 26). 
type collection from Coiba Island, Panama, in trop- 
ical moist forest with tropical-equatorial climate. It 
was collected in flower and fruit on August 26. 


Known only from the 


Psychotria fosteri may be recognized as a mag- 
nified version of P. chagrensis, having an inflo- 
rescence consisting of a fascicle of flowers and fruit 
ellipsoid with a conspicuous persistent calyx. Psy- 
chotria fosteri differs in having larger (sheath 8- 
vs. 4-7 mm long) biaristate (vs. aristate) stip- 
ules, larger mature leaves (usually 8-12 x 3-5 
vs. 4-7.5 x 1.5-2.5 cm), shorter calyx lobes 
(0.5-1 vs. 2-2.5 mm), and much shorter corolla 
tubes (3.5 vs. 6-7 mm). 

The lone flowering collection is a pin morph. 


— 
N 


32. Psychotria fruticetorum Standley, J. Ar- 
nold Arbor. 11: 42. 1930. TYPE: Honduras. 
Comayagua: vic. Siguatepeque, 1,080-1,400 
m, 14-27 Feb. 1928 (fr), Standley 56197 
(holotype, F; isotype, A). Figure 25. 


Shrub, 0.5-3 m tall; young stems glabrous, the 
bark red-brown or pale, smooth; stipules sheathing, 
biaristate with extensions arising from apical cor- 
ners of sheath, 1.5-2 x 1.5-2 mm, glabrous, 


406 


Annals o 
Missouri Botanical Garden 


caducous, leaving a pale or red-brown ridge with 
or without fringe. Leaves petiolate; petioles 2-5 
mm long, glabrous, flat above; blades membranous 
to subcoriaceous, elliptic to slightly obovate, the 
apex acute to acuminate, the base narrowly cu- 
neate to attenuate, the margin often inrolled, (3—)5— 
7(-9) x (1-)2-4(-4.5) cm, glabrous above and 
below, drying chalky yellow-green to red-brown; 
secondary veins (4-)5-7(-8) pairs, diverging 55°- 
65°, eucamptodromous, straight then constantly 
arcuate toward margin, barely elevated below, gla- 
brous, the axils often with minute tufts of white 
hairs below; tertiary veins inconspicuous, orthog- 
onal reticulate. Inflorescences terminal or pseu- 
doaxillary, panicles of cymes; panicle branched to 
3 degrees; main axis 1.5—4.5 cm long, the peduncle 
0.3-2.5 cm long or rarely lacking; secondary axes 
in (2-)3-4(-5) ranks, the first-rank axes 2 or 4, 
the longer pair 0.5-1.8 cm long, the shorter pair 
when present 0.5 cm long, the second-rank axes 
2, 0.1-1.1 cm long, the third-rank axes 2(4), the 
longer pair 0.2-0.7 cm long, the shorter pair when 
present 0.2 cm long, the fourth-rank axes 2, 0.4- 
0.5 cm long, the fifth-rank axes 2, 0.3 cm long; 
cymes branched to 2 degrees; bracts broad, irreg- 
ular, to 1 mm long, glabrous to puberulent, often 
minutely fringed. Flowers on pedicels 0.5-1 mm 
long; calyx cup-shaped, the tube 0.5 mm long, the 
lobes 5, triangular to barely evident, to 0.5 x 0.5 
mm, often minutely fringed; corolla white, the tube 
cylindrical, 2 x 1.5 mm, white pubescent in throat, 
the lobes 5, linear, 1.5-2 x 1 mm; stamens 5, 
the filaments 1.5-2 mm long in pins, 3-3.5 mm 
long in thrums, the anthers 0.8 mm long; style 
3.5-5 mm long in pins, 2.5 mm long in thrums, 
the branches short, linear, often recurved. Fruit 
when dry ellipsoid, 4-5 mm long, 3-3.5 mm diam., 
maturing red, drying red-brown; persistent calyx 
to 0.5 mm long; seed dorsal surface with eep 
longitudinal furrows, the ventral surface with 2 
deep longitudinal furrows. 


Distribution (Fig. 25). Widespread from 
Mexico through Panama, mostly on the Caribbean 
side, with greatest concentration in Belize and east- 
ern Petén, Guatemala, in tropical to premontane 


and parts of Guatemala. It has been collected in 
flower throughout the year, primarily March-July, 
and in fruit throughout the year. 


Selected specimens examined. | MEXICO. CAMPECHE: 
Tuxpeña, 13 Dec. 1931 (fr), Lundell 1062 (F). CHIAPAS: 
Mpio. Tenejapa, Ala Shashib River below Habenal, paraje 


of Mahben Chauk, 990 m, 15 July 1964 (fl), Breedlove 


Ocote 32 k cozocoautla, 6 A 
1972 (fr), Breedlove 27427 (MEXU, MO); Mpio. Oco- 
singo, nr cotal Grande, 800 n 21 


pio. de Juárez-Hidalgo, barrancas al N e Juárez-Hi- 
dalgo, 1,800 m, 10 May 1981 
al. 6114 (MEXU, MO i 


tumal- Felipe Carrillo oed a Nov. 198 
& Cortés rad (MEXU); 11 
N of Uni 100 m, 7? May 1982 (fl), Davidse et = 
20183 (MO); Mpio. je Carrillo Puerto, 21 Aug. 1 

(fr), G. Pérez S. 411 (ENCB, MEXU). TABASCO: Mpio. 
RaT km 15 de Huimanguillo a Francisco Rueda, 
12 June 1 979 b Cowan 2284 (ENCB, MEXU); Ba- 
a, 1-6 June 1939 (fr), Matuda 3266 
(A, F, MEXU, NY. Mpio. Huimanguillo, 1 km al N de 
Chontalpa, i n 1971 (fr), Puig 322 (ENCB, MEXU). 
VERACRUZ: Mpio. de Huayacocotla, camino a Rancho Nue- 
vo, 1,600 m, £i Mar. 1972 (fl, fr), R. Hernández M. & 
B. Rosales A. 1559 (MEXU). GUATEMALA. ALTA VERAPAZ: 
Chahal, El Mago, ca. 90 m E of Sebol Road, 15 Oct. 
1968 (fr), Contreras 7938 (F, NY); Rubelsanto, Balas- 
trera, 23 July 1975 (fl), Lundell & Contreras 19548 
(MO); SW of Lanquin, vic. caves, 600-1,000 m, 21 Feb. 
1942 (st), Steyermark 44128 (F, US); savanna N of 
Concepción, 3-5 mi. SE of Finca Yalpemech, nr. Alta 
Verapaz-Petén boundary, 100-110 m, 23 Mar. 1942 
(fl), Steyermark 45272 (E, NY). CHIQUIMULA: 3 mi. SE 


bollas, along Rio L m 

1939 (fr), Steyermark 31237 (F); Steyermark 31321 
(F). IZABAL: Cadenas/Puerto Méndez, 8 Sep. 1969 (fr), 
Contreras 9099 (MO); vic. Quiriguá, 75-225 m, 15-31 
May 1922 (fl), Standley 23922 (GH, US); Standley 
24248 (GH, US); seashore around Punta Palma, across 
bay from Puerto Barrios, 22 Apr. 1940 (st), Steyermark 
39830 (F). PETÉN: San Clemente to Dos Arroyos, 1 May 
1931 (fl), Bartlett 12819(F, US); Tikal, on top of Temple 
IV, 28 June 1960 (fl), Contreras 1192 (DUKE, F, NY); 
La Cumbre, W of km 139 on Cadenas Road, 23 Sep. 
1966 (fr), Contreras 6198 (F, NY); Carmelita, in thicket 
bordering aviation field, 2 July 1942 (fr), Egler 42-302 
(F); Tikal National Park, Bajo del Hormiguero, 6 July 
1959 (fl, fr), Lundell 16214 (DUKE, MEXU, NY); km 
62 of Flores-Poptün Rd., 16 Sep. 1976 (fr), Lundell 
& Contreras 20369 (MO); +35 km E de Sta. Elena, 
brecha El Remate-Tikal, 200 m, 9 Nov. 1965 (fr), A. 
Molina R. 15443 (F, NY, US); sitio arqueológico en el 
camino que conduce a cienaga “‘ 


S 
a 
zz 
Sm 
a 
O 
[a 
= 
= 
o 
A 
G 
p 
>% 
S 
e 
Led 
E 
Es 
to 
a 


2655 (ENCB, F, MO, NY). 
8 June 1974 (fl), Dwyer 12787 (MO, 
1933 (fr), Gentle 946 (A, F, MO, dd 9 Apr. 19 
Gentle 1188 (A, F, GH, K—2 sheets, MO, ); 
Isabella Pine Ridge, June 1933 (fl), Lundell 4243 (F); 
nr. Manatee Lagoon, pine ridge, 9 June 1905 (fl), Peck 
23 (GH, K). cayo: Duck Run, 11 May 1931 (fl), veo 
13122 (F, Ky vic. Cuevas S of O 600 m 
30 May 1973 (fl), Croat 23589 (F, MO); Moüntsin Pins 
idge, San Agustin, banks of Rio Frío, cod Aug. 1936 
Un Lundell 6702 (F, GH, NY, ro AL: 1.5 km 
f Buena Vista, 1.5 km W of Nor n Highwa ay on 
inu rd., 23 June 1973 (early s Di 11373A 


Volume 76, Number 2 
1989 


Hamilton 407 


Mesoamerican Psychotria, Part II 


(F, MO, NY); a EN m Aug. 1933 (fr), onn 
4913 (NY). o Northern Highwa 
to New River Toll | Bridge, 7 .. 1974 (fl), Dwyer 19746 


— 2 sheets); Honey Camp, 19 Oct. 1929 (fl, 
fr), Lundell 617 (F, GH, K, MO sheets, NY, US). 
STANN CREEK: Silk Grass Creek Reserve, 20 Sep. 1939 


(fr), Gentle 3002 (A, F, K, XU, NY); Stann Creek 
Ww Big Eddy Ridge, 6 Apr. 1941 (fl), Gentle 3546 
, NY); Mullins River Rd., swampy places, 30 m, 


OLEDO: San José, 6.7 mi. N of Columbia Paias 
Station, 13 June 1973 (fl), Dwyer 11150 (MO); Rio 
Temash, between 3 km upriver and mouth, 26 July 1979 
(fr), Dwyer 14779 (MO); Monkey River, - Aug. 1941 
(fr), Gentle 3600 (4, K, MO, 13 on rd. 
W from Punta Gorda, nr. junction of Sou kern Hwy 
and rd. to San Antonio, 30 m, 14 June 1973 (fl), eae 
8212 (DUKE, MO). HONDURAS. AE : 3 km S of 
i e 969 (fr), A. Molina R. 


Santa Rosa de Copán, 1,000 m, 15 Feb. 1982 (fr), Mejía 
124 (TEFH). FRANCISCO MORAZÁN: 1 km from Agalteca, 
800 m, 22 Nov. 1966 (fr), 4. Molina R. 18705 (F, 
NY). GRACIAS A DIOS: Puerto Lempira, orilla laguna de 
Caratusca, 5-8 Sep. 1978 (fr), = apie 4828 (TEFH); 
camino entre Dursuna y Mocorón, 70 m, 14-21 Feb. 
1979 (fr), C. Nelson & Pargas “4906 (TEFH). LA Paz: 
Agua Blanca River between Chinacla and Planes de Mulle, 
1,300 m, 22 Mar. 1969 (fr), d s R. & A. Molina 
24350 (F—2 sheets, bd d 
km de Gracias, 1,000 m 
et al. 91 (MO). oc OTEPEQU E: Deren El Agua Caliente 
and Machuca, hwy. to Guatemala-H border, 1,000 
m, 4 Sep. 1975 (fr), 4. Molina R. a A. Molina 31091 
(ENCB, F, MO). SANTA BARBARA: al N de Sta. Barbara, 
región La Cuesta, 500 m, 13 Dec. 1950 (fr), 4. Molina 
R. 3784 (F, GH). NICARAGUA. CHONTALES: Hda. Corpus, 
W of Juigalpa, 12%07'N, 85*28'W, 100 m, 5 Sep. 1982 
(early fr), W. D. Stevens 21782 (MO). MATAGALPA: 7 km 
al SO de Buenavista, carret. a cod 13°15'N, 
85?24'W, 300-440 m, 1 Mar. 1983 (fr), P 

& Robleto 21016 (MO); Hda. Sta. Maria á 
1,300-1,600 m, 17 Jan. 1977 (st), Tomlin 145 (MO). 
RIVAS: Isla de Ometepe, al lado N del Volcán Concepción, 
11%34'N, 85°37'W, 250-350 m, 12 Mar. 1981 (fr), 
Sandino 545 (MO). ZELaYa: Prinzapolka, 23 June 1978 
(fr), Neill 4551 (MO); Caño El Hormiguerro, on E slope 
of El Hormiguerro, 13?46'N, 84%59"W, 750-800 m, 17 
Mar. 1980 (fr), Pipoly 6104 (MO); nr. Tala Has and 
Puente Mango, over Rio Sd 14?41'N, 84°03'W, 
40-60 m, 18 Apr. 1978 (fl), W. D. ~n 7597 (MO); 


rd. to Panua, entrance ca. 7 f Sta. Marta, 
, 30-40 


8 

. 19 W. D. Stevens 7761 (M 
d Bluefields, 11%59' N, 83°46'W, 10-40 m, 2-3 Apr. 
1981 (fl), IW. D. Stevens 19836 (MO); vic. jet. of rd. to 
Alamikamba with rd. LH El Empalme and Limbaika, 
13*32'N, 84°30'W, 25 A July E (8, fr), W. D. 
je 21675 (MO); 4 rto Cabezas, 10- 
, 11 June 1978 (fr), Vincelli 533 (MO). PANAMA. 
CANAL AREA: Ancón Hill, 25 Feb. 1923 (fr), Piper 5554 
(US). PANAMÁ: nr. Calzada Larga, swampy area over 
exposed limestone, 27 Sep. 1970 (fl, fr), Croat 12389 
(F, GH, MO, NY); San José Island, Perlas vr 
13 June 1945 (fl Erlanson 279 (GH, NY, US); 8 Jan. 
1946 (fl, early fr), : Johnston 1071 (GH, MO, US); T 
Río Tocumen, N of Chepo rd., 0-30 m, 27 Jan. 1935 


(f), Hunter & Allen 240 (MO); southern tip of Isla de 
Rey, Punta de Cocos, 8?17'N, 78°55'W, 0-20 m, 19 
Jan. 1982 (fl, fr), Knapp & Mallet 2948( Wr SAN BLAS: 

mainland opposite Isla q 19 Oct. 1966 
(fr), Duke 8989 (MO—2 sheets). 


Psychotria fruticetorum may be recognized by 
its small leaves (usually 5-7 x 2-4 cm) drying 
chalky yellow-green to red-brown with barely el- 
evated secondary veins and inconspicuous tertia- 
ries, its small inflorescences (1.5-4.5 cm long) with 
secondary axes usually 2 per rank in 3-4 ranks, 
and small (4-5 x 3-3.5 mm) ellipsoid fruit. Mex- 
ican collections differ from those from the rest of 
the range in having generally smaller leaves, 5 vs. 
6-7(8) secondary veins, and a higher fruit length- 
to-diameter ratio. 

The exserted pistil of the pin morph is often 
longer than the exserted stamens of the thrum 


(3.5-5 vs. 3.5-4 m 


33. Psychotria jinotegensis Nelson, Molina & 
Standley, Phytologia 50: 1. 1981. TYPE: Nic- 
aragua. Jinotega: region of La Montanita and 
Las Mesitas, in sierra W of Jinotega, ca. 
1,100-1,400 m, 29 June 1947 (fl), Standley 
10314 (holotype, EAP, n.v.; isotype, F, n.v.). 
Figures 9d, 26. 


Shrub (0.5-)1-4(-6) m tall; young stems fer- 
rugineous-pubescent or very sparsely tomentose, 
the bark smooth; stipules sheathing, ovate to linear, 
15-23 x 4-6 mm, usually ferrugineous-pubes- 
cent, caducous, leaving a pale ridge with dark red- 
brown fringe. Leaves petiolate; petioles 0.5-1.5 
cm long, pubescent or glabrous, flat above; blades 
membranous, elliptic to oblanceolate, the apex acute 
or sometimes acuminate, the base attenuate, (7 —)9— 
14(-15.5) x (2-)2.5-4.5(-5.5) cm, pale tomen- 
tose or glabrous above, densely pubescent or gla- 
brous below, drying gray-green to red-brown; sec- 
ondary veins 9-14 pairs, diverging 40?-50 
eucamptodromous to brochidodromous, straight 
then arcuate near margin, elevated below, pubes- 
cent or glabrous below, the axils with tufts of hairs 
below; tertiary veins evident, orthogonal reticu- 
late to slightly percurrent. /nflorescences terminal 
or pseudoaxillary, few-branched panicles of con- 
densed cymes; panicle branched to 3-4 degrees; 
main axis (2.5-)4—6.5 cm long, the peduncle lack- 
ing or to 1(-3) cm long; secondary axes in (2-)3 
ranks, the first-rank axes 1-2, (1-)1.5-5 cm long, 
the second-rank axes 2, 0.5-2.5 cm long, the third- 
rank axes 2(4), the longer pair 0.2-0.8 cm long, 
the shorter pair when present 0.1 cm long; cymes 
branched to (1)2 degrees; bracts and bracteoles 


408 


Annals of the 
Missouri Botanical Garden 


linear, 1.5-3 mm long; usually ferrugineous-pu- 
bescent. Flowers sessile or on pedicels to 1.5 mm 
long; calyx cup-shaped, the tube cylindrical, 0.5- 
1 mm long, the lobes 5-6, triangular, 0.5-1 x 
0.7 mm, puberulent; corolla white, the tube cylin- 
drical to campanulate, 3.5-4 x 1.5-2 mm, white 
pubescent in throat, the lobes 5-6, 1.5-2.5 x 1.2 
mm; stamens 5-6, the filaments 3 mm long in pins, 
5-5.5 mm long in thrums, the anthers 1 mm long; 
style 5 mm long in pins, 3-4 mm long in thrums, 
the branches linear. Fruit when dry ellipsoid, (5—) 
5.5-6.5 mm long, 4-5 mm diam., maturing red, 
drying red-brown, ferrugineous-pubescent or gla- 
brous; persistent calyx conspicuous, 1-3 mm long 
(Fig. 9d); seed dorsal surface with 6-8 irregular 
longitudinal furrows, the ventral surface with 
deep regular plus several irregular longitudinal fur- 
rows. 


Distribution (Fig. 26). Known from south- 
eastern Guatemala through northwestern Nicara- 
gua in tropical moist forest with tropical to tropical- 


mountainous climate. 


Psychotria jinotegensis may be recognized by 
its usually ferrugineous aspect, elliptic to oblan- 
ceolate leaves with secondary veins diverging 40?— 
50°, and ellipsoid fruit with conspicuous (1-3 mm 
long) persistent calyx tube. 


KEY TO THE VARIETIES OF PSYCHOTRIA JINOTEGENSIS 


la. Ferrugineous-pubescent and/or tomentose on 
oung stems, both leaf surfaces, and o 
fruit; secondary vein axils with tufts of reddish 
airs a. var. Jinotegensis 
. Generally glabrous or only sparsely tomentose; 
secondary vein axils with tufts of white hairs .. 
30b. var. morazanensis 


- 
c 


33a. Psychotria jinotegensis Nelson, Molina 
tandley var. jinotegensis. Figure 


Shrub; young stems ferrugineous-pubescent. 
Leaves: petioles ferrugineous-pubescent; blades pale 
tomentose above, densely ferrugineous-pubescent 
below, drying green-brown to deep red-brown; sec- 
ondary veins (9-)10-14 pairs, ferrugineous-pu- 
bescent, the axils with tufts of reddish hairs below. 
Fruit when dry ferrugineous-pubescent. 


Distribution (Fig. 26). Known from south- 
eastern Guatemala eastward across southern Hon- 
duras and northern El Salvador to northwestern 
Nicaragua at elevations of 1,000-2,000 m. It has 
been collected in flower June-July and December— 
February and in fruit August- February. 


Selected specimens examined. GUATEMALA. BAJA 
VERAPAZ: Unión Barrios, 27 Feb. 1972 (fr), Contrereas 
11062 (MO); along hwy. to Cobán, ca. 1.5 km S of 
turnoff to Salamá, pine a 1,300 m, 16 July 1977 
(early fr), Croat 41138 (MO); nr. La Unión (Barrios), 
Sierra de las Minas, 1,800 m, 18 Jan. 1974 (fl, fr), > 
O. Williams et al. 43559 (F). CHIQUIMULA: 
Nonojá, 5-8 km E P pu 600-1,800 m 
1939 (fr), Steyermark 31675 (F). EL PARAISO: between 
Calera and middle slopes of quebradas of Volcán Siglo, 
2,000-2,200 m, 20 Jan. 1942 (fr), Steyermark 42992 
F, US). HONDURAS. COMAYAGUA: ravine nr. El Achote, 
hills above plains of Siguatepeque, 1,350 m, 30 July 1936 
(early fr), Yuncker et al. 6229 (F, GH, K, MO). DISTRITO 
CENTRAL: 2.5 km N of Zambrano along rd. to San Fran- 

cisco de Soroguara, 1,370 m, 1 July 1970 (fl), rus 
& Pohl 2245 (F, MO, NY); nr. Zambrano, 26 Sep. 
(fr), Lauvert & Barkley 39552 (MO, TEFH); San Matin 
NO de Tegucigalpa, 1,400 m, 25 Sep. 1983 (fr), Paredes 
97 (TEFH). EL PARAISO: drainage of Río Yeguare, már- 
genes del Rio Lizapa entre Guinope y Las Casitas, 1,300 
m, 1 Nov. 1950 (fr), 4. Molina R. 3357 (MO); Francisco 
Morazán, Cerro Canta Gallo, 14 km NE de Teguc e 
1,046 m, 2 Oct. 1983 (fr), M. Castro M. 83 (TEFH); 
along Río Rancho Quemado, S of Tegucigalpa, km 20- 
25 on rd. to Sabana Grande, 1,300 m, 9 Nov. 1966 (fr), 
A. Molina R. 18648 (GH, NY). INTIBUCÁ: 2 km de La 
e La Pozona, 28 June 1974 (fl), J. 
z& Bos 236 (TEFH). EL SALVADOR. C in 
ATENANGO: S del Rio Nomiapa, La Palma, 1,100 m, 6 
July 1976 (early fr), Montalvo 4695 (MO). NICARAGUA. 
JINOTEGA: sierra W of Jinotega, trail to Cerro de la Cruz, 
1,050-1,350 m, 27 June 1947 (fl), Standley 10214 (F). 


— 


33b. Psychotria jinotegensis Nelson, Molina 
& Standley var. morazanensis C. Hamilton, 
Phytologia. 64: 226. 1988. TYPE: El Salvador. 
Morazán: easternmost peak, Montes de Ca- 
caguatique, nr. summit on N side, 13?46'N, 
88?13'W, 1,500 m, 25 Dec. 1941 (fr), Tuck- 
er 610 (holotype, US: isotypes, F, K, NY). 
Figure 26 


Shrub; young stems very sparsely tomentose. 
Leaves: petioles glabrous; blades glabrous above 
and below, drying gray-green to pale red-brown; 
secondary veins 9-11 pairs, glabrous, the axils 
with tufts of whitish hairs below. Fruit glabrous 
when dry. 

Distribution (Fig. 26). 
type collection from eastern El Salvador, at ca. 
1,500 m. This variety was collected in fruit De- 


Known only from the 


cember 25. 


34. Psychotria mirandae C. Hamilton, Phy- 
tologia 64: 229. 1988. TYPE: Mexico. Chia- 
pas: Berriozábal a Las Vistas, 17 July 1949 
(fl), Miranda 5395 (holotype, US; isotype, 
MEXU). Figure 27. 


Shrub ca. 1 m tall; young stems red-brown 
tomentose, the bark pale, smooth; stipules sheath- 


Volume 76, Number 2 Hamilton 409 
1989 Mesoamerican Psychotria, Part II 
a 7 a -J- | TCs 
e 
| E f+ i 
MM ee 
z e 
^ e 
+ à 
F 15 4 
+ d P Pe s> Je 
? $$. q 
S 
Ox) 
L 10 4 
300 km + o la 
— e 
v ve 
105 100 95 90 85 a “ec 
FIGURE 27. Distributions of Psychotria mirandae (triangles) and P. nervosa (circles) in Mesoamerica. 


ing, rounded at apex, 4-8 x 3-4 mm, red-brown 
tomentose outside, caducous, leaving a pale ridge. 
Leaves subsessile to petiolate; petioles 1-4 mm 
long, red-brown tomentose, furrowed above; blades 
membranous, narrowly obovate, the pes caudate, 

2 m, glabrous 
above and below, drying dull bom secondary 


the base attenuate, 6-1 


veins 7-10 pairs, diverging 48?-55?, eucampto- 
dromous, slightly arcuate, prominent below, red- 
brown sparsely tomentose below, the axils with 
small tufts of hair below; tertiary veins evident, 
reticulate to slightly percurrent. /nflorescences ter- 
minal, fascicles of flowers or with several short 
axes, 1-1.5 cm long, 1-2.5 cm across; peduncle 
lacking; bracts triangular, 0.5 x 0.5 mm, glabrous 
to sparsely tomentose. Flowers subsessile, the ped- 
icels 1-2 m 
2.5 mm long, the lobes 5, linear, 2-3 x 1.2 mm, 
sparsely ciliate on margin; corolla white(?), the tube 
cylindrical, 3 X 1 mm, white pubescent in throat, 
the lobes 5, linear, 1.5-2 X 0.5 mm; stamens 5, 
the filaments not seen in pins, 4 mm long in thrums, 


m long; calyx cylindrical, the tube 2- 


the anthers 0.8 mm long; style not seen in pins, 
2.5-3 mm long in thrums, the branches club- 
shaped. Fruit ellipsoid when dry, 5-7 mm long, 
4-5 mm diam., maturing red, drying red-brown, 
red-brown tomentose; calyx persistent, the tube 2- 
2.5 mm long, the lobes ca. 2 mm long; seed dorsal 
surface with 5 longitudinal furrows, the ventral 
surface with 2 longitudinal furrows. 


Distribution (Fig. 27). Limited to eastern Ve- 
racruz and northwestern Chiapas, Mexico, at 700- 
1,000 m elevation in evergreen forest with equa- 
torial-tropical to tropical climate. It has been col- 
lected in flower in March and May-July and in 
fruit in August and December 


Additional specimens examined. MEXICO. CHIAPAS: 
Mpio. Berriozábal, 13 km N of Berriozábal nr. Pozo Turi- 
pache and Finca El Suspiro, 900 m, 15 May 1973 (fl), 
Breedlove 35316 (MEXU); Cerro a San Martín, Mar. 
1845 (fl), Galeotti 2684 (F—3 sheets, US); Berriozábal 
a Las Vistas, 4 Dec. 1949 (fr), Miranda 5834 (MEXU, 

; San Fernando, Predio del Rosario, 21 June 1951 
(fl), Miranda 7216 (MEXU). VERACRUZ: Mpio. Catemaco, 
cerro entre Zapoapan y San Juan Seco al SW de Lago 
Catemaco, 700 m, 7 June 1972 (fl), Beaman 6094 (F, 

U— 2 sheets); region of San Andrés Tuxtla, nr. 
Zapoapan, SE of Catemaco, 17 Aug. 1953 (fr), Dressler 
& Jones 93 (GH, NY, US); Catemaco, 25 Mar. 1956 
(fl), Paray 1946 (ENCB); km 156.5 of Veracruz-Coatza- 
coalcos rd., 13 July 1974 (fl), Sohmer 9409 (MEXU). 


Psychotria mirandae may be recognized first 
by its marked resemblance to P. nervosa and sec- 
ond by its contracted (sometimes contracted to 
fascicles of flowers) inflorescences and wider (4-5 
mm vs. 2.5-3 mm diam. in P. nervosa) fruit with 
persistent calyx having a conspicuous tube and 
lobes ca. 4 mm long. 

The seven flowering collections examined are all 
of the thrum morph, which suggests strongly that 
the species is thrum-monomorphic. 


Annals of the 
Missouri Botanical Garden 


35. Psychotria nervosa Swartz, Prodr., 43. 
1788. TYPE: Jamaica. (fl), Swartz s.n. (holo- 
type, S, n.v., photo, A). Cf. also Swartz, Fl. 

Ind. Occid., 403. 1797. Figure 27. 


Psychotria undata Jacquin, Pl. Hort. Schoenbr. 3: 5, pl. 
260. 17 Jragoga undata (Jacq.) Baill., Hist. 
Pl. 7: 371. 1880. Uragoga undata (Jacq.) Kuntze, 
Revis. Gen. Pl. 2: 958. 1891. Myrstiphyllum un- 
datum ssp Hitchc., Annual Rep. Missouri Bot. 
Gard. 4: 9 A dae TYPE: Bahamas. July- Aug. (fl), 


Coll. ign. i 

Psychotria pii. ens Kunth in Humboldt, Bonpland & 
Kunth, Nov. Gen. Sp. 3: 364. 1819. Uragoga ru- 
i (Kunth) Kuntze, Revis. Gen. P 
1891. Psychotria nervosa subsp. rufescens (Kunth) 
Steyerm., Mem. New York Bot. Gard. 23: 480 
1972. Non Psychotria rufescens Roem. & Schultes 


ich is a synonym of Psychotria micrantha 
5 YPE of - 


sp. rufescens var. 
hirta (Roem. & Schultes) cibis Mem. New York 

Bot. Gard. 23: 481. 1972. TYPE: Colombia. Quindiu, 
d & dL ud s. : oo B— Willde- 

w 4092, photo 

Poi oligotricha A. P de Candalle, Prodr. 4: 514. 
/ Kuntze, Revis. 

: : l. TYPE: Insular Caribbean. 
inn (fl), Balbis s.n. (holotype, C, n.v., photo, F 

). 


33534 
taa chimarrhoides A. P. de Candolle, Prodr. 
dd 1830. TYPE: Cuba. 1829 Au Sagra 289 he. 
type, G, n.v., photo, F neg. 33535). 
Pira portoricensis A. P. de Candolle, Prodr. 4: 
515. 1830. TYPE: Puerto Rico. 1820 (fl), Bertero 
holotype, G, n.v., photo, F neg. 33537). 
Psychotria elongata Ben tham in Oersted, Vidensk. Med- 
del. Dansk Naturhist. le Kjebenhavn 1852: 32. 
1853. TYPE: Nicaragua. Granada: Volcán Momba- 
cho, Tine 1846 mund fr), Oersted 11594 (holotype, 
, n.v., photo, F neg. 22829, photo & fragment, 


Psychotria quiinifolia a ~ dg ih Bot. Gard. 
53: 108. 1966. Pan Bocas del Toro: 
Almirante, Mei Canal. wr Blum 1402 (ho- 
lotype, MO). 


Shrub 0.5-2(-3) m tall; young stems ferrugin- 
eous-tomentose to sometimes glabrous, the bark 
smooth; stipules sheathing, rounded-ovate, some- 
times short-acuminate, 6-11 x 3-6 mm, ferrugin- 
eous-pubescent to glabrous, caducous, leaving a 
pale ridge with red-brown fringe. Leaves petiolate; 
petioles 0.3-1.2(-2.5) cm long, ferrugineous-pu- 
bescent to glabrous, grooved above; blades m 
branous, oblanceolate to rarely elliptic, the apex 

)ll- 


mem- 


long-acuminate, the base attenuate, (6- 

17(-18) x (2-)4-6.5(-7.5) cm, sparsely tomen- 
tose to sometimes glabrous above and below, usu- 
ally denser below, the midvein usually ferrugineous- 
pubescent, the margin often red-brown ciliate, 


drying pale red-brown to green-brown, the veins 
usually drying reddish; secondary veins (10-)12- 
1 7 pairs, diverging 55°-65°, eucamptodromous to 
brochidodromous, increasingly arcuate toward 
margin, elevated below, sparsely ferrugineous-pu- 
bescent, the axils lacking domatia or hairs or rarely 
with fascicles of several red-brown hairs; tertiary 
veins inconspicuous to evident, orthogonal reticu- 
late to percurrent. /nflorescences terminal or pseu- 
doaxillary, panicles of cymes, the cymes often con- 
tracted; panicle branched to 3 degrees; main axis 
1-4(-5) cm long, the peduncle lacking; secondary 
axes in 3(4) ranks, the first-rank axes 2(4), equal, 
0.8-2.5(-4) cm long, the second-rank axes (1)2(3), 
0.6-1.5 cm long, the third-rank axes 2, 
0.2-0.7 cm long, the fourth-rank axes 2, 
equal, 0.2 cm long; cymes branched to 2-3 de- 
grees; bracts and bracteoles triangular, inconspic- 
uous, to O long, ferrugineous-pubescent. 
Flowers sessile to subpedicellate, the pedicels to 
0.5 mm long; calyx cup-shaped, the tube 0.3-0.5 
mm long, the lobes 5, triangular to not evident, to 
0.2 mm long, puberulent; corolla white, the tube 
cylindrical, 2-3 x 1 mm, white pubescent in throat, 
often puberulent without, the lobes 5, lanceolate, 

5 mm; stamens 5, the filaments 1.5-2.5 


equal, 
equal, 


mm long in pins, 3.5-5 mm long in thrums, the 
anthers 0.7-1 mm long; style 4-5 mm long in 
pins, 2-2.5 mm long in thrums, the branches short, 
clublike. Fruit narrow ellipsoid, 5.5-7 mm long, 
2.5-3 mm diam., maturing red, drying red-brown, 
usually red-brown puberulent; persistent calyx in- 
conspicuous or a minute beak; seed dorsal surface 
with 4-5 longitudinal furrows, the ventral surface 
with 2 sometimes incompletely divided longitudinal 
furrows. 


Distribution (Fig. 27). Ubiquitous, from 
southern Mexico and the Yucatán peninsula through 
Panama, most common on the Caribbean coast, at 
0- 1,200 m elevation in tropical moist to premon- 
tane moist to sometimes tropical dry forest with 
equatorial to tropical (usually equatorial-tropical) 
climate. It occurs also in Florida, the Bahamas, 
throughout the Greater and Lesser Antilles, Colom- 
bia, and Venezuela. It has been collected in flower 
throughout the year, primarily February-June in 
Mexico-Costa Rica and throughout the year in 
Panama, and in fruit throughout the year. 


Selected specimens examined. MEXICO. CAMPECHE: 
28 km al N de Escárcega, rumbo a Champotón, 7 Mar. 
1982 (fl), Cabrera et al. 2037 (MEXU); Campo Experi- 
mental Forestal Tropica E roy km 5 carr. 
à i 1966 (st), Chavelas et 


dell 1373 (F, US). CHIAPAS: a lo largo de las cascadas 


Volume 76, Number 2 
1989 


Hamilton 411 


Mesoamerican Psychotria, Part Il 


de Agua Azul, a 60 km al N de Palenque, 3 Mar. 1982 
(fr), Cabrera et al. 1875 (ENCB, MEXU); along Hwy. 
199 at Chital, 9 mi. SW of Ocosingo, 1,190 m, 7 July 
1977 (fl), Croat 40371 (MO); 17 km SE of pgs on 
road to Bonampak, ejido León Brindis, 300 m, 12 May 
1982 (fl), Davidse et al. 20385 (MO); Mpio. de denied 
Centro Arqueológico Bonampak, 350 m, 24 Dec. 1981 
(fr), Meave et al. B- 128 (MEXU). QUINTANA ROO: 1 km 
W of Puerto Morelos, Campo Agropecuario del Centro 
de Investigaciones de Quintana Roo, 5 m, 4 May 1982 
(fl), Davidse et al. 20033 (MO); 5 km al W de Vigia 
Chico, 13 June 1983 (fl), Durán & Olmsted 126 (MEXU); 
Isla de Cozumel, 24 May 1977 (fr), L. A. Pérez J. & 
Ramos 1620 (ENCB, MEXU); 1 km al N de Playa del 
Carmen, 9 Mar. 1980 (fl), Tellez & Cabrera 1807 
(MEXU). TABASCO: Balancán, San Isidro, 7-11 June 1939 
(fl), Matuda 3366 (F, GH, K, MEXU, MO, NY). vera- 
CRUZ: Mpio. Minatitlán, 6.9 km al N de la terraceria Ti 
Laguna-Rio Grande, una depresión kárstica, 120 m, 
July 1980 (fl), Wendt et al. 2534 (ENCB, on 
YUCATÁN: Chichen Itza, 28 June 1929 (fl), Bequaert 101 
(F, GH); Izamal, (fl), Gaumer 974 (A, F sheets, GH, 
US). GUATEMALA. ALTA VERAPAZ: Semococh, 17 km from 
Sebol, on Coban Road, 17 May 1964 (fr), ie 
4708a (F); SW of Languin, vic. caves, 600- a 
2] Feb. 1942 (st), Steyermark 44127 (F, oH. 1.5-2 
mi. S of Cubilgüitz, 300-350 m, 1 Mar. 1942 (fr), Stey- 
ermark 44433 (F). ESCUINTLA: 8 km S of Palin, 8 July 
1970 (early fr), Harmon & Dwyer 2960 (MO); between 
Rio Jute and Rio Pantaleón, on rd. between Esquintla and 


Santa Lucia Cotz, 540-720 m, 24 Jan. 1939 (fr), vA 


dell 16253 (F, LL, NY); Dolores, e Arroyo ‘kcal, 
14 Apr. 1961 (fl), Contreras 2070 (DUKE, LL, NY); 
La Cumbre, km 139-140 of Cadenas Rd., in swamp, 20 


Sep. 1966 (fr), Contreras 6159 (DUKE); nr. archaeo- 


logical camp on N shore of Lake Yaxja, 150 m, 18 June 
1973 (t), Croat 24668 (GH, MO, NY); bordeando ) Lago 


para Poptún, a km 116, 6 Dec. 1970 (fr), 

(DUKE, F, MO); Santa Elena, en la ori 

para San Francisco, a km 14, 2 

Ortíz 1859 (ENCB, F, MO, NY); sitio arqueológico en 
uventud," km ‘5 


el camino que conduce a cienaga “La 


Y, US). SACATEPEQUEZ: nr. Ls Lajas, 
. 28 Nov. 1938 (fl), y Standley 58081 (F). 
BELIZE. BELIZE: along Northern Hwy., vic. Maskall River, 
6 June 1973 (early fr), Croat 23930 (MO); Sibun o 
1.5 km in from mi. 34.5 on Western Highway, 5 Jul 
1973 (early fr), Dwyer 11491 (DUKE, MO), Big Fall, 
21 Mar. 1933 (fl), Lundell 4159 (F, MEXU); forest nr. 
Manatee Lagoon, 25 Mar. 1906 (fl), Peck 385 (GH, K). 
CAYO: El Cayo, river bluffs, 13 Feb. 1931 (fl), Bartlett 
11435 (F, K, US); bordering Rio Mopan, Sucotz, 14 Oct 
1967 (fr), Contreras 7128 (MEXU, NY). COROZAL: Co- 
rozal- Consejo road, Aug. 1933 (fr), Gentle 4862 (A, GH, 
NY). ORANGE WALK: Indian Church, Sawmill Rd., 
12 May 1977 (A), Arnason & Lambert 17293 (MO); 
nr. Honey Camp, 1930 (fr), Meyer 98 (F). STANN CREEK: 
Sittee River, secondary forest, 5 m, 11 Sep. 1930 (fr), 


Schipp 644 (A, F, GH, K, MO, NY). TOLEDO: Columbia 
Forest Station, 24 June 1972 (fr), D 4 


Nicolas Cay, 5 July 1972 (early fr), Spellman i Sap el 
2341 (MO). HONDURAS. ATLANTIDA 

11 Mar. 1928 (fl), Standley 56733 (A. F, Tw r. Te » 
sandy soil nr. beach, 20 July 1934 (fl, early fr), de 
4664 (A, F). COLON: N bank of Rio Guaimoreto between 
old bridge and opening of Laguna Guaimoreto, 7.5 km 
NE of Trujillo, 1595 7'N, 85%54'W, 20 Feb. 1981 (fl), 
Saunders 1039 Le COMAYAGUA: aon 5 km NE 


Libertad, 400 m, 21 May 1956 (fl), 4. Molina R. 7068 
(F, GH, US); villa de Taulabé, qda. 7 Caliche, 600 m, 
21 Apr. 1978 (fl), Trochez 41 (MEXU, MO). DISTRITO 
CENTRAL: Tegucigalpa, Mont. de la Flor, 950 m, Dec. 
1937 (st), Hagen & Hagen 1194 (F, NY). EL PARAISO: 
matorrales de la Qda. El Coyol entre El Junquillo y El 
Robledal, 1,300 m, 12 June 1964 (fl), 4. Molina R. 
14185 (F, NY). FRANCISCO MORAZÁN: 5 km O de 
pinares, orillas de riachuelo Chimbo, 900-1,00 
30 May 19 B (A C. Nelson & Vargas 3456 (TEFH). 
GRACIAS A DIO ampusilpi, nr. Río Patuca, 30 Aug.- 
2 Sep. 1973 ti, “Cletdell bur dup TEFH); 

coastal thicket, O m, 9 Feb. 


OLANCHO: vaguada del rio de la población de € 500 
m, 17-22 July 1978 (fr), C. Nelson & Romero 4640 
(MO). vono: Santa Rita, orillas del Rio Hn 23 A pr. 
1971 (A), Mancías & J. Hernández 1138 (MO, TEFH); 
Aguan River valley, vic. Coyoles, 300 m, June-Aug. 
dae (early fr), Yuncker et al. 8173 (F, GH, K, MO, 
, US). NICARAGUA. BOACO: NE de Mombachito, 11 
n 1982 (fl, Sandino 2837 (MO). cHONTALES: 2 km 
al N de Santo Tomas, carret. a Santo Domingo, 12°04'N, 
85°06'W, 340-380 m, 30 Apr. 1982 (fl), P. P. Moreno 
16224 (MO); E of El Narajal, between Juigalpa and La 
Libertad, ca. 12?11'N, 85?15'W, 500 m, 6 Aug. 1983 
(fl, early fr), W. D. Stevens 22435 (MO). JINOTEGA: Salto 
Kayaska, Rio Bocay, 13%51'N, 85?22'W, 190-340 m, 
7 Mar. 1980 (fl, W. D. Stevens et al. 16569 (MO). 
MATAGALPA: entre las fincas La Castilla y Sta. Basilia, 
bordeando las márgenes del Rio Yasica, 13°01'N, 
85?46'W, 400 m, 19 Jan. 1982 (fr), D. Castro 2264 
(MO); Mt. Yasica, 26 km S of Matagalpa, rd. to El Tuma, 
400 m, 10 Mar. 1967 (fl), 4. Molina R. 20466 (F, NY, 
US). p SAN JUAN: San Miguelito, drenaje del Lago Gra- 
nada, 3 , 13 Nov. 1951 (fr), Shank & Molina 4564 


900 m, 28 Nov. 1982 (8), P. P. Moreno 18846 (MO); 
Isla de Ometepe, N de Volcán Concepción, entrada a Los 
Angeles, 11?33'N, 85°37'W, 4 


Sandino 4 


oe 12008 (MO); Cerro Kana Cop oo 
of Siuna, 250 m, 17 June 1978 (fl), Neill. 4520 (MO) 


(MO); El Falso Bluff a E de El Bluff, LUSON, 83°41'W, 
30 m, 2 Feb. 1982 (fl), Sandino 2180 (MO); costa N 
de Puerto Cabezas, 14?02'N, 83?22'W, 12 m, 5 Feb. 


T 


Annals of the 
Missouri Botanical Garden 


1983 (early fr), Sandino 3959 (MO); vic. Wani, including 
Rio Uli, 13?42'N, 84*51'W, 90-110 m, 22 Apr. 1978 
(A), W. D. Stevens 7978 (MO). Costa RICA. CARTAGO: 
Rio Reventazón, entre Florencia y Turrialba, 500-600 
m, 10 July 1965 (fl), 4. Jiménez M. 3269 (CR — 2 sheets, 
— 2 sheets). GUANACASTE: between Liberia and Bagaces, 
nr. Rio Potrero along Interamerican Hwy., 10?33'N, 
85°23'W, 80 m, 29 Dec. 1966 (fl), Burger & Ramírez 
4124 (CR, DUKE, F, MO); Sta. Rosa National Park, 
10%51'N, 85°37'W, 200-300 m, 22 Jan. 1978 (fl, fr), 
Liesner 4342 (CR, MO); 23 June 1977 (fl, fr), Liesner 
& Lockwood 2409 (CR, MO); 13-16 km N of Liberia 
along Camino Sta. María, 600-700 m, 9 July 1976 (fl), 
J. Utley & K. Utley 5394 (CR, DUKE); ca. 5 km 
La Cruz, 260 m, 10 June 1967 (f), Weston 5017 (CR, 
DUKE, MO, NY). LIMÓN: between Hone Creek and Ca- 
huita, 5 May 1983 A ian fr), L. D. Gómez et al. ere 
(MO—2 sheets). PANAMA. BOCAS DEL TORO: vic. Alm 
rante, Jan.- Mar. 1928 (fl, fr), Cooper 557 (F, NY, US) 
Bar Mouth, Changuinola Valley, 9 Mar. 1924 (fl, fr), 
Dunlap 538 (US). CANAL AREA: nr. beach at Fort Kobbe, 
12 Oct. 1961 (fr), Duke 4728 (GH, MO); Madden Dam, 
Boy Scout Rd., 23 July 1968 (fl, fr), Dwyer & Lallathin 
8813 (F, GH, MO, NY); forest nr. Curundu, 8?58'N, 
79°33'W, O m, 10 Mar. 1983 (early fr), Hamilton & 
Gentry 3272 (MO); Fort Sherman and nearby, 9?20'N, 
80°00'W, 0-100 m, 16 June 1983 (fl), Hamilton & 
Stockwell 3696 (MO); 6 km E of Gamboa, 160-190 m, 
24 Dec. 1973 (fl), Nee 8993 (F, MO). cocLÉ: Penonome, 
14 July 1962 (fl), Dwyer 2007 (MO); W of Rio Guias, 
10 Sep. 1972 (fr), Gentry pur (F, MO); El Espino de 
Antón- Rio Arenales, 7 Oct. 1973 (fr), E. Jaén 17 (DUKE); 


(MO); ‘coastline between Garrote and La 


betwee n Panama an a 
Dodge et i 7 (GH, MO. NY) 
zales, Perlas, 2 pu 1961 (fl), Duyer 1 
Goofy Lake, 26 6 No v. 1966 (fl, fr), Dure 7128 (CH 
MO, US); San José Island, Pearl dia 15 July 
1945 (fl), Erlanson 475 (GH, NY, US); 12 Oct. 1944 


e 
IE 


78?15'W, ca. 100 m, 12 Sep. 1981 (early fr), Knapp 
1185 (ENCB, MO); Rio Tocumen, 3 Jan. 1924 (fl), 
Standley 29433 (US). SAN BLAS: mainland in front of 
Ustupo, 9 Nov. 1975 (fl, fr), vd 9524 (MO, NY); 
Soskatupu, island 1.5 x 0.5-0.7 mi., 0-45 m, 15 Aug. 
1967 (fl, early fr), Elias ka (CH. 'MEXU, MO, hs 
nr. Puerto Obaldia, 8?40'N, 77°25'W, O m, 16 Apr 
1982 (fl, fr), Knapp & Maltei 4649 (MO). 


All the species in synonymy have been properly 
synonymized before, with the exception of P. 
quiinifolia, the morphology of which does not differ 
from that of P. nervosa. Degree of pubescence 
varies within populations, and infraspecific taxon- 


omy based on this character (e.g., subsp. rufescens) 
is not recognized. 

Psychotria nervosa may be recognized by its 
usually reddish pubescent aspect, leaves with usu- 
ally eucamptodromous secondary veins diverging 
55?-65^, small (1-4(-5) cm long) inflorescences 
lacking peduncles and with secondary axes usually 
2 per rank in usually 3 ranks, and narrow-ellip- 
soidal fruit (5.5-7 x 2.5-3 mm). There appears 
to be no geographic component to the variation 
between glabrous and densely tomentose aspect. 
Collections from Florida are noteworthy for small 
eaves and relatively large inflorescences. 


GROUP 6. THE PANAMENSIS GROUP 


Shrub or tree; young stems glabrous or (often 
red-brown) puberulent; stipules usually sheathing, 
lanceolate (Fig. 2d) or ovate, or less commonly 
linear or triangular with the apex sometimes bilobed 
or biacuminate, uniform in color, glabrous or fringed 
or puberulent, caducous. Leaf blades usually ellip- 
tic, or sometimes obovate, ovate, or lanceolate, 
drying green-brown to red-brown to brown, some 
times paler below; secondary veins (4—)6- 19(- 22) 
pairs, diverging (30?-)45?—75*(-809), eucampto- 
dromous to brochidodromous, the axils sometimes 
with domatia (sometimes P. cocosensis and P. lun- 
dellii) or with tufts of hair (sometimes P. chiri- 
quina, P. lundellii, and P. mexiae) or usually 
lacking; tertiary veins orthogonal reticulate to per- 
current. /nflorescences panicles of cymes (or of 
glomerules sometimes in P. panamensis or of in- 
dividual flowers in P. hornitensis), nonpeduncu- 
late (Fig. 7e) except in P. olgae, P. stockwellii, 
and sometimes in P. cascajalensis, P. chiriquina, 
P. dwyeri, P. lundellii, P. panamensis, P. sara- 
piquensis, and P. trichotoma; secondary axes 
usually in 1 pair per rank or less commonly 2 
equal pairs or 2 size-differentiated pairs per rank, 
diverging acutely except in P. stockwellii; bracts 
not conspicuously enlarged, to 3 mm long only in 
P. chiriquina. Corolla tubes (1.5-)2-5 mm long, 
the lobes without apical extensions. Fruit ellipsoid, 
obovoid, or spherical when dry; persistent calyx 
a beak or less commonly a conspicuous tube (P. 
cocosensis [Fig. 9c] and P. olgae and sometimes 
P. panamensis) or often not evident; seed dorsal 
surface with (3—) 4—10 often irregular longitu- 
dinal furrows, the ventral surface with 2 deep plus 
often several irregular longitudinal furrows (Figs. 


10g, 10h) 


Half of the 12 species of the group are assigned 
to the Psychotria panamensis complex; P. pana- 


Volume 76, Number 2 
1989 


Hamilton 413 
Mesoamerican Psychotria, Part II 


300 km 


105 100 95 


‘= L 


e A 
ee A 
« 9. 


90 ^ 85 80 


L i L 


E 28. Distributions of Psychotria cocosensis (open triangle), P. panamensis var. 


FIGUR 
Mes P. panamensis var. ixtlanensis (open cir 
r. panamensis (solid circles) in Mesoamerica. 


mensis is widespread (Fig. 28), the other five much 
less so: P. cascajalensis (Coclé, Panama; Fig. 29), 
P. cocosensis (Cocos Island, Costa Rica; Fig. 28), 
P. dwyeri (southern Mexico; Fig. 29), P. olgae 
(eastern Panamá Province; Fig. 29), and P. stock- 
wellii (Costa Rica and Panama; Fig. 30). Three 
other widespread species have character states that 
distinguish them from typical members of the group 
and are of uncertain affinities, although analysis 
might reveal they belong in the panamensis com- 
plex: P. trichotoma has obovoid fruits and con- 
spicuously brochidodromous venation; P. sarapi- 
quensis has usually narrowly oblanceolate leaves; 
and P. mexiae has 5-8 deep irregular furrows on 
the seed dorsal surface. The most cohesive complex 
is that of P. chiriquina, which includes that species 
(Nicaragua through Panama; Fig. 29), P. lundellii 
(Chiapas through El Salvador; Fig. 29), and P. 
hornitensis (western Panama; Fig. 30); they share 
ovate stipules and shallow furrows on the see 
ventral surface. 

Of the 15 taxa in the group, eight appear di- 
stylous; of those, Psychotria dwyeri, P. mexiae, 

olgae, P. stockwellii, and P. trichotoma show 
some between-morph asymmetry in floral part 
lengths. Psychotria lundellii appears to be pin- 
monomorphic in Belize, but recently I examined a 
thrum collection from Guatemala. Psychotria 


compressicaulis (solid 


rele), P. panamensis var. magna (solid squares), and P. panamensis 


panamensis var. compressicaulis, on the other 
hand, is thrum-monomor 
sensis and P. 
Psychotria cascajalensis, 
ixtlanensis, and P. panamensis var. magna do 


hic. Psychotria coco- 
hornitensis are long-homostylous. 
P. panamensis var. 


not have sufficient flowering material for analysis. 


36. Psychotria cascajalensis C. Hamilton, 
Phytologia 64: 220. 1988. TYPE: Panama. 
Coclé: N of Penonomé, between Llano Grande 
and Cascajal, rd. to Coclesito, 480 m, 2 May 
1979 (fi), Hammel 7227 (holotype, MO- 
2901097; isotypes, ENCB, MO). Figure 29. 


Tree 10 m tall; young stems glabrous to sparsely 
puberulent, the bark smooth; stipules oval, round- 
ed, 6-7 x 3-3.5 mm, fringed, glabrous, caducous, 
leaving a pale ridge with red-brown fringe. Leaves 
petiolate; petioles 2-4 mm long, glabrous, flat above; 
blades coriaceous, elliptic, the apex acute, the base 
attenuate, 3.5-6(-7) x (1.5-)2-2.7 cm, glabrous 
above and below, drying dull red-brown to green- 
brown; secondary veins 4-6 pairs, diverging 55°- 
65°, brochidodromous with secondary loops incon- 
spicuous, straight then arcuate near margin, not 
elevated below, glabrous, the axils lacking domatia 
or hairs; tertiary veins inconspicuous. /nflores- 
cences terminal, panicles of cymes or glomerules; 


414 


Annals of the 
Missouri Botanical Garden 


300 km 


105 100 95 


T.C.4 
n 
^ Y 204 
e? ° 
15 4 


à 
90 85 80 


FIGURE 29. 


Distributions of Psychotria cascajalensis (open triangle), P. chiriquina (solid triangles), P. dwyeri 
e). 


(open circles), P. lundellii (solid circles), and P. olgae (solid square 


panicle branched to 3 degrees; main axis 6 cm 
long, the peduncle 2-3.5 cm long; secondary axes 
in 3 ranks, the first-rank axes 2 or 4, the longer 
pair 1-2 cm long, the shorter pair 0.4-1.2 cm 
long when present, the second-rank axes 2, 0.5- 
0.8 cm long, the third-rank axes 2, 0.1-0.3 cm 
long; cymes branched to 1-2 degrees; bracts and 
bracteoles broadly triangular, 1-2 mm long, red- 
brown pubescent. Flowers subsessile, the pedicels 
0.5 mm long; calyx cup-shaped, the tube 0.8 mm 
long, the lobes 5, triangular to barely evident, to 
O. 0.8 mm, puberulent, fringed; corolla green- 
white, the tube cylindrical, 1. mm, thickly 
white pubescent in throat, the lobes 5, triangular, 
1.5 x mm; stamens 5, the filaments not seen 
in pins, 1.5 mm long in thrums, the anthers 1 mm 
long; style not seen in pins, 1 mm long in thrums, 
the branches linear. Fruit not seen. 


Distribution (Fig. 29). Known only from the 
type collection from the foothills south of the divide 
in Coclé, Panama, at 480 m in a region of tropical 
moist to premontane wet forest with equatorial- 
mountainous climate. It was collected in flower on 


Psychotria cascajalensis may be recognized by 
its small (usually 3.5-6 x 2-2.7 cm), coriaceous, 
elliptic leaves and robust pedunculate inflores- 
cences with secondary axes 2(4) per rank in three 
ranks. 


The only flowering collection appears to be a 
thrum morph. 


37. Psychotria chiriquina Standley, Contr. 
U.S. Natl. Herb. 18: 129. 1916. TYPE: Pan- 
ama. Chiriqui: Alto de la Cuesta, E slope of 
Chiriqui Volcano, 2,100-2,200 m, 10-13 
Mar. 1911 (fl), Pittier 3118 (holotype, US). 
Figure 29. 


Shrub or small tree, 1-6 m tall; young stems 
glabrous, the bark gray, deeply fissured longitu- 
dinally; stipules sheathing, ovate, sometimes bi- 
lobed, 8-10 x 3-5 mm, glabrous to red-brown 
puberulent, fringed, caducous, leaving a pale ridge 
with red-brown fringe. Leaves petiolate; petioles 
0.6-1.5 cm long, glabrous to red-brown puberu- 
lent, flat above; blades membranous, elliptic, the 
apex long-acuminate, the base attenuate, (4—)6.5— 
9(-12) x (1-)1.5-2.5(-3) cm, glabrous above and 
below, drying deep red-brown; secondary veins 5- 
11(-12) pairs, diverging 30?-65?, eucamptodro- 
mous, constantly arcuate, prominulous to not el- 
evated below, glabrous, the axils with or without 
minute tufts of hair; tertiary veins evident to in- 
conspicuous, orthogonal reticulate to percurrent. 
Inflorescences terminal, panicles of cymes; panicle 
branched to 2-4 degrees; main axis (1-)2- 7 cm 
long, the peduncle lacking or 1-4.5 cm long; 
secondary axes in 3(4) ranks, the first-rank axes 


Volume 76, Number 2 
1989 


Hamilton 415 
Mesoamerican Psychotria, Part Il 


2, (0.4-)1-3.2 cm long, the second-rank axes 2, 
(0.3-)0.5-1.4 cm long, the third-rank axes 2, 
(0.1-)0.3-0.5 cm long, the fourth-rank axes 2, 
0.2 cm long; cymes branched to 1-2 degrees; 
bracts linear, 2-3 mm long, glabrous to ciliate; 
bracteoles irregular, to 0.8 mm long, sparsely cil- 
iate. Flowers sessile to pedicellate, the pedicels to 
3 mm long; calyx cup-shaped, the tube 1 mm long, 
the lobes 5, triangular to barely evident, 0-0.6 x 
0.8 mm, glabrous to minute-puberulent; corolla 
white, the tube cylindrical, 4-6 x -2 mm, 
white pubescent in throat, the lobes 5, linear, 1.5- 

X 1 mm; stamens 5, the filaments 4-4.5 mm 
long in pins, 5.5-6 mm long in thrums, the anthers 
1 mm long; style 6 mm long in pins, 4-5 mm long 
in thrums, the branches minute, clublike. Fruit 
when dry ellipsoid, 5-6 mm long, 4-4.5 mm diam., 
maturing red, drying deep red-brown; persistent 
calyx a minute beak to 1 mm long; seed dorsal 
surface with 4 shallow longitudinal furrows, the 
ventral surface with 2 shallow longitudinal furrows. 


Distribution (Fig. 29). 
Nicaragua and from both sides of the Costa Rica- 
Panama border at elevations of 900 to 2,500 m 
in premontane to lower montane moist forest to 


Known from northern 


rainforest with equatorial-mountainous to tropical- 
mountainous climate. Psychotria chiriquina has 
been collected in flower July-August in Nicaragua 
and primarily February-May in Costa Rica and 
Panama, and in fruit November-May in Nicaragua 
and August- April in Costa Rica and Panama. 


Selected specimens examined. NICARAGUA. JINOTE- 


*Filas el Portal," 13?37'N, 
600-900 m, 26 Mar. 1981 (fr), P. P. Morena & Sandino 
7583 (MO). MADRIZ: Cerro Pataste, 1,600 m, 8 May 
1975 (fr), Neill 129 (MO). MATAGALPA: Cerro El Picacho, 
N del hotel Selva Negra, 13?00'N, 85?55'W, 1,500 m, 
7 n és ne "i P. d 21678 (MO); Hda. Santa 

ria de O of Matagalpa, 1,500 m, 12 
[s 1977 i, “Neill 2304 e Macizos de Penas Blan- 
El Quebradon, vo N 
a. San ag 1905 N, 85°38'W, 


1,4 > 

21145b yea Sta. María de Ostuma, Cordillera Central 
between Matagalpa and Jinotega, 1,300-1,500 m, 8-15 
Jan. 1963 (fr), L. O. Williams et al. 23342 (F, GH, NY, 
US). Rivas: Isla Ometepe, faldas del Volcán Maderas al 
S de la Hda. Magdalena, 11?27'N, 85?*30'W, 300-900 
m, 28 Nov. 1982 (fr), P. P. Moreno 18854 (MO). ZELAYA: 
Cerro La Pimienta, N Pe facing La Garrapata, 13?45'N, 
84°59'W, 900-1,180 16 Mar. 1980 (fr), Pipoly 
6050 (MO). Costa en PUNTARENAS: valley of Rio Cotón 
between Sitio Cotón (Cotonsito) and Sitio Tablas, 895 7'N, 
82?46'W, 1,500-1,600 m, 2 Sep. 1983 (early fr), Da- 


vidse 24415 (CR, MO); trail between Tres Colinas and 
Cerro Bekom, 9°08'N, 83%04'"W, 2,300 m, 28 Mar. 1984 


1,200 m, 29 Mar. 1984 (fl), Davidse & Herrera 26219 
(MO); Cerro Buri, 9%00'N, 82°49'W, 2,000-2,300 m, 
20 Aug. 1983 (fr), Davidse et al. 23825 (MO); upper 
Rio Burú, 2,010 m, 19 Aug. 1983 (fr), Gómez et al. 
21509 (MO). PANAMA. BOCAS DEL TORO: Valle de Silencio, 
9*05'N, 82%56'W, 8-10 Aug. 1979 (fr), gra 1586 
(MO). COE teg of Las Cumbres n o Punta, 
20 Feb. 1971 (fl, fr), Croat 13737 (F, MO “o sheets, 
NY); E slope a Mola Barú, 10 mi. from Boquete, 2,500 
m, 18 May 1976 (fl), Croat 34982 (MO, NY); Caldera- 
Chiriquicito trail, 21 Apr. 1968 (fr), Kirkbride & Duke 
1007 ); Cerro Hornito, above Los Planes de Hornito, 
9*42'N, 82°06’W, 2,100 m, 14 Mar. 1982 (fl), Knapp 
et al. 4196 (MO); slopes Aix Cerro Horqueta, 
,200-1,800 m, 5 Jan. 1975 (fl), Wilbur & Luteyn 
19316 (DUKE); E of eri along Rio Chiriqui Viejo 
ca. 2 mi. NE of Cerro Punta, Cerro Respinga ridge, 13 
Jan. 1971 (fl, fr), Wilbur et al. 13106 (DUKE, MO). 


= 


Psychotria chiriquina may be recognized by its 
small (usually 6.5-9 x 1.5-2.5 cm) narrow-elliptic 
leaves drying deep red-brown, deeply longitudinally 
fissured bark, long (4-6 mm) corolla tube, and 
robust (5-6 x 4-4.5 mm) ellipsoid fruits drying 
deep red-brown. There are several differences not- 
ed in collections from Nicaragua as opposed to 
those from Panama. Nicaraguan material shows 
fewer (5-7 vs. usually 8-1 1[-12]) secondary veins 
and more acute (30°-50° versus 55?-65?) angles 
of divergence of secondaries. Panamanian material 
shows often pedicellate inflorescences and fruits 
with a more prominent persistent calyx. 


38. Psychotria cocosensis C. Hamilton, Phy- 
tologia 64: 222. 1988. TYPE: Costa Rica: Co- 
cos Island, 18 Tue 1898 (fr), Pittier 12375 
(holotype, US). Figures 9c, 28. 


Shrub; young stems glabrous, the bark furrowed 
longitudinally; stipules lanceolate, 12-35 x 2-5 
mm, glabrous, caducous, leaving a pale ridge with 
red-brown fringe. Leaves petiolate; petioles 7-14 
mm long, glabrous, grooved or flat above; blades 
subcoriaceous to coriaceous, elliptic, the apex acu- 
minate to subcaudate, the base attenuate, (12-) 
14-17 x 5.5-7 cm, glabrous above and below, 
drying red-brown to green-brown; secondary veins 
(9-)10-12 pairs, diverging 45?-50?, eucampto- 
dromous to brochidodromous, straight then arcuate 
near margin, elevate 
leaves, the axils with small domatia below; tertiary 


w in less coriaceous 


veins inconspicuous to evident, percurrent to re- 
ticulate. Inflorescences terminal, panicles of cymes; 
panicle branched to 2-3 degrees; main axis 1 cm 
long, the peduncle lacking; secondary axes in 2 
ranks, the first-rank axes 2, 0.6 cm long, the 
second-rank axes 2, reduced; cymes branched to 


416 


Annals of the 
Missouri Botanical Garden 


l degree; bracts triangular, 0.7-1.5 mm long, 
glabrous. Flowers pedicellate, the pedicels 2-3 mm 
long; calyx cup-shaped, the tube 1 mm long, the 
lobes 5, broadly triangular to barely evident, gla- 
brous; corolla color unknown, the tube cylindrical, 

mm, white pubescent in throat, the lobes 
5, linear, 4 X 2 mm; stamens 5, the filaments 7 
mm long, the anthers 2 mm long; style 9-10 mm 
long, the branches minute, linear. Fruit when dry 
spherical to ellipsoid, 6 mm long, 5.5-6 mm diam., 
drying dark red-brown; persistent calyx cuplike, 
coriaceous, 1.5 mm long (Fig. 9c); seed dorsal 
surface with 4 shallow, irregular, longitudinal fur- 
rows, the ventral surface with 2 deep, irregular, 
longitudinal furrows. 

Distribution (Fig. 28). 
type locality, Cocos Island, at 50 m elevation in 
premontane rainforest. Psychotria cocosensis has 
been collected in fruit in January and June. 


Known only from the 


Additional specimens examined. Costa RICA, COCOS 
ISLAND: La vallée de Chatham, 50 m, Jan. 1902 (fl, fr), 
Pittier 16279 (GH, US). 


Psychotria cocosensis may be recognized by its 
close resemblance to P. panamensis and by its 
coriaceous leaves and large (6 x 5.5-6 mm) spher- 
ical to ellipsoid fruits with calyx persisting as a 
coriaceous cup ca. 1.5 mm long. 

Style and stamens are both exserted in the one 
flowering collection seen, suggesting strongly that 
the species is long-homostylous. 


39. Psychotria dwyeri C. Hamilton, Phytolo- 
gia 64: 223. 1988. TYPE: Mexico. Oaxaca: 
Dto. Choapan (Santiago Choapan), Sta. María, 
Montana Sta. Maria, 1,500 m, 7 Apr. 1938 
(fr), Mexia 9265 (holotype, NY; isotypes, B, 
F, GH, K, MO, US). Figure 29. 


Shrub or small tree, 3-6 m tall; young stems 
glabrous, the bark smooth; stipules sheathing, tri- 
angular, 8-13 x 3-5 mm, glabrous, caducous, 
leaving a pale ridge often with red-brown fringe. 
Leaves petiolate; petioles (S—)15—40(—45) mm long, 
glabrous, furrowed above; blades membranous, el- 
liptic, the apex cuspidate, the base cuneate to at- 
tenuate, (12 -)15-23 x 5-10 cm, glabrous above, 
glabrous to minutely white puberulent below, drying 
green-brown, paler below; secondary veins 
(12-)15-18 pairs, diverging 55?-70*(-80?), eu- 
camptodromous, straight to slightly arcuate, ele- 
vated below, often white puberulent below, the axils 
lacking domatia or hairs; tertiary veins evident, 
orthogonal reticulate, the loops near margin evi- 
dent. /nflorescences terminal, panicles of cymes; 


panicle branched to 4 degrees; main axis 5.5-10.5 
cm long, the peduncle lacking or to 2 cm long; 
secondary axes in 4-5(-6) ranks, the first-rank 
axes 2, (2.5-)7-9 cm long, the second-rank axes 
2(4), (2-)3-5.5 cm long, the third-rank axes 2, 
(0.5-)1.5-4 cm long, the fourth-rank axes 2(4), 
0.5-)1-2 cm long, the fifth-rank axes 2, 0.6-0.8 
cm long, the sixth-rank axes 2, 0.3 cm long; cymes 
branched 1-2 degrees; bracts inconspicuous. Flow- 
ers pedicellate, the pedicels 0.5-1.5 mm long; 
calyx cup-shaped, the tube 0.5 mm long, the lobes 
5, broadly triangular, minute, glabrous to minutely 
fringed; corolla white, the tube cylindrical, 2-2.5 
X 2 mm, white pubescent in throat, the lobes 5, 
mm; stamens 5, the fila- 


— 


triangular, 1.5-2 x 
ments 2-2.5 mm long in pins, 2.5-3 mm long in 
thrums, the anthers 1 mm long; style 4-4.5 mm 
long in pins, 1.5-2 mm long in thrums, the branch- 
es linear in pins, clublike in thrums. Fruit when 
dry spherical, 4.5-5 mm long, 4.5-5 mm diam., 
maturing red, drying deep red-brown; calyx per- 
sistent as a beak, to 0.8 mm long; seed dorsal 
surface with 4 irregular longitudinal furrows, the 
ventral surface with 2 deep longitudinal furrows. 


Distribution (Fig. 29). Known from Veracruz 
and Oaxaca, Mexico, at elevations of 50-1,500 
m in a region of subevergreen forest with tropical 
to tropical-mountainous climate. This species has 
been collected in flower in April and May and in 
fruit April, June, and December. 


ecimens examined. MEXICO. OAXACA: Ubero, 30- 


a al. ES-2832 (MEXU); El Mirador. 1853 (8). F. Mulier 
up B 1932 (fl), iud 14124 (A, F); 

apan, 5 (fl), Purpus 7525 (MO, US); May 1926 | (A), 

s s (US); Jalapa, 1, 200- 1,350 m, 1894 (fl), 

C. Smith 1844 (F, US). 

Psychotria dwyeri resembles P. panamensis in 
inflorescence and vegetative characters and is dis- 
tinguished most easily in fruit. Psychotria dwyeri 
has spherical (vs. ellipsoid to obovoid) fruit 4.5-5 
mm (vs. 4.5-8 mm) long. 

Reproductive organs at both levels appear to be 
longer in the pin morph than in the thrum. 


40. Psychotria hornitensis Dwyer & C. Ham- 
ilton in C. Hamilton, Phytologia 64: 225. 
1988. TYPE: Panama. Chiriqui: ridge and sum- 
mit of Cerro Hornito, above Los Planes de 
Hornito, 8%42'N, 82%06'W, 2,100 m, 14 Mar. 
1982 (fl, early fr), Knapp et al. 4198 (ho- 
lotype, MO). Figure 30. 


Shrub 2 m tall; young stems glabrous, the bark 
deeply furrowed longitudinally; stipules sheathing, 


Volume 76, Number 2 Hamilton 417 
1989 Mesoamerican Psychotria, Part II 
d ds A TC.4 
e =p 
1 204 
F E: 
© 
e "n. e he 4 
ds + + Ee m p 
e 
cs 
So 
300 km + "E "| 
D 
105 90 85 a ag 


100 95 


L L L 


SURE 30. 
in Mesoamerica. 


ovate to lanceolate, 3-4 x 1-1.5 mm, glabrous, 
caducous, leaving a pale ridge with red-brown fringe. 
Leaves subsessile; petioles to 2 mm long, glabrous, 
flat above; blades membranous, narrowly elliptic, 
the apex long acuminate, the base attenuate, 2.5- 
3.5 x 0.5-0.7 cm, glabrous above and below, 
drying dull green-brown above, dull green below; 
secondary veins 5-6 pairs, diverging ca. 50°, eu- 
camptodromous, constantly arcuate, not evident 
above, barely evident below, glabrous, the axils 
lacking domatia or hairs; tertiary veins inconspic- 
uous. Inflorescences terminal, panicles of 3-5 in- 
dividual flowers or cymes; panicle branched to 2 
degrees; main axis 2.5-3 cm long, the peduncle 
1.5-2 cm long; secondary axes in 1-2 ranks, the 
first-rank axes 1-2, 0.3-0.4 cm long, the second- 
rank axes 1-2, 0.1 cm long; cymes branched to 
l degree; bracts and bracteoles linear, 1.5-2 x 
0.7 mm, glabrous to puberulent within near base. 
Flowers sessile to pedicellate, the pedicels to 1 mm 
long; calyx cup-shaped, the tube ca. 1 mm long; 
the lobes 5, triangular, 0.8-1 x 0.8 mm, glabrous; 
corolla white, the tube cylindrical, 3.5-4 x 1.5 
mm, white pubescent in throat, the lobes 5, ovate, 
1-1.5 x 0.8 mm; stamens 5, 3-4 mm long, the 
anthers 1 mm long; style 4-6 mm long, the branch- 
es short, linear. Mature fruit not seen. 


Distribution (Fig. 30). 
type locality, Cerro Hornito, Chiriqui, Panama, at 


Known only from the 


Distributions of Psychotria hornitensis (square), P. mexiae (circles), and P. stockwellii (triangles) 


ca. 2,000 m elevation in low montane rainforest 
with equatorial-mountainous climate. Psychotria 
hornitensis has been collected in flower in Feb- 
ruary and March and with immature fruit in March. 


PANAMA. CHIRIQUÍ: 
979 


Additional specimen examined. 
ridge nr. top of Cerro Hornito, 1,950 m, 15 Feb. 1 
(fl), Hammel 6189 (MO). 


Psychotria hornitensis may be recognized 
readily by its small, narrow (2.5-3.5 x 0.5-0.7 
cm) leaves and small (2.5-3 cm long) few-flowered 
inflorescences. 

The one flower morph found does not show 
significant differentiation in length between the 
exserted style and stamens, so the species is pre- 
sumed to be long-homostylous. 


41. Psychotria lundellii Standley in Lundell, 
Contr. Univ. Michigan Herb. 4: 29. TYPE: 
Belize. Cayo: Valentin, 26 June 1936 (fl), 
Lundell 6260 (holotype, F; isotypes, GH, 
MICH, n.v., NY, US). Figure 29. 


Tree or shrub, (1-)2-15 m tall; young stems 
glabrous or red-brown puberulent, the bark pale, 
cracked and furrowed longitudinally; stipules ovate, 
rounded, 5-6 x 2-4 mm, glabrous, often ciliate, 
caducous, leaving a pale ridge with red-brown fringe. 
Leaves petiolate; petioles 0.8-2 cm long, glabrous, 
grooved above; blades subcoriaceous, elliptic, the 


Annals of the 
Missouri Botanical Garden 


apex acuminate, the base attenuate, (5-)7.5-15 
x (1.8-)2.5-5 cm, glabrous above and below, 
drying pale to glossy red-brown; secondary veins 
(5-)6-8(-12) pairs, diverging (30?-)60?-65?, bro- 
chidodromous, constantly arcuate, prominulous be- 
low, glabrous, the axils usually with minute tufts 
of red-brown hair and/or domatia below; tertiary 
veins not evident. /nflorescences terminal or pseu- 
doaxillary, panicles of cymes; panicle branched to 
(3)4 degrees; main axis 4-7.5 cm long, the pe- 
duncle (0-)1-3 cm long; secondary axes in 4(5) 
ranks, the first-rank axes 2(4), the longer pair 1.5- 
3.5 cm long, the shorter pair when present 1.6 
cm long, the second-rank axes 2, 0.8-1.7 cm long, 
the third-rank axes 2, 0.5-0.7 cm long, the fourth- 
rank axes 2, 0.2-0.3 cm long, the fifth-rank axes 
2, 0.2 cm long; cymes branched to 1-2 degrees; 
bracts sheathing, irregular, to 1.5 mm long, ciliate; 
bracteoles triangular, 0.3-0.5 mm long, ciliate. 
Flowers pedicellate, the pedicels 0.7-1 mm long; 
calyx cup-shaped, the tube 0.5 mm long, the lobes 
5, triangular, 0.3 X 0.6 mm, minutely ciliate; 
corolla green-white, the tube campanulate, 2-2.5 
X ].5 mm, white pubescent in throat, the lobes 
5, ovate-acuminate, 1.5-2 x ] mm; stamens 5, 
the filaments 1.5-2 mm long in pins, 4 mm long 
in thrums, the anthers 0.7 mm long; style 3-4 mm 
long in pins, 2 mm long in thrums, the branches 
linear. Fruit spherical when dry, (4-)4.5-5(-5.5) 
mm long, (4-)4.5-5(-5.5) mm diam., maturing 
red, drying red-brown; persistent calyx not con- 
spicuous or rarely to mm long; seed dorsal 
surface with 3-6 shallow irregular longitudinal fur- 
rows, the ventral surface with 2 deep and often 
2-4 additional irregular longitudinal furrows. 


Distribution (Fig. 29). Eastern Chiapas, Gua- 
temala, western Belize, and El Salvador, at ele- 
vations below 500 m except in El Salvador, where 
this species occurs at 1,800-2,000 m. It is found 
in evergreen forest with equatorial-tropical or 
sometimes tropical to tropical-mountainous cli- 
mate. Psychotria lundellii has been collected in 
flower April-July and in fruit July-April, primarily 
January- March. 

Selected specimens examined. MEXICO. CHIAPAS: 23 
mi. SE of Palenque, rd. to ee 160 m, y 1977 
(fl), Croat 40294 (MO); Mpio. Ocosingo, 3 km al S del 
Centro P npo Bonampak, orillas del des Lacanja, 
2 (fr), Meave et al. B-4 


F : elsa 
Pozo #4 and Laguneta Los Lagartos, 29 July 197 
Lundell £ Contreras 19575 p 
Barrios, 12 Mar. 1972 (fr), Contreras 11268 (MO); 
Perdido, Qda. Seca, 31 May 1977 (fl), une Re Con- 
treras 21037 (MO). PETÉN: Uaxactün, 7 Apr. 1 (fr), 
Bartlett 12533 (F, GH, K, NY, US); Tikal Ve: Park, 


in ramonal on Temple 27, 16 Mar. 1966 (fr), Contreras 
5609 (ENCB, F, MO, NY); Chinchila, 8 km from San 
Luis, on Sebol Rd., 


"Ce m SE de Say 
fa), "Durs 2681 y Ta CAYO: Valentin, in advanced 
forest, limestone valley, June-July 1936 (fl), Lundell 
6194 (F, GH, 


r forest, Cerro 
m, 23 Feb. 1968 (fr), vi “Molino R. & 
Montalvo 21607 (F, NY). 

Psychotria lundellii may be recognized by its 
narrow (length/width — 3) leaves drying glossy 
red-brown with few (usually 6-8) secondary veins 
and tertiary veins not evident, and its spherical 
fruit drying often glossy red-brown and appearing 
smooth outside due to the seed dorsal surface hav- 
ing 3-6 shallow irregular longitudinal furrows. Psy- 
chotria lundellii differs from the similar P. chi- 
riquina in having smaller (5-6 vs. 8-10 mm long) 
stipules, brochidodromous (vs. eucamptodromous) 
secondary venation, shorter (2-2.5 vs. 4-6 mm 
corolla tubes, and spherical (vs. ellipsoid) fruits. 
Material from El Salvador shows generally more 
secondary veins in the leaves than material gath- 
ered elsewhere. 

Eight flowering specimens from Belize have been 
examined, and only the long-style morph has been 
found, suggesting that the short-style morph has 
been lost in populations in that country. One thrum 
has been found in Baja Verapaz, Guatemala (Lun- 
dell & Contreras 21037) 


42. Psychotria mexiae Standley, Publ. den 
Columbian Mus., Bot. Ser. 4: 6. 192 
TYPE: Mexico. Jalisco: Santa Cruz de Ein 
wooded ravine on mountain side, 300 m, 9 
Dec. 1926 (fr), Mexia 1262 (holotype, F; 
isotypes, A, GH, MO, US-2 sheets). Figures 
10h, 30. 


Palicourea nigrescens M. Martens & Galeotti, Bull. Acad. 
s ci. Bruxelles 11(50): 136. 1844. TYPE: Mex- 
RE eracruz: Cordillera, June- Oct. 1840 (fl), Gal- 
eotti 2653 os C). Not Baeta nigrescens 

De Wild., 1924 


il . Bequaert. 2: 393. 
Psychotria schippii Standley & Steyermark, Publ. Field 
Mus . His xa ot. Ser. 23 43. TYPE: 


Belize Cayo: V n, limestone val ley, June-July 
1936 (i T ondell 6193 (holotype, F; isotypes, GH, 
Y, 


Tree or shrub, 2-5(-10) m tall; young stems 
sparsely puberulent, the bark pale, mottled, with 


Volume 76, Number 2 
1989 


Hamilton 419 


Mesoamerican Psychotria, Part Il 


irregular ridges and raised lenticels; stipules sheath- 
ing, linear, 12-20(-40 mm, gla- 
brous, caducous, leaving a pale ridge with or with- 
out red-brown fringe. Leaves petiolate; petioles 3- 
20 mm long, glabrous or sometimes sparsely to- 
mentose below, flat above; blades membranous to 
subcoriaceous, elliptic, the apex long-acuminate, 
the base cuneate, (8-)10-20 x (2.5-)3-8.5 cm, 
glabrous above and below, sometimes sparsely to- 
mentose along midvein below, drying green-brown 
to sometimes red-brown, the veins often reddish 
below; secondary veins 12-16(-18) pairs, diverg- 
ing (55?-)60?- 70*(-75?), brochidodromous to eu- 
camptodromous, constantly arcuate, prominent be- 
low, glabrous, the axils sometimes with minute tufts 
of hairs below; tertiary veins inconspicuous to ev- 
ident, orthogonal reticulate to rarely random re- 
ticulate. /nflorescences terminal or pseudoaxillary, 
globose panicles of cymes; panicle branched to 4 
degrees; main axis (1-)2-4.5 cm long, the pedun- 
cle lacking; secondary axes in (3)4(5) ranks, the 
first-rank axes 2 or 4, equal, 1-3 cm long, the 
second-rank axes 2 or 4, equal, 0.5-1.5 cm long, 
the third-rank axes 2, 0.2-1 cm long, the fourth- 
rank axes 2, 0.2-0.7 cm long, the fifth-rank axes 
2, 0.1 cm long; cymes branched to (1)2 degrees; 
bracts and bracteoles not evident. Flowers on ped- 
icels 0.5-1 mm long; calyx cup-shaped, the tube 
0.3 mm long, sometimes minutely puberulent; co- 
rolla white, often drying pink, the tube cylindrical, 
2.5-4.5 X 1.5 mm, white ey pid in throat, the 
lobes 5, lanceolate, 1.5- l mm; stamens 5, 
the filaments 2-3.5 mm bs in pins, 3-5 mm 
long in thrums, the anthers 0.8-1 mm long; style 
4—7 mm long in pins, 2-3 mm long in thrums, the 
branches spathulate. Fruit when dry ellipsoid to 
spherical, 4.5-5(-6) mm long, 3.5-4.5(-5) mm 
diam., maturing red, drying often shiny red-brown; 
persistent calyx a tubular beak to 0.5 mm long; 
seed dorsal surface with 5-8 deep irregular lon- 
gitudinal furrows, the ventral surface with 2 deep 
regular plus usually 2-4 additional irregular lon- 
gitudinal furrows (Fig. 10h). 


Distribution (Fig. 30). Scattered from Mexico 
through northern Costa Rica, most common in a 
band from Veracruz and Chiapas, Mexico, through 
Petén, Guatemala, into Belize, at elevations of 400— 
1,800 m in regions of premontane moist forest 
with equatorial to tropical climate. Psychotria 
mexiae has been collected in flower February- July 
and in fruit September- April. 


elected specimens examained. EXICO. CHIAPAS: 
Mpio. Jitotol, adjacent to large double waterfall, 6-8 km 
W of Jitotol, 1,450 m, 16 Dec. 1971 (fr), Breedlove 


23269 (ENCB, F, MO, NY); Mpio. Ocosingo, nr 
anja, 350 m, 2 Apr. 1973 (fl), de no 
NY); Mpio. atic de Espinosa, 18-20 km 
of Ocomscouusls along rd. to Mal Paso, 800 m, 20 
Oct. 1971 (fr), S uris " Thorne 20960 (DUKE, 
ENCB, F, MO, NY); Mpio. La Trinitaria, along Comitan 
River at its iaca. Lago de m NE of 
La Trinitaria, 1,3 


ontebello, 


ntes de Oca, San 

(o) Hinton et re 10295 (B, K, US); Dto. Min 
1 15 Apr. 1939 (fl), Hinton et al. 14 

: . OA XA CA: Dto. voa Sierra de Juárez, s 

175 Mon a Map a, 0.5 de La Esperanza, 

0 m 983 (fl), Cedillo 2: Lorence 2345 
(MEXU); Lu a Cedillo 4180 (MEXU). 

Mpio. Soteapan, Sierra de Sta. 

9 


50 
Beaman 5513 (F, MEXU — 2 sheets, 1 Mpio. San 
Andres Tuxtla, Cerro Mantagaga, 13 km 
Andres Tuxtla, 1,400 m, l Feb. 1972 (fr), na 5594 
MEX 


an Andres 


n comatepec a C 
Mar. 1982 (st), Lorence et al. 3891 (MEXU); Mpio. Es 
Yecuatla, Loma Santa Rita, 1,480 m, 12 Jan. 1972 (fr), 
Ventura 4763 (ENCB, MEXU); Mpio. Banderilla, Ban- 
derilla, 1,500 m, 29 Nov. (fr), Ventura 13658 
(ENCB); Mpio. Atzalán, Alseseca, 1,100 m, 6 June 1980 
(fl), Ventura 17258 (MO). ro ALTA VERAPAZ: 
16 Apr. 1964 (fl), 
aa 16 July 


emple 4 ad’ " group, 13 


Oy c cuevas at Little John's 
Camp, 10 July 1973 (early fr), Dwyer 11577 (GH, ig 
2 sheets). CAYO: vic. Cuevas S of Millionario, 570 m, 
30 Mr 1973 (fl), Croat 23581 (DUKE, MO—2 beh 
; Macal River, bridge between Augustine & Cuevas, 
10 July 1972 (A), Dwyer & Pippin 10193 (ENCB, MO, 
NY). TOLEDO: 2.3 km S of Mayan village of San Jose, 
ca. 8 km W of Columbia Forest Station, river banks on 
limestone, 12 June 1973 (fl), Croat 24302 (F, MO, NY). 


HONDURAS. S 


ojoa, 1,000 m, 7 Aug. 1948 (early "i 
L. O. Williams & Molina 14515 (F, GH, MO). Nica 
GUA. ESTELÍ: Laguna de Miraflor, 13?15'N, 86?15'W, ca. 
1,300 m, 21 Feb. 1982 (fr), P. P. Moreno 15476, 15487 
o JINOTEGA: Laguna Miraflores, 13?15'N, 86?15'W, 
1,250-1,300 m, 10-11 June 1981 (8), Henrich & Ste- 
vens 325 (MO). MATAGALPA: 6-10 km NE of Matagalpa, 
rd. to El Tuma, 1,000 m, 14-16 Jan. 1963 (fr), L. O. 
Williams et al. 23844 (NY, US). CosrA RICA. GUANA- 
CASTE: La Tejona, N of Tilarán, 600-700 m, 25 Jan. 
1926 (fr), Standley & Valerio 45770 (US); Naranjos 
Agrios, 600-700 m, 29 Jan. 1926 (fr), Standley & 
Valerio 46375 (US). 


420 


Annals of the 
Missouri Botanical Garden 


Palicourea nigrescens and Psychotria schippii 
are morphologically identical to P. mexiae and are 
therefore included. The existing name Psychotria 
nigrescens De Wild. (dating to 1924) leaves ““mex- 
iae" as the oldest available epithet. 

Psychotria mexiae may be recognized by its 
elliptic long-acuminate leaves drying green-brown 
to red-brown with veins drying reddish below; glo- 
bose panicles without peduncle and with 4 equal 
secondary axes in each of the first 2 ranks; long 
(2.5-4.5 mm) corolla tube often drying pink; and 
ellipsoid to spherical fruit drying often shiny red- 
brown and appearing smooth outside due to the 
irregularity of the furrows on the dorsal surface of 
the seed. Corolla tubes are shorter in specimens 
from Belize (2.5 mm) than in those from Mexico 
(4-4.5 mm), and the stamens and styles are scaled 
down accordingly. Fruits are larger (5-6 mm long) 
in specimens from Veracruz and Guatemala than 
from elsewhere (4.5-5 mm long). 

The exserted pistil of the pin morph is usually 
longer than the exserted stamens of the thrum (4— 
7 vs. 3.5-5.5 mm). 


43. Psychotria olgae Dwyer & Hayden, Ann. 
Missouri Bot. Gard. 55: 42, fig. 3. 1968. TYPE: 
Panama. Panamá: between Cerro Jefe and 
Eneida, 630-870 m, 17 Jan. 1968 (fr), Dwyer 
et al. 8193 (holotype, MO; isotypes, DUKE, 
F, NY). Figures 10g, 29. 


Tree 2-8(-10) m tall; young stems puberulent, 
the bark pale, slightly ridged longitudinally; stipules 
lanceolate, sometimes biacuminate, 3-5 x 1.5 mm, 
ciliate at apex, caducous, leaving a pale ridge with 
red-brown fringe. Leaves petiolate; petioles 2-3 
mm long, glabrous, flat above, robust; blades co- 
riaceous, elliptic to obovate, the apex obtuse to 
rounded, the base subcordate, the margins inrolled, 
(5-)6-10 x (1.5-)2.5-4.5 cm, glabrous above 
and below, drying green-brown to red-brown; sec- 
ondary veins 5-7 pairs, diverging 60°-65°, eu- 
camptodromous to brochidodromous, constantly 
arcuate, prominulous below, glabrous, the axils 
lacking domatia or hairs; tertiary veins not evident. 
Inflorescences terminal or pseudoaxillary, panicles 
of glomerules; panicle branched to 3 degrees; main 
axis 2.5-9 cm long, the peduncle 1.5-5.5 cm long; 
secondary axes in 2 ranks, the first-rank axes 2, 
1-3 cm long, the second-rank axes 2, 0.4-1 cm 
long; bracts and bracteoles irregular, 1 mm long, 
ciliate. Flowers pedicellate, the pedicels 2-3 mm 
long; calyx cup-shaped, the tube 1 mm long, the 
lobes 5, broad triangular to not evident, to 0.5 mm 
long, minutely ciliate; corolla green, leathery, the 


tube cylindrical, 2.5-3.5 x 2 mm, white pubescent 
in throat, the lobes 5, ovate, 2.5-3 x 1.5 mm; 
stamens 5, the filaments 3-3.5 mm long in pins, 
4 mm long in thrums, the anthers 1—1.5 mm long; 
style 6 mm long in pins, 2.5-3 mm long in thrums, 
the branches linear. Fruit when dry ellipsoid to 
slightly obovoid, 9-10 mm long, 5.5-7 mm diam., 
maturing red, drying red-brown; persistent calyx 
a tubular beak to 1 mm long; seed dorsal surface 
with 4 deep regular and numerous irregular lon- 
gitudinal furrows, the ventral surface with 2 deep 
regular and several irregular longitudinal furrows 


(Fig. 10g). 
Distribution (Fig. 29). Known only from the 


region of Cerro Jefe, eastern Panamá Province, 
Panama, at 800-1,200 m elevation in premontane 
wet forest with equatorial-mountainous climate. This 
species has been collected in flower in March, June, 
and July and in fruit in October and January. 
Additional specimens examined. PANAMA. PANAMÁ: 
Cerro Jefe, to 930 
an 
na N 
Falsom 3817 (CR, Mo. foem Jefe, on top nr. iso 
ca. 1,200 m, 19 Jan. 1977 (fr), om & Harp 1368 
CR, MO); Cerro Jefe region, 1,000 m, 8 Jan. 1975 (fr), 
Luteyn & Wilbur 4655 (DUKE); ps A ca. 1,000 
ns July 1975 (fl), Mori 7122 (M 
5 (fl), Mori 7133 (MO); : June Ys Mori & 
ds 6507 (MO— 2 sheet 
before weather station, 850-9 
Sytsma 1504 (MO); Cerro Jefe, 850- 
1980 (fr), Sytsma 1990 (MO). 


— 


0 m, 29 Oct 


Psychotria olgae may be recognized by its co- 
riaceous leaves with subcordate base and inrolled 
margins and by its large (9-10 x 5.5-7 mm) 
ellipsoid to slightly obovoid fruit. 

e reproductive organs are longer at both levels 
in the pin morph that in the thrum 


44. Psychotria panamensis Standley, Contr. 
U.S. Natl. Herb. 18: 132. 1916. TYPE: Pan- 
ama. Chiriquí: S slope of Cerro Horqueta, Los 
Siguas Camp, ca. 1,700 m, 17-19 Mar. 1911 
(fl, fr), Pittier 3194 (holotype, US). Figures 
2d, 7e, 28 


Shrub or tree, (1-)2-13 m tall; young stems 
minutely puberulent to short-ferrugineous pubes- 
cent, the bark usually furrowed longitudinally; sti- 
pules usually sheathing, lanceolate, (10-)20-80 x 

.5-)3.5- 7 mm, glabrous, minutely puberulent or 


[c 


red-brown ciliate, caducous, leaving a pale ridge 
with red-brown fringe (Fig. 2d). Leaves petiolate; 
petioles (0.5-)2-4.5(-5) cm long, glabrous or pu- 
berulent, flat or grooved above; blades membranous 


Volume 76, Number 2 
1989 


Hamilton 421 
Mesoamerican Psychotria, Part II 


to subcoriaceous, elliptic to obovate, the apex acu- 
minate, the base subcordate to cuneate, (7-)9-28 
x (2.5-)4-11(-13) cm, glabrous above and below, 
the midvein sometimes puberulent below, drying 
red-brown to green-brown; secondary veins (6—)8— 
19(-22) pairs, diverging 45?-70?, eucamptodro- 
mous to sometimes brochidodromous, constantly 
arcuate or straight then arcuate near margin, el- 
evated below, glabrous or minute-puberulent, the 
axils lacking domatia or hairs, or having segments 
of red-brown hairs along midvein and secondary 
vein; tertiary veins inconspicuous, orthogonal re- 
ticulate to percurrent. /nflorescences terminal or 
pseudoaxillary, panicles of cymes or glomerules 
(Fig. 7e); panicle branched to 3-5 degrees; main 
axis 1.5-12 cm long, the peduncle usually lacking 
or 1.5-3 cm long; secondary axes in 3-7(-8) 
ranks, the first-rank axes 2 or 4, equal or subequal, 
0.8-7 cm long, the second-rank axes 2, 0.4-4 cm 
long, the third-rank axes 2, 0.2-3 cm long, the 
fourth-rank axes 2, 0.2-2 cm long, the fifth-rank 
axes 2, 0.1-1 cm long, the sixth-rank axes 2, 0.2- 
0.6 cm long, the seventh-rank axes 2, 0.1-0.2 cm 
long, the eighth-rank axes 2, 0.1 cm long; cymes 
branched to 1-3 degrees; bracts and bracteoles 
irregular or not evident, to 1 mm long, ciliate. 
Flowers pedicellate, the pedicels 0.5-1 mm long, 
sometimes (var. ixtlanensis) with minute sphe- 
roidal glands; calyx cup-shaped, the tube 0.3-0. 

mm long, the lobes 0 or 5, triangular to not evident, 
to 0.3 mm long, minutely ciliate; corolla white or 
cream, the tube cylindrical, 2-3 x 1.5-2 mm 
white pubescent in throat, the lobes 5, lanceolate 
or triangular, 1.5-2.5 x 0.8-1 mm; stamens 5, 
the filaments 2-3 mm long in pins, 3.5-5 mm 
long in thrums, the anthers 0.7-1 mm long; style 
4-6 mm long in pins, 2.5-3 mm long in thrums, 
the branches clublike, spathulate, or linear. Fruit 
when dry ellipsoid to obovoid, 4.5-8 mm long, 
3.5-6.5(- 7) mm diam., maturing red, drying red- 
brown to sometimes (var. ixtlanensis) red-black; 
persistent calyx inconspicuous or a beak to 1 mm 
long; seed dorsal surface with 4 deep irregular or 
7-10 irregular longitudinal furrows, the ventral 
surface with 2 central furrows often plus 2 deep 
or several irregular lateral longitudinal furrows. 


Distribution (Fig. 28). Southern Mexico 
through Panama, at elevations of 0-2,100 m. Va- 
rieties compressicaulis and magna occur at lower 
elevations, while varieties ixtlanensis and pana- 
mensis occur mostly above 1,000 m. 

Psychotria panamensis may be recognized b 
its large (usually 20-80 mm long) lanceolate 
sheathing stipules, leaves drying usually dark red- 


brown, inflorescences with robust secondary axes 
usually 2 per rank in 3-7(-8) ranks and bier 
with no peduncle, and large (usually 4.5-8 

6.5 mm) ellipsoid fruit drying almost always red- 
brown and without a conspicuous persistent calyx. 


KEY TO VARIETIES OF PSYCHOTRIA PANAMENSIS 


la. ee main axis 1.5-3 cm long, t e 
ondary axes in 3(4) ranks; fruit obovo 
o 44b. var. ixtlanensis 
lb. Inflorescence main axis 5-12 cm long, the se 
ondary axes in 5-7 ranks; fruit usually ellipsoid, 
2a. Fruit 4.5-5 x 3.5-4 mm; Caribbean east- 
ern Panama Õun 44c. var. magna 
b. Fruit 5.5 x 4.5 mm and larger. 
3a. Leaf blades 15-28 x 5.5-13 cm, the 
secondary veins 16-19; fruit 5.5-7 
x 4.5-6 mm; lowland Caribbean Nic- 
aragua and — ig WA IA 
var. Compr east eats 
Leaf denn == 9-15 
cm, the secondary veins 2 16; fruit 
generally 7. x 5.5-6.5 mm; Mex- 
ico through Panama, mostly ins 500 
d. var. panamensis 


N 


Qo 
a 


44a. Psychotria panamensis Standley var. 
compressicaulis (K. Krause) C. Hamilton, 
Phytologia 64: 233. 1988. Psychotria com- 
pressicaulis K. Krause, Bot. Jahrb. Syst. 54: 
Beibl. 119: 44. 1916. TYPE: Costa Rica. Car- 
tago: Tuis, 600 m, July 1898 (fl), Pittier 
12412 (holotype, B, M photo, GH; 
isotype, US). Figure 


Shrub or tree (1-)2-10 m tall; stipules sheath- 
ing, lanceolate, (10-)20-35 x (3-)3.5-5 mm, 
glabrous. Leaves: blades membranous, the base 
cuneate to rarely subcordate, (8-)15-28 x 
(3-)5.5-13 cm, drying dark red-brown; secondary 
veins (13-)16-19(-20) pairs, the axils lacking 
domatia or hairs; tertiary veins inconspicuous, or- 
thogonal reticulate to percurrent. Inflorescences: 
panicle branched to 4-5 degrees; main axis 7-9 
cm long, the peduncle usually lacking or ca. 1.5 
cm long; secondary axes in 6-7 ranks, the first- 
rank axes 2, 4.5-7 cm long, the second-rank axes 
2, 2.2-4 cm long, the third-rank axes 2, 1-2 cm 
long, the fourth-rank axes 2, 0.8-1.5 cm long, the 
fifth-rank axes 2, 0.4-0.9 cm long, the sixth-rank 
axes 2, 0.2-0.6 cm long, the seventh-rank axes 
2, 0.1-0.2 cm long; cymes branched to 2 degrees. 
Flowers: corolla white, the tube 2-2.5 x 1.5-2 
mm, the lobes lanceolate, 1.5-2 x 1 mm; stamens 
5, the filaments not seen in pins, 3.5-4 mm long 
in thrums; style not seen in pins, 2.5-3 mm long 
in thrums, the branches linear. Fruit when dry 
ellipsoid, 5.5-7 mm long, 4.5-6 mm diam., drying 


422 


Annals of the 
Missouri Botanical Garden 


red-brown; persistent calyx not evident; seed dorsal 
surface with 4 deep irregular longitudinal furrows, 
the ventral surface with 2 deep incompletely di- 
vided plus several irregular longitudinal furrows. 


Distribution (Fig. 28). Caribbean lowlands of 
Nicaragua and Costa Rica, at 0-200 m elevation 
in tropical moist to wet (to premontane wet) forest 
with equatorial climate. This variety has been col- 
lected in flower July-August and in fruit Novem- 
ber-July. 


Selected specimens examined. NICARAGUA. RÍO SAN 
JUAN: nr. Caño Chontaleño, 20 km NE of El Castillo, Río 
Indio watershed, 200 m, 7-9 Mar. 1978 (fr), Neill 3402 
(MO). zELAYA: Mpio. de Nueva Guinea, Bocas de Piedra, 
2 Nov. 1982 (st), Laguna 131 (MO); logging camp nr. 
Qda. La Talolinga, 11°51'N, 84°26'W, 170 m, 19 Aug. 
1983 (fl), J. Miller & Sandino 1165 (MO); Monkey 
Point, 3 km al S, 11%37'N, 83*40'W, 10 m, 24 Oct. 
1981 (early fr), P. P. Moreno & Sandino 12259 (MO); 
Rio Punta Gorda, Atlanta, “La Richard," 200 m al SE, 
11*32'N, 84°05'W, 20 m, 13 Nov. 1981 (fr), P. P. 
Moreno & Sandino 13027a (MO); new rd. to Mina 
Nueva America, leading more or less W TE 14.3 km 
N of El Empalme on main rd. to Rosita, 7.7 km from 
main rd., 23 Feb. 1979 (fr), W. D. Stevens 12708 (MO); 
vic. junction of rd. to Alamikamba with rd. between El 
Empalme and Limbaika, 13°32'N, 84°30'W, 25 m, 24 
Feb. 1979 (fr), W. D. Stevens 12740 (MO). Costa Rica. 
ALAJUELA: 2-3 km NW of Bajo, Rodriguez, ca. 30 km 
N of La Balsa de San Ramón, 300 m, 8 June 1976 (fr), 
J. Utley & K. Utley 5137 (DUKE). HEREDIA: Finca La 
Selva, the OTS field station on the Río Puerto Viejo, 100 
m, 5 Apr. 1979 (fr), Beach 1418 (CR, DUKE, MO); 4 
Aug. 1980 (fl), diete i 1092 (DUKE); vrbi 
Tirimbina, Istarú Farm, 220 m, 26 Nov. 1971 (fr), Lent 
2246 (CR, DUKE, MO, NY, US). PUNTARENAS: Corco- 
vado National Park, slopes above Llorona, 8?36'N, 
83%42'W, 13 July 1977 (fl), Liesner 3263 (CR, MO). 


All eleven flowering collections are thrum morphs, 
suggesting that the variety is thrum-monomorphic. 


44b. Psychotria panamensis Standley var. 
ixtlanensis C. Hamilton, Phytologia 64: 233. 
1988. TvPE: Mexico. Oaxaca: Dto. de Ixtlán, 
21.4 km al S de Valle Nacional, 17?41'N, 
96?18'W, 1,140 m, 28 Nov. 1979 (fr), Wendt 
et al. 2258 (holotype, MEXU — 317322; iso- 
type, ENCB). Figure 28. 


Shrub 2—4 m tall; stipules lanceolate-acuminate, 
(10-)30-60 x (2.5-)5-7 mm, minutely puberu- 
lent. Leaves: blades membranous, the base cuneate 
to attenuate, (9-)16-21 x (3-)5-7.5 cm, drying 
green-brown; secondary veins 12-17 pairs, the 
axils lacking domatia or hairs; tertiary veins in- 
conspicuous, orthogonal reticulate. Inflorescences: 
panicle branched to 3-4 degrees; main axis 1.5- 
3 cm long, the peduncle lacking; secondary axes 


in 3(4) ranks, the first-rank axes (2)4, subequal, 
0.8-2.2 cm long, the second-rank axes 2, 

1.0 cm long, the third-rank axes 2, 0.2-0.6 cm 
long, the fourth-rank axes 2, 0.2 cm long; cymes 
branched to 1-2 degrees. Flowers: corolla cream, 
the tube 3 x 1.5 mm, the lobes triangular, 1.5 x 
l mm; stamens 5, the filaments 2-2.5 mm long 
in pins, not seen in thrums; style 4 mm long in 
pins, not seen in thrums, the branches short, club- 
like. Fruit when dry obovoid, 5-6 mm long, 4- 
4.5 mm diam., drying dark red-brown to red-black; 
persistent calyx inconspicuous or a minute beak to 
0.5 mm long; seed dorsal surface with 4 deep 
irregular longitudinal furrows, the ventral surface 
with 2 incompletely divided central plus 2 deep 
lateral longitudinal furrows 


Distribution (Fig. 28). | Known only from type 
region of Ixtlàn, Oaxaca, Mexico, at ca. 900- 
1,140 m elevation in a region of evergreen to 
subevergreen forest with tropical-mountainous cli- 
mate. This variety has been collected in flower 
April-June and in fruit September, November, and 
April. 

Selected baie examined. MEXICO. OAXACA: Dto. 
de Ixtlán, Sierra de Juárez, a 2.5 km al NE de Puerto 
Eligio, 900 m, 2 T 1983 (fl), Cedillo & Lorence 2397 
EXU); entre Vista Hermosa y Comaltepec, a 1.5 km 
al 5 de Vista Hermosa, Sierra Juárez, 16 Sep. 1965 (fr), 
G. Martínez C. 296 (ENCB, MO); Dto. i Ixtlán, 5.3 
km al N de Vista Hermosa, carr. a Oaxaca- Tuxtepec, 
277 Sep. 1982 (fr), Torres & Cedillo 1461 (ENCB, MEXU). 


~ 


This variety, with its leaves drying greenish and 
fruit obovoid, may be the result of local introgres- 
sion involving Psychotria trichotoma in the Ixtlán 
region of Oaxaca. 

Four flowering collections are all of the pin 
morph, suggesting (albeit weakly; p = 0.125) that 
the variety may be pin-monomorphic. 


44c. Psychotria panamensis Standley var. 
magna (Standley) C. Hamilton, Phytologia 
64: 234. 1988. Psychotria magna Standl., 
Contr. U.S. Natl. Herb. 18: 131. 1916. TYPE: 
Panama. Colón: Loma de la Gloria, nr. Fató 
(Nombre de Dios), 10-104 m, Aug. 1911 (fl), 
Pittier 4092 (holotype, US-679188; isotype, 
US-693188). Figure 28. 


Shrub; stipules not seen. Leaves: blades mem- 
branous, the base cuneate, 17-24 x 7-11 cm, 
drying red-brown; secondary veins 14-22 pairs, 
the axils lacking domatia or hairs; tertiary veins 
inconspicuous, percurrent. Inflorescences: panicle 
branched to 5 degrees; main axis 10-12 cm long, 
the peduncle lacking; secondary axes in 6 ranks, 


Volume 76, Number 2 
1989 


Hamilton 423 
Mesoamerican Psychotria, Part Il 


the first-rank axes 2, 6 cm long, the second-rank 
axes 2, 4 cm long, the third-rank axes 2, 3 cm 
long, the fourth-rank axes 2, 2 cm long, the fifth- 
rank axes 2, 1 cm long, the sixth-rank axes 2, 0.6 
cm long; cymes branched to 1-3 degrees. Flowers: 
corolla color unknown, the tube 2-2.5 x 1.5 mm, 
the lobes lanceolate, 1.5-2 x 0.8 mm; stamens 
5, the filaments not seen in pins, 3.5 mm long in 
thrums; style not seen in pins, 2.5 mm long in 
thrums, the branches clublike or linear. Fruit when 
dry ellipsoid to slightly obovoid, 4.5-5 mm long, 
3.5-4 mm diam., drying dark red-brown; persis- 
tent calyx a minute beak; seed dorsal surface with 
4 deep irregular longitudinal furrows, the ventral 
surface with 2 incompletely divided central plus 2 
deep lateral longitudinal furrows. 


Distribution (Fig. 28). Known only from the 
Caribbean coast of Panama just east of the Panama 
Canal in tropical moist forest with equatorial-trop- 
ical climate. This variety has been collected in 
flower in August and in fruit in October. 

Additional specimens examined. bird SAN BLAS: 


trail E of Cangandi- Mandinga airport rd., 2-5 mi. S o 
Mandinga airport, 27 Oct. 1967 (fr), Duke 14782 (MO, 
US). 


lac] 


The only flowering collection is of a thrum flower 


44d. Psychotria panamensis Standley var. 
panamensis. Figure 


— ee ei Standley, J. Wash. Acad. Sci. 
1928, not PRT E Merr., 
J. As t. Soc. Mal. P. gran- 
vin d Standl.: eue Rica. San José: vic. Sta. 
Maria de Dota, ca. 1,600 m, 26 Dec. 1925 (fr), 
Standley & Valerio 43268 (holotype, US). 
Psychotria fere Standley in Yuncker, Publ. Field 
Mus. Nat gh Bot. Ser. 17: 397. 1938. TYPE: 
Honduras. Comayagua: nr. summit of y above 
El Achote, Ku plains of Siguatepeque, 1,850 m 
21 July 1936 (fl), Yuncker et al. 6013 (holotype, 
F; isotypes, G 0). 
Psychotria molinae Standley, Ceiba 1: 46. 1950. TYPE: 


O. Williams & Molina 10390 (holotype, F— 
64188; isotypes, F, GH). 

Psychotria durilancifolia Dwyer, Ann. cam Bot. Gard. 
67:3 1980. TYPE: Panama. Pana l Llano- 
red Rd., ME Arie Lab Mosquito eee Project 

at 2, 1 Aug. 1974 (fl), Croat 26028 
i po 2240525). 

Tree (2-)4-13 m tall; stipules sheathing, lan- 
ceolate, (10-)30-80 x (3.5-)5-7 mm, red-brown 
ciliate. Leaves: blades membranous to subcoria- 
ceous, the base cuneate to subcordate, (7-)9- 


15(-22) x (2.5-)4-6(-10) cm, drying deep red- 
brown or sometimes green-brown; secondary veins 
(6-)8-16(-20) pairs, the axils lacking domatia or 
hairs or sometimes with segments of red-brown 
hairs along midvein and secondary veins below; 
tertiary veins inconspicuous, orthogonal reticulate. 
Inflorescences: panicle branched to 3-4 degrees; 
main axis (2.5-)5-9 cm long, the peduncle usually 
lacking, or 1.5-3 cm long; secondary axes in (4-) 
5-7(-8) ranks, the first-rank axes 2, (1-)2-6.5 
cm long, the second-rank axes 2, (0.4-)1-3.5 cm 
long, the third-rank axes 2, (0.2-)0.6-2 cm long, 
the fourth-rank axes 2, (0.1-)0.2-1.5 cm long, 
the fifth-rank axes 2, 0.1-0.6 cm long, the sixth- 
rank axes 2, 0.4 cm long, the seventh-rank axes 
2, 0.2 cm long, the eighth-rank axes 2, 0.1 cm 
long; cymes branched to 1-2 degrees. Flowers: 
corolla white, the tube 2-3 x 1.5-2 mm, the lobes 
lanceolate, 2-2.5 x 1 mm; stamens 5, the fila- 
ments 2.5-3 mm long in pins, 4-5 mm long in 
thrums; style 4-6 mm long in pins, 2.5 mm long 
in thrums, the branches spathulate. Fruit when dry 
ellipsoid, (6—)7—8 mm long, (4.5-)5.5-6.5(- 7) mm 
diam., drying red-brown; persistent calyx usually 
a beak to 1 mm long; seed dorsal surface with 7- 
10 irregular longitudinal furrows, the ventral sur- 
face with 2 deep plus several irregular longitudinal 
furrows. 


Distribution (Fig. 28). Continuous along the 
central cordillera from Veracruz, Mexico, to east- 
ern Panama, at 200-2,100 m elevation, almost 
all above 500 m, in premontane to low montane 
wet forest to rainforest with usually equatorial to 
tropical-mountainous climate. It has been collected 
in flower throughout the year, primarily March- 
August, and in fruit throughout the year, primarily 
November- April. 


cted specimens examined. MEXICO. CHIAPAS: 
Mpio Berriozábal. 13 km N of Berriozábal nr. Pozo Tur 
ipache and Finca El Suspiro, 1,000 m, 24 July 1972 
(fr), Breedlove 26311 (F, MEXU U, MO); Mpio. La Trin- 


itaria, E of Laguna Tzikaw, Mente Bello National Park, 
1,300 m, 13 May 1973 (fl), Breedlove 35141 (ENCB, 


EXU, MO, NY); 
ens) 29565 (MEXU, MO, NY); Mpio. R 
gra 10 km above Rayón Mezcalapa along rd. to Jitotol, 
1.700 m, 25 Jan. 1973 (fr), Breedlove & Smith 32414 


of Valle Nacional bridge, 1,500 m, 19 Feb. 1979 (fl), 
Croat 48055 (CR, MO); Mpio. Matias Romero, 9.5 km 


17%03'N, 94?43'W, 400 m, 2 
1981 (fl), Wendt et al. 3307 (MEXU); Mpio. Matias 
Romero, orilla N del Rio Verde, 6.4 km al SE de Aser- 


radero La Floresta sobre camino a Arroyo Amaca, 1 7?03'N, 


424 Annals of the 

Missouri Botanical Garden 
94?45'W, 200 m, 30 Nov. hes (fr), Wendt et e 3555  Oluma, 750 m, 4 Jan. 1984 (st), Gentry et al. 43930 
(MEXU). PUEBLA pio. ziutlan, Rio 5450 (MOQ). ESTELÍ: Laguna de Miraflor, 13?15'N, 86?15'W, 
m, 29 Mar. 1973 n Puis 8094 (EN CB). VERACRUZ: 1,260-1,300 m, 29 Aug. 1982 (fl), Grijalva 947 (MO); 
Mpio. San pesa uxtla, lado W de Cerro A eae 29 Mar. 1983 (fr), P. P. Moreno 21132 (MO); Cerro 
a. 13 km al Quiabu, ca. 8 km NO de Esteli, 1,500-1,600 m, 19 Oct. 


diede Tuxtla, 1,200 m 
EXU 


cima del Cerro Vigia, 950 m, 22 July 
1972 (fr), Beaman 6391 (MEXU); Mpio. Atzalán, la 
Calavera, carret. Altotonga a Tlapacoyan, 1,500 m 
), Chazaro & Don 110 (MEXU); Mpio. 
oteapan, camino de Tebanca a Bastonal, 6-8 km al S 
de Tebanca, 1,000- 1,100 m, 26 Apr. 1982 (fl), pasen 
et al. 4238 (MEXU); Mpio. Teocelo, 19°23'N, 96°57'W, 
17 Feb. 1973 e Menéndez 67 (MEXU); Mpio. Cho- 
camán, ] km hocamán, gorge of river upstream 
from Chocamán- aR hwy., 19%02'N, 97°01'W, 
. 1981 (fr), Nee 23880 (MO); Mpio. 


de Santiago Tuxtla, 


ar 
km 154, 8 June 1975 (fl), Lundell & ea 
(LL); 3 km is of Purulha, Sierra de los Minas, 
4 Jan. 1974 (fr), L. O. Williams 
et al. 43149 (F). oa ae 0: lower and middle 
slopes ui Volcán Fuego, above Finca Montevideo, along 
Barranco Espinoza er tributary of Rio Pantaleon, 1,200- 
1,600 m, 20 Sep. 1 942 (early fr), Steyermark 52052 


. 1941 (st), Steyermark 
rina Ts JALAPA: Volen Jumay, N of Ja lapa, 1,300- 
2,200 m, 1 Dec. 1939 (f), Steyermark 32472 (F). 
QUEZALTENANGO: Volcán Zunil, 1,700 m, 31 July 1934 
(fl), Skutch 890 (A, I , NY, US); lower S-facing slopes 
à aria de Jesüs 


e DN 1940 (fr), 
Volcán Tues evel 


fl), Steyermark 37948 (F). 
HONDURAS. COMAYAGUA: bosque de Montana La Choca 


y 1962 v 4. 
Molina R. 10823 (F, NY); 10 km We of Sigua atepeque, 
pine forest, 1,200 m, 29 July 1974 (early fr), A. Molina 
R. 30578 (ENCB, F, MO). DISTRITO CENTRAL: ca. 10 km 
N of Tegucigalpa, cloud forest, La Tigra, 16 Feb. 1972 
(fr), Clewell & M. Hernández 3039 (TEFH). FRANCISCO 
MORAZÁN: Mt. Uyuca, nr. Zamorano, 1,500 m, 21 Feb. 
1952 (fr), Carlson 2449 (F); Cerro de e trail from 
Las Flores to La Labranza, 1,600-1 ; 
1948 (fl), Standley 13377 (F). INTIBUCA: Rastrojos del 
Pelón de Guise a lo largo de la quebrada, 1 m, 

Apr. 1956 (fr), 4. Molina R. 6370 (GH, US); Cordillera 

í La 


22615 (NY— 2 sheets). La Paz: Cordillera Guajiquiro 5 
km a Sabanetas, 2,100 m, 21 Mar. 1964 (fl), 4. Molina 
R. & A. Molina 13882 (NY); Cordillera Guajiquiro, 
Montaña Verde, 1,900 m, 23 Mar. 1969 (fr), 4. Molina 
R. & A. coms oe (F, NY). LEMPIRA: Montaña de 
Celaque, 1,900 8-22 Nov. 1974 (fr), Hazlett s.n. 
(MO). NICARAGUA. ' BOACO: Cerro Alegre, San José de Los 
Remates, 12°26'N, 85°44'W, 1,100-1,180 m, 11 Feb. 
1983 (fr), P. P. Moreno 20240 (MO). CHONTALES: Cerro 


1979 (fr), Grijalva & Araquistain 656 (MO — 2 sheets). 
y Volcán Mombacho, rd. and trail 
a, from reservoir to somewhat above 
Plan del Flores, DEN 85°58'W, 950-1,150 m, 1 
Oct. 1977 (A), W. D. Stevens 4327 (MO); Volcán Mom- 
bacho, Plan del Flores to W rim, cloud forest, 11%50'N, 
85*58'W, 1,100-1,220 m, 14 Nov. 1978 (fr), W. D. 
Stevens 10830 (MO). JINOTEGA: 4 km al SE del Cerro 
Kilambé, 13?35'N, 85?40'W, 800-1,000 m, 25 Mar. 
1981 (fr), P. P. Moreno 7459 (MO). MATAGALPA: El 
Comején, 1 km W de la carret. a Waslala, 13%15'N, 
85°34'W, 600 m, 23 Feb. 1983 (fr), P. P. Moreno & 
Robleto 20569 (MO). rivas: Isla de Ometepe, N slope 
of Volcán Maderas on trail from Balgue to Laguna Made- 
ras, 11?27'N, 85?32'W, 1,200 m, 23 Jan. 1981 (fr), W. 
Hahn 510 (MO); Isla de Ometepe, N de Volcán Con- 
cepción, 11°33’N, 85%37'W, 1,100-1,150 m, 11 Mar. 
1981 (fl, Sandino 492 (MO). CosrA RICA. ALAJUELA: 
Sta. Maria National Park, 10%48'N, 85°16'W, 600 m, 
8 Feb. 1978 (fr), Liesner 5218 (CR, MO); 15 km NW 
of San Ramón by air, Cerro Azahar, headwaters of Río 
San Pedro, 10%09'N, 84*34'W, 1,400-1,500 m, 14 May 
1983 (fr), eed et al. 15563 (CR, MO); Zapote, 1,400 
m, 17 Aug. 1938 (fl), 4. Smith 1103 (A, F, MO); region 
of Zarcero, 1, n m, 6 July 1937 (A), A. Smith 4224 
F). GUANACASTE: Los Ayotes, nr. Tilarán, 600-700 m, 
21 Jan. 1926 (fr), Standley & Valerio 45433 (US). 
HEREDIA: Volcán Barba, nr. San José de la Montana, 
1,600 m, 19 June 1965 (fl), Hatheway 1460 (US). 
PUNTARENAS: rd. to Las Alturas, 8*56'N, 82%51'W, 1,400 
m 10 July 1972 (fl, early dus Lent 2727 (CR, F, MO). 
N JOSÉ: Pacific slope of Cerro Chirripó massif, 2,000 

m, 5.7 Apr. 1969 (fl), Davidse & Pohl 1668 (F, MO, NY); 
ic. Sta. María de Dota, 1,500-1,800 m, 26 Dec.- Jan. 
1926 (fr), Standley & Valerio 44098 (US). PANAMA. 
CHIRIQUÍ: trail from Bambito to Cerro Punta, along Rio 
Chiriqui Viejo, 6 Apr. 1937 (fl, fr), Allen 308 (A, F, 
— 2 sheets, MO— 2 sheets, US); vic. “New Switzer- 
land," central valley of Rio Chiriqui Viejo, 1,800-2,000 
m, 6-14 Jan. 1939 (fl), Allen 1387 (F, GH, MO, NY, 
US); SO del campamento Fortuna, propriedad del IRHE, 
8°45'N, 82°15'W, 1,000-1,200 m, 25 Sep. 1976 (early 
fr), M. Correa A. et al. 2796 (CR, MO); on NW side of 
Cerro Pando cloud forest, 21 July 1971 (fl), Croat 15990 
(MO— 2 sheets); 21 July 1971 (early fr), Croat 15993 


~ 


along Rio Colorado, 8°50'N, 82°4 
11 July 1983 (st), Hamilton 

Nueva Suiza, 8%52'N, 82 
1982 (early fr), Hamilton et ol 763 (CR, 
Clara region, 27 km NW of El Hato del Volcán, finca ol 
R. Hartman, 1,500-1,600 m, 18 July 1975 (early ES 
Mori & Bolten 7190 (MO). cocLé: foothills of Cer 

Pilón, nr. El Valle, ca. 900 m, 5 Oct. 1967 (early fr), 
p & Correa 14672 (MO—2 sheets); 7 km N of E 
Copé, lumber camp, ca. 900 m, 11 Jan. 1977 (fl, fr), 
Folsom 1202 (MO); Coclesito rd., 20 Apr. 1978 (fr), 
Hammel 2558 (MO); 15-20 km NE of La Pintada to- 
wards Toabre, 600-1,000 m, 15 Feb. 1981 (fl), Sytsma 
& D'Arcy 3643 (MO). COLÓN: Cerro Santa Rita, ca. 6 
mi. from the Transisthmian Hwy., 250 m, 13 Sep. 1979 


Volume 76, Number 2 
1989 


Hamilton 425 
Mesoamerican Psychotria, Part II 


(early fr), pos 1804 (ENCB, MO); Santa Rita Ridge 

.1 A. & Dressler 

CH. MO); Santa Rita Ridge, 23 Jan. 1968 

DU ". sheets, NY). 

of Rio Tuqueza, betw da. Venado 

= Peje Swamp, 28 June 1967 m Dian 1051 (MO). 
68 (fr), Góm 


— 


ANAMÁ: Cerro Jefe, Feb. 19 ez-Pompa et 
di 3065 (MEXU, MO); El e Can Rd. -11 
from Inter-American Hwy. E , 13 Aug. 1975 


(A), Mori 7734 (MO, US) Cerro a ca. 10 km 
SE of Capira, trail to summit, 870-1,000 m, 7 Dec 
1974 (fr), Mori & Kallunki 3556 (MO —2 FM VER- 
AGUAS: rd. between Escuela Agricola Alto Piedra and Rio 
Dos Bocas, ca. m from escuela, 530-620 m, 26 
a 1974 oe Croat 25878 (DUKE, F—2 sheets, GH, 

O, NY); “Cerro Tute” ridge up from former Escuela 
pan Santa Fe, 8?35'N, 81°05’W, 800-1,000 m, 
20 Feb. 1983 (fr), Hamilton & Dressler 2986 (CR, MO); 
NW of Sante Fe, 1 km from Escuela Agricola Alto de 
Piedra, 24 Feb. 1975 (fr), Mori & Kallunki 4785 (ENCB, 
MO). 

The morphologies of P. grandistipula Standl., 
P. yunckeri, P. molinae, and P. durilancifolia 
fall well within the broad range of P. panamensis 
var. panamensis, so these former species are 
esed with var. panamensis. 

Psychotria panamensis var. panamensis shows 
great variation within its range, including the fol- 
lowing: leaves from Veraguas, Panama, are nar- 
rower and more coriaceous than those from else- 
where; leaves from Colón, Panama, are unusually 
large; secondary veins are more numerous in ma- 
terial from Guatemala, Honduras, and Colón and 
fewer in material from Chiapas, Mexico, and from 
Coclé, Panamá, and Veraguas provinces in Pan- 
ama; secondary vein angle of divergence is greatest 
in leaves from Chiriqui, Panama; secondary veins 
are brochidodromous in Nicaraguan specimens; in- 
florescences are largest in specimens from Panama 
and Costa Rica; flowers are generally larger in 
Costa Rican material; and the corolla tubes from 
Honduras are narrowest (1.5 mm) 


45. Psychotria sarapiquensis Standley, Publ. 
Field Mus. Nat. Hist., Bot. 18: 1360. 
1938. TYPE: Costa Rica. Heredia: Vara Blanca 
de Sarapiqui, 1,500 m, July-Sep. 1937 (fl), 
Skutch 3330 (holotype, US; isotypes, GH, K, 
MO). Figure 31. 


Tree 2-10 m tall; young stems glabrous, the 
bark smooth; stipules sheathing, lanceolate, 11- 
45 x 2-5 mm, glabrous, caducous, leaving a pale 
ridge with red-brown fringe. Leaves petiolate; pet- 
ioles (0.5-)1-2.5(-3.5) mm, glabrous, grooved 
above; blades membranous to subcoriaceous, ob- 
lanceolate to narrowly (or rarely broadly) elliptic 
to lanceolate, the apex short-acuminate, the base 


attenuate to cuneate, 6.5-15(-18) x 2-4.5(-8.5) 
cm, glabrous above and below, drying green-brown 
to red-brown; pig! veins 6—9(—11) pairs, di- 
verging 30?- 70°), eucamptodromous to 
brochidodromous, constantly arcuate to straight, 
prominutous below, glabrous, the axils acking do- 
matia or hairs; tertiary , orthog- 
onal reticulate. Inflorescences terminal or pseu- 
doaxillary, panicles of cymes; panicle branched to 

-4 degrees; main axis (2-)5-13 cm long, the 
peduncle lacking or 3-4.5 cm long; secondary axes 
in (3-)4-5 ranks, the first-rank axes 2, (1-)1.5- 
5.5 cm long, the second-rank axes 2, (0.3-)1-2.5 
cm long, the third-rank axes 2, (0.1-)0.5-1.4 cm 
long, the fourth-rank axes 2, 0.3-0.8 mm long. 
the fifth-rank axes 2, 0.1 mm long; cymes branched 
to 1-2 degrees; bracts not evident; bracteoles ir- 
regular, ca. 0.5 mm long, fringed. Flowers pedi- 
cellate, the pedicels 1-3 mm long; calyx often 
drying paler than the pedicel, cup-shaped, the tube 
0.3-0.5 mm long, the lobes not evident, glabrous; 
corolla green-white, the tube cylindrical, 2.5-3 x 
2 mm, white pubescent in throat, the lobes 5, 
lanceolate, 2-3.5 x 1.2 mm; stamens 5, the fil- 
aments 2.5-3 mm long in pins, 4 mm long in 
thrums, the anthers 1.2-1.5 mm long; style 4- 
4.5 mm long in pins, 2.5 mm long in thrums, the 
branches short. Fruit when dry ellipsoid to slightly 
obovoid, 7-8 mm long, 4-5.5 mm diam., maturing 
red, drying black to red-brown; persistent calyx 
often a minute beak; seed dorsal surface with 4— 
5 irregular often plus several shallow irregular lon- 
gitudinal furrows, the ventral surface with 2 deep 


plus several shallow irregular longitudinal furrows. 


Distribution (Fig. 31).  Disjunct, occurring in 
Veracruz, Mexico, at 400-900 m elevation, and 
along the cordillera in Costa Rica and Panama, at 
600-1,600 m, in tropical moist to premontane wet 
forest with equatorial-mountainous to usually trop- 
ical-equatorial climate. Psychotria sarapiquensis 
has been collected in flower in January, February, 
May, and August and in fruit July-March. 


Selected specimens examined. MEXICO. d a 
Mpio. Catemaco, cerro al E de Coyame, 900 m, 13 Dec. 
1971 (fr), Beaman 5307 (MEXU); Estación de dr ía 
Tropical Los Tuxtlas, 400 m, July 1975 (fr), CES 
417 (ENCB, MEXU); Estación de Biología Tropical Los 
Tuxtlas, Cerro “El Vigia,” 18 Mar. 1971 (fr), Flores 50 


14 Jan. 1981 (fr), Schatz & Nee 233 (F). ig Rica. 
GUANACASTE: Los Ayotes, nr. Tilarán, 600-700 m, 21 
Jan. 1926 (fr), Standley & Valerio 45344 ee HEREDIA: 
Vara Blanca de Sarapiqui, 1,500-1,750 m, July-Sep. 
1937 (A), Skutch 3282 (A, K, MO, US). PUNTARENAS: 
Monteverde, 1,600 m, 20 May 1981 (fl), Haber 531 


426 


Annals of the 
Missouri Botanical Garden 


-H- > | | T.C. 
e 
| + as E 
e 
e o 
e 
- e Bs 15 4 
o. e 
"y 
3% 
t 
à 10 + 
A 
200 km + pr 
A b 
a 
105 100 95 90 85 80 
FIGURE 31. Distributions of Psychotria sarapiquensis (triangles) and P. trichotoma (circles) in Mesoamerica. 


(MO). pre CHIRIQUÍ: Fortuna dam site, 1,200-1,400 
77 (fr), Folsom et al. 5505 (MO — 2 sheets); 
977 (fr), Folsom et al. 5569 


IRHE Fortuna Hydroelectric Project, 8%45'N, 82°12'W, 
1,200 m, 13 Mar. 1982 (early fr), Knapp et al. 4153 
(MO). cocLÉ: base of Cerro Pilón above El Valle, 9 Mad 
1972 (early fr), Gentry 3648 (MO, NY). PANAMÁ: Cerr 
ampana above Su Lin Motel, 25 May 1971 (fl), eS 
14780 (GH, MO, NY); 800-1,000 m, 11 Mar. 1981 
(fr), Sytsma & D'Arcy 3725 (MO). VERAGUAS: 5 mi. W 
of Santa Fe on rd. past Escuela Agricola Alto Piedra on 
Pacific side of divide, 800-1,200 m, 18 Mar. 1973 (fr), 
Croat 23037 (MO); above Santa Fe on slopes of Cerro 
Tute, 1,200-1,400 m, 28 Sep. 1972 (early fr), 
5 (F, GH, MO, NY); Alto Piedra, Santa Fe, 1,200 
m, 27 Sep. 1972 (fr), Lao 519 (MO) N of Santa ds 
-2 km 


i 4868 (MO —2 sheets); 
29 Mar. 1975 (fr), Mori & Kallunki 5325 (MO). 


Psychotria sarapiquensis may be recognized 
by its usually narrow (length/width — 3) leaves 
with few (6-9[-11 ]) secondary veins and its large 
(7-8 mm long) ellipsoid to often obovoid fruit drying 
black or dark red-brown. 

Mexican material is noteworthy for having lan- 
ceolate leaves with secondary veins diverging 60?- 
70°. Collections from the Fortuna and El Valle 
areas in Panama have large (to 18 cm long) broadly 
elliptic leaves with ca. 11 secondary veins, but 


they fit this species concept in other respects. Three 
of the Costa Rican collections— Dryer 1539 (not 
cited above), Skutch 3282, and Haber 53 1 —have 
subcoriaceous leaves drying red-brown and pedun- 
culate inflorescences. Psychotria sarapiquensis is 
the most variable and/or the most problematic 
species in this treatment; it is conceivable that 
additional material from throughout its range will 
reveal it to be two or perhaps three different taxa. 


46. Psychotria stockwellii C. Hamilton, Phy- 
tologia 64: 235. 1988. TYPE: Costa Rica. Ala- 
juela: region of Zarcero, 1,800 m, 18 Oct. 
1937 (fl, early fr), 4. Smith 543 (holotype, 
US; isotype, F). Figures 7i, 30. 


Tree or shrub, (1-)2-10 m tall; young stems 
puberulent, the bark pale, ridged longitudinally; 
stipules sheathing, ovate, 9-1 mm, gla- 
brous, caducous, leaving a pale ridge with red- 
brown fringe. Leaves petiolate; petioles 0.5-2(-2.5) 
cm long, glabrous, terete; blades membranous, ob- 
ovate or elliptic, the apex acuminate, the base 
attenuate, (8.5-)9-19 x (2-)3-7 cm, glabrous 
above and below, the midvein sometimes minutely 
puberulent below, drying dark brown above, slate 
rown or pale brown below; secondary veins (11-) 
14-17 pairs, diverging 60?-75*(-80*), eucampto- 
dromous to sometimes brochidodromous, constantly 
arcuate, prominulous below, glabrous or minutely 


Volume 76, Number 2 
1989 


Hamilton 
Mesoamerican Psychotria, Part II 


427 


puberulent below, the axils lacking domatia or hairs; 
tertiary veins conspicuous to evident, orthogonal 
reticulate. Inflorescences terminal or pseudoaxil- 
lary, spreading panicles of cymes (Fig. 7i); panicle 
branched to 3-4 degrees; main axis (7-)9-12 cm 
long, the peduncle (4-)5-9 cm long; secondary 
axes in 4—5 ranks, usually diverging over 90°, the 
first-rank axes 2, 1.8-5 cm long, the second-rank 
axes 2(4), 1-2.5 cm long, the shorter pair when 
present 0.4 cm long, the third-rank axes 2, 0.7- 
1.5 cm long, the fourth-rank axes 2, 0.4-0.6 cm 
long, the fifth-rank axes 2, 0.3 cm lon 
branched to 1-2 degrees; bracts and bracteoles 
triangular, 0.5-2 mm long, ciliate. Flowers on 
pedicels 0.5-1.5 mm long; calyx cup-shaped, the 
tube 0.8 mm long, the lobes 5, triangular, often 
reflexed, 0.7 mm long, puberulent; corolla cream, 
the tube cylindrical, 4-5 x 1.5-2 mm, white pu- 
bescent in throat, minutely puberulent without, the 
lobes 5, ovate, 2 X 1 mm; stamens 5, the filaments 
3.5-4 mm long in pins, 4.5-5.5 mm long in thrums, 
the anthers 1-1.2 mm long; style 6-7 mm long 
in pins, 3-4 mm long in thrums, the branches 
clublike or linear. Fruit when dry ellipsoid, 5-6 
mm long, 4.5-5 mm diam., maturing red, drying 
dark red-brown; persistent calyx not evident or a 
beak or ring drying pale brown; seed dorsal surface 
with 6-10 irregular longitudinal furrows, the ven- 
tral surface with 2 deep regular plus sometimes 
several irregular longitudinal furrows. 


Distribution (Fig. 30). Known from Alajuela, 
Costa Rica, and western Chiriqui, Panama, at 
1,000-2,200 m elevation in regions of low mon- 
tane rainforest with equatorial-mountainous cli- 
mate. Psychotria stockwellii has been collected 
in flower January-October, primarily January- 
March, with immature fruit July-October, and in 
fruit January-March. 


8; cymes 


Additional specimens examined. Costa Rica. 
ALAJUELA: region of Zarcero, 1,800 m, 18 Jan. 1937 (fl), 
A. Smith 165 (F, MO). PANAMA. CHIRIQUÍ: Las Nubes, 
ca. 2,000 m, 7 Aug. 1974 (fl, ve hes Croat 26450 
(MO); r5 Chorro, 1,800 m, 22 Jan. 1938 (fl), Davidson 
172 (F, MO); 2 Mar. 1938 (fl, fr), xe = 363 
(A, F, MO); end of rd. to Bajo Mono, 21 Mar. 1977 (fl), 


Knapp 1646 (MO); vic. Las Nubes i. NW of Rio 
Chiriquí Viejo W of Cerro ms 2, i m, P Feb. 1973 
e a Liesner 293 (F, —2 sheets, NY); Las Nubes, 
of Cerro Due 1,800-1,950 m, 19 July 

I (early fr), Mori & Bolten 7240 (MO—2 sheet) 
Cerro Pando, nr. Panama-Costa Rica border, 2, 
2,482 m, 21 July 1975 (fl, early fr), Mori & Bolten 
7328 (MO, US); NW side of Cerro Punta beyond Las 
Nubes, 2,250 m, 15 Jan. 1971 (fl, fr), Wilbur et al. 
13211 (DUKE-— 2 sheets, MO); above San Ramón nr 


ajo Mono, 4 mi. NW of Boquete, 1,800 m, 22 Jan. 
1971 (fl, fr), Wilbur et al. 13544 (DUKE). 


Psychotria stockwellii may be recognized readily 
by its inflorescence, whose secondary axes diverge 
over 90? from the main axis and whose tertiary 
axes diverge likewise from the secondary axes, and 
so forth, a character state unique in the subgenus. 
In addition, P. stockwellii differs from P. pana- 
mensis in having a larger corolla tube (4-5 vs. 2- 
3 mm long), long-pedunculate inflorescences, and 
conspicuous to evident (vs. inconspicuous) tertiary 
veins. 

Reproductive organs at both levels are longer 
in the pin morph than in the thrum. 


47. Psychotria trichotoma M. Martens & 

Me e e Acad. Roy. Sci. Bruxelles 11: 

221 4. Uragoga trichotoma (M. Mar- 

tens P Con Kuntze, Revis. Gen. Pl. 2: 

963. 1891. TYPE: Mexico. Veracruz: Jalapa 

et Mirador, 900-1,200 m, Aug. 1840 (fl), 

Galeotti 7092 (holotype, G, n.v., photo, MO). 
Figure 31. 


Shrub or small tree 1.5-6 m tall; young stems 
glabrous or rarely minutely puberulent, the bark 
smooth or longitudinally furrowed; hie sheath- 
ing, lanceolate, (10-)20-30 mm, glabrous, 
caducous, leaving a pale ridge with red-brown fringe. 
Leaves petiolate; petioles 1-4(-5.5) cm long, gla- 
brous, flat above; blades membranous, ovate to 
elliptic, the apex acuminate, the base attenuate to 
cuneate, (10.5-)13-23(-26) x (5.5-)6.5-12 
(-15) cm, glabrous above and below, drying usually 
green-brown or sometimes red-brown, the veins 
usually drying paler; secondary veins (12-)14— 
18(-20) pairs, diverging 45°-60°, brochidodro- 
mous with secondary loops near margin, straight 
or sometimes slightly arcuate, elevated below, gla- 
brous, the axils lacking domatia or hairs; tertiary 
veins evident to inconspicuous, orthogonal reticu- 
late to percurrent. /nflorescences terminal or pseu- 
doaxillary, panicles of cymes: panicle branched to 
4—5 degrees; main axis (4-)7-16 cm long, the 
peduncle usually absent or 5-9 cm long; secondary 
axes in 5-7 ranks, the first-rank axes 2(4), equal, 
4—8.5 cm long, the second-rank axes 2, 2-4.5 cm 
long, the third-rank axes 2, 1-3 cm long, the 
fourth-rank axes 2, 0.5-1.5 cm long, the fifth- 
rank axes 2, 0.2-1 cm long, the sixth-rank axes 
2, 0.3-0.4 cm long, the seventh-rank axes 2, 0.2 
cm long; cymes branched to (1)2 degrees; bracts 
and bracteoles not evident. Flowers pedicellate, 
the pedicels 0.5-1 mm long; calyx cup-shaped, 
the tube 0.2-0.5 mm long, the lobes (4)5, trian- 


428 


Annals of the 
Missouri Botanical Garden 


gular to barely evident, glabrous; corolla white, the 
tube cylindrical, 2.5 x 1.5 mm, white pubescent 
in throat, the lobes (4)5, lanceolate, 2-2.5 x 
mm; stamens (4)5, the filaments 2-2.5 mm long 
in pins, 3-3.5 mm long in thrums, the anthers 
0.8-1.3 mm long; style 4 mm long in pins, 2 mm 
long in thrums, the branches short. Fruit when dry 
obovoid, (5-)6-8 mm long, (4-)4.5-5.5 mm diam., 
maturing red, drying usually black or sometimes 
red-brown; persistent calyx inconspicuous or a broad 
beak ca. mm long; seed dorsal surface with 
4-5 irregular often plus several irregular shallow 
longitudinal furrows, the ventral surface with 2 
deep regular and often 2-4 irregular longitudinal 
furrows. 


Distribution (Fig. 31). Southern Mexico 
through Nicaragua, at 100-1,600 m elevation in 
regions of mostly evergreen forest with usually 
tropical-mountainous climate. Psychotria tricho- 
toma occurs also in Venezuela and Ecuador. It has 
been collected in flower April-July and December— 
January, and in fruit throughout the year, primarily 
August- March. 


Selected end d ES MEXICO. CHIAPAS: 
Mpio. Solusuchiapa, 3 above Solusuchiapa along 
rd. to Tapilula, 450 m, 26 July 1972 (fl), Breedlove 
26461 (ENCB, MEXU, MO); Mpio. A Mn 
to Laguna Ocotal Grande, 800 m, 6 Feb. 1973 (fr), 
Breedlove 33093 (MEXU, MO); Mpio. Cina de P 
Mina, 12 km S 
Oro, 1,000 m, 16 Oct. 1971 Ln 
fisesdlou e & Thorne 20640 (MEXU, MO, NY); M 
Mapastepec, ped Rio pias 10 km SE of Mapas. 
tepec, , 24 Dec. 1972 (fl), Breedlove & Thorne 
30660 (MEXU); Escuintla, Esperanza, 14 Apr. 1947 (fl), 
Matuda 16461 (F, MEXU); nr. Tumbala, 1,200-1,650 
m, 20 Oct. 1895 (fr), E. Nelson 3299 (F, US). JALISCO: 

of San Sebastian, Hda. n Ototal, Arroyo de los Palos 
Blancos, 1,500 m, 9 Mar. ] 
F, pu ue NY, US). 
N Putla, sobre el Rio Cuchara, 970 m, 20 June 
s (8, Cedillo & Torres 1487 (ENCB, MEXU, MO); 
Dto. Cuicatlan, De La R 


Schultes & Reko 757 (F). TABASCO: eii. 
carret. a Tecoluta, 26 m, 9 Oct. 8 (ir), Calzada 
4965 (ENCB, MEXU); Mpio. Huizangaille km 12.6 de 
la desviación de Huimanguillo hacia Francisco e 
Oct. 1980 (fr), Cowan & Magaña 3267 (ENCB, MEXU); 
pio. Macuspana, along Arroyo Hular, S of Macuspana, 
28 Sep. EU (fr), Gilly & E. Hernández X. 339 (GH, 
MEXU); a 11.5 km de Villahermosa, por la carret. a 
Escárcega, is Jan. 1966 (fr), González & Pérez P-546 
(ENCB). VERACRUZ: Mpio. ue Sk "La D d 
y Chavaxtla, 19%9'N, 96*53'W, 650 m, 3 Oct. 1979 
(fr), Avendaño & Calzada 516 ENCB, F, MEXT) 


Teocelo, El Olmo, 11 Oct. 1980 (fr), Barrera et al. 298 
MEXU); Mpio. Hidalgotitlán, Benito Juárez segundo, 
17°47'N, 100 m, 2 Nov. 1978 (fr), Castillo 
358 (F); km 3 carret. Playa Vicente a Santa Cecilia, 6 
Oct. 1971 (fr), Chavelas et al. ES-4245 (MEXU); orillas 
de la Laguna de Catemaco, camino a Coyame, 14 Apr. 
1969 (fr), Nevling & Gómez-Pompa 903 (F, GH, MEXU); 
swamps nr. Jalapa, 1,200 m, 19 May 1899 (fl, fr), Prin- 
gle 8198 (A, F, GH, K, MEXU—2 sheets, MO, NY, 
US— 2 sheets); Mpio. Hidalgotitlán, brecha Hermanos 
Cedillo-La Escuadra, HAD, 94 s W,150m,3 June 


— 


94°39'W, 


. Vazquez T. 5 

pio. Atzalan, Napoala, 950 m, He Nov. 1969 
(fr), Pod 103 (ENCB); Mpio. de Totutla, el Mirador, 
1,000 m, 3 May 1976 (fl, fr), Ventura 12723 (MEXU). 
nas ALTA VERAPAZ: nr. 


Guacalata, 600 m, 16 Dec. 1938 n Pd 60207 


(F). SUCHITEPÉQUEZ: nr. Santo Dom azatenan- 
go, 300 5 Mar. 1941 (fr), Standley 88882 (F). 
Doom. COPÁN: cerca de Dulce re, 1,200 m, 30 


Mar. 1963 (fr), 4. Molina R. 11760 (F—2 sheets). 
OLANCHO: Montaña del Chifiringó, 20 km S de Campa- 
mento, 1,000 m, 6 Sep. 1979 (fr), C. Nelson 5339 
TEFH). os BOACO: entre Cerro Alegre y el Rob- 
lar, San José de los Remates, 12%36'N, 85?43'W, ca. 
1,000 m, IL al 1983 (fr), P. P. Moreno 20193 (MO); 
seh and upper SW slope of Cerro Mombachito, 
85°33'W, 950-1,020 m, 18 Jan. 1981 (fr), 


: a Fundadora, 13%03'N, 85°54 
1,200-1,400 m, n Oct. 1979 (fr), W. D. Stevens & 
Grijalva 15443 (MO). MANAGUA: ca. 5.4 km NE of El 
Crucero, ridge of Sierra de Managua nr. Hda. Palmira, 
12°01'N, 86°16'W, 800-900 m, 25 Aug. 1977 (fr), W. 
D. Stevens 3466 sea MATAGALPA: camino al Cerro La 
Carlota, a 2k carret. al Tuma, 12%58'N, 85%52'W, 
0 m, 5 Mar. 1982 (fr), P. P. Moreno 15666 


Jan. 1979 (fr), W. D. Stevens 11801 (MO); Macizos de 
Peñas Blancas, drainage of 
and W of Hda ds 
18-20 Jan. 1982 (fr), W. D. Stevens 
et al. 21144a (MO, rd. to La Fundadora, N of Sta 
Maria de Ostu 1,300-1,500 m, Feb. 1963 (fr), L. 
O. Williams et a , 24803 ay ALT US). zELAYA: El 
Hormiguero 45'N, 84*59'W, 800- 
1,000 m, 17 Mar. 1980 (fr), joe 6182 (MO). 


The name Psychotria trichotoma has been used 
to misidentify specimens of most of the species in 
Group 6 plus P. tenuifolia, P. limonensis, P. car- 
thagenensis, P. quinqueradiata, and P. chagren- 
sis. 

Psychotria trichotoma may be recognized by 
its fairly wide (length/ width = 2) leaf blades drying 
usually green-brown with veins drying paler; sec- 
ondaries usually 14-18 pairs, straight and making 
arches near margin; large inflorescences (usually 
7-16 cm long) with no peduncle; and large (usually 


Volume 76, Number 2 Hamilton 429 
1989 Mesoamerican Psychotria, Part II 


Specimens from Oaxaca show also minute puber- 
ulence on the inflorescence axes. 

Reproductive organs at both levels are longer 
in the pin morph that in the thrum. 


6-8 mm long) obovoid fruit often drying black. 
The pedunculate inflorescences are found in some 
material from Oaxaca and Veracruz, Mexico, where 
the basal secondary axes may have been aborted. 


NEW SPECIES AND OTHER 
TAXONOMIC MATTERS IN THE 
NEW WORLD MEMECYLEAE 
(MELASTOMATACEAE) 


Thomas Morley! 


ABSTRACT 


r new species of Mouriri and one of (dial are described and illustrated. Mouriri beider is a to 


Fou 
a variety of / - gleasoniana. The relation of Votom 
seed structure; no change 
drawings of M. longifolia and V. pubescens are provide 


to Mouriri is discussed in the light of new on 


in generic boundaries is inibi ah for the present. Supplementary dl jur: 
d. 


Recent collections and older ones recently iden- 
tified correctly to genus have yielded five new species 
of the tribe Memecyleae DC. (Melastomataceae). 
Four of these are members of the genus Mouriri: 
one from the Osa Peninsula of Costa Rica, one 
from western Venezuela in the state of Táchira, a 
third from southern Venezuela in Amazonas, and 
the fourth from northeastern Peru and adjacent 
Brazil. Also in Mouriri, new collections of M. glea- 
soniana from the Panama mainland, from Costa 
Rica, and from the island of Coiba revealed much 
intraspecific variation and require reassessment of 
the status of its close ally, M. coibensis. A recent 
flowering collection of M. longifolia permits the 
completion of its description and illustrations and 
allows its sectional placement. The fifth new species 
is a Votomita from Roraima of Brazil, the eighth 
known member of that small genus. The specimens 
of this species, as well as those of another collection 
of V. pubescens, bear ripe fruit, so that it is now 
possible to compare more accurately the seeds of 
Votomita and Mouriri. Previously, mature seeds 
were known only from V. monantha of Cuba. A 
discussion of ovule and seed structure and of ge- 
neric relations therefore follows the description of 
V. roraimensis. More collections of V. pubescens 
make possible the completion of the descriptions 
and illustrations for that species. 

References in the descriptions to the "type" of 
stomatal crypt refer to the forms described on 
pages 6 and 7 of Morley (1976). 


Mouriri osaensis Morley, sp. nov. TYPE: Costa 
ica. Peninsula de Osa, Llorona. Arbol co- 


mün; solamente con frutos; con aprox. 7 m 
de alt. y 30 cm DAP. Madera pesada y muy 
resistente, usada para timones y ejes de car- 
retas, macetas para pilar arroz, plumas o pa- 
lancas etc., al rajarla al acha se va en capas 
y al quebrarla quedan chuzos muy punzantes. 
Nombre regional: Huesillo & Quiebracho. 19 
Sep. 1975, L. J. Poveda & Chrys y Adelaida 
Vaughan s.n. (holotype, MIN). Figure 1. 


Arbor usque 7 m alta trunco 30 cm diametro; petioli 
3-5 mm longi; laminae grandes, basi rotundata usque 
truncata usque subcordata incisura usque 1 mm profunda; 


fructus atropurpureus, globosus vel depresso- sb sine 
lobis; cicatrix hypanthii 7-11 mm diametro, 1.5-2 mm 
alto; loculi 2-5; semina 2-5. 


Tree to 7 m tall and 30 cm in diam., glabrous 
except for the inflorescence; crown at times dense 
and elongate; young twigs terete or with 4 rounded 
angles; bark light gray, exfoliating in very small 
sheets; wood heavy and very resistant. Petioles 3— 
5 mm long; blades somewhat shiny on the face, 
16-27 cm long, 5.6-9 cm wide, elliptic to ovate- 
elliptic or with an oblong tendency, abruptly acu- 
minate at the apex, rounded to truncate to shal- 
lowly cordate at base, when cordate the notch to 
l mm deep; midrib grooved above at least in the 
lower half, prominent and rounded below; lateral 
nerves conspicuous when dry. Midrib xylem tu- 
bular, depressed in the center above; stomatal crypts 
none; upper epidermis mostly 1 cell thick but oc- 


' Botany Department, 220 Bioscience Center, University of Minnesota, St. Paul, Minnesota 55108, U.S.A. 
ANN. MISSOURI Bor. GARD. 76: 430-443. 1989. 


Volume 76, Number 2 Morley 431 
1989 New World Memecyleae 
A B 
S //& 
ON S 
300 
pm 
ND 
9 // € 
20 © 9) 
LD 


Mouriri osaensis Adan & C. & A. Vaughan s.n.).—A. Leav 


FiGUR - 
showing v and terminal sclereids. — C. 
Cross section n of leaf midrib. TE Fruit cluster. — F. Seed 


casionally 2 cells thick at scattered places in the 
same leaf, mucilaginous walls occasional to fre- 
quent in the single cells and in the inner of the 
doubled cells; hypodermis none; terminal sclereids 
irregularly stellate with a strong columnar tenden- 
cy, most with columnar arms, many entirely co- 
lumnar. Flowers borne at leafless nodes of twigs 
3.5-4 mm thick, in clusters of 1-3 per side, the 
peduncles 1-flowered, ca. 4.5 mm long to base of 
pedicel with 2 internodes in that length; bracts 
triangular, 2 mm long. True pedicels 7-9 mm long 
in fruit, very minutely puberulent. Flowers un- 
known. Fruits dark purple at maturity, globose to 
depressed-globose, not lobed, 12.2-16.5 mm high 


including hypanthium rim, 12-17 mm diam. when 


s. — B. Cleared portion of leaf blade 
—D. 


Cross section of leaf blade showing omes and upper epidermis. 
(perhaps not fully mature 


dry, bearing a circular rim of the circumscissile 
hypanthium 7-11 mm diam. and 1.5-2 mm high, 
2-5-locular, 2-5-seeded. Seeds 8-8.7 mm high, 
6.5-7.7 mm wide, 5-5.2 mm thick, irregularly 
ovoid, with a light brown slightly thickened portion 
6-6.5 mm high in the lower half, the rest dark 
brown; hilum basal, whitish, rough, 1.5-3 x 3 


mm. 


Distribution. Known only from the type lo- 
cality in the western part of the Osa Peninsula on 
the southwest coast of Costa Rica. 

The new species appears to belong in subg. 
Mouriri sect. Olisbea. The characteristics sug- 
gesting this alignment are the large leaves, absence 


432 


Annals of the 
Missouri Botanical Garden 


of stomatal crypts, simple upper epidermis with 
mucilaginous walls, stellate terminal sclereids, and 
lack of a hypodermis. Within the section, the new 
species is closest to M. completens, on the basis 
of the former's similar leaves with a cordate ten- 
dency, very similar upper epidermis and foliar 
sclereids, and above all, the circumscissile hypan- 
thium as seen on the fruit. The absence of flower 
characters of course detracts from understanding 
the relationships, but I still regard this alignment 
as reasonably certain. Mouriri osaensis differs from 
M. completens in leaf, fruit, and seed characters. 
The leaf blade is less cordate than in M. comple- 
tens; the fruit is purple instead of greenish, lacks 
a broad stipe at base, and is not abruptly narrowed 
to the hypanthium at the apex, nor is it lobed 
according to the seed number; the hypanthium on 
the fruit is 7-11 mm in diameter rather than 4.5- 
7 as in M. completens and is only 1.5-2 mm deep 
instead of 3.8-4.5 mm. The seeds are a bit smaller 
and differ in shape, best compared in the illustra- 
tions. 


Mouriri papillosa Morley, sp. nov. TYPE: Ven- 
ezuela. Tachira: 10 airline km E of La Fun- 
dacion, 13-23 km by road, around Represa 
Dorada, primary evergreen forest with patches 
of secondary vegetation. Soils tending to be 
sandy, derived from metamorphic rock (schist 
or gneiss), 600-900 m, 7?47-48'N, 71°46- 
47'W, 4 m tree, petals and anthers pinkish. 
30 Apr. 1981, R. Liesner & M. Guariglia 
11616 (holotype, US; isotypes, MIN, MO). 
Figure 2 


Arbor usque 15 m alta; laminae 4.7-11.5 cm lon 

.3-5.5 em altae, elliptico-ovatae usque lipticas, Las 
truncata vel subcordata; pagina inferior costae mediae 
rotundata ad basem, 2-angulata 44-14 pipe versus 
apicem; cryptae stomatophorae Typi II, tecto 
presso; epidermis supera stratiis 2-3 sellularom, la 
interiorae plerumque parietibus mucilaginis aureis, pari- 
etes lamellis 5-9; calyx ovarium inferum includens 6-7 
mm longus, ovarium papillosum; calyx 1 .8 mm fin- 
dens inter lobos sub anthesi; antherae 3.6-4 mm longae; 
ovarium 4-5-loculare; ovula 12-16 


Tree to 15 m high, glabrous except for the 
inflorescence; young twigs terete to 2-channeled 
to 4-angled. Petioles 1.5-3 mm long; blades 4.7— 
11.5 cm long, 2.3-5.5 cm wide, elliptic-ovate to 
seldom elliptic, abruptly short- to medium-acumi- 
nate at apex, rounded to a truncate or slightly 
cordate base, the basal notch up to 1 mm deep; 
midrib plane above, prominent below, rounded at 
base, becoming 2-angled 14-14 of the way to the 
apex, then becoming plane near the apex; lateral 
nerves slightly visible above and below when dry. 


Midrib xylem tubular; stomatal crypts Type II, 
averaging in a leaf ca. 45 um diam., 35 um high, 
57 per mm? (extremes 35-55 um diam., 30-40 
um high, 45-69 per mm"), the roof often depressed 
in the center; upper epidermis 2 or 3 cells thick, 
the upper 1 or 2 layers without mucilaginous walls, 
the cells of the inner layer mostly with golden 
mucilaginous walls /,—'/ the height of the cell and 
containing 5-9 lamellae, these cells enlarged and 
together resembling a hypodermis, aligned or not 
with the cells of the outer layers; terminal sclereids 
stellate-columnar, crowded. Inflorescences in the 
lower leaf axils and at leafless nodes of twigs up 
to 5 mm thick, 1-3 per side, each 1-3-flowered, 
1.5-3.5 mm long to base of farthest pedicel mea- 
sured along the axes with 2 internodes in that 
length, the internodes very minutely puberulent; 
bracts 1-1.5 mm long, ovate-triangular, often de- 
ciduous by anthesis. True pedicels 1-4 mm long; 
calyx including inferior ovary 6-7 mm long, cam- 
panulate, it and the pedicels very minutely puber- 
ulent, the ovary papillose, the papillae enlarging 
into rounded wartlike bumps crowded on the im- 
mature fruit; free hypanthium 2.5-2.8 mm long; 
calyx lobes before anthesis low-triangular, 0.7-0.8 
mm long, 2-2.1 mm wide, 1.8-2.4(- 2.8) mm long 
when measured from the stamen attachment, the 
calyx splitting between the lobes at anthesis a dis- 
tance o mm. Petals pinkish, 5.5-6 mm 
long, 5.5-6 mm wide, triangular, erose, clawed at 
base. Mature filament length unknown; anthers 
pinkish, 3.6-4 mm long; thecae 3-3.5 mm long, 
dehiscing by apical slits; gland 0.4-0.6 mm long, 
2.6-2.9 mm from apex of anther when measured 
from center of gland; cauda ca. 0.45 mm long, 
scarcely noticeable externally. Ovary 4—5-locular; 
ovules axile-basal, produced only outwardly from 
each placenta, 12-16 in all; style ca. 15 mm long. 
Persistent calyx red on immature green fruit. Ripe 
fruit and seed unknown. 


VENEZUELA. TÁCHIRA: Distr. Uribante. In 
forest along road from La Siberia to entrance to Las 
Cuevas Represa. Evergreen forest with much bamboo and 
Scleria secans, € 1,100 m (immature fr), 8 July 1983, 
H. van der Werff & A. González 5123 (US, MO). 


Paratype. 


Distribution. Evergreen forests of far western 
Venezuela in south-central Táchira State SE of San 
Cristobal at 600-1,100 m. 


This species is best placed in the sect. Mouriri 
(Morley, 1976: 36) on the basis of its tubular 
midrib xylem, small Type II stomatal crypts, hy- 
podermis, acuminate leaf apex, axile-basal placen- 
tation, and relatively short thecae. Section Lito- 
phyllum is very similar, differing only in its lack 
of a hypoderms. The hypodermis of sect. Mouriri 


Volume 76, Number 2 
1989 


o 433 


Morley 
New World Memecyleae 


E 2. Mouriri acia e - d iglia 


—C. Cro 


appears to be derived from the inner layer of a 
double or triple epidermis, judging by the presence 
in its cells of mucilage walls and the occasional 
alignment of the hypodermal cells with cells or 
groups of cells in the epidermis. In M. papillosa 
the alignment is rather frequent, as seen in section 
view; however, as seen in face view, the cells are 
most often unaligned. Thus the inner layer of cells 
appears to be sufficiently modified to justify con- 


6).— 
section of bd epidermis of leaf blade. — 


11616). —A. Leaves. — B. Cleared a of leaf blade 
ss section of leaf 
Cross section of leaf midrib. —F. meme 


sidering it a hypodermis like those of the other 
species of sect. Mouriri, and not simply as the 
inner layer of a multiple epidermis. 

Section Mouriri contains three species i in ad- 


panthium, anthers, thecae, and splitting distance 


434 


Annals of the 
Missouri Botanical Garden 


between calyx lobes. It further differs from M. 
barinensis in having short petioles. 

The curious multiple lamellations in the mucilage 
walls of M. papillosa I have noted elsewhere only 
in two collections of M. barinensis. Since this char- 
acter varies in the latter species, it is possible that 
it will also prove to be a variable in M. papillosa. 

Although plants of sect. Litophyllum lack a 
hypodermis, in other respects some of them re- 
semble rather closely the members of sect. Mouriri. 
Most similar is M. gleasoniana, including its var. 
coibensis. From this, M. papillosa differs in its 
lack of a two-winged lower midrib and in its pa- 
pillose ovary and longer ovary plus calyx, longer 
anthers and thecae, and greater splitting distance 
between the calyx lobes. 


Mouriri latihila Morley, sp. nov. TYPE: Vene- 
zuela. Amazonas: Selva alta pluvial, a 7-9 km 
de Yavita hacia Pimichin, lat. 2°55’N, long. 
67?25-30'W. Altura: 125 metros. Tree 3 m 
tall; leaves subchartaceous, dark green above, 
pale green below; fruit orange red. 22 Apr. 
1970, Julian A. Steyermark & George Bunt- 
ing 102927 (holotype, VEN; isotype, MICH). 
Figure 3. 


Arbor 3 m alta; cryptae stomatophorae Typi II; epi- 
dermis supera strata uno cellularum; sclerides terminales 
m plerumque columnares; flores tetrameri, axil- 
laron, solitarii; calyx omnino clausus usque ad anthesem, 

ad lineas conjunctionis loborum rumpens; semen l, 
lateribus 2, ips latissimum, partem majoram lateris uni 
seminis occupan 


Tree 3 m high, glabrous; young twigs narrowly 
4-winged. Leaves subchartaceous, dark green 
above, pale green below; petioles 1-2.5 mm long; 
blades 6.5-11.5 cm long, 3.3-5.7 cm wide, ovate- 
elliptic or ovate-oblong to ovate, abruptly acute to 
abruptly acuminate at apex, broadly acute to 
broadly rounded at base; midrib slightly grooved 
above, prominent and rounded below; lateral nerves 
prominent above and below when dry. Midrib xy- 
lem tubular; stomatal crypts Type II, averaging in 
a leaf ca. 55 um diam., 31 um high, 50 per mm? 
(extremes 40-75 um diam., 25-38 um high, 33- 
66 per mm?); upper epidermis 1 cell thick, the 
cells with mucilage walls 34-34 of the cell height; 
hypodermis none; upper palisade layer clear, not 
filled with tannin compounds, but most of the other 
mesophyll cells so filled; terminal sclereids mostly 
columnar, narrow, often slanting and weaving 
somewhat, sometimes branching in the spongy 
parenchyma, the ends there sometimes horizontal. 
Flowers single in the leaf axils, borne on peduncles 
0.2-1 mm long bearing 2-3 pairs of bracts, the 


bracts 0.7-1.2 mm long, triangular. True pedicels 

.9-1.3 mm long; flowers tetramerous; calyx in- 
cluding inferior ovary 3 mm long when immature, 
mature length unknown, ellipsoid or slightly ob- 
ovoid, conical at each end; calyx lobes fused to the 
apex of the flower in bud, splitting apart regularly 
at anthesis, each then triangular and acute, 1.8- 
2.4 mm long, 1.2-1.4 mm wide, 2-2.4 mm long 
when measured from the stamen attachment; free 
hypanthium ca. 1 mm long. Petals, stamens, and 
ovary at anthesis unknown. Fruit orange-red, de- 
pressed-globose, ca. 9 mm long, mm 
bearing the calyx lobes and the remains of the 
style. Seed 1, ca. 7.5 mm high, 10.5 mm wide, 
4.6 mm thick, oblongish with an apical notch as 
seen in face view, one face and most of the edges 
with polished surfaces, these curved over onto the 
opposite side at 4 places, 2 above, 2 below, the 
irregularly roughened hilum occupying most of the 
opposite side of the seed. 


— 
N 


diam., 


Distribution. Known only from the type lo- 
cality in tropical rainforest near the west-central 
border of Amazonas, Venezuela, between the head- 
waters of the Guainía and Atabapo rivers. 


Mouriri latihila clearly falls within subg. Mou- 
riri on the basis of its tubular midrib xylem and 
its seed with a large polished area. The new species 
is very distinctive, standing apart on the combi- 
nation of its tetramery, closed calyx, and small 
Type II stomatal crypts. Its sectional relation is 
uncertain; it is here placed tentatively in sect. 
Litophyllum because of its tubular midrib xylem, 
small crypts, leaf shape, small flower, and lack of 
a hypodermis (Morley, 1976: 36). When placed 
here, it weakens further the already weak sectional 
boundaries. From this position the new species ap- 
pears to have ties with sect. Megacrypta, whose 
two species also have tetramery, closed calyces, 
and columnar sclereids. The new plant differs widely 
from these two, however, in its much smaller leaves 
and small Type II stomatal crypts, as well as in its 
shorter pedicels and calyces. Within sect. Lito- 
phyllum, no species appear very similar to M. 
latihila, although two are tetramerous and have 
short one-flowered peduncles like the new species: 
M. exadenia and M. lancifolia. Mouriri latihila 
differs from M. exadenia in having Type II crypts, 
columnar sclereids, a closed calyx, and a different 
seed shape; from M. lancifolia it differs in its 
broader leaves, presence of stomatal crypts, co- 
lumnar sclereids, and closed calyx. 


Mouriri retentipetala Morley, sp. nov. TYPE: 
Brazil. Amazonas: Rio Javari, behind Palmei- 


Volume 76, Number 2 
1989 


Morle 


y 435 
New World Memecyleae 


Mouriri latihila (Steyermark & Bunting 102927). 


IGURE 
dd veins and terminal sclereids. — C. 


epidermi 
side; b, bium s 


ras Army Post, 72°49’W, 5?8'S. Forest on 
terra firme. Treelet, 3 m tall. Fruit green, 1 
Aug. 1973, E Lleras, W. C. Steward, J. C. 
Ongley, D. F. Coelho, J. F. Ramos & J. F. 
Lima P16980 (holotype, NY; isotypes, CAS, 
G, INPA, MIN). Figure 4. 


bens Lenis laminae 7.8-16 cm longae; cryptae sto 
tophorae simplices; sclerides terminales foliorum stel- 
Eus flores axillares, solitarii, tetrameri; calyx ovarium 
inferum includens 2.5-3.4 mm longus; lobi calycis in 


—D. = section of leaf midrib. — E. Deflorescent flower and flower bu 


—A. Leaves.—B. Clea 


red portion of leaf blade 


Cross section of leaf blade showing sclereids, aba crypts, and upper 


d. —F. Fruit. — C. Seed: a, polished 


alabastro separati; petala persistentes in fructu immaturo 
thecae antherae ambae poro unico rd deliscenes 
ard nulla; ovarium 1-loculare, ovula 4 en l. 
Small tree, 3 m the only cited height, the plants 
glabrous except for the flower; young twigs round- 
ed. Petioles 1.5-4 mm long; blades 7.8-16 cm 
ng, 3.1-7 cm wide, ovate to elliptic or these 
oblong, abruptly acuminate to short caudate at 
apex, medium acute at base; midrib moderately 
grooved above, prominent and rounded below; sec- 


436 Annals of the 
Missouri Botanical Garden 


A] 


© 
da 
3 


Mouriri retentipetala. —A. Leaves dp collections). — B. Cleared portion of leaf blade showing veins 
and tenanik sclereids.— C. Cross section of leaf blade owing sclereids, stomatal crypts, and upper epidermis. — D. 
Cross section of leaf midrib. —E. Flower before anthesis. — ee at anthesis. — C. Longisection of flower before 
anthesis. — H. Petal. — I. Anther. — J. Fruit t. — K. Seed: a, ales side; b, micropyle side. (B-I, Lleras et al. P16980; 
J, K, Bernardi s.n.) 


Volume 76, Number 2 
1989 


Morley 
New World Memecyleae 


ondary and smaller nerves prominent above and 
below when dry. Midrib xylem tubular; stomatal 
crypts Type II, averaging in a leaf 31-50 um 
diam., 18-25 um high, 27-54 per mm’; superficial 
stomata numerous; upper epidermis 1 cell thick, 
nearly all the cells with thick mucilaginous walls; 
hypodermis none; terminal sclereids stellate, the 
central body sometimes isodiametric with 4—5 arms, 
more often 2-6 times as long as wide with 1-3 
large arms. Flowers tetramerous, single in the leaf 
axils on sessile pedicels 2-2.5 mm long with 1 or 
2 pairs of ovate-triangular bracts 1-1.6 mm long 
at base of pedicel. Calyx including inferior ovary 
2.5-3.4 mm long, campanulate, the surface scat- 
tered-mealy with loose inflated hairs, these white 
on the outer surface of the calyx lobes, golden on 
the inner surface; free hypanthium ca. 0.7 mm 
long; calyx lobes 0.2-0.4 mm long, 1.1-1.4 mm 
wide, broadly triangular or slightly apiculate, split- 
ting apart a distance of 0.5- m at anthesis. 
Petals protruding from the calyx early in the bud, 
narrowly ovate-elliptic, 3-3.6 mm long, 1.1-1.5 
mm wide, minutely glandular-hairy mostly on the 
apical third, persistent on the unripe fruit. Ante- 
sepalous filaments 1.2-1.4 mm long, antepetalous 
ones 1.7-1.9 mm long; anthers 1.6-2.2 mm long; 
thecae 0.9-1.2 mm long, dehiscing by a single 
apical pore; gland none; cauda 0.8-1 mm long. 
Ovary 1-locular with 4 ovules arranged around a 
short central and basal placenta; style ca. 5 mm 
long, sharply curved at tip, with a sharply demar- 
cated 4-lobed stigma. Fruits subglobose, ca. 6.5— 
7.5 mm high, 8-9 mm thick, crowned with the 
persistent calyx. Seed 1, ca. 5.5-5.8 mm high, 7- 

mm wide, 4—5 mm thick, medium brown and 
polished except for a basal hilum as wide as the 
seed, the funiculus visible as a hardened and pol- 
ished irregular stalk extending below the main pol- 
ished part of the seed. 


LORETO: Arboreti Jenaro Herrera 
vire ad flumen Uca- 


Paratype. PERU. 


(Amazoniae Peruv vianae, g 


a 
iqui 
a Ca IX. 1976. 6-R-83 (Parcela Marmillod) (G). 


Distribution. Tropical forests in the drainages 
of the Ucayali and Javari rivers in NE Peru and 
adjacent Brazil at about the latitude of Requena, 

eru. 


This species, like the foregoing one, falls in subg. 
Mouriri, its tubular midrib xylem, anatropous 
ovules, and polished seed being the determining 
features. The tetramery, monoporous and eglan- 
dular anthers, and persistent petals of this entity 


make it very distinctive. This is only the second 
species of the genus to be found with monoporous 
anthers, and persistent petals have not been noted 
before. The petals were persistent on the green but 
nearly ripe fruits of the type collection; they were 
gone from the ripe fruit of the paratype. It is 
impossible to know if they had dropped naturally 
or were broken off during collection in the latter 
case. Within its subgenus the new species is most 
similar to three species in sect. Litophyllum: M. 
exadenia, M. latihila, and M. monopora. Prob- 
ably M. retentipetala is most closely related to 
these three and belongs with them in sect. Lito- 
phyllum 

The chararieriiiks of M. retentipetala sug- 
gesting the three species named are: small simple 
stomatal crypts, a one-layered upper epidermis, 
small flowers, tetramery, monoporous eglandular 
anthers, a one-locular ovary with few ovules, and 
a large hilum on the seed. Some of these characters 
are not found in all three of the presumed allies, 
and several have not yet been investigated in M. 
latihila owing to lack of material. 

Mouriri retentipetala differs from M. exadenia 
in having higher, more numerous stomatal crypts, 
pedicellate flowers, persistent petals, monoporous 
caudate anthers, and a large hilum on the s 
From M. monopora the new species departs in its 
tetramery, triangular calyx lobes, and eglandular 
anthers with a longer cauda and straight thecae. 
From M. latihila, M. retentipetala varies in hav- 
ing stellate foliar sclereids, short calyx lobes so that 
the petals protrude from the calyx early in the 
bud, and persistent petals. 


Mouriri gleasoniana Standley ex Standley & 
Steyermark var. coibensis (Morley) Morley. 
Mouriri coibensis Morley, Fl. Neotropica 15: 
143-145. 1976 


Recent collections of M. gleasoniana from Cos- 
ta Rica and Panama require reassessment of the 
status of M. coibensis, a close relative of the former 
species from the island of Coiba off the southwest 
coast of Panama. When M. coibensis was de- 
scribed, M. gleasoniana was known only from 
Honduras to Mexico, and the differences between 
the two taxa as then known were sharp. The new 
material partly bridges the gap between them and 
seriously weakens their separation. I believe real- 
ities are best served viewing both as part of a 
single variable species. As presently known, the 
island plants are sl pasan and are re- 
tained as a variety. Six major variables are in- 
volved, as shown in Table 1. Plants of the complex 
are yet to be found in Nicaragua. 


438 Annals of the 
Missouri Botanical Garden 
TABLE 1. Geographic variation in six characters of Mouriri gleasoniana. 
Anthers 
Distance 
from 
Theca 
Base to 
Anther 
Theca Gland Base on 
Stomatal Length ^ Length Gland Side Ovule 
Foliar Sclereids Crypts (mm) (mm) mm) Numbers 
Central Panama roundish Type II 2.7-3.0 0.8-0.9 1.0-1.2 29-33 
Coiba Island roundish mostly Type II 1.9-2.2 0.2-0.5 1.2-1.4 18-20 
Northeastern Costa Rica roundish to columnar mostly Type II 2.2-2.7 0.5-0.7 0.9-1.2 23-33 
Honduras-Mexic columnar tendency mostly Type II 2.0-3.0 0.4-0.9 0.7-0.8 15-24 


Ovule numbers are relatively large in plants of 
Costa Rica and the Panama mainland. In the one 
flowering collection from Panama (Knapp & Huft 
4424) and one of the two Costa Rican collections 
(Hammel 9317), the large number of ovules ap- 
parently results in an unusual placentation in that 
the ovules in each locule are arranged on all sides 
of the axile- ine placenta. These are the only 

nown pl such an arrangement in sections 
4 dur. 9 of subg. Mouriri. 

A transition from a large number of ovules to 

a small one raises the question of which state, if 


either, represents the primitive condition. A similar 
trend was found in M. guianensis, where the con- 
clusion was that the greater number was primitive 
(Morley, 1983). In M. gleasoniana the presence 
of the larger numbers close to South America, 
where the genus appears to have originated, with 
the smaller numbers to the north, where the en- 
vironment gradually becomes more extreme, sug- 
gests that the same interpretation should be made 
in this species. Throughout the range, the fruits 
are 1-4(-5)-seeded; as in M. guianensis, many 
ovules must therefore abort when a large number 
is present before fertilization. Accordingly, those 
plants with fewer ovules in the flower and thus less 
wastage would be regarded as better adapted and, 
in this case, more specialized. It would appear that 
in increasingly extreme environments the demand 
or floral economy becomes greater. 


Mouriri longifolia (HBK) Morley. Until re- 
cently, this species was known only from fruit- 
ing material. A flowering collection has now 
been made, so that the following addendum 
to the description of the species can be given, 
with illustrations (Fig. 5). 


Inflorescences at leafless nodes of twigs 4-7 mm 


thick, 1 or 2 per side, each 1-5-flowered, 2.5- 


Orinoco, 1-2 km 


(defl.), Berr 
l 


7.7 mm long to base of farthest pedicel measuring 
along the axes and with 2 or 3 internodes in that 
length; bracts 1.4-2.5 mm long, triangular or nar- 
rowly so, mostly present at anthesis, deciduous 
later. Axes of inflorescence, ovary, and calyx mi- 
nutely puberulent. True pedicels 7-15 mm long; 
calyx including inferior ovary 6.2-8.5 mm long, 
campanulate; free hypanthium 2.2-2.7 mm long; 
calyx lobes just before anthesis 2.1-2.4 mm long, 

.o-3 mm wide, 3.8-4.1 mm long from the stamen 
attachment, triangular with inverse-curved sides, 
the calyx splitting 0-0.5 mm between the lobes at 
anthesis. Petals ovate-oblong, ca. 10 mm long in- 
cluding a basal claw ca. 3 mm long, 7 mm wide, 
broadly acute at apex, erose toward the base. Fil- 
aments 11-14 mm long; anthers 3-3.3 mm long; 
thecae 2.8-3 mm long, dehiscing by apical slits; 
gland 0.8-1 mm long, 2.1-2.3 mm from apex of 
anther when measured from center of gland; cauda 
0.6-0.8 mm long. Ovary 5-locular; placentae basal 
in each locule, the ovules borne on all sides of a 
short basal column, ca. 5 per placenta, ca. 25 in 
all; style 17-22 mm long. 

With the flowering material in hand it now be- 
comes apparent that M. longifolia belongs in sect. 
Apirangia. Specifically, the characters that place 
it there are the tubular midrib xylem, filiform scler- 
eids, Type III crypts like those of M. apiranga, 
the lack of a hypodermis, the free calyx lobes with 
the petals protruding from them early in the bud, 
the anther shape which generally agrees with that 
of the other species, the proximate position of the 
placentae which are not widely separated, and the 
position of the ovules on all sides of each placenta. 

Additional collections examined. VENEZUELA. 
AMAZONAS: Estación Experimental de Sta. Bárbara s 
al sur de Trapichote, 26 Feb.-2 
1976 (st), Berry & ao 2083 (US); 28 Mar. 1976 
& Chesney 2179 (US); Tayari, 1 Nov. 

S 


975 (fl), Lissot s.n. (US) 


— 


Volume 76, Number 2 
1989 


Morley 439 
New World Memecyleae 


D 


FIGURE 5. — Mouriri longifolia (Lissot. s.n.). 
Anther, shown cleared except for the thecae. — D. Pet 


Votomita roraimensis Morley, sp. nov. TYPE: 
Brazil. Roraima: BR 174, Manaus- Caracarái, 
km 523, 1?20'N, 60?43'W. In damp water- 
logged woodland. Straggling tree to 5 m, 2 
June 1986, Hopkins, Pereira de Lima, 
Guedes de Oliveira & Lowy 502 (holotype, 
INPA; isotypes, K, MIN, NY, US). Figure 6. 


Arbor usque 5 m alta; costa media folii subta lateribus 
papillosa; cryptae stomatophorae unaquaeque cavitatibus 

2-5; ovarium 4-loculare dais axialibus; ovula 3-4 
ai d circum Me placentam genita, in recta ver 
ticilla; fructus 6.7-9.5 mm diametro, stylo Ber 
semen l, vitta angusta e 2% circumdatum 


Tree to 5 m, glabrous except for the leaf midrib; 
young twigs terete. Petioles 1.5-2.5 mm long; 
blades 6.1-10.2 cm long, 2.3-4 cm wide, elliptic 
to elliptic-ovate, acuminate at the apex, medium 
acute at base; midrib flat to slightly rounded above, 
narrowly winged below for its length, the sides with 
papillae 20-35 um long; lateral nerves not visible 
above or below when dry. Midrib xylem tubular; 
stomatal crypts multiple, each branched above the 
mouth into 2-5 small cavities, with prominent pa- 
pillae on the central roof, the multiples 60-125 
um total diam., 55-75 um high, averaging ca. 28 
per mm?; upper epidermis mostly 1(2) cell(s) thick, 
a deep hypodermis present with walls tending to 
align with the epidermal cells above, the hypoderm 
cells commonly with mucilaginous walls; foliar 
sclereids terminal, thick-columnar with large den- 
droid branches at each end. Peduncles terminal 
and at leafless nodes of twigs up to 5 mm thick, 


— A. Inflorescence. — B. Longisection of flower before anthesis. — C. 
al. 


1-2 per side, 1-flowered, 2-11 mm long to base 
of farthest pedicel including 1 or 2 internodes; 
bracts narrowly triangular, 0.7 mm long, deciduous 
before anthesis. True pedicels 3-5 mm long; flow- 
ers 4-merous; calyx including inferior ovary ca. 
4.5 mm long, campanulate; calyx lobes 0.4 mm 
long, 1.4 mm wide, rounded-triangular and mu- 
cronulate, stretching and splitting irregularly be- 
tween lobes as bud enlarges. Immature petals ovate, 
sessile, 4.7 mm long, 2 mm wide. Stamens lightly 
connate in the lower third, 2.9-4.2 mm long; fil- 
aments 0.3-0.4 mm long; anthers 2.6-2.8 mm 
long; thecae 1.2-1.3 mm long; gland terminal, 
concave, 0.6-0.7 mm long, Ovary 4-locular with 
axile placentation, each locule with 3-4 ovules in 
a vertical whorl, the ovules 15-16 in all; style 6 
mm long. Fruit medium brown when dry, spheroid 
with persistent style, 7-9.3 mm high excluding 
7.7-9.4 mm wide, 6.7-8.9 mm 
thick. Seed 1, spheroid, light brown with dark 
brown markings, 5.9-6.8 mm high, 7.4-8.2 mm 
wide, 7-7.9 mm thick, with a narrow, smooth but 
unpolished tan band encircling 25 of the seed, ex- 
tending from a small projection near the broken 
chalazal vascular strand where the band is narrow- 
est to the micropylar end where it is widest, the 
width here 2.7-3 mm; remainder of surface ru- 
gose. 


style and calyx, 


Distribution. Known only from the type lo- 
cality in tropical forest in SW central Roraima, 
Brazil, east of the Rio Branco. 


440 Annals of th 
Missouri Botanical Garden 
A B 
90 
m 300 
| , O (9 
FIGURE 6. Votom —A. Leaves. —B. Cleared portion of leaf blade showing 
veins and terminal s iere a. ss section ‘of leaf blade showing sclereido, dies aa and upper epidermis. 


— D. Cross o of leaf midri ib. — E. Flower before anthesis. —F. Longisect 
shown cleared.— I. Cluster of ovules in one locule as seen from abaxial side. - D 
abaxial view; b, view from top; c, underside. 


n of flower. — GC. Petal.—H. Anther, 


a ok Fruit.—L. Seed: a, 


Volume 76, Number 2 


Morley 
New World Memecyleae 


This species is identified more by its combination 
of features than by any unusual characters. It is 
allied to the other four species that have four locules 
or placentas, numerous ovules, and (with one ex- 
ception) hairs on the midrib. Its laterally fused 
stamens occur in two members of this group of 
four, V. plerocarpa and V. monadelpha. From all 
of these the new species differs in having much 
smaller leaves and flowers and only 15-16 ovules. 
Three of the other four, V. monadelpha, V. or- 
binaxia, and V. plerocarpa, have 36-48 ovules. 
The fourth, V. pubescens, is closest in ovule num- 
ber with 20-26, but the new species differs greatly 
from that species by having small nearly glabrous 
leaves, a nonindurated ovary wall, laterally fused 
stamens, four locules, and a persistent style. The 
remaining three species in the genus, V. guianen- 
sis, V. orinocensis, and V. monantha, have one- 
locular ovaries with 8-10 ovules and glabrous mid- 
ribs. Because of its small leaves, small flowers, and 
geographic proximity, a close relation of V. rorai- 
mensis to V. orinocensis and the related Cuban 
species V. monantha seems likely, although the 
floral structure mostly does not sustain this view. 
The small leaves, deep hypoderm, and persistent 
style of the new species in particular suggest NV. 
monantha, although the hypoderm of the former 
differs in having abundant mucilage walls. 

Ovules in Votomita and Mouriri vary from a 
campylotropous type in which the ovule is bent at 
a sharp angle from a short, broad placental at- 
tachment at the chalazal end to half-anatropous, 
and fully anatropous forms occur in Mouriri. In 
V. pubescens and V. roraimensis, the ovules are 
half-anatropous and are borne on an axile placenta, 
the only known examples of such a combination 
in the two genera. In Mouriri, half-anatropous 
ovules occur only on axile-basal, free central, or 

asal placentas. In V. pubescens, the ovules are 
sessile laterally with extruded inner integuments 
that appear to serve as internal stigmas (Fig. 7D). 

Seeds are known in three species of Votomita: 
V. monantha, V. pubescens, and V. roraimensis. 
In V. monantha, which has free-central placen- 
tation, the angular distance on the seed between 
the micropyle and the broken chalazal strand is 
about 94°, while in V. roraimensis and V. pubes 
cens, with axile placentation, the angles are 60? 
and 50? respectively. 

In V. monantha the whole abaxial (outer) sur- 
face of the seed is smooth, whereas in V. rorai- 
mensis and V. pubescens the smooth part is narrow 
and is broadest at or near the micropylar end (Figs. 
6L, 7F). In none of these seeds is the smooth part 
polished. Seeds of the latter two species resemble 


superficially seeds found in Mouriri sect. Brevi- 
pedillus; however, in that section the smooth area 
is narrowest at the micropylar end and is usually 
conspicuously widened at the opposite end nearest 
the broken chalazal strand. I have no explanation 
for the difference in shape. 

The reason for the smooth area is unknown; it 
may be that this is the part of the ovule first pushed 
against the ovary wall as enlargement begins. 


THE GENUS QUESTION 


Partial generic differences between Votomita 
and Mouriri in ovule and seed structure are sug- 
gested by the above and may be summarized thusly: 
(1) half-anatropous ovules on an axile placenta 
occur only in some species of Votomita; (2) the 
smooth area of the seed is unpolished in Votomita 
but is polished in almost all species of Mouriri; (3) 
in no known species of Votomita does the smooth 
area cover all of the seed except for the hilum, as 
it does in seeds of the subgenus Mouriri; and (4) 
seeds of Votomita most closely resembling those 
of Mouriri are found only in sections Acutiflos and 
Brevipedillus of Mouriri subg. Taphroxylon. Seeds 
of V. monantha somewhat resemble those of sect. 
Acutiflos but lack the polished face and rugose 
surrounding area of the seeds of that section. Seeds 
of V. pubescens and V. roraimensis have narrow 
smooth areas with surrounding rugose surfaces as 
in Mouriri sect. Brevipedillus, but the smooth 
parts differ in shape as described above. 

e above considerations may affect present 
generic divisions. The seed structure in V. rorai- 
mensis and V. pubescens maintains the difference 
between Votomita and Mouriri subg. Mouriri but 
increases the resemblance between Votomita an 
Mouriri subg. Taphroxylon. 

It may be that the New World Memecyleae 
should be regarded as consisting of three genera 
or subgenera: Votomita, Mouriri, and Taphrox- 
ylon. Absolute differences exist between all three 
groups, the most between Votomita and subg. 
Taphroxylon, the fewest between subgenera Ta- 
phroxylon and Mouriri, thus supporting the pres- 
ent disposition of these groups. Between Votomita 
and subg. Taphroxylon the differences are those 
of midrib xylem form, flower mery, hypanthium 
development with accompanying features of sta- 
men form, and the presence or absence of free 
stone cells in the petals. Between Votomita and 
subg. Mouriri the differences are the presence or 
absence of free stone cells in the petals, and seed 
structure; between subgenera Taphroxylon and 
Mouriri the single absolute difference is that of 


442 Annals of the 
Missouri Botanical Garden 
15 
FIGURE 7.  Votomita pubescens. —A. Longisection of flower (ovules copied from Morley, 1985, Lao 22).— 


Petal. — C. Anther 


r, shown cleared (Bernardi s.n., tree 5-145). 


— D. Ovule (Lao 22). — E. Immature fruit. — F. E 
168.) 


a, lengthwise view showing smooth face; b, underside. (A, B, E, F, Bernardi s.n., tree 2/ 68. 


seed structure. On this basis Votomita is overall 
more like subg. Mouriri than subg. Taphroxylon, 
even though seed structure in Votomita is more 
like that of the latter. 

Because the above differences do not negate the 
present arrangement of genera, I think any serious 
effort to reconsider the generic divisions must await 
more evidence. Seeds of more species of Votomita 
should be examined, and detailed comparative stud- 
ies of wood anatomy, karyotypes, and phytochem- 
istry may be expected to aid in the evaluation. 
When the similarities and dissimilarities between 
the three are well known it can then be decided 
how many of them should be assigned generic 
status. 


Votomita pubescens Morley. Several more col- 
lections of this species have been made from 
the same area as the type. Among these are 
specimens with petals, stamens, fruit, and 
seed —all lacking from the type. An extended 
and illustrated (Fig. 7) description of the species 
is therefore presented, omitting the anatomical 
features, which are not modified. 


Tree to 16 m high. Petiole 4-6.5 mm long; leaf 
blades 10.5-15.5 cm long, 3.6-6.5 cm wide, el- 
liptic or slightly elliptic-ovate, abruptly caudate at 
the apex with a cauda 11-13 mm long, medium 
acute at base, glabrous above, pubescent beneath; 
midrib nearly plane above but with a slight groove 
down the center, prominent below, the underside 
rounded at the petiole but abaxially soon 2-angled 


with 2 very narrow wings for its length, pubescent 
on the sides; lateral nerves not visible above or 
below when dry. Peduncles axillary, 1 or 2 per 
side, each 2-9-flowered, 19-53 mm long to base 
of farthest pedicel measured along the axes and 
with 1-3 internodes in that length; bracts un- 
known, deciduous before anthesis. True pedicels 
8-11 mm long; flowers 4-merous, white or yellow; 
calyx including inferior ovary 10.3-13.3 mm long, 
7.8-9 mm thick, glabrous, shrinking only slightly 
on drying, when dry with a hard, smooth, somewhat 
polished olive-green to brown outer layer, the ovary 
widest 14-23 of the way above the base; calyx lobes 
separate, 0.9-2 mm long, 2.4-2.7 mm wide, tri- 
angular and slightly apiculate, not separating fur- 
ther at anthesis. Immature petals oblong-ovate, 6.5 
mm long, 3.5-3.7 mm wide. Stamens 4—5.5 mm 
long; filaments 1.2-1.5 mm long; anthers coherent, 
apparently not truly fused, 3.6-4.5 mm long; the- 
cae 2-2.5 mm long; gland terminal, concave, 0.8- 
1.2 mm long. Ovary 2-locular, each locule with 2 
axile placentas and an irregular very incomplete 
ridge running down the outer wall representing a 
missing partition; ovules sessile with an extruded 
inner integument, 5-7 per placenta, 20-26 in all; 
style ca. 5 mm long. Fruit maturing red, ellipsoid, 
ca. 20-24 mm long, 14-18 mm wide. Seed 1, 
light brown, ovoid-ellipsoid, 12-15 mm high, 8- 
10 mm wide, 7.5-8 mm thick, with a narrowly 
ovate, slightly elevated, rather smooth, slightly peb- 
bled, unpolished face (the enlarged outer face of 
the ovule) extending 54 of the length of the seed 


Volume 76, Number 2 
1989 


Morley 443 
New World Memecyleae 


from the base, the widest end of the face down and 
turned under opposite the broken chalazal strand, 
the micropyle in the underturned end; remainder 
of seed surface rugose. 

Vernacular name. Lanza caspi. 

Additional collections examined. PERU. LORETO: Prov. 
Requena, Ucayali River, Arboreti Jenaro Herrera, Ber- 
nardi L. 15608 (G), s.n. trees 2/39, 2/168, 5/145 (G); 
Diaz M. 59.A, 76-A (G); slightly disturbed moist forest 

W of Arboretum Jenaro Herrera, D. C. Daly et al. 
5641 (MIN, NY); Jenaro Herrera, 73?45'W, 4?50'S, ca. 


O m, bosque primario, R. Vasquez, K. Young & N. 
Jaramillo 2057 (US). 


LITERATURE CITED 


MonLEY, T. 1976. Memecyleae (Melastomataceae). Fl. 
Neo ut 15: 1-295. 

——. 1983. A directional cline in Mouriri gui- 
anensis (Melastomataceae). Acta Amazonica 13: 
799-811. 

Five new taxa of New World Meme- 

cyleae (Melastomataceae). Ann. Missouri Bot. Gard. 

72: 548-557. 


STUDY OF THE 
FLORA OF RUCAMANQUE, 
CAUTIN PROVINCE, CHILE: 


Carlos Ramirez,’ Enrique Hauenstein,’ 


José San Martín* and Domingo Contreras? 


ABSTRACT 


The flora of forest, scrub, and grass communities was studied in Rucamanque, located near the city of Temuco, 


Cautin Chile, in the central valley of south central Chile. Forty-six vegetation samples were made, an 


collected intensively outside the sample areas. A 


importance of each in the different vegetational communities, and their uses in industry, crafts, and medicine 


ucamanque, ““The House of the Condor" in 
Mapudungun, the language of the indigenous peo- 
ple of this region (Ramirez-Sanchez, 1985), is a 
valley located 12 km northwest of the city of Te- 
muco, capital of the IX Region of the Araucania, 
Chile (Fig. 1). The primitive forest vegetation of 
this valley escaped destruction during last century's 
process of colonization of central southern Chile 
(Donoso, 1983) because it was protected as an 
early watershed area for the drinking water for 
Temuco (Magofke, 1985). 

With the increase in population, water for the 
city had to be drawn from other sources, principally 
subterranean, so the state lost its interest in Ru- 
camanque and offered it for sale in March of 1986. 
Because of its value as a forest relict and its sci- 
entific interest, several state and private institutions 
and ecological organizations intervened to preserve 
it from exploitation. Finally, the University of the 
Frontera in Tem ie the legal owner, saving 
the relict forest for science, education, and the 


general public. 

The present catalog of the flora of Rucamanque 
was made during the time that the land was for 
sale. 


STUDY AREA 


camanque is a small river valley oriented 
southeast-northwest to the south of the Ñielol- 
Huimpil hills (Magofke et al., 1986). Through it 
runs Chivilcán Creek, which receives the water of 
two small unnamed streamlets. Its elevation ranges 


from 300 to 530 m. The area occupies about 525 
ha, including some of the adjacent forests that 
belong to private owners. The basin and slopes of 
Rucamanque Valley are covered by native forest, 
which is evergreen in the lower areas and partially 
deciduous at higher elevations. In several places 
the forest has been destroyed and secondary scrub 
replaces it. In the higher slopes, originally of de- 
ciduous forest, permanent grasslands of human 
origin can be found. Small bogs, which we include 
in the grassland formation, appear in open areas 
on the edges of streams at the bottom of the valley. 

The climate of the area is humid and temperate 
with a Mediterranean influence (Di Castri & Hajek, 
1976). The average annual rainfall is 1,400 mm 
and the median annual temperature is 12°C. Rain- 
fall is abundant in winter, and summers often have 
one to two dry months, as shown in the climatic 
diagram in Figure 1. Winter and spring frosts are 
frequent. To this climate corresponds a subtropical 
semideciduous forest type as natural vegetation 
(Schmithüsen, 1956). 

The Nielol- Huimpil hills comprise a tectonic unit 
characteristic of the central valley of Chile and do 
not connect with either the Coastal Ranges or with 
the Andes. The red clay soil does show similarity 
to the Coastal Range soils (Weinberger & Binsack, 
1970). When cultivated or grazed, this soil is sub- 
ject to erosion. 


METHODS 


The catalog of plant species of Rucamanque was 
made from 46 vegetation samples taken using the 


E 


l "x Ae by Grant RS-84-22 from the Research Fund of Universidad Austral de Chile, Valdivia, and the Projecto 
88. 7 of the research fund of the Universidad Católica de Chile, Sede Te 


emuco, 


* Instituto de Botánica, Universidad Austral de Chile, Casilla 567, Valdivi via, Chile. 


* Departamento de Ciencias Nat 


urales, Universidad tollen de Chile, Casilla 


15-D, Temuco, Chile. 


* Area de Ciencia, Universidad Católica de Chile, Casilla 617, Talco, Chile. 
ANN. Missouni Bor. Garb. 76: 444-453. 1989. 


Volume 76, Number 2 
1989 


Ramírez et al. 
Flora of Rucamanque 


b 
TEMUCO (114m) 
oc 33 y 12° 

a 


FIGURE 1. 
of observation, b = altitude, c = year average temperature, d = year average precipitation, p = precipitation, t = 
temperature. Black areas = surplus of precipitation. 


Braun-Blanquet (1964) phytosociological method 
for the study of the local vegetation. Other plants, 
collected outside the study quadrats, were added 
to the collection. Eighteen samples were taken in 
the forest, 20 in scrub vegetation, 6 in pastures, 
and 2 in bogs. The vegetation contains the following 
associations: Lapagerio—Aextoxiconetum punc- 
tatii, Nothofago—Perseetum linguae, Fuchsio— 
Chusqueetum quilae, Rhaphithamno- Aristote- 
lietum chilensis, Chusqueetum coleu, Aristotelio— 
Rubetum  constrictae, Hyperico-Agrostidetum 
castellanae, and Juncetum procerii (Oberdorfer, 
1960; Ramirez, 1982; Hilldebrand, 1983). 

The frequency of each species was estimated in 
each plant association. The relative importance 
value was calculated adding this frequency and the 
average cover expressed in relative percentage of 
each species in the different associations according 
to the Wikum & Shanholtzer (1978) method. This 
value was not determined for species collected out- 
side the quadrats. 

Nomenclature and phytogeographic origin of the 
taxa were based on Marticorena & Quezada (1985) 
and the family nomenclature on Cronquist (1981). 
The common names are according to Munoz (1966). 
Introduced species were classified using the Ger- 
man flora by Rauh & Senghas (1968). The col- 
lected species were kept in the herbarium of the 
Institute of Botany, Austral University of Chile 
(VALD). 


Ninth Region of Chile. Climate diagram of Temuco according to Hajek & Di Castri (1975): a = years 


Life forms follow the Raunkiaer system (Ellen- 
berg & Mueller-Dombois, 1966). Vines, epiphytes, 
and parasites were included under phanerophytes. 
The interpretation of the biological spectrum was 
based on Cain (1950) 

Information was added on the present and po- 
tential uses of the plants. 


RESULTS AND DISCUSSION 


The flora of the study area is made up of 203 
species, of which 190 were inventoried in the 46 
vegetation samples taken in the forest, scrub, and 
grassland formations (Appendix I). The other 13 
species were collected outside the sample quadrats. 
The 203 species are distributed among 161 genera, 
84 families, and 6 classes (Table 1). The class with 
the greatest number of taxa was Magnoliopsida 
(dicotyledons), which include almost 69% of the 
species present in Rucamanque. Following these 
are the Liliopsida (monocotyledons) with 42 species 
(21%), and the Polypodiopsida (Filicinae) with 16 
species (8%). 

The best-represented family of the Magnoliop- 
sida was the Asteraceae, with 18 genera and 25 
species, amounting to 12% of the total. Next came 
the Fabaceae with 4 genera and 9 species, the 
Apiaceae with 6 genera and species, the Lamiaceae 
and Rosaceae with 5 species each, and the Myr- 
taceae, Proteaceae, and Scrophulariaceae with 4 


446 Annals of the 
Missouri Botanical Garden 
TABLE l. T ic distribution of the flora of Ruca- TABLE 2. Phytogeographic origin of the flora of Ruca- 
manque. manque, distributed by class. Percentages in parentheses. 
Fami Classes Native Introduced Total 
Classes lies Genera Species (%) pila 3 0 3 
Bryopsida 3 3 3 (1.47) Equisetopsida l 0 l 
Equisetopsida l l 1 (0.49) Polypodiopsida 16 0 16 
Polypodiopsida ri 9 16 (7.88) Pinopsida 1 0 1 
Pinopsida l l . Magnoliopsida 96 (68.6) 44 (31.4) 140 
Magnoliopsida 60 115 140 (68.96) Liliopsida 30 (71.4) 12 (28.6) 42 
Liliopsida 12 32 42 (20-68) Total 147 (72.41) 56 (27.58) 203 


species each. Outstanding among the Liliopsida are 
the Poaceae, with 16 genera and 18 species, the 
Cyperaceae with 5, and the Juncaceae with 4 
species. Among the ferns, the Hymenophyllaceae 
presented 6 species, all of which were epiphytes 
in the forest formation. 


ORIGIN 


Of the 203 species, 147 (72.41%) are native 
and 56 (27.58%) are introduced (Table 2). This 
high percentage of introduced weeds indicates a 
strong human influence on the vegetation (Sukopp, 
1969). The introduced species only belong to the 
classes Magnoliopsida and Liliopsida, and make up 
31.4% and 28.6% of each group, respectively. Of 
course the greatest percentage of introduced species 
is found in the shrub grassland formations, of an- 
thropic and therefore secondary, origin (Ramirez, 
1982). The evergreen forest of Aextoxicon punc- 
tatum is the association least invaded by weeds. 
The majority of the introduced species are origi- 
nally European and of little grazing value (Mon- 
taldo, 1975; Anazco et al., 1981). 


LIFE FORMS 


The biological spectrum of the study area was 
constructed from 200 species; the remaining three 
were mosses. In the epiphytic synusia that were 
not considered, mosses are very abundant (Riveros 
& Ramirez, 1978). The most-represented life form 
was the phanerophytes with 80 species, repre- 
senting 40% of the total. This life form abounds 
in forest and scrub formations, occupying different 
strata in the vegetation. Chamaephytes and cryp- 
tophytes were the scarcest, with eight species (4%) 
each. Hemicryptophytes appeared in about the same 
numbers as phanerophytes, with 78 species, or 
39%. Therophytes were represented by 26 species, 
or 13%. The latter two herbaceous life forms were 
abundant in grassy and scrub formations (Table 3 

The biological spectrum here described (Fig. 2) 
corresponds to a phanerophytic phytoclimate, typ- 


x 


ical of intermediate latitudes in temperate zones 
(Cain, 1950). The relative abundance of hemi- 
cryptophytes and therophytes is related to the de- 
gree of anthropic intervention suffered by the orig- 
inal vegetation. In the original vegetation, these 
ife forms certainly were not as important. 

This is confirmed by separating the native ele- 
ments from the introduced for each life form (Table 
4, Fig. 3). Only one of 80 phanerophytes is exotic. 
The cryptophytes are all native. Among the cha- 
maephytes and hemicryptophytes, the percentage 
of introduced species is much higher, reaching 
37.5% and 38.5%, respectively. Cryptophytes are 
not very frequent, unlike the hemicryptophytes, 
which contribute 30 exotic species. These perennial 
cespitose herbs and rosettes are plants that accom- 
pany man and do not characterize any particular 
phytoclimate (Le Blanc, 1963). Their abundance 
is due to destruction of forest and to grazing, which 
impedes their replacement by more aggressive 
woody species (Pessot & Montaldo, 1974). 

The therophytes constitute a special case of 
plants typical of dry climate or weeds of cultivated 
areas. They act as pioneers in uncovered soil (Cain, 
1950). Only 15% of the therophytes present in 
the study area are native. This life form was scarce 
in the original vegetation but invaded anthropo- 
genic grasslands formed on eroded soils, where the 
vegetational cover never reaches 100%. Thus, 
therophytes are not typical of Rucamanque. 

The phanerophytes, therefore, make up the most 


TABLE 3. Biological spectrum of the flora of Ruca- 
manque. 

Life Forms Species Percentage 
Phanerophytes 80 40% 
Chamaephytes 8 4% 
Hemicryptophytes 78 39% 
Cryptophytes 8 4% 
Therophytes 26 13% 

Total 200 100% 


Volume 76, Number 2 
1989 


Ramírez et al. 
Flora of Rucamanque 


e 


FIGURE 2. — a. Biological spectrum of Rucamanque. F = phanerophytes, C = chaemaephytes, H = hemicryptophytes, 


Cr = cryptophytes, m ther 
epiphytes, P — paras 


abundant and representative life form of the re- 
search area, including trees, shrubs, epiphytes, and 
Es (Table 5). Shrubs were very abundant, 

28 species, 35% of all the phanerophytes. 
This life form dominates the secondary scrub and 
tends to invade degraded and abandoned pastures 
or grassy areas (Ramirez et al., 1984a, 1985). 
Twenty-two species of trees were present, or 27% 
of the total. 

Sixteen climbing plants flourish in Rucamanque, 
making up 20% of the total. The most robust of 
these, Hydrangea serratifolia, can have a stem 
diameter of up to 15 cm. Climbers are typical of 
forest and proliferate in secondary scrub where 


dE I 


Cc H Cr T 
Native vs. introduced 1 aperies in the bio- 


y ypto 
s = therophytes. White entage of 
native Le Hatched bars — percentage of la 


Spec 


rophytes. —b. Distribution of phanerophytes: 


S — shrubs, V — vines, E = 


T = trees, 


more light penetrates. The epiphytes, principally 
filmy ferns of the Hymenophyllaceae, presented 
11 species, or 13.5%. Only three parasitic species 
were present. 


FREQUENCY IN THE PLANT ASSOCIATION 


Only two species were present in each of the 
eight studied plant associations: Rubus constrictus 
and Prunella vulgaris (Table 6). These are Eu- 
ropean weeds, the first a spiny, semiclimbing shrub, 
and the second, a chamaephytic herb. Both have 
wide ecological ranges, colonizing any disturbed 
site in central-southern Chile. Another European 
species, Holcus lanatus, was found in seven of the 
eight associations studied. In six associations, Plan- 
tago lanceolata, another European herb, was found 
along with Aristotelia chilensis and Eucryphia 
cordifolia, two native woody species. 

Native species with a frequency of five, that is, 
which occur in five associations, are: Aextoxicon 
punctatum, Luma apiculata, Baccharis race- 
mosa, Lapageria rosea, Cissus striata, Boquila 
trifoliolata, and Blechnum hastatum. The first 
two are arboreal species of forests and scrubland; 
the third is a small shrub fairly frequent in the 
south of Chile. The remaining three are vines that 


TABLE 4.  Phytogeographic origin of the plant species 
of Rucamanque distributed by life form. Percentages in 
parentheses. 


Life Forms Native Introduced Total 
Phanerophytes 79 (98.75) 1 (1.25) 80 
Chamaephytes 5 (62.50) 3 (37.50) 8 
Hemicryptophytes 48 (61.53) 30 (38.47) 78 
Cryptophytes 8 (100) 0 (0) 8 
Therophytes 4(15.38) 22(84.62) 26 


448 


Annals of the 
Missouri Botanical Garden 


Hahit T 1 po. 


L p phytes p the flora TABLE 6. Frequencies in the plant associations, and 
of Rucamanque. importance values of the main species of Rucamanque. 
Habits of predi of cc edd 
Phanerophytes Speci Percentage Species Frequencies Value 
Trees 22 27.50 Chusquea quila 4 11.69 
Shrubs 28 35.00 Rubus constrictus 8 9.33 
Vines 16 20.00 Aextoxicon punctatum 5 7.92 
Epiphytes 11 13.75 Aristotelia chilensis 6 6.96 
Parasites 3 3.75 Nothofagus obliqua 4 6.66 
Tota] 80 100.00 Agrostis castellana 5 5.61 
Chusquea coleu 2 5.03 
Juncus procerus l 2.93 
form large populations in secondary brush, and the Blechnum chilense l 4 2.81 
last is a fern tolerant of a wide range of luminosity, Eucryphia cor difolia 6 2.46 
as Godoy et al. (1981) demonstrated. The exotic Prunella vulgaris a 2.39 
. : Holcus lanatus 7 2.30 
hemicryptophyte herbs having the same frequency 
Persea lingue 4 2.25 
) are Agrostis castellana and Rumex acetosella. m igi; 
i e rifolium repens 4 2.17 
The former dominates the grassy vegetation in the Senecio yegua 4 2.00 
study area (Oberdorfer, 1960) and the latter thrives Plantago lanceolata 6 1.95 
on worn-out soils. Leontodon taraxacoides 2 1.95 
Peumus boldus 3 1.91 
IMPORTANCE VALUES A us echinatus : x 
accharis racemosa E 
Chusquea quila, a climbing bamboo, was found Conium maculatum 2 1.88 
to be the most important species in the vegetation — Lapageria rosea 5 1,19 
of Rucamanque (Table 6). It grows in areas of Vulpia bromoides 3 1.77 
moist soil (Ramírez et al., 1984b) and prefers Baccharis concava 3 1.74 
light in forest clearings. It forms extensive and Cissus striata 7 ds 
PM Leucanthemum vulgare 3 1.62 
mpenetrable secondary scrub, locally called **qui- 
| Following t ith dob» leai Blechnum hastatum 5 1.49 
antales ollowing 1t with slight y ess Hopertanee Luma apiculata 5 1.49 
is the blackberry (Rubus constrictus), which col- Boquila trifoliolata 5 1.49 
onizes all of the vegetational communities in Ru- Rumex acetosella 5 1.49 


camanque. This blackberry forms scrubby areas 
called **murrales" when it invades degraded mead- 
ows and hedges (Hildebrand, 1983). It is an ag- 
gressive, woody plant whose control poses a serious 
problem in the study region (Ramirez, 1973). The 
importance of these two shrubs, which belong to 
disclimax communities, indicates the degree of an- 
thropic intervention found in the vegetation of Ru- 
camanque. They invade areas when the forest has 

een cut, and in this case make difficult the re- 
generation of the original forest community (Ra- 
mirez et al., 

Third in order of importance is “olivillo”” (4ex- 
toxicon punctatum), a tall tree that forms ever- 
green forests in the lowest and deepest parts of 
Rucamanque Valley. Aristotelia chilensis and 
Nothofagus obliqua show similar importance val- 
ues. The former dominates in shrub communities 
and the latter in partially deciduous forests of the 
Nothofago—Perseetum. The next-lowest values 
were found for Agrostis Bases, Chusquea co- 
leu, and Juncus procerus, these 
areas, scrub, and bogs, A aS 


tin grassy 


APPLICATIONS 


The flora of Rucamanque offers many uses, some 
of which are employed by the Araucarian Indians 
(Mapuche) who populate Cautin Province. Chus- 
quea coleu and C. quila are used in crafts and 
for the construction of furniture and household 
articles. To fasten pieces of these bamboos, “vo- 
quis” (Boquila trifoliolata and Campsidium val- 
divianum) serve as ties. Juncus procerus is used 
to make mats. From Luma apiculata, tool handles 
are cut, and the wood of Embothrium coccineum 
is carved. 

Applications as construction materials are im- 


mannia trichosperma. Tannins for the leather tan- 
ning industry are obtained from the barks of Persea 
lingue and Weinmannia trichosperma. Many in- 
dividuals of the latter species can presently be 


Volume 76, Number 2 
1989 


Ramírez et al. 449 


Flora of Rucamanque 


eyreceyve 


found with the bark cut off, resulting 
exploitation. Eucryphia cordifolia is valued for 
domestic firewood, and for this reason the tree is 
being cut so extensively that it risks local extir- 
pation. Edible fruits grow on Lapageria rosea, 
Chile's national flower. “Maqui cider" is made from 
the berries of Aristotelia chilensis. Edible fruits 
also grow on Gevuina avellana, Ribes trilobum, 
Fragaria chiloensis, Ugni molinae, Fuchsia ma- 
gellanica, and Peumus boldus. The fruits of Ru- 
bus constrictus are used to make ja 

The bark of Dasyphyllum e al- 

legedly regulates blood sugar; Boquila trifoliolata 
is applied for eye infections; Peumus boldus is 
used for stomachaches and liver illnesses; an in- 
fusion of Tristerix tetrandrus is ipe di to reduce 
cholesterol in the blood. Solanum gay a 
reputed febrifuge; Buddleja Borsa allegedly | helps 

eal ulcers; and Equisetum bogotense is used as 
a diuretic. The following introduced plants have 
applications in popular medicine: Urtica urens as 
an antirheumatic, Hypericum perforatum as a vul- 
nerary, and Mentha rotundifolia and M. pule- 
gium for the stomach. 

The rhizomes of Dioscorea brachybotrya are 

edible and high in starch. Maytenus boaria and 
Chusquea quila provide excellent winter forage. 


LITERATURE CITED 
ANazco, N., M. Moraca & C. Ramirez. 1981. Dis- 


tribucion de comunidades pratenses antropogénicas 
en un gradiente de inclinación en Valdivia, Chile. 
Agro Sur 9: 14-27. 
BRAUN-BLANQUET, J. . Pflanzensoziologie — Grund- 
er Vegetationskunde, 3rd edition. Springer, 


; . Life forms and phytoclimate. Bot Rev. 
(Lancaster) 16: 1-31. 
e A. 1981. An Integrated System of Classi- 
n of Flowering Plants. Columbia Univ. Press, 
New ork. 
Di Castri, F. & E. Hajek. 1976. Bioclimatologia de 
Chile. Universidad MES de Chile, Vice-Rectoria 
Académica, Santiag 
Donoso, C. 1983 Modificaciones del paisaje forestal 
chileno a lo largo de la historia. Versiones Abreviadas 
I encuentro EE sobre el medio ambiente chi- 
leno. Talca 1 9-113. 
Eus H. & p MUELLER-DOMBOIS. oon A key 
jog mu: plant life forms with revised subdivi- 
Ber. Geobot. Inst. ETH Stiftung Rübel 37: 
56-73. 
Gopoy, R., C. Ramírez, H. FIGUEROA & E. HAUENSTEIN. 
1981. Estudios ecosociológicos en pteridófitos de 
comunidades boscosas Valdivianas, Chile. Bosque 4: 


12-24. 
Hajek, E. € F. Di Castri. 1975. Bioclimatografia de 
Chile. Universidad ^ ai de Chile, Vice-Rectoria 


Académica, Santia 


HILDEBRAND, R. Die Vegetation der Tieflandsge- 

us sche des südchilenischen Loorbeerwaldgebiets un- 
r besonderer Berücksichtigung der Neophyten- 
iid ACH Phytocoenologia 11: 145-223. 

LE BLanc, F. 1963. The life forms of the flora of Mount 
Yamaska, pri County, Quebec. Canad. J. Bot. 
41: 1425-14 

MAGOFKE, J. 1985.  Rucamanque: un relicto de bosque 

nativo en Temuco, Chile. Rev. Universidad de la 
pit i d 

o & H. Lop 1986. Rucamanque: 
un dieto de bosque Valdiviano tipo costero. Ver- 
siones abreviadas II encuentro científico sobre el 

edio ambiente chileno. Talca 1: -187. 

MARTICORENA, C. & 1985. Catálogo de 

. Gayana, Bot. 42: 5-157. 

MoNTALDO, P. 1975. Soest de las praderas an- 
Hg per en la provincia e Valdivia, Chile. Agro 
Sur 3: 16-24. 

Muñoz, C. 1966. Sinopsis de la Flora Chilena, 2* Edi- 
ción. Universidad de Chile, Santiago. 

OBERDORFER, E. 1960. Pflanzensoziologische Studien in 
Chile— ein Vergleich mit Europa. Flora et Vegetatio 
Mundi 2: 1-208. 

Pessor, R. & P. MONTALDO. 1974. Cambios sinecológi- 
cos en una pradera permanente bajo la influencia de 
talajeo, fertilización, quema y competencia interes- 
pecifica. Turrialba 24: 265-273. 

RAMÍREZ, C. 73. Germinación, crecimiento juvenil y 
relaciones de competencia de Rubus constrictus Lei 
et M. y Ulex europaeus L. Rev. Agric. Técnica 33: 


90-93. 


. 1982. La vegetación nativa del Sur de Chile. 
Creces 3: e 5. 
, M. Moraca & H. FIGUEROA. 


1984a. La simili- 


del bosque Med 

. COLIQUEO, H. Fic 

1985. Estudio Sastry sche estadistico de las pra- 

deras SL PP de la cordillera pelada, Chile. 

Agro Sur 13: 114-130. 

, H. da TN R. CARRILLO & D. CONTRERAS. 
1984b. E 

res en un bosque de pino (Valdivia, Chile). Bosque 


Raisins: SÁNCHEZ, C. 1985. Voces Mapuches. Uni- 

versidad Austral de Chile, Valdivia. 

Raun, W. & K. SencHas. 1968. Flora von Deutsch- 
land, 81. Edic. Quelle & Meyer, Heidelberg. 
RIVEROS, M. & C. RAMÍREZ. 1978. Fitocenosis epífitas 
de la asociación Lapagerio-Aextoxiconetum en el 
fundo San Martín (Valdivia, Chile). Acta Ci. Venez. 

29: 163-169 

ScHMITHÜSEN, J. 19 Die ráumliche Ordnung a 
chilenischen Vegetation. Bonner Geogr. Abh. 17: 

SUKOPP, H. 1969. Der Einfluss des oo auf die 

-37 

R. Binsack. 1970. a Entstehung 

und Verbreitung der Aschenbóden in Südchile. Der 
Tropenlandwirt 71: 19-31. 

Wikum, D. € F. SHANHOLTZER. 1978. Application of 
the Braun-Blanquet cover-abundance scale for vege- 
tation analysis in land development studies. Environ. 
Managem. 2: 323-320. 


450 Annals of th 
Missouri Sener Garden 
APPENDIX I. Scientific names, families, local names, life forms (Lf), origins (O), frequencies (F), and importance 


values (I.V.) of the flora of Rucamanque (Cautín, Chile). 


Scientific Name Family Local Name If O F LV. 
BRYOPHYTAE 
Dendrolygotrichum dendroides (Hedw.) Brot. Polytrichaceae Musgo pinito n 1 0,29 
Hypopterigium thouinii Bridges ypopterygiaceae Paragua de sapo n 1 0,29 
Rygodium implexum Schwaegr. Brachytheciaceae Lana del pobre n 2 0,59 
EQUISETATAE 
Equisetum bogotense H.B.K. Equisetaceae Limpiaplata C n 1 0,29 
POLYPODIATAE 
Adiantum chilense Kaulf. Adiantaceae Patita negra H n 3 049 
Asplenium dareoides A. N. Desv. Aspleniaceae pio del monte F n 2 0,59 
Asplenium trilobum Cav. Calahuala F n 2 059 
Blechnum blechnoides (Bory) Keyserl. Blechnaceae Helecho H n 2 0,9 
Blechnum chilense (Kaulf.) Mett. Quil-Quil H n 4 281 
Blechnum hastatum Kaulf. Palmilla H n 5 1,9 
Ctenitis spectabilis (Kaulf.) Kunkel Aspidiaceae Unknown H n 2 0,9 
Hymenoglossum cruentum (Cav.) K. Presl Hymenophyllaceae ^ Helecho pelicula F n ] 029 
MPG eA caudiculatum Mart. F n 1 0,29 
Hymenophyllum dentatum F n col 
Hymenophyllum ae Phil. F n 1 0,29 
Hymenophyllum pectinatum Cav. F n col 
Hymenophyllum plicatum Kaulf. F n 2 0,9 
Hypolepis rugosula (Labill.) J. E. Sm. Dennstaedtiaceae Unknown H n 2 0,59 
Piscina. feuillei Bert. Polypodiaceae Hierba del lagarto F n 3 049 
Polystichum chilense (Christ) Diels Aspidiaceae Helecho palmita H n 1 0,9 
PINATAE 
Podocarpus salignus D. Don Podocarpaceae Mario hoja larga F n 1 0,90 
MAGNOLIATAE 
Acaena ovalifolia R. et P. Rosaceae Cadillo H n 3 0,9 
Aextoxicon punctatum R. et P. Aextoxicaceae Olivillo F n 5 7,92 
Anagallis arvensis L. Primulaceae Pimpinela escarlata T f 1 0,29 
Anmannia coccinea Roth Lythraceae Unknown T f 1 0,29 
Aristotelia chilensis (Mol.) Stuntz Elaeocarpaceae Maqui F n 6 6,9 
Aster bellidiastrum Nees ex Walp. Asteraceae Unknown H n 1] 0,0 
Azara integrifolia R. et P Flacourtiaceae Aromo F n 2 0,66 
Azara lanceolata Hook. f. Corcolén F n 3 0,89 
Azara microphylla Hook. f. Chin-chín F n 1 0,9 
Baccharis concava (R. et P.) Pers. Asteraceae Chilca F n 3 1474 
Baccharis confertifolia Bert. ex Colla F n col 
Baccharis linearis (R. et P.) Pers. F n 3 0,9 
Baccharis racemosa (R. et P.) DC. F n 5 31,88 
Baccharis rosmarinifolia Hook. et Arn. Romerillo F n col 
Boquila trifoliolata (DC.) Dcne Lardizabalaceae Pil-pil voqui F n 5 149 
Buddleja globosa Hope Buddlejaceae Palguín, matico F n 1] 029 
Calcluvia paniculata (Cav.) D. Don Cunoniaceae Tiaca F n 1 029 
Callitriche stagnalis Scop. Callitrichaceae Estrella de agua H f 1 0,81 
Carduus pycnocephalus L. Asteraceae Cardo T f 1 0,29 
Centaurea calcitrapa L. Unknown T f 1 0,29 
Centella triflora (R. et P.) Nannf. Apiaceae Centella H n 1 0,29 
Cerastium arvense L. Caryophyllaceae Cerastio C f 3 0,89 
Chevreulia sarmentosa (Pers.) Blake Asteraceae Unknown H n 1] 0,4 
Chrysosplenium valdivicum Hook Saxifragaceae H n 1] 0,29 
Cichorium intybus Asteraceae Achicoria H f 1 0,29 
Cirsium vulgare (Savi) Ten. Cardo negro T f 4 1,19 
Cissus striata R. et P. Vitaceae Voqui-naranjillo F n 5 1604 
Citronella mucronata (R. et P.) D. Don Icacinaceae Huilli-patagua F n 3 0,98 
Conium maculatum L Apiaceae Cicuta T f 2 1,88 


Volume 76, Number 2 
1989 


Ramírez et al. 
Flora of Rucamanque 


451 


APPENDIX I. Continued. 

Scientific Name Family Local Name If O F LV. 
Coniza bonariensis (L.) Cronq. Asteraceae Coniza T f 4 1,19 
Crepis capillaris (L.) Wallr. Crepis T f 2 0,59 
Cryptocarya alba (Mol.) Looser Lauraceae Peumo F n 1 0,9 
Cynanchum pachyphyllum (Dcne.) Schum. Asclepiadaceae Voqui F n 2 0,59 
Dasyphyllum diacanthoides (Less.) Cabr. Asteraceae alo sant F n 2 0,59 
Daucus carota Apiaceae Zanahoria silvestre T f 4 132 
Dichondra sericea Sw. Convolvulaceae Oreja de ratón H n 1] 0,29 
Digitalis L Scrophulariaceae Cartucho T f 1 0,29 
Discaria serratifolia (Vent.) B. et H. Rhamnaceae Chacay F n l 029 
Disopsis glechomoides (A. Rich.) Muell.-Arg. | Euphorbiaceae Unknown H n 1 0,60 
Echium vulgare L. Boraginaceae Viborera T f 1 0,29 
Ecremocarpus scaber R. : Bignoniaceae Chuplin F n L 0,29 
Elytropus chilensis (A. DC. ) Muell.-Arg. Apocynaceae Quilmay F n 2 059 
Embothrium coccineum J. R. et G. Forster Proteaceae Notro F n 1 0,29 
Epilobium puberulum Hook. et Arn. Onagraceae Epilobio H n 1 0,29 
Ercilla volubilis A. H. L. Juss. Phytolaccaceae oqui F n 1. 0,29 
Eryngium paniculatum Cav. et Domb. Apiaceae Cardoncillo H n 1 0,29 
Escallonia alpina Poepp. ex Escalloniaceae Siete camisas F n 3 1,19 
Eucryphia cordifolia Cav. Eucryphiaceae o F n 6 246 
Flourensia thurifera (Mol.) DC. Asteraceae Incienso F n col 
Fragaria chiloensis (L.) Duch. osaceae Frutilla H n 1 0,29 
Francoa appendiculata Cav. Saxifragaceae Llaupangue H n 1 0,29 
Fuchsia magellanica Lam. Onagraceae Chilco F n 3 133 
Gamochaeta americana (Mill.) Wedd. Asteraceae Vira-vira H n 2 0,59 
Gamochaeta spicata (Lam.) Cabr. Unknown H n 1 040 
Geranium robertianum L. Geraniaceae Core-core T f 4 1,19 
Gevuina avellana Mol. Proteaceae Avellano F n 3 0,89 
Hydrangea pisi (H. et A.) F. Phil. Hydrangeaceae Pehuelden F n 4 121 
Hydrocotyle poeppi Hydrocotylaceae Tembladerilla H n 3 049 
Hypericum derit Choisy Hypericaceae N H n 1 O19 
Hypericum perforatum L Hierba de San Juan H f 5 1,63 
Hypochaeris radicata L. Cichoriaceae Hierba del chancho H f 4 1,19 
Lardizabala biternata R. et P. Lardizabalaceae Voqui-có F n 2 0,59 
Lathyrus hookeri D. Don Fabaceae einde. T n 1 0,29 
Laurelia philippiana Looser Monimiaceae Tepa F n 2 1,23 
Laurelia sempervirens (R. et P.) Tul Laurel F n 4 1,32 
Leontodon taraxacoides (Vill.) Merat Cichoriaceae Chinilla H f 2 1,95 
Leptocarpha rivularis DC. Palito negro F n 3 0,89 
Leucanthemum vulgare Lam. Margarita H f 3 1,62 
Linum usitatissimum L. Linaceae Lino T f 4 1,19 
Loasa acanthifolia Desr. oasac Ortiga caballuna H n 3 0,89 
Lobelia tupa L. Lobeliaceae Tabaco del diablo H n 1 0,29 
Lomatia dentata (R. et P.) R. Br. Proteaceae inol F n 4 1,19 
Lomatia ferruginea (Cav.) R. Br. Huinque F n 1 0429 
Lotus corniculatus L. Fabaceae Lotera H f 1 0,29 
Lotus uliginosus Schkuhr Alfalfa chilota H f 2 1,2 
Luma apiculata (DC.) Burret Myrtaceae Arrayán F n 5 149 
Margyrycarpus pinnatus (Lam.) O.K. Rosac Perlilla C n 1 0,29 
Matricaria perforata Merat Asteraceae Manzanilla H f 1 029 
Maytenus boaria Mol. Celastraceae Maitén F n 1 0,56 
Mentha pulegium L. Lamiaceae C f 3 0,89 
Mentha Bur e (L.) Hudson Menta alemana H f 1 029 
Mimulus bridgessii (Benth.) Clos Scrophulariaceae Berro H n 1 0,29 
Misodendrum Aer ic iE DC. Misodendraceae Injerto F n col 
Mitraria coccine Gesneriaceae Botellita F n 3 1,16 
Muehlenbeckia siete (J. E. Sm.) Johnst. Polygonaceae Voqui F n 4 1,34 
Mutisia retusa Remy Asteraceae Clavel del campo F n 1 0,29 


452 Annals of the 
V Botanical Garden 

APPENDIX I. Continued. 
Scientific Name Family Local Name Lf O F LV. 
i ah oblonga R. et P. Santalaceae Orocoipo F n 2 0,59 
enia exsucca (DC.) Berg Myrtaceae Pitra F n col 
Ma p (H. et A.) Berg icha F n 2 1,06 
Nertera granadensis (Mutis ex L. f.) Druce Rubiaceae Rucachucao H n ] 0,29 
Nothanthera hetereph lla (R. et P.) D. Don Loranthaceae Quintral del boldo F n L 029 
Nothofagus dombeyi (Mirb.) Oerst. Fagaceae igü F n 1 0,36 
Nothofagus obliqua aki ^ Oerst. oble F n 4 60,6 
Osmorhiza chilensis H. e Apiaceae Asta de cabra H n 2 0539 
Oxalis araucana Reiche Oxalidaceae Vinagrillo T n 4 1,19 
Persea lingue (R. et P.) Nees Lauraceae Lingue F n 4 2325 
Pe ldus Monimiaceae Boldo F n 3 1,91 
Plantago hirtella H.B.K Plantaginaceae Llantencillo H f 1 0,29 
Plantago lanceolata L Siete venas H f 6 1,95 
Prunella vulgaris L. Lamiaceae Hierba mora C f 8 2,39 
Pseudopanax valdiviensis (Gay) Seem. ex Araliaceae Curaco F n 3 0,1 

Reiche 
Ranunculus minutiflorus Bert. ex Phil. Ranunculaceae Botón de oro H n 4 1,19 
Relbunium ee (L.) Hemsl. Rubiaceae Relbún C n 4 1,19 
Rhammus diffusu Rhamnaceae Murta negra F n 2 0,59 
Rhaphanus sativu Brassicaceae Rabanito silvestre T f 1 029 
Rhaphithamnus spinosus (A. L. Juss.) Mold. Verbenaceae uayún F n 4 1,22 
Ribes trilobum Meyen Grossulariaceae Zarzaparrilla F n 3 0,89 
Rubus constrictus Muell. et Lef. Rosaceae Zarzamora F f 8 933 
umex acetosella L Polygonaceae Romacilla H f 5 1,9 
Rumex sanguineus L Romaza H f 2 0,59 
Sanguisorba minor Scop. Rosaceae Pinpinela H f 1 0,29 
Sanicula crassicaulis Poepp. ex DC. Apiaceae Pata de guanaco H n 1 0,59 
Sarmienta repens R. et P. Gesneriaceae Medallita F n 2 0,59 
Satureja multiflora (R. et P.) Briq. Lamiaceae Menta de árbol F n 1 0,29 
Senecio chilensis Less Asteraceae Palo de yegua F n 3 0,89 
Senecio otites Kunze ex DC. Tutuco H n 3 1,08 
Senecio yegua (Colla) Cabr. Palpalén F n 4 2,00 
Sherardia arvensis L. Rubiaceae Unknown T f 1 0,29 
Solanum gayanum (Remy) Reiche Solanaceae Natri F n 3 0,89 
Solanum nigrum L. Tomatillo T f 3 0,89 
Solanum Quse Dunal Yaguecillo F n 3 0,89 
Soliva sessilis R. e Asteraceae Dicha H n col 
Stachys pudo ans Lindl. Lamiaceae Unknown C n 1 0,29 
Stellaria cuspidata Willd. Caryophyllaceae Quilloi-quilloi C n 2 0,9 
Taraxacum officinale Weber Cichoriaceae Diente de león H f 1 0,9 
Trifolium dubium Sibth. Fabaceae Trébol enano T f 3 0,99 
Trifolium polymorphum Poir. Trébo H f 1 0329 
Trifolium repens L. Trébol blanco H f 4 2,17 
Tristerix tetrandrus (R. et P.) Mart. Loranthaceae Quintral del álamo F n 2 0,59 
Tropaeolum ciliatum R. et P. Tropaeolaceae Espuela de galán Cr n 2 0,59 
Ugni molinae Turcz Myrtaceae Murtilla F n 2 1,44 
Urtica urens Urticaceae rtiga H f 3 0,89 
Valeriana Joihanda Phil. Valerianaceae Valeriana H n 3 0,9 
Verbascum thapsus Scrophulariaceae Hierba de paño H f 1 0329 
Veronica anagallis-aquatica L. No me olvides del H f 1 029 
campo 

Vestia foetida (R. et P.) Hoffmanns. Solanaceae Huevil F n col 
cia macraei H. et A. Fabaceae Arvejilla T n col 
Vicia sativa L. T f 1 0,29 
Vicia vicina Clos T n 2 0,59 
Weinmannia trichosperma Cav. Cunoniaceae Tineo F n 2 1,28 


Volume 76, Number 2 Ramírez et al. 453 

1989 Flora of Rucamanque 
APPENDIX I. Continued. 

Scientific Name Family Local Name Lf O F LV. 

LILIATAE 
Agrostis castellana Boiss. et Reuter Poaceae Chépica H f 5 5,61 
Aira caryophyllea L. Air T f 1 1,08 
Alstroemeria pulchra Sims Alstroemeriaceae Amancay Cr n 2 0,59 
Arachnitis uniflora Phil. Corciaceae Flor de la araña Cr n 1 0,29 
Arrhenatherum elatius (L.) P. Beauv. oaceae Pasto cebolla H f 3 0,9 
Bomarea salsilla (L.) Herb. Amaryllidaceae Copihuito Cr n 1 0,29 
Briza minor L Poaceae emblequ T f 1 0,29 
Bromus unioloides H.B.K. Pasto lanco H n 1] 0,59 
Carex acutata Boott Cyperaceae Cortadera H n 1 0,29 
Carex fuscula D H n 1 0,29 
Chloraea aca Poepp. Orchidaceae Pico de loro Cr n 1 029 
Chusquea culeou Desv Poaceae Colihue F n 2 5,03 
Chusquea quila Kun Quila F n 4 11,69 
Codonorchis lessonii (D'Urv.) Lindl. Orchidaceae Azucena - ice Cr n col 
Cynosurus echinatus Poaceae Cola de ra T f 4 190 
Dactylis glomerata L. Pasto s H f 3 1,02 
Dioscorea andin Dioscoreaceae Papa del monte Cr n col 
Dioscorea brachybothrya Poepp. Papa cimarrona F n 2 0,59 
Eragrostis lugens Nee Poaceae Unknown H f 1 0,29 
Fascicularia bicolor (R. et P.) Mez Bromeliaceae Chupalla F n 1 0,29 
Gavilea odoratissima Poepp. Orchidaceae Orqui Cr n col 
Holcus lanatus Poaceae Pasto dulce H f 7 237 
Juncus capillaceus Lam. Juncaceae Unknown H n 1 0,29 
Juncus dombeyanus J. Gay. ex Lah. Junquillo H n 1 0,29 
Juncus microcephalus H.B.K. H n 1 0,9 
Juncus procerus E. Mey. H n 1 2,93 
Lapageria rosea R. et P. Philesiaceae Copihue F n 5 L79 
Libertia chilensis (Mol.) Gunckel Iridaceae Calle-Calle H n col 
Libertia coerulescens Kunth et Bouché Calle-Calle azul H n ] 0,29 
Libertia ixioides Gay H n 3 0,9 
Lolium multiflorum Lam. Poaceae Ballica italiana H f 1 0,29 
Lolium perenne L Ballica inglesa H f 1 029 
Luzuriaga radicans R. et P. Philesiaceae oralito C n 2 0,9 
Nasella excerta Phil. Poaceae Pasto quila H n 1 0,29 
Paspalum distichum L. Unknown H f 2 0,94 
Polypogon chilense (Kunth) Pilger Cola de zorro H n 1 043 
Sagittaria montevidensis Cham. et Schlecht. Alismataceae Rosa de agua H n 1 0,29 
Scirpus cernuus Vahl Cyperaceae Can-cán H n 1 0,43 
Stipa poeppigiana Trin. et Rupr. Poaceae Quilmén H n 1 0,29 
Uncinia erinacea (Cav.) Pers. Cyperaceae Unknown H n 3 0,9 
Uncinia phleoides (Cav.) Pers. Clin-clin H n 4 1,19 
Vulpia bromoides (L.) S. F. Gray Poaceae Cepilla T T 8 LiT 


NOVELTIES IN THE 
ORCHID FLORA OF 
SOUTHERN VENEZUELA! 


Gustavo A. Romero?? 
and Germán Carnevali?^ 


ABSTRACT 


Six new orchids from Venezuelan Guayana are described and illustrated, including three species, Coryanthes 
at 


cataniapoensis, Catasetu 


zii, and Catasetum parguaz 


ybrid, Catasetum X dunstervillei, 


three hybrids involve four sympatric Catasetum species with overlapping flowering 


seasons that share Eulaema cingulata (Fabricius) as their main pollinator 


This paper and others in preparation precede 
the publication of a comprehensive treatment ‘a 
the Orchidaceae of southern Venezuela for J. 
Steyermark’s Flora of the Venezuelan oe 
(Carnevali et al., in prep.). The taxa described here 
belong to two of the most complex genera in the 
Orchidaceae, Coryanthes Hook. and Catasetum 
Rich. Species in these two genera are extremely 
variable, and in the Venezuelan Guayana many 
closely related species are sympatric or syntopic, 

e pollinators, and have overlapping flowering 
seasons; natural hybrids are common. 

alf of the taxa proposed here are natural hy- 
brids. The parentage of these nothospecies was 
inferred from pollination data, phenological data, 
and morphological characters. This approach has 
its limitations, but one can postulate safely parent 
species if a large number of specimens of the sus- 
pected hybrid are examined, because, as Hurst 
(1902) indicated, “In orchid hybrids of the first 
generation single specific characters are inherited 
in all degrees of blending, forming, on the whole, 
a perfect series between the respective characters 
of the two parents." In Catasetum x wendlingeri 
Foldats, the parentage was confirmed by artificial 
hybridization experiments (A. Pardo, pers. comm., 


Coryanthes cataniapoénsis G. Romero et Car- 
nevali, sp. nov. TYPE: Venezuela. Territorio 


Federal Amazonas: Departamento Atures, Rio 
Gavilan, alrededores de Cucurital, 9 Ma 
1987, G. A. Romero 1307 (holotype, VEN; 
isotype, K). Figure 1A-C. 


Planta C. macranthae (Hook.) Hook. affinis sed hy- 
pochilo antice eee campanulato et mesochilo subae- 


quilongo differ 


Plant epiphytic in ant gardens. Rhizomes short; 
pseudobulbs ovoid to narrowly ovoid, grooved, bi- 
foliate at the apex, to 11 cm long, surrounded at 
the base by scarious sheaths. Leaves oblong-elliptic, 
acuminate, to 34 cm long, including the short, 
sulcate petiole; lamina to 9 cm wide and 31 cm 
long, plicate, with 3 conspicuous nerves beneath. 
Inflorescence from the base of the pseudobulbs, 
pendulous, elongate, 1- or 2-flowered at the apex; 
peduncle slender, to 18 cm long, with short remote 
tubular-infundibuliform sheaths. Ovary pedicellate, 
elongate, slender, ca. cm long, almost 4 times 
as long as the concave, scarious subtending bract. 
Flowers white, densely spotted and blotched with 
dark red-maroon. Sepals and petals membrana- 
ceous; dorsal sepal ovate, obtuse, ca. 4.3 cm long 
and 5 cm wide; lateral sepals obliquely oblong- 
ovate, retrorse, ca. 9 cm long an cm wide, 

margins involute; petals narrowly oblong-lanceo- 
late, falcate, obtuse, ca. 5 cm long and 1.3 cm 
wide near the base, margins shallowly undulate. 


' We are grateful to J. A. Steyermark and B. J. 
E. Melgueiro, and J. Moreno for field assistance. We t 
collection of euglossine bees; the curators of AM 
and A. Pardo for providing p and materi 


Indiana University, Sigma Xi, and The Am merican Socie 


al on artificial hybrids. Thi 
ety of Plant Taxonomists to G. A. Romero. Venezuela’s “Fondo 


per for the line drawings and to P. Beer-Romero, C. Gom 


. L. Dressler for having kindly identified a tolera 


Nacional de Investigacion nes in opecuarias" provided logistic support. 


* Jard in Botánico de Caracas, INPARQUES, Apartado 


2156, Caracas 1010-A, Venezuela. 


urrent address: Oakes Ames Orchid Herbarium, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, 


Pai 02138, U.S.A. 


* Current address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. 
ANN. Missouni Bor. GARD. 76: 454-461. 1989. 


Volume 76, Number 2 Romero & Carnevali 455 
1989 Orchid Flora of Southern Venezuela 
GURE l.— A-C. — cataniapoensis. A. Side view of labellum. B. Front view of hypochile. C. Side view 


of bue n after removing hypochile, showing the labellum claw (Cl), the transverse ridges (Lm, and Lm,), and the 


projections (Pr) near the base (Lm, and 
(H Hook. shown for comparison. A, B from 
s.n. (VEN). 


Lip fleshy, divided into 3 parts, the clawed, semi- 
spherical hypochile 4.5-5 cm 
bular mesochile ca. 4-4.5 cm long, with 5-6 trans- 
verse ridges, bilobed to multilobed, decreasing in 
size from the base, with 3 toothlike projections 
near the base (these seen only after removing the 
hypochile), 8-13 mm high; the deeply saccate 
epichile bucketlike, with a 3-lobed apex, the lateral 
apex, the middle lobe lin- 
gulate, shallowly obcordate. Column gradually di- 
lated toward the apex, recurved near the apex, 
bialate, ca. 2.5 cm long, with a pair of shallowly 
falcate, fluid-producing glands at the base. Pollinia 


wide; the semitu- 


lobes uncinate at the 


2, waxy, yellow. 


atypes. VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: Departamento Atures, Rio Gavilán, alrede- 
es de Cucurital, June 1985, Romero 1273 
(AMES) Rio Cataniapo, entre la boca del Río Gavilán y 
los Raudales de Rabipelado, Jan. 1985, C. Gomez s.n. 
(TFAV). 


Pr not visible unless the hypochile is removed). — D. 
G. A. Romero 1307; C fro 


Coryanthes macrantha 


m C. Gomez s.n.; D from G. Bergold 


Etymology. Named for the Rio Cataniapo, 
where the species was first found. 


This species differs from Coryanthes macran- 
tha (Hook.) Hook. in the globose hypochile with 
the anterior margin that almost reaches the epi- 
chile, and in the toothlike projections near the base 
of the mesochile. It flowers from January to June 
and is pollinated around Puerto Ayacucho, T. F. 
Amazonas, Venezuela, by Eulaema cingulata (Fa- 
bricius) and, less frequently, by E. meriana (Oliv- 
ier) (G. A. Romero, unpubl. data) 


Catasetum gomezii G. Romero et Carnevali, sp. 
nov. TYPE: Venezuela. Territorio Federal Ama- 
zonas: Departamento Atures, Rio Cataniapo, 
cerca de la desembocadura del Rio Gavilán, 
30 June 1987, G. A. Romero & C. Gomez 
1333 (holotype, VEN; isotypes, K, TFAV). 
Figure 2A-B. 


456 


Annals of th 
Missouri Bond Garden 


FIGURE 2.- 


: rawn from the 
s and J from Saunders, 


Planta inter C. barbatum Lindley et C. ochrac 
Lindley quasi intermedia, sed E yei labello nde. 
simo marginibus dentatis differ 

Plant epiphytic, indistinguishable vegetatively 
from other members of the genus. Staminate in- 
florescence lightly arcuate to pendent, racemose, 


e —ÀÁ 
-A, B. Catasetum gomezii. A. Staminate flower. B. Detail of staminate flower 
paraguazense. —D. Catasetum callosum 
setum pileatum. — 
Catasetum discolor. Scale bars — 


O = Foldats, 1970: 59; E from 


—C. Catasetum 


plates accompanying the original descriptions (Hooker, 1840, and Lindley, 1834, 
1882. 


to 9-flowered, originating from the base of the fully 
developed pseudobulb, to 18 cm long. Staminate 
flowers resupinate. Tepals light green, densely spot- 
ted with maroon on their inner surface, less so on 
their outer surface; dorsal sepal slightly concave, 
oblanceolate, acute, margins entire, 11-12 mm 


Volume 76, Number 2 
1989 


Romero & Carnevali 457 


Orchid Flora of Southern Venezuela 


wide, 25-26 mm long; lateral sepals slightly con- 
cave, oblong-lanceolate, acute or shortly apiculate, 
slightly oblique, margins entire, 12-13 mm wide, 
2 m long; petals narrowly ovate, acute, 
margins entire or finely serrate with 1 indentation 
and a flush falcate tooth on each side at the widest 
point, 10-11 mm wide, 24-25 mm long. Labellum 
fleshy, rigid, yellowish green, spotted with maroon 
in the margins, deeply concave, deltate in general 
outline, with 2 lateral indentations midway between 
the base and the apex, thus slightly trilobulate, 12 
mm deep, 22 mm long, 22 mm wide between the 
spread basal lobes, margins crenate to dentate in 
the middle lobe, dentate to fimbriate in the lateral 
lobes, the base with a transverse, tricuspid callus, 
the apex with a V-shaped, verru l 

yellowish green, with maroon dots in the inner 


3 


altus. oium 


surface near the base, densely spotted with maroon 
on the outer surface, semiterete, stout, straight, 
apically thickened, conspicuously rostrate, the cli- 
nandrium apiculate and slightly incurved at the 

ex; antennae bilaterally symmetrical, slightly 
convergent, 8 mm long. Anther white, the margins 
spotted with maroon. Pollinia 2, cleft, waxy, com- 
pressed, more than twice as long as wide, attached 
to a light yellowish green stipe with a white, ad- 
hesive viscidium. Pistillate inflorescence unknown. 


Etymology. | Named in honor of Carlos Gomez, 
the discoverer of this species. 


Par VENEZUELA. TERRITORIO FEDERAL 
AMAZON ui ea say Rio Cataniapo, entre el 
Milagro y San Pablo, 1 1987, . Romero, C. 
G Beer- E. 1365 (AMES, TFAV). 


This showy species resembles Catasetum ochra- 
ceum Lindley (Lindley, 1844) but differs in the 
epiphytic habit, pendulous inflorescence and la- 
bellum with dentate to fimbriate margins. It is also 
similar to C. cornutum Lindley (Lindley, 1840) 
and the recently described C. semicirculatum Mi- 
randa (Miranda, 1986), but the new species is 
distinguished by the deeply concave labellum with 
a frontal, V-shaped callus. It flowers from May to 
September. 


Catasetum parguazense C. Romero et Carne- 
vali sp. nov. TYPE: Venezuela. Bolivar: Distrito 
Cedeño, Rio Parguaza, 12 May 1967, E. Rut- 
kis 1985 (holotype, VEN). Figure 2C. 

Planta Cataseto calloso Lindley et eius affinitatis prox- 
ima, sed labello basi subcordato et callo basali truncato 
differt 

Plant epiphytic, indistinguishable vegetatively 
from other members of the genus. Staminate in- 


florescence lightly arcuate to pendulous, racemose, 
to 8-flowered, originating from the developing leaf 
growth, to 15 cm long. Staminate flowers resupi- 
nate. Tepals tinged green with maroon, or maroon 
entirely; dorsal sepal narrowly elliptic or narrowly 
obovate, acute or acuminate, somewhat concave, 

g tire or slightly ciliolate, 29-31 mm long, 
Te 10 mm wide; lateral sepals similar to dorsal sepal 
but slightly oblique, 9-10 mm wide; petals nar- 
rowly elliptic to narrowly obovate-elliptic, acute or 
shortly acuminate, margins entire or finely dentic- 
ulate to ciliolate, 26-31 mm long, 7-9 mm wide. 
Labellum green with maroon dots or maroon en- 
tirely. Labellum fleshy, in general outline narrowly 
ovate, obtuse, rounded to truncate, basal margins 
extended, making the labellum subtrilobate, 2-2.6 
cm long, 1.8-2 cm wide between the spread basal 
lobes, margins ciliate and glandular papillose or 
coarsely verrucose, the ventral face glandular ver- 
rucose, especially toward the margins, leaving an 
almost longitudinally smooth central zone, basally 
with a transverse, truncate low callus having a 
verrucose surface; the labellum excavate in front 
of the callus, forming a shallow, conical rounde 
sack; the apex of the labellum callus thickened. 
Column green or maroon or maculated with green 
and maroon, straight, apically thickened, conspic- 
uously rostrate, 16-22 mm long; clinandrium apic- 
ulate and slightly incurved apically; antennae bi- 
laterally symmetrical, subparallel or somewhat 
divergent, smooth or finely glandular verrucose, 
6-8 mm long. Anther yellowish green, maroon at 
the base and apex. Pollinia 2, cleft, waxy, yellow, 
compressed, more than twice as long as wide, at- 
tached to a light yellow-green stipe with a white, 
adhesive viscidium. 


Paratypes. | VENEZUELA. BOLÍVAR: Distrito Roscio, 60 
km al oeste de Santa Elena de Uairén, por la via de 
Icabarü, G. Carnevali 2614 (VEN); Distrito Piar, on 
tributary to Río Carrao, ca. 10 mi. above Canaima and 
Rio Churün, G. C. K. Dunsterville 259 (line drawing, 
AMES). 


Etymology. Named after the Rio Parguaza, 
where the type of the species was collected. 


This species has been collected several times in 
the Venezuelan Guayana and in Brazil (see color 
photograph in Miranda, 1987). In the past it had 
been assigned incorrectly to C. bicallosum Cogn. 
(Dunsterville & Garay, 1979: 78; Foldats, 1970: 
58; Miranda, 1987), a species from the upper Rio 
Orinoco closely related to C. barbatum Lindley. 
The type specimen of C. bicallosum Cogn. in the 
Cogniaux Herbarium (BR, photo) and the original 
description clearly show this affinity. Catasetum 


458 


Annals of the 
Missouri Botanical Garden 


parguazense belongs in the C. callosum Lindley 
alliance (Pabst & Dungs, 1975: 172). It differs 
from C. callosum by having a subcordate labellum 
and truncated basal callus, and it differs from C. 
poriferum Lindley (see color photo in Fergusson, 
1967) by having long, fully developed antennae 
and a labellum with erect-patent lateral lobes. It is 
further distinguished by flowering almost invariably 
from the developing leaf shoot. 


Catasetum x dunstervillei G. Romero et Car- 
nevali, hybr. nat. nov. inter Catasetum pilea- 
tum Reichb. f. et C. discolor Lindley. TYPE: 
Venezuela. Territorio Federal Amazonas: De- 
partamento Atures, Los Oripopos, 7 km al 
norte de Puerto Ayacucho, 23 Nov. 1982, G. 
A. Romero & A. Cortez 914 (holotype, VEN; 
isotype, TFAV). Figure 2H. 


Planta inter parentes ad C. discolorem Lindley ver- 
gens, sed inflorescencia arcuata vel pendula, floribus re- 
supinatis majoribus, labello leviter saccato, callo V-formi 
praedito, antennis longioribus, columnae clinandrio api- 
culo brevi recedit. 


Plant epiphytic, indistinguishable vegetatively 
from other members of the genus. Staminate in- 
florescence lightly arcuate to pendent, racemose, 
to 7-flowered, originating from the base of the fully 
grown pseudobulb, to 16 cm long. Staminate flow- 
ers resupinate. Tepals white, maroon, light green 
or greenish yellow with maroon dots. Dorsal sepal 
concave, lanceolate, acuminate, erect, 23-25 mm 
long, 9-10 mm wide; lateral sepals concave, ob- 
dae slightly s apiculate, recurved, 22-25 
mm long, 9-10 mm wide. Petals convex, obovate, 
slightly oblique, ale 25-27 mm long, 11- 
abellum pearl white, green, light 
green or waxy yellow, saccate, the sack 15-18 
mm deep, simple or with 2 shallow indentations 
alongside the apex, thus slightly trilobulate, and 
with a V-shaped callus near the inner apical margin; 
margins patent or reflexed, finely denticulate to 
fimbriate. Column light greenish white or yellow, 
semiterete, straight, apically thickened, shortly ros- 
trate; clinandrium shortly apiculate, incurved api- 
cally; antennae bilaterally symmetrical, parallel to 
slightly convergent, 6-8 mm long. Anther greenish 
white. Pollinia 2, cleft, waxy, yellow, compressed, 
shorter than twice as long as wide, attached to a 
light yellowish white stipe with a white, adhesive 
viscidium. 


13 mm wide. 


Paratypes. | VENEZUELA. m FEDERAL 
AMAZONAS: upper Orinoco, (P. Anduze) G. C. K. Dun- 
"id e 748 (line drawing, AMES); quias Atures, 

o Pavoni, cerca de su confluencia con el Orinoco, 3 


Nov. 1983, G. A. Romero & J. Moreno 1153 (VEN, 
TFAV); same locality, 10 Aug. 1987, G. A. Romero, P. 
L. Beer-Romero & J. Moreno 1336 (VEN, TFAV); Cano 
Rueda, 6 Dec. 1983, G. A. Romero & L. Rojas 1158 
VEN) 


— 


Etymology. Named in honor of G. C. K. 
Dunsterville, who first illustrated this hybrid in his 
series coauthored by L. A. Garay, Venezuelan 


Orchids Illustrated. 


This natural hybrid has been previously assigned 
to Catasetum fimbriatum (Morren) Lindley (Dun- 
qi & Garay, 1966: 42; 1979: 85; Foldats, 
1970: 80). However, three lines of evidence reveal 
C. x dunstervillei as distinct. First, northern South 
America lies outside the known geographical range 
of Catasetum fimbriatum: southeastern Brazil, 
southwestern Bolivia, Paraguay, and northern Ar- 
gentina (Bicalho, 1965; Dodson, 1978). Second, 
typical C. fimbriatum from southern Brazil differs 
considerably from Catasetum X dunstervillei in 
having a column with long antennae; a long, apic- 
ulate, acuminate clinandrium; an apiculate, fleshy 
anther; and a labellum with the fleshy mid-lobe 
projected forward (see Vasquez & Dodson, 1982). 
Third, based on fragrance and pollinator data, 
Catasetum Xdunstervillei cannot be assigned to 
C. fimbriatum. Hills et al. (1972) showed that C. 
fimbriatum is a poor fragrance producer with a 
fragrance spectrum unlike any other species in the 
genus. Moreover, the only known C. fimbriatum 
pollinator is Eufriesea auriceps (Friese) (Pijl € 
Dodson, 1966: 182). Catasetum X dunstervillei, 
in contrast, produces a fragrance that attracts Eu- 
laema cingulata, one of the pollinators of Catase- 
tum discolor Lindley, C. longifolium Lindley, and 
in general the C. maculatum Kunth complex (Wil- 
liams & Whitten, 1983). Catasetum X dunster- 
villei was also compared with C. fimbriatum var. 
cogniauxii L. Linden, described from material pre- 
sumably collected in Venezuela (Linden, 18954). 
A plate in Lindenia (Linden, 1895b) and Cog- 
niaux's sketches in the type sheet (BR, photo) 
reveal Catasetum fimbriatum differing from C. 
x dunstervillei in the characters presented above. 

Dressler (1976) pointed out that Dunsterville 
748, assigned here to Catasetum X dunstervillei 
peia in Dunsterville & Garay, 1966: 42; 

9: 85), was a hybrid between Catasetum pi- 
d and C. discolor, but he never proposed a 
binomial. Dressler's hypothesis is supported by two 
lines of evidence. The flowers of C. x dunstervillei 
are intermediate between Catastum pileatum and 
C. discolor, and these two species are by far more 
abundant than any other possible parent species 


Volume 76, Number 2 
1989 


Romero & Carnevali 459 


Orchid Flora of Southern Venezuela 


found in the general area where C. X dunstervillei 
has been collected, as indicated by plant sampling 
and by the frequency of their pollinaria on euglos- 
sine bees (G. A. Romero, unpubl. data). Further- 
more, Catasetum pileatum and C. discolor share 
euglossine bee pu and have overlapping 
flowering season 

Dodson d that Dunsterville 748 was a 
hybrid between Catasetum longifolium Lindley and 
C. macrocarpum Rich. ex Kunth. (Dodson, 1978), 
based on plants collected in Dawa, Guyana (also 
pictured in Tan, 1971, fig. 8). However, the plant 
illustrated in Dodson (1978) differs from C. x dun- 
stervillei in the pendent habit, the long and narrow 
leaves, and the labellum morphology. 

Catasetum X dunstervillei is similar to Cata- 
setum Xissanensis Pabst (a natural hybrid be- 
tween C. pileatum and C. longifolium, Pabst, 1975) 
but is distinguished by the shorter and wider leaves 
and the labellum with a V-shaped callus. 

Catasetum X dunstervillei flowers from the ma- 
ture pseudobulb from August to December. The 
pollen donor in C. X dunstervillei is most likely C. 
discolor: the large pollinia of C. pileatum could 
not fit in the narrow stigmatic cleft of C. discolor. 


Catasetum Xroseo-album (Hook.) Lindley (pro. 
sp.), hybr. nat. nov. inter Catasetum discolor 
Lindley et C. longifolium Lindley. Mona- 
chanthus roseo-albus Hook., Bot. Mag. 66: 
t. 3736. 1839. Catasetum discolor var. ro- 
seo-album (Hook.) Mansf., Repert. Spec. Nov. 
Regni Veg. 30: 264. 1932. TYPE: Brazil. Para: 
sent by Mr. Campbell to Mr. Murray at the 
Glasgow Botanical Garden, collector unknown 


(holotype, K, photo). 


Selected specimens examined. VENEZUELA. BOLÍ. 
vAR: Distrito Cedeño, Agua Mena, epifita sobre Mauritia 
.M 


Pavoni, epifita sobre árbol seco, 10 Nov. 1987 (male fl), 
. A. Romero . Moreno 1421 (TFAV); Rio Autana, 
Caño Cabez Manteco a sobre Bactris sp., 8 


(male fl), G s ero & J. 
E 

19 Apr. 1970 kadie fl), 

10257 1 (VEN); same locality and date (female fl), J. A. 
Steyermark & G. Bunting 102649 (VEN). 


Etymology. The specific epithet probably re- 
fers to the rose and white flowers in the original 
description. 


Natural hybrids between Catasetum discolor 
and C. longifolium were expected in the Vene- 
zuelan Guayana: the two species are separated by 
less than the flying range of euglossine bees (1-5 
km), they share male euglossine bee pollinators, 
Eulaema meriana (Olivier), E. bombiformes 
(Packard), and E. cingulata (Fabricius), and they 
have overlapping flowering seasons. Around Puerto 
Ayacucho, Venezuela, the same species of Eulae- 
ma bees have been caught carrying the very dis- 
tinct C. discolor and C. longifolium pollinaria in 
late February through May. The pollinaria of both 


» 


species are placed on the ventrum of the bees 
thorax, but C. discolor pollinaria have a dark brown 
stipe and a very small viscidium (narrower than 
the involute stipe), whereas C. longifolium polli- 
naria have a light brown stipe and a large viscidium 
(wider than the involute stipe). Plants bearing flow- 
ers intermediate between the two presumed parents 
had been collected in the Venezuelan Guayana, 
Guyana (Dodson, 1978), and Brazil (Mansfeld, 
1932) and were assigned previously to Catasetum 
discolor var. roseo-album (Hook.) Mansf. Dodson 
(1978) suggested that this taxon may have resulted 
from a natural cross between Catasetum discolor 
and C. longifolium, but he never changed its no- 
menclatural status to that of nothotaxon. 

Catasetum X roseo-album is widespread in low- 
land "morichales" (gallery forest communities 
dominated by Mauritia flexuosa L.f.) in the north- 
western Venezuelan Guayana, where it occupies 
typical C. longifolium habitat, growing near the 
crown on the stems of Mauritia flexuosa. It is also 
found on Bactris sp., and on dead trees in sea- 
sonally flooded forest. 

The hybrid differs from C. longifolium by hav- 
ing generally erect (vs. pendulous) habit; leaves 6— 
20 cm long, 4-7 cm wide and generally 5-nerved 
(vs. 15-150 cm long, 1-3 cm wide, and 3-nerved), 
shortly apiculate, apically incurved (vs. obtuse and 
flat) clinandrium of the staminate flower, and the 
C. discolor-like pollinarium. 

The hybrid differs from C. discolor in the epi- 
phytic habit, generally arcuate to pendent stami- 
nate inflorescence (always erect in C. discolor), 
and deeply galeate, dorsally compressed labellum 
of the staminate flower (shallowly concave and 
globose in C. discolor). Pistillate flowers of the 
hybrid are often distinguished by the deeply galeate 
labellum (globose in C. discolor) but can vary be- 
tween C. discolor-like to C. longifolium-like (deep- 
ly saccate), ranging in color from light green (yel- 
low-green in old flowers) to deep maroon. The 
hybrid staminate flowers vary in color between 
pinkish white to green, purple, orange, or maroon. 


460 Annals of the 
Missouri Botanical Garden 
Catasetum X roseo-album flowers from the ma- — dulate-dentate. The hybrid differs from C. pilea- 


ture pseudobulb from June to January. Pollen flow 
between either parent is likely, as the pollinia of 
both species are of similar size. 


Catasetum x wendlingeri Foldats (p ), hybr. 
nat. nov. inter Catasetum pileatum Reichb. 
f. et C. planiceps Lindley. TYPE: Venezuela. 
Territorio Federal Amazonas: Departamento 
Atabapo, alrededores de San Fernando de Ata- 
bapo, July 1957, E. Foldats 2890 (holotype, 
VEN). Figure 2E. 


Selected specimens examined. VENEZUELA. BOLÍ. 
VAR: Distrito Cedeño, Caripo, June 1980, (I. Sucre) G. 
C. K. vog dns 1413 (line drawing, AMES); same 
re) G. Carnevali 2633 (VEN, TFAV). 


~ 
ho 
e 
~ 


VEN); carretera a Gavilin. ca. del Fundo Dona 
Juana, 23 June 1985, G. A. Romero & J. Taine 1263 

TFAV); same locality, 17 May 1987, G. A. Romero 
1310 (TFAV). 


Etymology. Named after K. Wendlinger, who 
cultivated the plant featured in the original de- 
scription. 


C. H. Dodson first proposed this natural hybrid 
between Catasetum pileatum and C. planiceps 
(quoted in Dunsterville & Uzcategui, 1981) based 
on the floral morphology of the G. C. K. Dun- 
sterville 1413 specimen cited above. Dodson, how- 
ever, never proposed a binomial. Since then, plants 
resulting from the artificial cross between the pre- 
sumed parents flowered in the collection of A. Par- 
do in Venezuela, and three more specimens of the 
presumed natural hybrid were collected around 
Puerto Ayacucho. Comnarison of the staminate 
flower of the artificial hybrid with the holotype of 
Catasetum wendlingeri and C. planiceps (K, pho- 
to) showed it to be more similar to C. wendlingeri; 
in fact, some of the hybrids were identical to C. 
wendlingeri. Catasetum wendlingeri had been 
treated previously as a synonym of C. planiceps 
(Foldats, 1959; 1970: 97; Schweinfurth, 1967). 

Catasetum X wendlingeri differs from C. plani- 
ceps in the epiphytic habit and in the generally 
larger staminate flower with the labellum apex 
broadly patent or reflexed and the column with a 
long, apicular clinandrium. Further, the staminate 
flowers of the hybrid are sometimes resupinate (as 
in G. C. K. Dunsterville 1413 and G. A. Romero 
& L. Rojas 1201), or the margins of the labellum 
are fimbriate (as in G. C. K. Dunsterville 1413, 
illustrated in Dunsterville & Uzcategui, 1981), 
whereas the staminate flowers of C. planiceps are 
never resupinate and the margins crenate or un- 


tum in the hooded labellum and the bilaterally 
symmetrical antennae of the staminate flowers. 

Catasetum X wendlingeri flowers from the de- 
veloping leaf shoot from April to July. This hybrid, 
as is its proposed parent species, is pollinated by 
Fulaema cingulata. The pollen donor of the hybrid 
is most likely Catasetum planiceps; C. pileatum 
pollinia could not be inserted in the narrow stig- 
matic cleft of C. planiceps. 


LITERATURE CITED 


BicaLHo, H. D. 1965. Estudos sistemáticos no género 
Suid tum L. C. Rich. (Orchidaceae). I. — Catase- 
um fimbriatum (Morren) Ldl. Loefgrenia 20: 1-28. 
Donson. C. H. 1978. The catasetums (Orchidaceae) of 
apakuma, Guyana. Selbyana 2: 159-168. 
me R.L. 1 Venezuelan Orchids Illustrat- 
monumental work. Orquidea 6: 129-131 
Dien . K. & L. A. Garay. Ven- 
ezuelan Orchids Illustrated IV. Andre Deutsch, Lon- 
don 


979. Orchids of Venezuela. Bo- 

tanical Museum of Harvard University, Cambridge, 

Massachusetts 

& E. Uzc. ATEGUI. 1981. ¿Una nueva especie 

de Catasetum? Bol. Comité dad Soc. Venez. 
7-31. 


l. t 
FERGUSSON, M. 
Orchid Dig. 31: 109- 
Forpars, E. 8 PA a la oe 
de Venezuela. Acta Biol. Venez. 2: 369-40 
. Orchidaceae, m L. E Rich. 
Flora de Meus XV(4): 48-109. 
Hiis, H. G., N. H. C HD Dopson. 1972. 
Floral eee ir isolating mechanisms in the 
genus Catasetum (Orchidaceae). Biotropica 4: 61- 


iA to the genus Catasetum. 


HOOKER, W. 1840. Monachanthus roseo-albus. Bot. 
67: t. 3796. 
. Mendel's principles applied to or- 
id hybrids. J. Roy. Hort. Soc 27: 614-624. 
822 


895b. Catasetum enu ie iri 
Lin a var. Cogniauxi. Lindenia 11: 41, 
Livin, J. 1834. Move landat: doa Edvard's 
t. Reg. 20: t. 1735. 
Pts g cornutum. Edward's Bot. 
Reg. 26: misc. p. 77. 
1844. Catasetum ochraceum. Edward's Bot. 


Re eg. 30: misc. p. 
MANSFELD, R. 1932. “Dio Gattung Catasetum L. C. 
Rich. Rep. Spec. Nov. Regni Veg. 30: 257-275. 
MIRANDA, F. E. 1986. New orchids from Brazil — 1. 


ag ri 1: 156. 
————, tasetums in Brazil — notes on habi- 


tats x aabt: Amer. Orchid Soc. Bull. 56: 473- 
482 


Passt, G. F. 1975. New or critical orchids from Bra- 
zil— VII. Catasetum x issanensis Pabst. Orchid Rev. 
48: 405-406. 
F. DuNcs. 1975. Orchidaceae Brasiliensis, 
Volume I. Kurt Schmersow, Hildesheim 


Volume 76, Number 2 
1989 


Romero & Carnevali 461 


Orchid Flora of Southern Venezuela 


PiL, L. VAN DER & C. H. Dopson. 1966. Orchid Flow- 
olut 


ution. Univ. Miami 

— ERS, W. : Refugium Botanicum II. John 

rst, London. 

SCHWEINFURTH, C. 1967. Orchidaceae of the Guayana 
Highlands. Mem. New York Bot. Gard. 14: 69-214. 

Tan, K. W. Orchids of "un Guyana, Amer. 
Orchid Soc. Bull. 40: 585-59 


Vasquez, R. & C. H. Dopson. 1982. Catasetum "ad 
briatum epah Lindl. Icon. Pl. Trop. t. 
WiLLIAMS, N. H. & W. M. WHITTEN. 1983. Orchid 
floral fragrances and male euglossine bees: methods 
and advances in the last sesquidecade. Biol. Bull. 
pedis 164: 355-395. 


NOVELTIES IN THE 
LAURACEAE FROM 
VENEZUELAN GUAYANA! 


Henk van der Werff* 


ABSTRACT 


Eleven new species (Licaria n Ocotea flavantha, O. liesneri, O. glabra 
"roatii, P. croizatii, and P. fluviatilis) are described. Ocotea julianii is 
for Phoebe peint Phoebe areolatocostae is transferred to Persea, and Ocotea 


o ulata, O. tomentosa, 
published as a new name 


steyermarkiana is transferred to Rhodostemonodaphne 


, O. huberi, O. megacarpa, O. 


During the preparation of the treatment of the 
Lauraceae for the Flora of the Venezuelan Guaya- 
na, a number of collections were found that ap- 
peared to represent undescribed taxa. Most of these 
collections were gathered during recent expeditions 
and are of high quality. As can be expected, there 
are, after completion of the family treatment, still 
several sterile or fruiting collections that do not 
match any of the known species from Guayana 
and that represent additional or even undescribed 
species. These species could not be included in the 
treatment for lack of adequate collections. 

In general, the Lauraceae in Venezuelan Guaya- 
na are still undercollected. Of the 128 species 
(Nectandra excluded; this genus will be contributed 
by Dr. J. Rohwer), 55 are known only from one 
or two collections, while an additional 13 are known 
from three or four collections. This large number 
of rarely collected species suggests that there are 
still several additional species to be collected and 
that our knowledge about Lauraceae in Venezuelan 
Guayana is far from complete. 

The available data show that about one-third of 
the species (41 out of 128) are endemic to Guaya- 
na. Of these 41, 31 species occur on the slopes 
and summits of the tepuis, while 10 are lowland 
species. The number of endemic lowland species 
will likely diminish when adjoining Colombia and 
Brazil are better explored, because I expect that 
lowland species such as Ocotea tomentosa, Persea 
fluviatilis, and Phoebe semecarpifolia, which oc- 
cur near the borders with Colombia and Brazil, are 
not restricted to the Venezuelan side of the borders. 

w species, combinations, and the new 
name are arranged in alphabetical sequence. 


Licaria tomentosa van der Werff, sp. nov. TYPE: 
Venezuela. Bolivar: Chimantá Massif, along 
trail from Base Camp to Bluff Camp, western 
slopes of Chimantá-tepui, 1,100-1,700 m, 
Steyermark 75607 (holotype, NY; isotypes, 
COL, F). Figure 1. 


Arbor, ad 10 m. Ramuli teretes, ferrugineo-tomentosi, 
vetustiores glabrescentes. Folia chartacea, elliptica, 8-11 
x 3-4.5 cm, opposita, basi acuta, apice acuminata; supra 
secus costam nervosque aliquant um tomentosa; subtus 
ferrugineo-tomentosa. Nervi utroque costae latere 3- d 
versus marginem arcuati et conjuncti. Petioli 0.5-1 c 
longi, tomentosi, supra plani. Inflorescentiae Tien ig- 
noti. rune ellipsoideus, 1. dl cm. Cupula 1.2 cm 
alta ata, extus la margine leviter duplicata, 
3 stam wk bilocellatis et 3s stamin odiis praedita. Stamina 
vetera triangularia, locellis extrorso-apicalibus versus api- 
cem aperientibus. 


Tree, 10 m tall. Twigs terete, ferruginous to- 
mentose, becoming glabrous with age. Leaves firm- 
ly membranaceous, elliptic, 8-11 x 3-4.5 cm, 
opposite, the base acute, apex acuminate, upper 
surface with some tomentose pubescence along the 
midvein and lateral veins, otherwise glabrous, the 
lower surface ferruginous tomentose, the tomen- 
tum wearing off with age; lateral veins 3-5 pairs, 
immersed above, raised on lower surface, the lat- 
eral veins arching upwards and loop-connected. 
Petioles 0.5-1 cm long, tomentose, not canalicu- 
late. Inflorescence and flowers unknown. Infruc- 
tescence unknown. Fruit ellipsoid, 1.2 x 0.6 cm 
when dry. Cupule deeply cup-shaped, 1.2 cm high 
and 1.4 cm broad, the outside smooth, without 
lenticels; the margin weakly double-margined, with 
traces of tomentum and bearing on the inner mar- 
gin 3 2-celled stamens and 3 staminodia. Old, dried 


! [ thank the curators of F, G, ME 


Dwyer kindly checked the Latin descriptions, and John My 
9, St. Louis, Missouri 63166-0299, U.S.A. 


? Missouri Botanical Garden, P.O. Box 29 


¿RF, MY, MYF, NY, and VEN for making their specimens available. Dr. J. 


ers drew the fine illustrations 


ANN. MISSOURI Bor. GARD. 76: 462-475. 1989. 


Volume 76, Number 2 
1989 


van der Werff 
Lauraceae from Venezuelan Guyana 


463 


FIGURE 1. 
Leaf. — E. Detail of lower surface of leaf 


stamens triangular, the locelli extrorse-apical and 
opening towards the tips of the anthers. Staminodia 
similar in size and shape to stamens but without 


i. 

Although the type collection of Licaria tomen- 
tosa is in fruit, the combination of characters found 
on the holotype leaves no doubt about the generic 
identity. Indicative for Licaria are the three 2-celled 


Licaria tomentosa. — A. Habit. — B. Cupule.— C. Detail of rim of cupule, with old 


ERRE 3 
ASÍS 3 
MER EIC RS 


stamens, the double-rimmed margin of the cupule 
and the opposite leaves with few, loop-connected 
lateral veins which stay well away from the margin. 

he ferruginous-tomentose pubescence of this 
species readily separates it from the other Licaria 
species with opposite leaves, because these all have 
appressed pubescence (if any at all), usually with 
a lighter color. 


uoc 


stamen. — D. 


464 


Annals of the 
Missouri Botanical Garden 


I am not certain if Licaria tomentosa has only 
three staminodia; the staminodia present on the 
cupule represent whorl II and are only present 
when the tepals facing them have not fallen off. 
The tepals in front of the stamens (whorl III) are 
absent, and it is likely that staminodia representing 
whorl I would have fallen off as well. 

At present, Licaria tomentosa is known only 
from the holotype in NY (with detached cupules) 
and sterile isotypes in COL and F. Allen (1964) 
identified and cited these specimens as Ocotea ro- 
raimae Mez, a species with alternate leaves, four- 
celled anthers, a canaliculate petiole, without to- 
mentose pubescence, and lacking loop-connected 
lateral veins. I consider O. roraimae now part of 


O. aciphylla (Nees) Mez sensu lato. 


Ocotea flaventha van der Werff, sp. nov. TYPE: 
Venezuela. T. F. Amazonas: Dept. Atures, val- 
ley of Río Coro-Coro, W of Serranía de Yutaje. 
Tree, 28 m, flowers dull golden-yellow, Holst 
& Liesner 3443 (holotype, MO; isotypes, 
HBG, NY, VEN). Figure 2. 

Arbor, ad 30 m alta. dead hornotini angulati dense 


brunneo- anto losi, vetustiores teretes, indumento ad- 


neo-tomentella ir 


3-5 c cm, coriacea, alterna, supra n 


acuta margine 


s antheris paulo 
lo ongis, locellis Iac did 3 in- 


teriora longa, filamentis ca. m longis, fere 
será antheram aequantibus dubi liberis globos- 
is; sta ia non visa. Pistillodium ca. 2.4 mm longum, 


uA basi pubescens. Tubus floralis pubescens. Flos 
femineus: (partes florales super fructu juvenali prae- 
sentes): coda ca sh mm longa, filamento anthera 
perangustiore ongo; locellis stab Stylus 
triangularis, pubescens Due ctescentiae ad 8 cm longae. 
Cupula immatu PEU. lenticellata, ad 1. 5 cm lata, 
in pedicellum attenuata 


Tree, to 30 m tall. Twigs (especially when young) 
angled, becoming terete with age; young twigs with 
a dense, short, brown tomentum, this soon becom- 
ing appressed and finally disappearing on older 


twigs; inflorescence-bearing parts of twigs with con- 
spicuous scars from fallen inflorescences. Terminal 
bud brown-tomentellous to appressed pubescent. 
Leaves elliptic, 8-12 x 
ternate, smooth and lustrous above, pale green 


—5 cm, coriaceous, al- 


below, apex acute to shortly acuminate, the base 
acute with slightly revolute margin; glabrous above, 
with some appressed hairs below (especially near 
the base of the leaves and along the costa). Ve- 
nation pinnate, lateral veins 6-9 pairs, costa and 
lateral veins impressed above, slightly raised below, 
a fine reticulation rarely apparent (mostly on young 
leaves). Petioles appressed-brown-pubescent when 
young, becoming glabrous and black. 
cences brown-tomentellous to appressed pubescent, 
to 10 cm long, paniculate, the branchlets usually 
with several cymes near their tips; inflorescences 
frequently arranged on leafless apical parts of twigs, 
these to 15 cm long and with large scars of old, 


Inflores- 


fallen inflorescences, the entire complex appearing 
as one terminal inflorescence. Flowers unisexual, 
sessile or nearly so, yellow. Tepals 6, equal, ap- 
pressed-brown-pubescent, broadly ovate, 2.5 x 2.2 
mm, the inner surface pubescent, erect at anthesis. 
Male flower: stamens 9, all 4-celled, the cells ar- 
ranged in 2 rows. Outer 6 stamens ca. 1.6 mm 
long, the filaments ca. 0.8 mm long, slightly nar- 
rower than the anthers, the cells introrse; inner 3 
stamens ca. 2 mm long, the filament ca. 

long and about as wide as the anther; glands free, 
globose, basally attached; staminodia not seen. Pis- 
tillode ca. 2.4 mm long, pubescent, especially near 
the base; ovary and style poorly differentiated. 
Floral tube pubescent inside. Pistillate flower (based 
on remnants found on young fruits): staminodia 
ca. 0.8 mm long, filament and anther ca. 0.4 mm 
long, filament much narrower than anther, anther 
cells difficult to see and not opening. Filament 
dorsally pubescent near the base. Style deciduous, 
triangular, pubescent; stigma platelike. Top of young 
fruit pubescent. Infructescences to 8 cm long. Flo- 
ral parts persisting on young fruits, tepals slightly 
fused near the base and falling as a unit. Cupule 
deeply cup-shaped, enclosing ca. % of the young 
fruit, lenticellate, to 1.5 cm wide, narrowed into 
the swollen pedicel at the base. 


Paratypes. VENEZUELA. T. F. AMAZONAS: Dept. Atures, 
reds of Rio Coro-Coro, fr, Holst & Liesner 3402, 3421 
(MO, VEN). 


Ocotea flavantha is known only from the three 
type collections in T. F. Amazonas, Venezuela. The 
nearly sessile flowers, angled twigs, deep cupules, 
and erect tepals that tend to persist on young fruits 
place it in the group of O. glomerata. Three species 


Volume 76, Number 2 van der Werff 465 
1989 Lauraceae from Venezuelan Guyana 


C S nd 
RM 


qo 


Sim 
QS 
AB d 


Mp 


[t 


NSI deck 


M 

M “4 = 
TAY T 
DE X, AAA B 3 
NE M1 WT 

Lf 

| 


FIGURE 2. Ocotea flavantha. —A. Habit. —B. Flowers.— C. Young fruits. —D. Outer stamen.— E. Inner sta- 
men. —F. Pistillode. —G. Style and stigma from young fruit. — H. Staminode.—I, J. Leaf base, seen from below and 
above.— K. Stem showing scars from fallen inflorescences. — L. Leaf apex 


466 


Annals of the 
Missouri Botanical Garden 


of this group in the Venezuelan Guayana have a 
wide distribution (O. glomerata (Meissner) Mez, 
O. guianensis Aublet, and O. longifolia HBK), 
but these species have light-colored or very dense 
pubescence. The color of the indument and the 
smooth upper leaf surfaces suggest a relationship 
with the more restricted O. bracteosa (Meissner) 
Mez, O. cujumary Martius, and O. canaliculata 
(Rich.) Mez. These three species in Venezuela are 
known only from the eastern parts of T. F. Delta 
Amacuro and Estado Bolivar. Ocotea cujumary 
can be readily identified by its double-rimmed cu- 
pules; the other two species differ from O. flavan- 
tha in their glabrous ovary and style and in the 
absence of leafless, but inflorescences-carrying ter- 
minal parts of the twigs. The swollen pedicel sup- 
porting the cupule and the rather large (and still 
immature) cupules are additional distinctive char- 
acteristics. Differences in leaf shape are rather 
subtle. Ocotea canaliculata has the petiole poorly 
differentiated and lacks the weakly recurved leaf 
base, while O. flavantha has an obvious petiole 
and a slightly recurved leaf base. Ocotea bracteosa 
has larger and wider (to 15 x 6 cm) and more 
glaucous leaves than O. flavantha. 


Ocotea glabra van der Werff, sp. nov. TYPE: 
Venezuela. Bolivar: summit of Carrao-tepui, 
2,470-2,500 m, shrub 4-8 ft. tall, Steyer- 
mark 60897 (fl) (holotype, VEN; isotype, F). 
Figure 3E. 


Frutex ad 2.5 m altus. Ramuli ne glabri, teretes. 
Gemma terminalis glabra. Folia obovata vel anguste ob- 
ovata, 5-8 x 1.5-3 cm, alterna, sa n. aggregata, co- 
riacea, basi acuta, apice subacuta vel rotundata, glabra, 
supra polita. Venatio pinnata; nervi laterales 
r sub angulo 70-80? p 


spicatae vel anguste paniculatae, pauciflorae, ex axillis 
bractearum persistentium vel infrequenter foliorum ortae. 
glabri; tepala 8, et late elliptica, 
2 mm longa, 1. .7 mm lata; flos culinus: stamina 

9, 4-locellata, glabra, 6 ennt n locellis introrsis, fila- 
ento 3 interiora locellis inferioribus 
extrorsis, locellis superioribus lateralibus, filamenti base 
glandulis duabus connatis praedita; staminodia non visa; 
partilodnum lineare, ca. 0.8 mm lon PEE in flore femineo: 
staminodia ca. 0.7 mm longa, ovarium glabrum, ellip- 
soideum, ca. 2 mm longum, Minds capitata. Fructus 


ignotus 

Shrub, to 2.5 m tall. Twigs thick, glabrous, 
terete, with conspicuous leaf scars. Terminal bud 
glabrous, usually hidden by the leaves. Leaves ob- 
ovate to narrowly obovate, widest above the middle, 
5-8 x 1.5-3 cm, alternate, somewhat clustered 
along twigs, coriaceous, the base acute, the apex 


o 


3cm 


FIGURE 3. Leaf sha apes of Ocotea species with gla- 
brous PE: leaves, terminal buds, n: orescences, and 
flowers. —A. Ocotea cowaniana (Cowan 31106).— B. 
Oc i venosa (tate c 6).— 
ermark 97 bos 
10127).— E bs ae (Huber 12003) .—F. Oco- 
tea huberi (Huber 11008). 


very shortly acute to rounded, glabrous, the upper 
surface shiny, especially on young leaves, the lower 
surface less shiny. Venation pinnate, lateral veins 
6-9 pairs, departing from the midvein under an 
angle of 70-80", raised on the upper surface and 
immersed or nearly so on the lower surface; margin 
often slightly revolute. Petiole glabrous, to 2 mm 
long, drying dark. Inflorescences glabrous, ca. 6 
cm long, few-flowered, mostly in the axils of per- 
sistent bracts, these 5-7 mm long, narrowly ellip- 
tic, glabrous. Flowers unisexual, glabrous; stami- 
nate flowers with tepals erect at anthesis; tepals 6, 
equal, broadly elliptic, ca. 2 mm long, 1.7 mm 
wide; stamens 9, 4-celled, glabrous, the outer 6 
stamens with introrse cells arranged in 2 rows, 
filament and anther both ca. 0.6 mm long; inner 


Volume 76, Number 2 
1989 


van der Werff 467 


Lauraceae from Venezuelan Guyana 


stamens with lower cells extrorse, the upper ones 
lateral, the base of the filament with 2 large, fused 
glands; staminodia not seen; pistillode present, lin- 
ear, ca. 0.8 mm long; pistillate flowers with stami- 
nodia ca. 0.7 mm long, the ovary ellipsoid, ca. 2 
mm long, glabrous, style lacking, stigma capitate. 
Fruit unknown. 


VENEZUELA. BOLÍVAR: Municipio Gran Sa- 
O m, fl, Huber & Picon 


Paratype. 
bana, Caraurén-tepui, ca. 1,95 
12003 (MYF). 

Ocotea glabra is known only from two collec- 
tions from the summits of Carrao- and Caraurén- 
tepuis. It is characterized by being completely gla- 
brous, its (narrowly) obovate leaves, and its ve- 
nation with lateral veins departing at almost 90? 
from the midvein. This last-mentioned character 
occurs also in O. wurdackiana Allen, a species 
restricted to the tepuis of Estado Bolivar. Allen 
(1964) cited the type collection of O. glabra as a 
paratype of O. wurdackiana and described O. 
wurdackiana as having hermaphrodite flowers. 
Recent collections have shown that O. wurdack- 
iana has unisexual flowers, like O. glabra. The 
two species, although superficially similar, can be 
separated as follows: O. glabra is entirely glabrous 
and has the lower leaf surfaces smooth, while 
wurdackiana always has a pubescent terminal bud, 
the inner surface of tho tepals densely pubescent, 


Ilat 
mate. 


and the lower leaf 
Ocotea glabra has its closest bie amon 
the ten unisexual Ocotea species in the Venezuelan 
Guayana that share glabrous twigs, leaves, and 
terminal buds. Two of these have pubescent flowers 
or inflorescences (O. myriantha (Meissner) Mez 
and O. perrobusta (Allen) Rohwer) and are not 
very closely related. Ocotea myriantha is a low- 
land species, rather common in southern T. F. 
Amazonas, whereas O. perrobusta is restricted to 
Auyan-tepui, Estado Bolívar. The remaining species 
in this group have a restricted distribution and are 
known only from the upper slopes or summits of 
one or two sandstone mountains. Three have rather 
large, many-flowered inflorescences: O. crassifolia 
(Sarven-tepui, Roraima), O. neblinae (Neblina), 
and O. basirecurva (Serranía de Yutaje, Cerro 
Coro-Coro). Short, almost spicate, and few-flowered 
inflorescences, also strongly coriaceous leaves, 
characterize the last large group of species. Several 
t not all species in this group have glaucous, 
clustered, almost sessile leaves, shiny upper leaf 
surfaces, and lobed or toothed cupules. This group 
can be subdivided into (1) species from Estado 
Bolivar, characterized by immersed or elevated 
costa and lateral veins but without a raised reticula- 


tion (O. erectifolia (Allen) van der Werff, Cerro 
Jaua; O. glabra van der Werff, Carrao-tepui and 
Caraurén-tepui; O. roseopedunculata van der 
Werff, Acopan-tepui), and (2) species from T. F. 
Amazonas with weakly developed lateral veins and 
raised reticulation on upper and/or lower leaf sur- 
face (O. venosa Gleason, Cerro Duida, Marahuaca; 
O. cowaniana Allen, Serrania Paru; O. huberi van 
der Werff, Cerro Coro-Coro, Serrania Guanay). 
Ocotea glaucophylla Moldenke, based on material 
from Cerro Duida, is not well understood; to this 
small-leaved species also belong collections from 
Chimantá, revealing an unusual distribution pat- 
tern. With the exception of O. glaucophylla, these 
species are well defined by their inflorescence and 
leaf characters (Fig. 3); they are also geographi- 
cally separated, and the paucity of collections is 
probably due to occurrence in inaccessible places. 
The collection Maguire 33026 (NY) from Cerro 
Guaiquinima belongs in this species group (it has 
glabrous leaves, twigs, and terminal bud, and clus- 
tered, glaucous, coriaceous, subsessile leaves) and 
differs from the other species in its smooth upper 
leaf surface; it is almost certainly undescribed, but 
because it is sterile, I do not wish to describe it. 


Ocotea huberi van der Werff, sp. nov. TYPE: 
Venezuela. Bolivar: Distr. Cedeno, Serrania 
Guanay, ca. 1,700 m, Huber 11008 (holo- 
type, MO; isotype, MYF n.v.). Figure 3F. 


rutex, Bas 3m Mi Ramuli glabri, juniores suban- 
li 


gu ulares, vetustior re emma terminalis non visa, 
n. necta, Folia aplico: -oblonga vel leviter obovata, 
-13 -8 cm, coriacea, alterna, basi rotunda vel 


m apice rotunda vel apiculata, supra polita, prae- 
sertim in foliis novellis, subtus subpolita, glauca. Venatio 
pinnata, nervis lateralibus s utroque costae Aes 6-8, prope 

margi m versus apicem curvatis coalitisque; costa utrin- 
que immersa; nervis lateralibus et reticulatione elevata, 
sed foliis veteribus paulo elevatis. Petioli glabri, crassi, ad 


m glabris; staminibus 9, 4-locellatis, eis seriei 
primae ca. 1.6 mm longis, antheris ca. 0.9 mm lo 
locellis introrsis, filamentis ca. 0.5 mm latis, eis 
secundae ca. 1.0 mm longis, Iouis introrsis, filamentis 
ca. 0.2 mm latis, eis seriel tertiae ca mm longis, 
antheris rectangularibus, ca. 0.8 x 0.6 mm, locellis la- 
teralibus, filamentis basi 2 glandulis magnis praeditis; 
staminodiis 3, ca. 0.5 mm longis, linearibus, basi pubes 
centibus; ovario clipsvideo, ca. 1.3 mm longo, iae in 
evem poi rum: stigmate capitato. Cupula non 
ata, margine 6-lobato vel 6- dans 
sensim in catalan attenuata. Fructus ovaloideus, 1.5 
X ] cm. 


Shrub, 1-3 m tall. Twigs glabrous, subangular 
when young, becoming terete. Leaves elliptic-ob- 


468 


Annals of the 
Missouri Botanical Garden 


long to slightly obovate, 8-13 x 5-8 cm, alter- 
nate, + evenly distributed along twigs, coriaceous, 
the base rounded to cordate, the apex rounded to 
apiculate, the upper surface shiny, especially the 
young leaves, the lower surface less so, glaucous. 
Venation pinnate, the lateral veins 6-8 pairs, curv- 
ing upward towards the margin and becoming con- 
nected; midrib immersed on upper and lower sur- 
faces, lateral veins and tertiary venation raised and 
forming a coarse reticulum, this less visible on older 
leaves. Petioles glabrous, thick, to 3 mm long, 
blackish when dry. Inflorescences glabrous, in the 
axils of persistent leaves, 12-1 
ing the leaves, narrowly paniculate. Flowers gla- 
brous, 5-6 mm tepals 6, equal, ovate, 2- 
2.2 mm long, 1.4 mm wide, the inner 3 with a 
few hairs on the inner surface, otherwise glabrous, 
reflexed in old flowers; stamens 9, the anthers 
4-celled; outer 3 (whorl I) ca. 1.6 mm long, the 
anther ca. 0.9 mm long, cells introrse, the filament 
ca. 0.5 mm wide; stamens of whorl II ca. 1.0 mm 
long, cells introrse, the filaments ca. 0.2 mm wide; 
stamens of whorl III ca. 1.4 mm long, the anther 
rectangular, ca. 0.8 mm long, 0.6 mm wide, cells 
lateral; glands at base of filaments free, ca. 0.5 
mm long; staminodes 3, linear, ca. 0.5 mm long, 
pubescent near base; ovary ellipsoid, ca. 1.3 mm 
long, gradually narrowed into short style, 0.3 mm 
long; stigma large, capitate. Cupule a shallow cup, 
6 teeth or lobes at the margin 
and gradually narrowed into the pedicel. Fruit oval- 


cm long, exceed- 


diam.; 


ca. 1 cm wide, with 


, 1.5 X 1 cm fide collector. 
Paratypes. VENEZUELA. T. F. AMAZONAS: Depto. 


Atures, cumbre del Cerro Coro-Coro, 2,200 m, Huber 
12298 (MO, MYF). 


Ocotea huberi is restricted to the Departamento 
Atures and Distrito Cedeño, along the border of 
Estado Bolivar and T. F. Amazonas. Its closest 
relative is O. venosa Gleason, a species only known 
from the type collection from Cerro Duida and a 
fruiting specimen from Cerro Marahuaca, both in 

. Amazonas. The two species have similarly 
id reticulation, toothed or lobed cupules, and 
swollen fruiting pedicels. They differ in their leaf 
bases, which are obtuse in O. venosa vs. rounded 
to cordate in O. huberi, and in the much shorter 
inflorescences of O. venosa. The types of O. venosa 
(NY, US) do not have open flowers, and I have 
not compared their floral details; I accept O. huberi 
as a distinct species based on its striking leaf shape 
and long inflorescences. 

The group of species to which Ocotea huberi 
belongs is characterized by unisexual flowers. The 
MO sheet of Huber 11008 has two twigs, one in 


young fruit, the other in flower. Dissection showed 
the flowers to be staminate (anthers with well-de- 
veloped locelli; pistil hollow). Thus, the flowering 
and fruiting twigs on the type sheet probably came 
from different plants, and I select the flowering 
twig as the holotype. ipsias ee of O. huberi 
are further discussed under O. gla 


Ocotea julianii van der Werff, nom. nov. Based 
on Phoebe steyermarkiana Allen, Mem. New 
York. Bot Gard. 10(5): 74. 1964. TYPE: Stey- 
ermark & Wurdack 921 (holotype, NY; iso- 
types, F, MO). 


Ocotea julianii was erroneously described as 
belonging to the genus Phoebe by Allen. The large, 
cup-shaped cupule excludes this species at once 
from Phoebe, a genus in which the fruit is sub- 
tended by a small cupule with the tepals usually 
persisting. Leaf venation, the large cupule, and the 
deep floral tube in old flowers all point to the Ocotea 
aciphylla group (I have not seen good flowering 
material). Because Allen already described Ocotea 
steyermarkiana, a new epithet is needed when 
Phoebe steyermarkiana is transferred to Ocotea. 

The similarity between Ocotea julianii and 
Phoebe tetragona (Meissner) Mez pointed out by 
Allen (1964) is superficial. Fruiting specimens of 
the latter species have a cupule typical of Phoebe, 
and the leaves and flowers are glabrous. Distri- 
bution also argues against a close relationship, since 
P. tetragona is a cerrado species from Minas 
rais, Brazil. I regard O. julianii as closely related 
to O. revoluta Moldenke, a poorly known species 
from Cerro Duida. This species differs in its nar- 
rower, generally smaller leaves with revolute mar- 
gins and in its smaller, funnel-shaped cupules; both 
species belong to the O. aciphylla group. 


Ocotea liesneri van der Werff, sp. nov. TYPE: 
Venezuela. T. F. Amazonas: Dept. Río Negro, 
Cerro Neblina Camp IV, 780 m, Liesner 
16687 (holotype, MO; isotypes, HBG, NY, 
VEN) 


Arbuscula, ad 5 m altus. Ramuli teretes (novelli paulo 
angulati), glabri. Gemma terminalis glabra. Folia late el- 
liptica vel late ovata, 10-17 x 7-11 


s L subtus Aisa 


Inflorescentiae glabrae, ad 10 cm 
longae, foliis breviores, paniculatae, ramulis basalibus 2- 
5-floris, apicalibus unifloris. Pedicelli glabri vel paucis pilis 


Volume 76, Number 2 
1989 


van der Werff 469 


Lauraceae from Venezuelan Guyana 


praediti, 3-4 mm longi. Flores albidi, ad 4 mm diametro, 
tubo florale infundibuliforme, conspicuo. Tepala 6, ae 
qualia, elliptico-ovata, ca. 1.7 mm longa, intus parte in- 
feriore pubescentia. Stamina 9, 4-locellata, locellis 2 se- 
riebus dispositis; 6 exteriora ca. 1.4 mm longa, filamento 
0.7 mm longo, locellis introrsis, filamento pubescente; 3 
interiora 1.3 mm longa, filamento 0.7 mm longo, pubes- 
cente; glandulis Breviter dmca 0.3 mm diametro, e 
DETI oribus la 
nodia 3, linearia, bbdd apice incrassata et reni 
Ovarium globosum, 1.1-1.2 mm in diametro, in tubo 
florale intus aine inclusum; style 0.8 mm longo, 
stigmate non incrassato. Fructus ignotus. 


teralibus Stam 


Small tree, to 5 m tall. Twigs terete or slightly 
angled when young, glabrous. Terminal bud gla- 
brous. Leaves broadly elliptic to broadly ovate, 10— 
17 x 7- cm, coriaceous, glabrous, alternate, 
evenly distributed along twigs, the tip rounded or 
acute, the base obtuse or rounded, the margin 
slightly revolute, both surfaces dull. Venation pin- 
nate, lateral veins 5-7 pairs, arching upwards and 
disappearing near the margin, midvein and lateral 
veins immersed on upper surface, raised on lower 
surface, the lower surface with a slightly raised, 
fine reticulation, this less visible on the upper sur- 
face. Petioles glabrous, 1.5-2 cm long, somewhat 
canaliculate. Inflorescences glabrous, ca. 10 cm 
long, shorter than the leaves, paniculate, the lower 
branchlets with 2-5 flowers, otherwise flowers sin- 
gle, the inflorescences few-flowered. Pedicels gla- 
brous or with very few hairs, 3-4 mm long. Flowers 
whitish, with a few appressed hairs on the outside, 
4 mm diam., floral tube funnel-shaped, conspicu- 
ous. Tepals 6, equal, elliptic-ovate, ca. 1.7 mm 
long, the inner surface pubescent in the lower half. 
Stamens 9, all 4-celled, the cells arranged in 2 
rows; outer 6 ca. 1.4 mm long, both filament and 
anther 0.7 mm long, the cells introrse, filament 
rather densely pubescent; inner 3 stamens 1.3 mm 
long, the filament 0.7 mm long, pubescent; glands 
shortly stipitate, 0.3 mm diam.; lower pair of cells 
extrorse, the upper pair lateral. Staminodia 3, lin- 
ear, ca. 0.6 mm long, pubescent, but the tip gla- 
brous and slightly swollen. Ovary globose, 1.1-1. 
mm diam., enclosed in the well-developed floral 
tube, this with pubescent walls; style distinct, ca. 
0.8 mm long, not widened into a distinct stigma. 
Fruits unknown. 


Although Ocotea liesneri, known only from the 
type collection, is in all vegetative parts glabrous 
like Ocotea erectifolia and allies, these species are 
probably not closely related. Ocotea liesneri differs 
from the O. erectifolia group, for instance, by 
having well-developed petioles, a rather lax inflo- 
rescence (comparable to but smaller than that of 
O. basirecurva), relatively long and pubescent fil- 


aments, and flowers apparently hermaphroditic. 
The last-mentioned character, difficult to ascertain, 
needs verification from additional collections. Oco- 
tea liesneri seems quite isolated. In the Venezuelan 
Guayana, the number of hermaphroditic Ocotea 
species is not great; most belong to the O. aci- 
phylla group, which always has pubescent flowers 
and a quite different venation, and the O. oblonga 
group, which has entirely different leaves. The 
remaining species are also isolated and possess some 
striking characters; they include O. rubra Mez, O. 
cymbarum esmeraldana Moldenke, and 
O. fasciculata (Nees) Mez. 


Ocotea megacarpa van der Werff, sp. nov. TYPE: 
Venezuela. T. F. Amazonas: Depto. Atabapo, 
Cerro Marahuaca, 1,220-1,350 m, tree, 15 
m tall, Steyermark & Holst 130610 (holo- 
type, MO; isotype, VEN) 


Arbor, ad 15 m alta. Ramuli juniores cinnamomeo- 
tomentosi, parum angulosi, vetustiores glabrescentes 
teretesque. Folia coriacea, 15-22 x 5-9 cm, alterna, 


rescentiae floresque ignoti. Fructus ellipsoideus, 5-6 x 
3.5-4 cm. Cupula ad 4 cm lata, 1.5 cm profunda, ver- 
rucosa. 

Tree, 10-15 m tall. Twigs brown-tomentose and 
weakly angled when young, becoming glabrous and 
terete with age. Leaves stiff, leathery, 15-22 x 
5-9 cm, alternate, elliptic, frequently condupli- 
cate, the base obtuse or acute, the apex acuminate, 
the upper surface glabrous and with the venation 
immersed, the lower surface brown-tomentose when 
young, glabrous with age, the midvein prominent, 
lateral veins slightly raised, in 5-8 pairs, arching 
upwards and fading away close to the margin. 
Petioles 1.5-2 cm long, canaliculate, brown-to- 
mentose when young, glabrescent with age. Inflo- 
rescences and flowers unknown. Infructescences 
with few fruits, glabrous, to ca. 7 cm long. Fruit 
ellipsoid, 5-6 cm long, 3.5-4 cm wide. Cupule 
(immature) a cup ca. 3 cm wide and 1.5 cm deep, 
the outside ridged and warty, the inside glabrous 
or with a few appressed hairs; mature cupules 
shallower, 3.5-4 cm wide and to 1.5 cm deep, the 
outside ridged and warty. 

Paratypes. | VENEZUELA. a F. AMAZONAS: Depto. Ata 
bapo, Cerro Marahuaca, 1,140 m, Steyermark & Holst 
130555 (MO, VEN); same ae 1,225 m, Liesner 
17732, 17733 (MO, VEN). 

tea megacarpa is known only from three 
fruiting collections from Cerro Marahuaca. Leaf 


470 


Annals of the 
Missouri Botanical Garden 


shape, venation, canaliculate petioles, and the ini- 
tially deep cups all point to the Ocotea aciphylla 
group. Within this group (and within the genus 
Ocotea), the large fruits and large, shallow cupules 
are unmatched, and O. megacarpa is solely de- 
scribed on the distinctiveness of its fruits and cu- 
pules. In the Ocotea aciphylla group, O. julianii 
also has large fruits (3.5 x 3 cm) and cupules (ca. 
3 cm wide). However, in addition to its smaller 
fruits and cupules, O. julianii differs in its rounded 
to subcordate leaf bases, short, thick petioles, and 
blunt to rounded leaf apices. Ocotea megacarpa 
is in leaf shape (but not in indumentum) very similar 
to O. aciphylla (Nees) Mez s.l., 
lowland populations in T. F. Amazonas, which have 
been identified as O. costulata (Nees) Mez in the 
past, but this species has appressed pubescence, 
cupules to 2 cm wide, and fruits to 2.5 cm long. 


especially to the 


Ocotea roseopedunculata van der Werff, sp. 

YPE: Venezuela. Bolivar: Distr. Piar, 

summit area of Acopan-tepui, ca. 1,950 m 

fl, Huber, Ahti & Pipoly 10127 (holotype, 
MO; isotype, MYF n.v.). Figure 3D. 


ad 2.5 m altus. Ramuli glabri, cristati petiolis 
cb foliorum cicatricibus conspicuis praediti. 
emma terminalis glabra. Folia elliptica vel anguste ellip- 
tica, raro subovata vel subobovata, 7-11 x 2.5-4.0 cm, 
coriacea, glabra, basi apiceque acuta, alterna sed plerum- 
ibus inter folios con- 


m longi, in sicco nigri. Inflorescentiae gla- 
brae, roseae, folios aequantes vel eis paulo longiores, ex 
axillis foliorum vel bractearum ortae. Pedicelli glabri, ad 
5 mm longi, basi bractea lineare ad 0.5 mm longa praediti. 
Flores vindi flavi. Tepala aequalia, ovata, 2.5 x 1.8 
mm. Stamina 9, 4- locellata, locellis i in 2 seriebus dispositis; 


.1 mm, locellis introrsis; 3 interiora ca. 1.5 
mm longa, anthera ca. 1 mm longa, locellis lateralibus 
vel laterali-extrorsis, glandulis magnis, liberis, ca. 0.6 mm 
in diametro; staminodia non visa; pistillodium parvulum, 
lineare, ad 0.3 mm longum. Flores feminei et fructus 
ignoti. 


Shrub, to 2.5 m tall. Twigs ridged due to de- 
current leaf bases and with conspicuous scars from 
fallen leaves. Terminal bud glabrous. Leaves elliptic 
to narrowly elliptic, or rarely somewhat ovate or 
obovate, 7-11 O cm, coriaceous, gla- 
brous, base and tip acute, alternate, mostly clus- 
tered, the few leaves between the leaf clusters 
usually smaller, the upper surface shiny, the lower 
surface dull, glaucous. Venation pinnate, lateral 


veins 6-10 pairs, the basal ones ascending, the 
middle and distal ones leaving the midvein under 
blunter angles, near the margin curving upwards 
and becoming loop-connected, midvein and lateral 
veins raised on the upper surface but not on lower, 
tertiary venation immersed or nearly so on both 
sides. Petioles thick, glabrous, to 5 mm long, black 
when dry. Inflorescences glabrous, pink, about as 
long as the leaves or slightly longer, spicate or the 
lower branches with 2 or 3 flowers, in the axils of 
leaves or bracts. Pedicels glabrous, to 5 mm long, 
with a linear bract ca. 0.5 mm long at the base. 
Flowers yellow, glabrous. Tepals 6, equal, ovate, 
2.5 x 1 tamens 9, all 4-celled and the 
cells arranged in 2 rows; outer Ó stamens sessile 
or nearly so, the filament to 0.3 mm long, the 
anther 1.3 x 1.1 mm, cells introrse; inner 3 sta- 
mens ca. 1.5 mm long, anther ca. 1 mm long, the 
cells lateral or lateral-extrorse, glands large, free, 
ca. 0.6 mm diam.; staminodia not seen; pistillode 
very small, linear, ca. 0.3 mm long, completely 
idden by stamens and glands, stigma not devel- 
oped. Pistillate flowers and fruits not known. 


VENEZUELA. BOLÍVAR: Distr. Piar, summit 
950 m, fl, Huber, Ahti & 


Paratype. 

ea of Acopán-Tepui, ca. 
Pipoly 10120 (MO). 

Ocotea roseopedunculata is restricted to Aco- 
pán-tepui and is currently known from only two 
collections. Its nearest relative is Ocotea glabra, 
known from Carrao and Caraurén-tepuis in Estado 
Bolivar. These two species are best separated by 
their leaf characters: O. roseopedunculata has 
elliptic leaves with mostly ascending lateral veins, 
while O. glabra has (narrowly) obovate leaves with 
the lateral veins leaving the midrib under a blunter 
angle and more rounded leaftips. In sme ae there 
are some small floral differences: in O. glabra the 
filaments and anthers of the outer stamens are 
about equally long and the pistillode is ca. 0.8 mm 
ong; in O. roseopedunculata the anthers are sub- 
sessile and the pistillode is smaller, 0.3 mm long. 
The two species share ridged stems, conspicuous 
leaf scars, almost spicate, few-flowered inflores- 
cences, and a tendency toward clustered leaves. 
See also discussion under O. glabra. 


Ocotea tomentosa van der Werff, sp. nov. TYPE: 
Venezuela. T. F. Amazonas: San Carlos de Rio 
Negro, IVIC main study site, km 4.3 on Solano 
Road, small tree 3 m high, flowers white, Clark 
& Maquirino 6802 (holotype, MO; isotypes, 
HBG, NY). 


rutex vel arbor parva, ad 3 m alta. Ramuli angulati, 
hornotini brunneo-tomentosi, vetustiores tomentelli, de- 


Volume 76, Number 2 
1989 


van der Werff 471 


Lauraceae from Venezuelan Guyana 


nique glabrescentes. Gemma terminalis brunneo-tomen- 
x 6- , a aie seg Os 
subtus 


e 

o superiore consociatis; venatione ue 

immersa, costa die nervis subtus elevatis, indumento 

neo venationis tomento albido laminae anya Petioli 
.5 em longi, brunneo-tomentelli vel subglabri. ien flo- 

rescentiae axillares, brunneo- tomentosae, ad 10 cm lon- 

ongi, tomentosi. 


et minus pubescentia quam inflorescentia, intus subglabra; 
staminodia 9, ca. 0.3 m m longa, glabra; ovarium ca. 
mm diametro, de inconspicuum, stigma capitatum 
Fructus globosus, ca. 1 cm diametro, cupula non profun- 
da, pedicello un. petalis persistentibus. 


Shrub or small tree, to 3 m tall. Twigs angular, 
brown-tomentose when , becoming tomen- 
tellous and finally glabrous. Terminal bud brown- 
tomentose. Leaves elliptic, 15-25 x 6-8 cm, al- 
ternate, coriaceous, glabrous above except some 
pubescence on midrib, the lower surface with brown 
curled hairs along midrib and main veins, the leaf 
tissue white-tomentose, turning grayish with age, 
base and apex acute, lateral veins 6-8 pairs, these 
arching upwards and becoming loop-connected wit 
the distal vein; venation immersed on the upper 
surface, costa and lateral veins raised on lower 
surface, the venation with brown indumentum con- 
trasting with the white tomentum on the laminar 
tissue. Petioles ca. 1.5 cm long, brown-tomentel- 
lous to glabrous. Inflorescences axillary, brown- 
tomentose, to 10 cm long, paniculate. Pedicels ca. 
1 mm long, tomentose. Flowers white (only pistillate 
seen). Tepals 6, equal, ca. 1 mm long, ovate, with 
some gray hairs on the outside and pubescence on 
tepals much less than on inflorescence or pedicels; 
inside of tepals with a few hairs or glabrous; sta- 
minodia 9, ca. 0.3 mm long, glabrous; ovary ca. 
] mm diam., style not evident, stigma capitate. 
Fruit a globose berry, ca. 1 cm diam., the cupule 
shallow, pedicel swollen and the petals persisting 
as lobes on the margin of the cupule. 


Paratypes. | VENEZUELA. T. F. AMAZONAS: San Carlos 

e Río Negro, IVIC main study site, fl, Clark & Ma- 
quirino 8216 (MO); Río Guainía, ca. 7 of Maroa, 
frequent in shrub savanna, fr, Maguire et al. 41755 
(NY). 


Ocotea tomentosa is known only from the upper 
drainage of the Río Negro between Maroa and San 
Carlos de Rio Negro. Although the area around 
San Carlos de Río Negro has been reasonably well 
explored, O. tomentosa has only been collected 
there twice (both collections from the same marked 


tree) and is apparently rare. It is very likely present 
in Colombia and can be expected in Brazil. 

The angled twigs, somewhat clustered flowers 
and, above all, scarce pubescence on the tepals in 
comparison with that on the inflorescence point to 
the Ocotea leucoxylon group as the closest rela- 
tives. None of the species in that group has leaves 
with dense abaxial tomentum or such persistent 
tepals in the fruiting stage; moreover, O. tomentosa 
lacks the white lenticels on the cupule that so 
readily identify the O. leucoxylon group. 


Persea areolatocostae (Allen) van der Werff, 
comb. nov. BASIONYM: Phoebe areolatocostae 
C. K. Allen, Mem. N.Y. Bot. Gard. 10: 75 
1964. 


The type collection of Phoebe areolatocostae 
has buds and immature fruits. However, the fruits 
are diseased; the relatively long petioles, leaf shape, 
and leaf reticulation all point to the genus Persea. 
The equal tepals are a character infrequently found 
in Persea; P. americana and a few related species 
share this character, but P. areolatocostae is prob- 
ably not related to those species. In my opinion, 
the relationship is with a small group of species 
characterized by equal tepals, a small, round fruit, 
and persistent tepals in the fruiting stage. This 
group includes the neotropical P. steyermarkii, P. 
rigens, P. silvestris, and a few additional, as yet 
undescribed, species. It is not yet clear to which 
degree this group is related to the Asian Persea 
subg. Machilus. 


Persea croatii van der Werff, sp. nov. TYPE: 
enezuela. T. F. Amazonas: Cerro Neblina, 
near Canon Grande at Camp No. 7, ca. 1,800 

m, Croat 59519 (holotype, MO). 


Arbor, ad 4 m. Ramuli teretes, sparse adpresse pu- 

escentes, vetustiores glabri. Folia pde subcoriacea, 
elliptica, 7-10 cm, basi acuta, apice obtusa; supra 
glabra, die adpre esse  Pubescen tia, sed superficie folio- 
non o dg a, costa nervisque supra im- 
mersis, sibus costa eleva a sed nervis immersis vel per- 
atis. Petioli escis 1-2 cm longi, glabri 
vel iler adpresse pu entes. Inflorescentia axillaris, 
subterminalis, paniculata m longa, adpresse ci- 
nereo-pubescens. Pedicelli ca. 4 mm longi. Tepala 6, 
inaequalia, 3 exteriora ca. 4 mm longa, (anguste) ovata, 


cellis introrsis, famen tis Wai antheris glabris, 
a. m longis; 3 interiora locellis lateralibus vel la- 
ee A ubescentibus, ca. 3 mm longis, 
ig glandulis praeditis, antheris glabris, ca. 1.5 mm 
longis; ed , ca. 1 mm longa, sagittata, pubes- 
centia. Ovarium glabrum vel apice paucis pilis praeditum, 


472 


Annals of the 
Missouri Botanical Garden 


globosum, ca. 1.5 mm diametro; stylo ca. 3 mm longo, 
pubescente. Fructus ignotus. 


Tree, 4 m tall. Twigs terete, sparsely appressed 
pubescent, becoming glabrous with age. Leaves 
alternate, subcoriaceous, elliptic, 7-10 x 2-3 cm, 
evenly distributed along twigs, the base acute, apex 
obtuse; glabrous on the upper surface, appressed 
pubescent below, but pubescence not obscuring the 
leaf surface; costa and lateral veins (6-9 pairs) 
immersed on upper surface; costa raised on lower 
slightly 
raised on lower surface. Petioles canaliculate, 1— 
2 cm long, glabrous or with some appressed hairs. 
Inflorescence axillary, pseudoterminal, paniculate, 
ca. 12 cmlong, appressed gray-pubescent. Pedicels 
ca. 4 mm long. Tepals 6, unequal; outer 3 ca. 4 
mm long, ovate to narrowly ovate, appressed pu- 


surface, lateral veins immersed or ver 


bescent outside, glabrous inside; inner 3 ca. 6 mm 
long, narrowly ovate, appressed pubescent on both 
surfaces. Fertile stamens 9, 4-locellate, the outer 
6 with introrse cells, 4.5-5 mm long, filaments 
densely pubescent, anthers glabrous, ca. 1.6 mm 
long; inner 3 stamens with lateral or lateral-ex- 
trorse cells, 4.5-5 mm long, the filaments ca. 3 
mm long, densely pubescent, each with 2 glands 
attached near the base, anthers glabrous, ca. 1.5 
mm long. Filaments of all stamens about as wide 
as anthers or slightly narrower. Staminodia 3, ca. 
| mm long, sagittate, pubescent. Ovary globose, 
ca. 1.5 mm diam., glabrous except for some hairs 
near the tip; style ca. 3 mm long, pubescent. 


Persea croatii is known from only one collection 
from the upper slope of Cerro Neblina. The pu- 
bescent style and the few apical hairs on the ovary 
place it in Kopp's (1966) section Aurataea, which 
includes two other species from the Venezuelan 
Guayana. Persea grandiflora, known from the 
lower slopes of Cerro Neblina, differs in having 
larger flowers, prominently raised lateral veins on 
the lower leaf surface, ovate (vs. elliptic) leaves, 
and much denser pubescence. Persea maguirei 
differs in having smaller leaves, denser pubescence 


croatii differs in its sparser, noncupreous pubes- 
cence on the lower leaf surface and its noncupreous 
pubescence on flowers and inflorescence. 


Persea croizatii van der Werff, sp. nov. TYPE: 
Venezuela. mazonas: upper Rio Ori- 


noco, Croizat 918 (holotype, F). Figure 4. 


Arbor vel frutex. Ramuli teretes, juvenales sparse ad- 
presse pubescentes, glabrescentes. Folia alterna, elliptica, 
9-14 x 4-6 cm, firme chartacea, margine paulo reflexa, 
base apiceque acuta, supra glabra, subtus pilis adpressis 

u 


sve gla- 

bri, piu gi. Inflorescentiae anguste paniculatae, 

10-15 cm longae, adpresse pubescentes. Pedicelli ca. 2 

mm longi, eso se pubescentes. Tepala 6, inaequalia; 3 
exteriora late pue intus glabra, ca long : 

nteriora ovata, intus adpresse pubescent - 


pa Stamina 9, 4 locellata; 6 leeis locellis ostia 
. 2 mm longa, filamentis pubescentibu us, ca. 1.1 mm 
lo ongis, ide glabris, c ca. 0.9 mm longis, filamentis quam 
antheris angustioribus; 3 interiora locellis lateralibus vel 
extrorsis, ca. 2. 5m m longa, filamentis pubescentibus, basi 
i and  preeditis, ca. l. i 
0.6 mm 
1 mm 
varium glabrum, globosum; E gracile, ca. 1.6 mm 
dia Fructus globos sus, ca. 8 mm dapes tepala per- 
sistentia, paten 
Woody plant of unknown size. Twigs terete, with 
some appressed pubescence when young, soon be- 
coming glabrous. Leaves alternate, elliptic, 9-14 
X 4—6 cm, firmly chartaceous, the margin slightly 
reflexed, base and apex acute, glabrous above, with 
some appressed hairs below, especially along mid- 
vein and lateral veins, or almost glabrous, lateral 
veins 5-6 pairs; midvein and lateral veins im- 
mersed above, raised on lower surface; final retic- 
ulation not or very slightly raised. Petioles glabrous 
or with a few appressed hairs especially near the 
base, 2-3 cm long. Inflorescences narrowly panic- 
ulate, 10-15 cm long, about as long as the sub- 
tending leaves, appressed pubescent. Pedicels ca. 
2 mm long, pubescent. Tepals 6, unequal; outer 
3 broadly ovate, appressed pubescent outside, gla- 
brous inside, ca. 3 mm long, inner 3 appressed 
pubescent on both surfaces, ovate, 5-6 mm long; 
tips of inner tepals often breaking off in old flowers 
or in fruiting stage. Fertile stamens 9, all 4-celled; 
those of the outer 2 whorls with introrse cells, ca. 
2 mm long, the filaments pubescent, 1.1 mm long, 
the anther glabrous, ca. 0.9 mm long, the filaments 
distinctly narrower than the anthers; inner 3 sta- 
mens linear, ca. 2.5 mm long, the filaments pu- 
bescent, ca. 1.9 mm long, with 2 globose glands 
attached slightly above the base, the anther gla- 
brous, ca. 0.6 mm long, slightly wider than the 
filament; staminodia 3, pubescent, ca. 1.1 mm 
long, sagittate. Ovary glabrous, style slender, ca. 
1.6 mm long, stigma platelike. Fruit globose, ca. 
8 mm diam., with glaucous bloom; tepals persistent, 
spreading. 
Persea croizatii is known only from the type 
collection, made in July-October 1951, along the 


Volume 76, Number 2 
1989 


van der Werff 473 


Lauraceae from Venezuelan Guyana 


IGURE 4. Persea croizatii. — A. Habit. — B. Old flowers. — C. Fruit. — D. Inner stamen. — E. Arrangement of 
anther cells of inner stamen. — F. Outer stamen.—G. Staminode. — H. Pistil. 


upper Rio Orinoco, probably at low elevation. There 
are no label data indicating habit, size, habitat, or 
other information. In Kopp's (1966) revision of 
the neotropical Persea species, P. croizatii keys 
to her section Kriodaphne because of its glabrous 


ovary. It does not seem closely related to any of 
the species of that section with appressed pubes- 
cence, consisting mostly of P. caerulea and its 
relatives, but instead resembles more closely species 
of section Aurataea, especially P. nivea Mez and 


474 


Annals of th 
Missouri BON Garden 


P. benthamiana Meissner. These species both have, 
however, whitish and more pubescent lower leaf 
surfaces and smaller flowers. The inflorescences of 

benthamiana are also only half as long as the 
leaves. In all three species, the inner stamens are 
longer than the outer ones ich is an unusual 
feature in Persea. Unfortunately, all three species 
are rarely collected, and I have only seen one 
collection of each. More collections are needed to 
elucidate the relationships between these three 
species, whose 
rated from each other. 


istributions are not widely sepa- 


Persea fluviatilis van der Werff, sp. nov. TYPE: 
Venezuela. T. F. Amazonas: Depto. Rio Ne- 
gro, inundated forest along lower Rio Baria, 


G. Davidse 27717 (holotype, MO). 


Arbor, ad 10 m. Ramuli crassi, fistulosi, Seins pu- 
bescentes annotini glabri. Folia firme chartacea vel co 
riacea, elliptica vel anguste elliptica, 15- 20 4 x 3.5-7.5 

m, basi acuta vel obtusa, apice acuta; plus minusve 
aggregata; nervis lateralibus 8-10, supra immersis, subtus 
elevatis; margine paulo recurvata. Petioli 2-3 cm longi, 
juvenales Mies scentes, vetustiores glabri. Inflorescentiae 
axillares, folis perbreviores, dense pubescentes. Flores 
viridi-lutei, fere sessiles. Tepala 6, subaequalia, extus dense 
pubescentia; tepala exteriora late ovata, intus glabra, ad 


centi 

quadrilocellata, ca. 2.2 mm longa; filamentis pubescen- 
tibus, sensim in antheris glabris dilatatis; staminodia serei 
III ca. 1.9 mm longa, dense pubescentia, ad basim 2 
sandalia praedita; staminodia serei IV sag ittata, ca. ] 


mm longo. Fructus globosus; tepala persistentia, patentia; 
pedicellum at maturitatem fructus incrassatum. 


Tree, to 10 m. Twigs stout, terete, hollow, the 
leaf scars clustered, not evenly distributed along 
the twigs, older twigs glabrous, recent growth with 
rather dense, spreading pubescence. Leaves firmly 

chartaceous or coriaceous, elliptic or narrowly el- 
liptic, 15-20 x 3.5-7.5 cm, apex acute, base 
acute or obtuse, + clustered along the twigs; lateral 
veins 8-10 pairs, immersed on the upper surface, 
prominently raised on the lower surface, arching 
towards the tip of the leaf, not connected to the 
upper lateral vein; margin slightly recurved; upper 
leaf surface glabrous, lower leaf surface with some 
erect, gray hairs, these denser along midvein and 
lateral veins. Petioles 2-3 cm long, gray pubescent 
when young, glabrous in age. Inflorescences axil- 
lary, much shorter than the leaves, to 6 cm long, 
densely gray pubescent. Flowers greenish yellow 
or tawny, nearly sessile. Tepals 6, their outside 
surfaces completely covered by dense, gray pu- 
bescence, outer 3 tepals broadly ovate, ca. 2.5 


mm long, glabrous inside, inner 3 broadly ovate, 
ca. 3.2 mm long, pubescent inside. Fertile stamens 
6, representing the outer 2 whorls, ca. 

long, the filament densely pubescent, ca. 1.2 mm 
long, gradually widened into and poorly differen- 
tiated from the glabrous anther; staminodia of whorl 
III ca. 1.9 mm long, densely pubescent and with 
2 large, globose glands near the base, the tip not 
cordate or sagittate; staminodia of whorl IV ca. 1 
mm long, densely pubescent, sagittate. Ovary with 
a few hairs, 0.8 mm diam., style glabrous, slender, 
ca. 1.5 mm long. Fruit globose, ca. 1.3 cm 
with a glaucous bloom; tepals persisting in fruiting 
stage, spreading; pedicel in fruit strongly swollen, 
to 7 mm thick 


iam., 


Paratypes. VENEZUELA. T. F. AMAZONAS: along Rio 
Temi near Yavita, Ll. Williams 14122 (F); same alo, 
Bunting et al. 4041, 4050 (MY). 


Persea fluviatilis is known only from periodi- 
cally inundated forest in Territorio Federal Ama- 
zonas, Venezuela, at elevations of about 100 m. 
In Kopp's (1966) revision of the neotropical species 
of Persea, it keys to section Heteranthera because 
the stamens of whorl III are sterile. Within that 
section, it is closest to P. meridensis, a poorly 
known Andean species, which differs in its ferru- 
ginous pubescence, pedicelled flowers, and solid 
twigs. Unusual characters of P. fluviatilis are the 
subequal tepals, hollow twigs, clustered leaves, dif- 
ference in pubescence between the young and old 
twigs, and swollen pedicels in fruit. Because many 
of the neotropical Persea species are infrequently 
collected and are poorly known, close relatives of 
this distinctive species have not been pinpointed. 
The swollen pedicels in fruit are a very unusual 
feature, but because there are only two fruiting 
collections known, it is possible that this condition 
is caused by a disease and is not typical for the 
species. At the time of her revision Kopp had only 
a fruiting collection of this species at hand, which 
she cited as P. nivea Mez, a species that does not 
occur in Venezuela. 


Rhodostemonodaphne steyermarkiana (Al- 
en) van der Werff, comb. nov. BASIONYM: 
Ocotea steyermarkiana C. K. Allen, Acta 
Bot. Venez. 2: 216. 1967. 


Rhodostemonodaphne, as defined by Rohwer 
& Kubitzki (1985), includes a small number of 
species previously included in Nectandra or Oco- 
tea and differing from Nectandra in being dioe- 
cious and in a general lack of papillose pubescence 
on the floral parts; they differ from Ocotea in 
having almost sessile anthers and the locelli ar- 


Volume 76, Number 2 
1989 


van der Werff 475 


Lauraceae from Venezuelan Guyana 


ranged in an arc, not in two superposed rows. A 
study of the available collections of O. steyer- 
markiana shows that this species has all the char- 
acters of Rhodostemonodaphne, and therefore a 
new combination is made. I did not find that the 
outer stamens have distinct filaments and that the 
anther cells are arranged in two rows, as Allen 
described and illustrated. Pubescence and inflores- 
cence type suggest that RA. steyermarkiana is 
closely related to R. celiana (Allen) Rohwer and 
ndlicheria vinotincta Allen, both endemic to the 
Venezuelan Guayana. I include two collections from 
Cerro Sarisarinama, cited by Bernardi in Steyer- 
mark (1976) as O. celiana Allen and O. ferruginea 
(Meissner) Mez, in R. steyermarkiana. 


LITERATURE CITED 

C. K. 1964. Lauraceae. In: B. Maguire, J. J. 
urdack, et al., The Botany of the Guayana High- 
rui Part III. Mem. New York Bot. Gard. 10(1): 


ALLEN, 


Kore, z 1966. A taxonomic revision of the genus 
Persea in the Western Hemisphere. Mem. New York 
Bot. Gard. 14(1): oe 

ROHWER, twicklungslinien 
im Ocotea- Komplex da A Jahrb. Syst. 


SrevenvaRk, ] 1976. La vegetación de la cima del 
de Jaua. Soc. Ven. Ci. Nat. Bol. 132/133: 
"405. 


TAXONOMIC STUDIES 

IN THE MICONIEAE 
(MELASTOMATACEAE). III. 
CLADISTIC ANALYSIS OF 
AXILLARY-FLOWERED TAXA' 


Walter S. Judd? 


ABSTRACT 


The axillary- flowered members of the Miconieae are a to be a 
e been biota la ted by m 
iettea is ia D including Henriettella and Llew 
cia and Myriasp 
ophyso a and Maieta 


ithin this group hav 
g taxonomic po ora Hen 
Mon to include ee elluc 
di Henricten sensu lato. Clidemia sect. Calo 
congene Mecranium (inclu oe 
ecies of Üssaea all are considered gene 
a along w 
rues an autapo 


die ir 


ct. Sagraea 
distinct genus, morphy for 


Ekmaniocharis), Killip 
rically distinct P to their isolated cladistic positions. Species of Clidemia 
Necramium and the axillary-flowere 
this group was not consistently present in the generated cladograms, 


a monophyletic group, and the generic 
minary cladistic analysis, which resulted i in 


of a prelim 
f 
ia, Huilaea, and the -merous 


and 4-merous species of Ossaea are considered a 


e genus is considered metaphyletic. Clidemia sect. e peta io althoug ee considered axillary-flowered, 


and t 

actually - oth terminal and axillary 
closely to sect. Tamo 
iius Kirk idea, Loreya, M 


atet a, 


orm tw 
(Sometimes along with druses) occur in 


ing genera . Cladistic den id amo dm the g 
[xm is probably basal within ee plex, as ah ere d by 
Sagraea is likely most closely related to Meer 


inflorescences an 
nea). Thus, rd flowered groups recognized at the generic level include: Henriettea, Huila 
ecranium, Pentos 
major clades based primarily on bus crystal type present in le leaf mesophyll and stem. Megastyloids 
Lor ee 


era possessing only 


may congeneric Miconia (and likely is id. 


ossaea, and Sagraea. Axillary-flowered Miconieae E 


and Henriettea, while on 
ruse crystals 
its large flowers with well-developed pedicels, and 


m, the two being linked by the ds of 4-merous flowers 


i ae of axillary-flowered taxa to on eri Pese sn infloresences are discussed brie 


The Miconieae Triana (Melastomataceae) are a 
diverse tribe of ca. 30 neotropical genera, including 


characterized, often difficult to discern, and defined 
quite arbitrarily (Cogniaux, 1891; Macbride, 1941; 
Gleason, 1932, 1958; Wurdack, 1972). In order 
to clarify relationships within this large and com- 
plex group, a preliminary cladistic analysis of the 
axillary-flowered taxa of Miconieae was conducted 
as reported here 

Within the Miconieae, the inflorescence may be 
either terminal or lateral. Plants with terminal in- 
florescences have hapaxanthic shoots in which the 
apical meristem of the shoot becomes wholly trans- 


formed into a flowering axis after a period of veg- 
etative growth, while plants with axillary inflores- 
cences have pleonanthic shoots in which the apical 
meristem continues its vegetative activity while 
producing lateral flowers or inflorescence axes (see 
Judd, 1986a; a UN Inflorescence po- 
sition, along wit at t f growth 
architecture, has been shown to be a valuable taxo- 
nomic character in the tribe and is usually uniform 
within sections or genera (Judd, 1986a). 

The axillary-flowered condition is likely a de- 
rived state within the Miconieae because nearly all 
members of the Merianieae show terminal flowers. 
The latter tribe is a reasonable outgroup taxon 
considering possible tribal relationships suggested 


' This research was supported by NSF Grant BSR-8406760. I thank the curators of DUKE, F, GH, NY, MO, 


and US, from which specimens have been borrowed for this study, and Charles A. Woods, Thomas A 


Z noni, and 


. Za 
Lena Green for logistical assistance during fieldwork in Haiti, the Dominican Republic, and Jamaica. I am grateful 


to James D. Skean, Jr. and Susan 


e S. Renner for helpful — concerning the m 
A. Kron for assistance with rec Bele specimen data. I also thank Sus 


nuscript, and to Kathleen 
ne S. ho for sending me portions of her 


recent revisions of Loreya and ola Thanks are due Kent Perkins for processing specimen loans and to Gerald 


E. Masters for his fine illustratio 
? Department of Botany, 


ANN. Missouni Bor. GARD. 76: 476-495. 


220 Donum Hall, University of Florida, Gainesville, Florida 32611, U.S.A. 
1989. 


Volume 76, Number 2 
1989 


Judd 
Miconieae 


by J. Wurdack (Welle & Koek-Noorman, 1981). 
The same conclusion would be reached through 
the use of a generalized melastomataceous out- 
group and the tree counting algorithm of Frolich 
(1987), since terminal cymose inflorescences are 
the strongly predominant condition within non-Mi- 
conieae melastomes. Terminal inflorescences occur 
in over 80% of non-Miconieae genera, and strongly 
predominate in all tribes except the Blakeeae Benth. 
& Hook. f. In addition, Weberling (1988) noted 
that all melastomes show monotelic inflorescences, 
with the basic form being a pleiothyrsoid termi- 
nating a leafy shoot. In this study the monophyly 
of the axillary-flowered Miconieae was accepted as 
a useful preliminary hypothesis and a cladistic anal- 
ysis of these taxa was conducted. The results of 
this analysis are then used to suggest generic re- 
alignments and to reevaluate the preliminary hy- 
pothesis of monophyly of the axillary-flowered taxa. 


CLADISTIC ANALYSES 
METHODS 


The cladistic analyses were conducted using the 
Wagner parsimony algorithm (with global branch- 
swapping and multiple parsimony options) of the 
PAUP (i.e., Phylogenetic Analysis Using Parsi- 
mony, Version 2.4; Swofford, 1985) package of 
computer software. 

Fifteen Evolutionary Units (EUs) were used in 
most of the analyses (Table 1), based on a survey 
of ca. 370 herbarium specimens representing 150 
species (and supplemented, where possible, by the 
addition of information from recent taxonomic 
studies). Several taxa also were studied during the 
course of fieldwork in Haiti, the Dominican Re- 
public, Puerto Rico, and Jamaica. In some anal- 
yses, an additional EU representing either a gen- 
eralized terminal-flowered taxon or Clidemia sect. 
Miconiopsis Cogn. was used. Although terminal- 
flowered, Clidemia sect. Miconiopsis has been 
considered to be related to the other sections of 
Clidemia included in this study, and its inclusion 
in a few analyses was designed to test this hypoth- 
esis. The inclusion of a generalized terminal-flow- 
ered taxon in some analyses was designed, in a 
very preliminary way, to evaluate the hypothesis 
that the axillary-flowered Miconieae are monophy- 
letic. Placement of this generalized terminal-flow- 
ered taxon is considered very tentative because 
this EU is extremely heterogeneous, making the 
determination of its ancestral character states prob- 
lematic (see Table 1). 

Forty-two characters were used and assigned 
plesiomorphic (ancestral: scored as 0) and apo- 


morphic (derived: scored as 1) states (Table 1). 
Apomorphic features used in the computer analyses 
are listed below. Nearly all characters were polar- 
ized using the Merianieae as an outgroup, with 
character states uniformly found (or, for a few 
characters, those states most commonly repre- 
sented) in the out-group considered plesiomorphic 
(Stevens, 1980; Wheeler, 1981; Wiley, 1981; 
Maddison et al., 1984; Frolich, 1987). When EUs 
are polymorphic for a given character, an estimate 
of the ancestral condition was made. The states of 
a few characters were unknown for certain taxa, 
and these are scored as “missing data," indicated 
by a “?” in the data matrix (Table 1). Trees were 
rooted by using a hypothetical ancestor with all 
characters scored as plesiomorphic. 


APOMORPHIC CHARACTERS 


1. Stems quandrangular and 4-winged/ ridged; 
plesiomorphic condition: stems + terete. Quadran- 
gular stems are considered an autapomorphy of 
Killipia Gleason and occur in K. quadrangularis 
Gleason (the type species). However, two other 
species have terete stems, and the ancestral con- 
dition within Killipia is not known. A coding of 
"0" for this EU would not affect the analyses 
because the feature is autapom i 

2. Growth pleonanthic Lr plesio- 
morphic condition: growth hapaxanthic (determi- 
nate). (See character 12.) 

3. Megastyloid crystals present; plesiomorphic 
condition: only small styloids and/or druse crystals 
present. (See character 4.) 

4. Crystals exclusively druses; plesiomorphic 
condition: small or large styloids and druse crystals 
present. The polarity of characters 3 and 4 was 
determined by using the Crypteroniaceae (Dahl- 
gren & Thorne, 1984) as an outgroup (see dis- 
cussion in Johnson & Briggs, 1984; Vliet et al., 
1981). This family, which is closely related to the 
Melastomataceae, contains genera with druses and 
with small styloids (Baas, 1981), and this crystal 
complement is taken as the ancestral condition in 
melastomes. It is likely that megastyloids have 
evolved in a few lineages (sometimes with the loss 
of druses), while styloids have been lost in other 
lineages, resulting in druses being the only crystal 
type in those groups. That styloids and druses are 
known from more distantly related families of Myr- 
tales, such as Myrtaceae and Combretaceae (Vliet 
& Baas, 1984) supports this polarity. Variation in 
crystal complement within the Merianieae is poorly 
known. 

5. Elongate multicellular hairs absent; plesio- 


478 


Annals of the 
Missouri Botanical Garden 


TABLE 1. 
Henriettella 


Character divergence values for taxa used in cladistic analyses. Taxon names abbreviated by first four letters of genus name, except Henriettea (HENT), 


HENL), Clidemia juruensis (CJUR), generalized terminal-flowered taxon (TERM). An asterisk indicates that the opposite state occurs in some species 


a question mark indicates that the character state is in doubt or unknown. 


of that taxon; 


18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 


8 9 10 11 12 13 14 15 16 17 


1234567 


MECR 0101101000 0 


Taxon 


0 0 0 0 0 0 0*0 0 1000000 0 


l 


0 0 0 0 


l 


l 


l 


HENT 0110010000 0 


0 0 00 00 0 0 0 0 


— 


] 


OF 0* 0* OF OF 0 0*0 0*0 D 0*0 0 0*0 0*0 0 000 0 0 0 


l 


TERM 0 001 0*0 00 0*0* 0 0 0 


Volume 76, Number 2 
1989 


Judd 479 
Miconieae 


morphic condition: elongate multicellular hairs 
present, often strigose. Mecranium Hook. f. an 
Killipia are unique in lacking a well- da 
indumentum. A few species of Loreya DC., 
L. ovata Berg ex Triana, L. arborescens Gann 
and L. nigricans Triana, lack such hairs as 
wei je (in press) hypothesized the loss of 
strigose hairs to be a synapomorphy of a species 
group within Loreya. Thus, the presence of such 
hairs is considered the ancestral condition within 
the genus. 

6. Multicellular hairs elongate with radiate to 
stellate-globular basal portion (or clearly derived 
types); plesiomorphic condition: multicellular hairs 
various, but not as above. This distinctive hair 
morphology is much modified (see discussion, and 
Judd, 1986b) in a few species, but such extremes 
are connected by intermediate types, sometimes 
even on the same plant. 

7. Multicellular hairs minute, globular, and un- 
branched to irregularly branched and matted (not 
strigose); plesiomorphic condition: multicellular hairs 
various but not as above. This character is limited 
to Mecranium (including Ekmaniocharis Urban) 
and Ossaea acuminata The feature is im- 
portant in determining that Mecranium spp., 
saea acuminata, and Ekmaniocharis crassinervis 
Urban form a monophyletic group but does not 
help in inferring relationships among the EUs. 

8. Multicellular hairs conspicuously stellate (with 
stalk and elongated arms); plesiomorphic condition: 
multicellular hairs various but not as above. In 
these analyses this feature is considered an autapo- 
morphy of Clidemia sect. Miconiopsis. Very sim- 
ilar hairs occur in many terminal-flowered species, 

., Miconia sects. Jucunda (Cham.) Naudin and 
Tamonea (Aublet) Cogn. 

9. Leaves clearly anisophyllous; plesiomorphic 

condition: leaves isophyllous or nearly so ew 
specialized species of Clidemia sect. Sograt (DC.) 
ogn. have anisophyllous leaves. The isophyllous 
condition is considered ancestral within Clidemia 
sect. Sagraea, following a preliminary analysis of 
variation within the group. 

10. Leaves with pair of pouchlike formicaria at 
base of blade; plesiomorphic condition: leaves lack- 
ing formicaria. Similar ant-domatia that occur in 
terminal-flowered taxa, e.g., Tococa Aublet, Cli- 
demia D. Don, Myrmidone Mart., probably evolved 
independently (see discussion). Ossaea bullata (Pil- 
ger) Gleason, possibly a member of the Sagraea 
group, has been reported to possess similar for- 
micaria (Whiffin, 1972). 

11. Cuticle of abaxial leaf epidermis papillose; 
plesiomorphic condition: epidermis nonpapillose. 
Renner (in press) pointed out that this feature cor- 


relates well with cuticle thickness and suggested 
that the character has important ecological con- 
sequences. 

12. Inflorescences axillary (in leaf axils or at 
nodes on old wood); plesiomorphic condition: inflo- 
rescences terminal. Terminal cymose inflores- 
cences (as discussed above) occur in over 80% of 
genera within non-Miconieae melastomes. Thus, 
proliferating inflorescences (in which the terminal 
axis returns to vegetative growth, producing axil- 
lary buds that develop vegetative branches instead 
of flower-bearing paracladia—see Weberling, 1988) 
are considered derived. The process of proliferation 
results in a vegetative shoot in which the flowers 
are borne in axillary cymose paracladia. Delayed 
anthesis of the paracladia can even result in cau- 
liflory. 

13. Inflorescences 
condition: inflorescences cymose. Although fasci- 
cles occur in a few species of Loreya, Clidemia 


fasciculate; plesiomorphic 


sect. Sagraea (including axillary, 4-merous species 
of Ossaea DC.), and the 5-(6-)merous species of 
Ossaea, cymose inflorescences are considered to 
be the ancestral condition in these groups. Renner 
(in press) also considered fasciculate inflorescences 
to be derived within Loreya. 

14. Flowers more or less sessile; plesiomorphic 
condition: flowers pedicellate. 

15. Flowers 4-merous; plesiomorphic condition: 
flowers other than 4-merous. (See character 16.) 

16. Flower parts frequently more than 5; ple- 
siomorphic condition: flowers 4- or 5-merous. Gen- 
erally, 5-merous flowers are considered primitive, 
with higher and lower floral numbers treated as 
derived. Henriettea DC. and Loreya contain a few 
species with numerical plans greater than five, while 


Henriettella Naudin and Kirkbridea Wurdack 


sidered ancestral. Renner (in press) considered flo- 
ral numbers greater than five to be derived and 
used this feature to link some species of Loreya 
with Bellucia Raf. The ancestral condition in Kirk- 
bridea is in doubt; however, recoding the state of 
this EU as “1” for this character does not change 
the topology of the discovered cladograms. 

17. Flowers small; plesiomorphic condition: 
flowers moderate to large. 

18. Hypanthium strongly ribbed; plesiomorphic 
condition: hypanthium nonribbed. This character 
is an autapomorphy of Clidemia sect. Calophy- 
soides Cogn. 

19. External calyx lobes more or less conspic- 
uous and well developed; plesiomorphic condition: 
external calyx lobes absent or inconspicuous. 

20. External calyx lobes absent; plesiomorphic 


480 


Annals of the 
Missouri Botanical Garden 


condition: external calyx lobes present. Some species 
of Henriettella have lost external calyx lobes in- 
dependently of the loss in Loreya, Bellucia, and 
Myriaspora Is was determined because it 
is evident that Henriettella evolved out of a group 
within Henriettea, which has external calyx lobes. 

21. Internal calyx lobes very elongate and 
closely associated with external lobes; plesiomor- 
phic condition: internal calyx lobes not elongate, 
variously developed but not closely appressed to 
internal lobes. This feature is an autapomorphy of 
Killipia. 

22. Internal calyx lobes connate into a mem- 
branous, dome-shaped, apiculate cap that ruptures 
irregularly at anthesis; plesiomorphic condition: in- 
ternal calyx lobes free. This character is an aut- 

of Mecranium (including Ekmani- 

Although Ossaea acuminata has 
somewhat caplike calyx lobes, they are only par- 
tially connate. The internal calyx lobes are free 
and imbricate at their extreme apices. This species 
is included within the Mecranium EU since it has 
the other apomorphies of this clade (characters 7, 
19, and 32; see discussion). 

23. Internal calyx lobes circumscissilly dehis- 
cent (and noncalyptrate); plesiomorphic condition: 
internal calyx lobes more or less persistent, or if 
circumscissile, then calyptrate. This character is 
considered an autapomorphy of Clidemia sect. Mi- 


included within the generalized terminal-flowered 
taxon). 

24. Buds closed, with thick triangular calyx 
lobes that cover bud up to the final stage prior to 
anthesis (and sometimes fused into a cap); plesio- 
morphic condition: buds open, calyx lobes of vari- 
able shape and thin, the petals exposed much before 
anthesis. This feature is an autapomorphy of Bel- 
ucia 

25. Internal calyx lobes reduced to a more or 
less smooth rim; plesiomorphic condition: internal 
calyx lobes clearly evident. This is a noninformative 
character (see Table 1) because it is found only in 
a few derived Vll within Henriettella and Cli- 
demia sect. Sagraea. 

Teas de lobes fused into a conical, 
apically closed calyptra and conspicuously strigose; 
plesiomorphic condition: internal calyx lobes non- 
calyptrate (or if so, then not as above). This is an 
autapomorphy of Myriaspora. 

27. Internal calyx lobes fused into an apically 
open cap (opening into sepal-like segments or cir- 
cumscissilly); plesiomorphic condition: internal ca- 
yx lobes noncalyptrate (or if so, then not as above). 
This feature is limited to Llewelynia Pittier and a 


few (probably related) species of Henriettea (see 
discussion). 

. Calyx a cap open at apex and falling off 
by circumscissile slit; plesiomorphic condition: ca- 
lyx not open-calyptrate, or if so, then opening by 
longitudinal slits into triangular sepal-like segments. 
This character is an autapomorphy of Llewelynia 
and is a further specialization of character 27. 

29. Petals narrowly triangular and more or less 
acute at apex; plesiomorphic condition: petals ovate 
to elliptic (or obovate) with blunt to more or less 
acuminate apices. Preliminary analyses of variation 
within Mecranium (including Ekmaniocharis) and 
Clidemia sect. Sagraea (including axillary, 4-mer- 
ous species of Ossaea) have led to the conclusion 
that ovate to elliptic petals with more or less acu- 
minate to blunt apices constitute the ancestral con- 
dition within these groups. Obovate petals have 
evolved within a group of Mecranium species. It 
is noteworthy that Ossaea acuminata, a narrow- 
petaled species, is a sister species to Mecranium 
amygdalinum (Desr.) C. Wright in Sauv., a fairly 
blunt-petaled species (Skean, in prep.). 

30. Petals with a dorso-apical tooth; plesio- 
morphic condition: petals lacking a dorso-apical 
tooth. This feature occurs in a few species of Cli- 
demia sect. Sagraea (including the species of Os- 
saea with axillary, 4-merous flowers) and in the 

r 6-merous species of Ossaea. A scoring of 
«p is this character in these two EUs does not 
alter their placement in relation to the very dif- 
ferent Henriettea group. 

31. Petals usually pubescent, sometimes only 
with very few hairs near apex; plesiomorphic con- 
dition: petals glabrous (or with a single apical hair). 
Hairs on the petals have been lost in a few species 
of Loreya and Bellucia. 

2. Androecial fringe present; plesiomorphic 
condition: androecial fringe absent. This feature is 
a strong and consistent synapomorphy of Mecra- 
nium species, including Ekmaniocharis crassiner- 
vis and Ossaea acuminata. 

. Anthers with two well-developed dorso-bas- 
al aan: plesiomorphic condition: anthers 
various but lacking two well-developed dorso-basal 
projections. 

4. Anthers usually robust and fleshy, opening 
by one or two minute pores; plesiomorphic con- 
dition: anthers not robust and fleshy, various. 

35. Anther sacs basally bifurcate, i.e., sagittate; 
plesiomorphic condition: anther sacs not bifurcate 
at base. 

36. Anther sacs more or less wrinkled; plesio- 
morphic condition: anther sacs smooth. The best 
expression of this feature is in Maieta Aublet. 

37. Stamens numerous, i.e., more than twice 


Volume 76, Number 2 
1989 


Judd 481 


Miconieae 


the number of petals; plesiomorphic condition: sta- 
mens twice the number of petals. This is an aut- 
apomorphic character of Llewelynia. 

38. Stigma very conspicuously and radiately 
ridged; plesiomorphic condition: stigma not radiate- 
ly ridge 

39. Piscenis highly divided; plesiomorphic con- 
dition: placenta undivided or nearly so. This is an 
autapomorphy of Myriaspora. 

40. Fruit a berry; plesiomorphic condition: fruit 
a capsule. 

41. Chromosome number of n = 28; plesio- 
morphic condition: n — 17 (or 15, 20). (See char- 
acter 42.) 

42. Chromosome number of n — 15, 20; ple- 
siomorphic condition: n — 17 (or 28). The terminal- 
flowered Miconieae were used as a generalized out- 
group for the determination of polarity of the in- 
completely known characters 41 and 


RESULTS 


Initial computer runs using the entire data set 
for the 15 axillary-flowered EUs resulted in the 
discovery of 100-- equally parsimonious trees (but 
only 15 different topologies when multi-furcations 
are taken into account), all with a length of 28 
steps and a consistency index of 0.714 (or 48 steps 
and consistency index of 0.813 when autapomor- 
phies and uniform characters were included). The 
strict consensus tree resulting from this analysis is 
presented in Figure 1B. In addition a tree indicating 
character state changes is presented (Fig. 1A). This 
tree is preferred due to its pattern of character 
variation, especially the linking of Mecranium and 
Clidemia sect. Sagraea on the basis of 4-merous 
flowers (15), and the linking of Henriettea, Hen- 
riettella, and Llewelynia on the basis of fasciculate 
inflorescences (13). The relationships of the EUs 
representing Loreya, Bellucia, Myriaspora, Hen- 
riettea, Henriettella, and Llewelynia did not vary 
in any of the trees generated by PAUP. In addition, 
Kirkbridea consistently linked with Henriettea, 
Henriettella, and Llewelynia, in some trees as the 
cladistically basal member of this clade, in others 
as the sister group of Henriettella. In all trees, the 
EUs representing Maieta, Clidemia sect. Calo- 
physoides, and Clidemia juruensis (Pilger) Glea- 
son were always joined, and Huilaea Wurdack was 
always the basal member of the clade representing 
taxa with only druse crystals. Thus the large num- 
ber of generated trees is due to the numerous 
possible topologies among the remaining EUs in 
the analyses, i.e., Mecranium (including Ekmani- 
ocharis and Ossaea acuminata), Clidemia sect. 
Sagraea (including Necramium Britton and axil- 
lary 4-merous species of Ossaea), Killipia, and 


Pentossaea, here described to include the axillary- 
flowered, 5-merous (or less commonly 6-merous) 
species of Ossaea 

Thus, several analyses were conducted using just 
those EUs possessing only druse crystals (i.e., top 
eight EUs listed in Table 1) with characters not 
pertaining to this complex deleted. These analyses 
generated 45 equally parsimonious trees (but com- 
prising only seven different tree topologies) of 11 
steps and with a consistency index of 0.727 when 
all characters were weighted equally (or of 25 steps 
and a consistency index of 0.880 when autapo- 
morphies and uniform characters were included). 
However, when the apomorphy of 4-merous flowers 
was stressed (weighted by 2, as seems reasonable, 
since this character is quite uniform within EUs 
under investigation), only 15 trees (representing 
only two topologies) were produced, all with 12 
steps and a consistency index of 0.750 (or of 26 
steps and a consistency index of 0.889 when aut- 
apomorphies and uniform characters were includ- 
ed). The consensus tree resulting from the analysis 
in which all characters are weighted equally is 
identical to that produced in the analysis of all 
axillary-flowered taxa (Fig. 1B). Both cladograms 
resulting from the weighted-character analysis are 
given in Figure 1C and D 

When an EU representing a generalized ter- 
minal-flowered taxon is included in the analysis, 
over 100 equally parsimonious trees were found, 
all with a length of 32 steps and a consistency 
index of 0.688 (or of 50 steps and a consistency 
index of 0.780 when autapomorphies and uniform 
characters were included). These trees are very 
similar to those generated in the analyses involving 
only axillary-flowered taxa (Fig. 1), and the four 
different topologies involving their basal branches 
are presented in Figure 2, illustrating the various 
positions of the genus Huilaea and a generalized 
terminal-flowered taxon. When Clidemia sect. Mi- 
coniopsis (another taxon producing terminal inflo- 
rescences) is included in the analysis, many trees 
are again generated (of 55 steps and a consistency 
index of 0.745, including autapomorphies), all link- 
ing this entity with either Killipia or a clade con- 
taining Killipia, Clidemia sect. Sagraea, and 
Mecranium. 


DISCUSSION 


The cladograms presented in Figures 1 and 2 
indicate that the axillary-flowered Miconieae are 
divided into two major clades. The first is composed 
of those taxa containing megastyloids (3) in their 
leaf and stem tissues (often with druse crystals as 
well) and usually possessing slightly pubescent pet- 
als (31). The group also generally has distinctly 


482 Annals of the 
Missouri Botanical Garden 


ME SA KI PE CL MA CA HU LO MY BE KK HL HT LL 
5 417 4 26 11 20 
1 3 27 
19 29 35 18 is 17 17 28 
2 5 24 19 
32 21 37 
33 
9 2n 13 
33 34 
10 
15 f 16 38 he 
6 42 
41 
14 
17 
19 
3 
à . 
I (A) 

2 

12 
40 


ME A KI PE CL MA CA HU LO MY 


(B) 


Volume 76, Number 2 
1989 


Judd 483 
Miconieae 


FIGURE 2. 


"hone taxon, all of 50 steps, consistency index — 


Huilaea (HU), exclusively drase- ponents dnd (ee) megastyloid-containing group (Styloid), and Miser 
1 2:2 1 I ] tly 4 1 1 1 megas 


O e 


Basal portion of cladograms resulting from analysis of axillary-flowered taxa and generalized terminal- 


Reversals indicated by open bars. Taxon abbreviations: 


styloid- 


terminal-flowered taxon (TE). * 
and druse-containing lineages (see text). 


pedicellate flowers. This clade may be divided into 
two clearly marked subgroups based on vegetative 
and reproductive characters. Loreya, Bellucia, and 
Myriaspora are united by the synapomorphies of 
flowers lacking external calyx lobes (20), possessing 
robust fleshy anthers (34), an expanded and ra- 
diately ridged stigma (38), and a chromosome num- 
ber of n = 15 or 20 (42). In contrast, Henriettea, 
Henriettella, Llewelynia, and Kirkbridea form a 
clade based upon the presence of multicellular stri- 
gose hairs with a stellate, radiate, irregularly 
branched, or suprabasally expanded base, or hair 
types clearly modified from these conditions (6), 
petals with an apical apiculum on the abaxial sur- 
face (30), and a chromosome number of n — 28 
(where known, 41). Kirkbridea likely is cladisti- 
cally basal in this group, with Henriettea, Hen- 
riettella, and Llewelynia being united by the ad- 
ditional synapomorphy of fasciculate inflorescences 


(13). (It should be noted that parallelism in flower 
size (17) occurs within these genera, which is not 
surprising considering the variability of this feature 
within the tribe. Linking Kirkbridea and Henriet- 
tella on the basis of this character is considered 
implausible.) 

The remaining taxa included in the analysis form 
a second clade based on the following synapomor- 
phies: presence of only druse crystals in leaf and 
stem tissues (4), small more or less sessile flowers 
(17, 14), and usually well-developed external calyx 
lobes (19). The extremely isolated genus Huilaea 
probably is basal within this clade (Fig. 1A, B) 
because it shows only the synapomorphy of exclu- 
sive presence of druse crystals (4), having large 
and clearly pedicellate flowers. The relationships 
among the taxa comprising this clade are not well 
resolved. However, it is clear that Maieta, Cli- 
demia sect. Calophysoides, and C. juruensis form 


— 
FIGURE 1. 


flowered taxa, 48 steps, consistency index — 0.813. 


analyses of taxa containing only druse crystals —two 26-ste 
nalysis. Reversals indicated by open bars; ta 


topology C equals consensus tree of this a 


Cladograms resulting from PAUP analyses. — A. Preferred tree resulting from analysis of all axillary- 
— B. Consensus tree. — C, D. 


Cladograms resulting from PAU 
trees generated in Ta weighting character 15; 
a abbreviated by first two 


letters of generic or sectional name, except yo (HT), Henriettella (HL), and Kirkbridea (KK). 


Annals of the 
Missouri Botanical Garden 


a monophyletic group based on the presence of 
morphologically very similar pouchlike ant-domatia 
(formicaria) at the base of the leaf blade (10) and 
wrinkled anthers (36). In addition, Maieta and 
Clidemia sect. Calophysoides have anisophyllous 
leaves (9). If the character of 4-merous flowers is 
stressed, then Mecranium is probably most closely 
related to Clidemia sect. Sagraea, and the two 
may be linked by this feature (15). Linking Killipia 
with Sagraea and Mecranium in some trees on 
the basis of the shared loss of strigose hairs (5) 
probably has little phylogenetic significance. 

The terminal-flowered taxa of Miconieae may 
be a paraphyletic or monophyletic group basal to 
all the axillary-flowered taxa (Fig. 2A-C) or a pos- 
sible sister taxon to the taxa containing only druses 
(Fig. 2D). In the first alternative, the axillary- 
flowered condition evolved just once and serves as 
a synapomorphy of all axillary-flowered Miconieae; 
in the second, axillary arrangements are hypoth- 
esized to have evolved in the druse and megastyloid 
clades independently (or, less likely, evolved in the 
stem lineage of the entire tribe with a reversal to 
terminal cymes occurring in the ancestor of Mi- 
conia and relatives). The possible polyphyly of the 
axillary-flowered Miconieae clade will be investi- 
gated more thoroughly in a later paper. It is cer- 
tainly possible that Miconia and relatives are more 
closely related to the axillary-flowered ““druse-con- 
taining" genera than initially hypothesized. 

When an EU representing a generalized ter- 
minal-flowered taxon is included in the analysis, 
the position of Huilaea in the cladograms becomes 
more variable than in the initial analyses (compare 
Figs. 1 and 2). This variability may reflect the 
presumed phylogenetic isolation of this genus. It 
seems more reasonable to link Huilaea with 
Mecranium, Clidemia sect. Sagraea, Maieta, and 
Killipia on the basis of the presence of only druse 
crystals in its tissues (4) than with Loreya, Hen- 
riettea, and Kirkbridea on the basis of a reversal 
to pedicellate flowers (14) (Fig. 2). 

he presence of both terminal and axillary flow- 
ers in Clidemia sect. Miconiopsis (Cogniaux, 1888, 
1891) is not surprising (see Judd, 1986a, for dis- 
cussion of similar variation evident in Miconia, 
especially in sects. Octomeris (Naudin) Benth. & 
Hook. and Tamonea; and Troll, 1964, and We- 
berling, 1965, 1988, for more general discussion 
of inflorescence structure and position). Wurdack's 
suggestion (pers. comm.) that this group is closest 
to Miconia is likely correct. Note especially the 
similarity in stellate indumentum (8; Fig. 4) and 
circumscissilly dehiscent calyx (23). The section 
may be quite close to Miconia sects. Tamonea and 


Jucunda. The linking of Clidemia sect. Miconiop- 
sis with. Killipia or a clade containing Killipia, 
Clidemia sect. Sagraea, and Miconia in the cla- 
distic analysis probably is due to homoplasy in 
several fairly weak characters, e.g., loss of strigose 
hairs (5), or to several potential synapomorphies 
between the axillary-flowered taxa containing only 
druse crystals and Miconia (and relatives), e.g., 
more or less sessile flowers (14) or well-developed 
external calyx lobes (19). 

In spite of the necessarily preliminary nature of 
these cladistic analyses, their results help determine 
the previously confused generic limits within this 
difficult tribe (see also Judd, 1986a, b). In inter- 
preting these results, I have attempted to delimit 
monophyletic and phylogenetically equivalent gen- 
era within the tribe that are recognizable on the 
basis of several morphological characters. Thus 
Huilaea, Mecranium (including Ekmaniocharis 
and Ossaea acuminata), Killipia, Maieta (in- 
cluding Clidemia sect. Calophysoides and C. ju- 
ruensis), Loreya (including Bellucia and Myri- 
aspora), Henriettea (including Henriettella and 
Llewelynia), and Kirkbridea are considered to be 
generically distinct. In addition, Clidemia sect. 
Sagraea (including Necramium and the species of 
Ossaea with axillary 4-merous flowers) is recog- 
nized at the generic level. Finally, the species of 
Ossaea with axillary 5- (or 6-)merous flowers are 
in need of generic recognition and are described 
herein as Pentossaea. The systematics of each of 
these taxa is discussed briefly below. 


Discussion of axillary-flowered genera. 
Loreya is clearly delimited by its distinctive usually 
robust fleshy anthers (34), expanded and radiately 
ridged stigma (38), lack of external calyx lobes 
(20), and range of chromosome numbers (42, see 
Solt & Wurdack, 1980). The group possesses me- 
gastyloids in its leaf and stem tissues (pers. obs.; 
Baas, 1981) and is clearly monophyletic. However, 
if Bellucia and Myriaspora are segregated as sep- 
arate genera, the result is a paraphyletic Loreya 
(Fig. 1). Bellucia is characterized by papillose 
abaxial leaf cuticle (11), closed buds (24), i.e., the 
buds have thick calyx lobes that cover the buds 
up to the final stage prior to anthesis (and are even 
fused into a cap in a few species, e.g., Bellucia 
dichotoma Cogn. and B. acutata Pilger), and ten- 
dency for floral merosities greater than five (16) 
(see Renner, in press). In contrast, Myriaspora is 
characterized by its highly divided placentae (39), 
calyx lobes fused into an unusual cap (26, which 
is probably not homologous with the cap of some 
species of Bellucia), and long hairs on the abaxial 


Volume 76, Number 2 
1989 


Judd 
Miconieae 


485 


surface of its petals. In addition, Myriaspora pro- 
duces inflorescences in the leaf axils (as do several 
species of Bellucia), while the usual condition in 
Loreya is to have the inflorescences lateral on 
leafless nodes below the leaves; it should be noted, 
however, that this character is often variable within 
a genus (Judd, 1986a; Renner, in press). 

No autapomorphies are evident for Loreya when 
certain species are segregated as either Bellucia 
or Myriaspora (Fig. 1). The paraphyly of Loreya 
is made even more apparent when it is realized 
that Loreya subrotundifolia (Wurdack) S. Renner 
and L. riparia S. Renner link with Bellucia on 
the basis of higher numbers of floral parts and that 
L. mespiloides Miq., L. spruceana Benth. ex 
Triana, and L. klugii S. Renner link with Myrias- 
pora on the basis of testa characters and one- 
pored anthers (Renner, in press). 

hus, recognition of Bellucia and Myriaspora 
distorts the best estimate of phylogenetic structure 
currently available for this group of species. Their 
recognition is also doubtful on purely phenetic 
grounds, since the patristic distances separating 
Bellucia and Myriaspora from the cladistically 
related taxa within a narrowly delimited Loreya 
are comparable to those separating other clades 
within Loreya (Renner, in press). Thus, as is char- 
acteristic of traditional taxonomic procedure (see 
Stevens, 1984, 1986), recognition of Bellucia and 
Myriaspora can only be justified by selective 
weighting of certain features (in this case the char- 
acters of cuticular papillae, bud structure, placental 
form, and inflorescence position) due to their sup- 
posed greater ecological significance. I see no rea- 
son to weight these characters more than any oth- 
ers used in these analyses. The decision to recognize 
a broadly circumscribed Loreya is not new; Baillon 
(1877) also considered Bellucia and Loreya con- 
generic (although only few species were known to 
him), and Macbride (1941: 497) considered this 


. to 'straining at 


treatment “‘preferable it seems . . 
a gnat’ to keep them apart.” Gleason (1932) con- 
sidered the genera only “weakly separated"; how- 
ever, they were maintained as distinct by Wurdack 
(1973, 1980) and Renner (in press). 


Henriettea is easily recognized by its usually 


expanded, radiate to globular-stellate basal portion 
(or easily derived types, such as stellate-hairs, pel- 
tate-scales, elongate hairs with numerous small 
mainly basal projections, elongate hairs with a su- 
prabasal expanded region and a narrowed basal 
attachment, or hairs very reduced, e.g., Henriet- 


tella caudata (DC.) Naudin, H. glabra Cogn.; see 


Wurdack, 1986) (6), petals with a minute dorso- 
apical apiculum (30), and chromosome number of 
n — 28 (41, Solt € Wurdack, 1980). Fasciculate 
inflorescences have evolved independently in a few 
species of Loreya (Renner, in press), and the in- 
florescences of Henriettea squamulosa (Cogn.) W. 
Judd have reverted to a pedunculate condition (Judd, 
1986b). Henriettea is clearly monophyletic and 
shows the presence of calcium oxalate megastyloids 
(similar in structure to those of Loreya) in its leaf 
and stem tissues (pers. obs.; see also Welle & 
Mennega, 1977; Welle & Koek-Noorman, 1981; 
Baas, 1981). 

As with Loreya, problems of paraphyly occur 
when Llewelynia and Henriettella are recognized 
as distinct from Henriettea (Fig. 1). Llewelynia 
is phenetically distinctive (Pittier, 1939; Wurdack, 
1973) and easily recognized by its loss of external 
calyx lobes (20), open calyx cap that opens by a 
circumscissile slit (27, 28), and numerous stamens 
(37), while Henriettella is only marginally sepa- 
rable due to its smaller flowers (17) and tendency 
to have shorter anthers that open by larger, gaping 
pores (see Wurdack, 1973, 1980). A nearly ob- 
solete internal calyx has evolved in a few species 
of Henriettella, e.g., H. sessilifolia (L.) Triana, 
and in many species the internal calyx lobes are 
not well developed. Henriettea and Henriettella 
were considered congeneric by Macbride (1941: 
500), who stated that "there seems to be no sig- 
nificant concomitance of the characters of petals, 
anthers or pubescence that have been used to main- 
tain two similar but distinct genera.” He added 
that “the characters moreover often exist in de- 
grees of development.” Richard A. Howard also 
will consider Henriettea and Henriettella to be 
congeneric in his Lesser Antilles flora (pers. comm.). 
Wurdack (1972: 418) maintained these genera 
but noted that “a generic transition zone exists 
between Henriettella ovata Cogn. and Henriettea 
patrisiana DC., with Henriettella parviflora 
(Griseb.) Triana perhaps not distinct from the lat- 
ter.” 

The genera differ in flower size, anther char- 
acters, and indumentum, but intermediates exist. 
Not surprisingly, there are no autapomorphies for 
Henriettea when defined narrowly, and its species 
certainly intergrade with Henriettella. 

The floral buds of some other species of Hen- 
riettea (e.g., H. spruceana Cogn., H. succosa 
(Aublet) DC.. H. stellaris Berg ex Triana, and H. 
ramiflora (Sw.) DC.) possess an expanded and open 
cap formed by the inner calyx lobes very similar 
to (and likely homologous with) that of Llewelynia, 
although opening longitudinally rather than cir- 


Annals of the 
Missouri Botanical Garden 


cumscissilly. These species are likely basal mem- 
bers of the clade leading to Llewelynia. It is of 
interest that Wurdack (1973) suggested a similar- 
ity in general facies between Llewelynia and Hen- 
riettella, and Welle & Koek-Noorman (1981) re- 
ported similarity in wood anatomical characters 
between Llewelynia and Henriettella. It thus seems 
best to recognize a broadly defined, and monophy- 
letic Henriettea by including both Henriettella and 
Llewelynia within it. 

Kirkbridea clearly is related closely to Hen- 
riettea s.l. (Fig. 1). Kirkbridea (Wurdack, 1976) 
is quite distinctive due to its pedunculate inflores- 
cences and anthers with two well-developed dorso- 
basal projections (33). Kirkbridea and Henriettea 
show variation in number of floral parts. 

Huilaea is evidently very isolated, as Wurdack 
(1957, 1976) suggested at the time of his descrip- 
tion of the genus. It clearly is distinct generically 
and is recognizable due to its pendent, three-flow- 
ered cymose inflorescences with very long pedun- 
cles and large, clearly pedicellate, 6-merous flow- 
ers. 

Maieta, defined broadly, is a monophyletic group 
best recognized by its characteristic pouchlike for- 
micaria at the base of its leaf blades (10) (Fig. 3; 
see discussion of morphology, development, and 
ecological functions of these structures in Gleason, 
1931; Schnell, 1967; Whiffin, 1972; Neto € 
Asakawa, 1978; Benson, 1985). It should be noted 
that ant-domatia of various types undoubtedly have 
evolved independently in a few to many species of 
several other genera (e.g., Sagraea DC., Tococa, 
Clidemia, Henriettea, and Conostegia D. Don), 
and additional developmental studies are necessary 
in order to determine appropriate homologies of 
such structures within the tribe. The presence of 
structurally and positionally very similar pouchlike 
ant-domatia is accepted as a provisional synapo- 
morphy of Maieta s. od me Cudenia sect. t 
ophysoides until s 
are conducted. The group alo tends to Hadas e 
ly wrinkled anthers (36) and clearly anisophyllous 
leaves (9). Druses are the only crystals known in 
this group. The species of Maieta (defined nar- 
rowly—see Aublet, 1775; Wurdack, 1980) have 
sagittate anthers (35) and show a reversal toward 
large flowers (17), while those of Clidemia sect. 
Calophysoides have a distinctly ribbed hypan- 
thium (18). Both, along with possibly a few other 
species of Clidemia, such as C. juruensis, are 
considered congeneric. The close relationship be- 
tween these species of Clidemia with pouchlike 
formicaria and the phenetically similar species of 


Maieta has long been recognized. Macbride (1941: 


479) remarked that “Clidemia remains . . . a con- 
cept rather arbitrarily retained but conveniently so 
on account of Maieta (in which Baillon, Hist. Pl. 
7: 56. 1880 rightly merged it) being the older 
name." I agree and here consider Maieta to be 
congeneric with Clidemia sect. Calophysoides (and 
C. juruensis). 

Killipia (Fig. 4) is a distinctive genus not easily 
confused with any other in the complex of genera 
possessing only druse crystals. Noteworthy apo- 
morphies are the very long internal calyx lobes 
with closely associated external calyx lobes (21) 
and distinctive anthers with two dorso-basal pro- 
jections (33); see description of genus in Gleason 
(1925). The stems of Killipia quadrangularis are 
quadrangular and ridged-winged (1); however, two 
other species have terete stems. The abaxial leaf 
surface of K. quadrangularis is usually red, an 
apomorphy not included in the analysis, which has 
also evolved within several genera with terminal 
inflorescences. The cladistic relationships of Kil- 
lipia are obscure, although it is closest to Maieta, 
Mecranium, Sagraea, and Pentossaea (Fig. 1). 

The species formerly of Ossaea with axillary 
5-merous or occasionally 6-merous flowers warrant 
generic recognition (Fig. 5). The petals of this 
group are narrowly triangular with an acute apex 
(29), and the group is clearly separable from related 
taxa (see key). It is not surprising that a group of 
Ossaea species was found to be in need of generic 
recognition, since the genus has long been sus- 
pected of being polyphyletic (see Judd, 1986a). 
Wurdack (1962: 206), for example, stated that 
"the genus is an artificial assemblage of various 
species groups." The distinctive clade, with axillary 
5-(6-)merous flowers, is here described as the genus 
Pentossaea. 


Pentossaea W. Judd, gen. nov. 


Genus novum, Sagraea DC. affine, a quo floribus 5-meris 
vs. 4-meris), petalis semper anguste triangularibus, api- 
cibus acutis differt. TYPE SPECIES: Pentossaea brachy- 
stachya (DC.) W. Judd. 


— 


Evergreen subshrubs or shrubs with pleonanthic 
shoots; twigs + terete, nodose, glabrous to densely 
strigose, with a ridge encircling each node; druse 
crystals present. Indumentum of conspicuous, un- 
branched, slender to stout, strigose, multicellular 
hairs with or without glandular apices and of mi- 
nute, multicellular, pun hairs (or sometimes 
glandular-stellate hair urring on stems, leaves, 
inflorescence axes, and paniis or in some species 
+ restricted to reproductive parts. Buds narrowly 


ovoid, sylleptic, usually densely pubescent. Leaves 


Volume 76, Number 2 Judd 
1989 Miconieae 


FLUTE. 
eT, LA 
" Een 
WS 
Y SS 


A NEN Pone 
fs 


3cm 


T "my 


ay. 


FIGURE 3. Maieta poeppigii Mart. ex Cogn. — A. Habit.—B. Leaf, abaxial view showing formicaria.—C. Detail 
showing entrance to formicaria. — D. Stamen.— E. Flower. 


488 


Annals of the 
Missouri Botanical Garden 


(029 mm 


D 


LJ. S. J. 


Imm 


j —— 


FIGURE 4. —A-F. Killipia quadrangularis. A. 


Ste 
a e, H. Clidemia kappleri Cogn., stellate es rs 


opposite, decussate, petiolate to nearly sessile, 
estipulate, herbaceous to coriaceous, + plane; 
adaxial surface essentially glabrous (at maturity) 
or sparsely to densely covered with unbranched, 
elongate, multicellular hairs and sometimes also 
with minute glandular hairs (especially in vein 
impressions or at petiole junction); abaxial surface 
usually sparsely to densely covered with slender to 
stout, unbranched, elongate, multicellular hairs 
along with minute glandular to glandular-stellate, 
multicellular hairs (i.e., glandular hairs unbranched 
or sparsely branched with one branch glandular 
and the others nonglandular), sometimes only with 
glandular hairs and becoming + glabrous with age; 
domatia not observed in vein axils; margin serrate 
in distal ca. ?4 of lamina to entire, plane to slightly 
revolute near base, the teeth ending in an elongate 
multicellular hair (and often associated with addi- 
tional elongate hairs); venation acrodromous, slightly 
suprabasal, with a prominent midvein, 1—4 pairs 
of secondary veins (1 of these often intramarginal 


zd 


, section. B. Flower bud. C. Calyx lobes. D. Petal. E, 


and inconspicuous), numerous percurrent tertiary 
veins * perpendicular to midvein and reticulate to 
orthogonal quaternary and higher-order veinlets; 
petiole caniculate, essentially glabrous to densely 
strigose-pubescent. Inflorescences axillary cymes 
(occasionally reduced and + fasciculate) borne in 
the leaf axils and in a few species on leafless nodes 
below the leaves, usually borne singly, rarely 2 per 
axil, bracteate, with opposite, narrowly triangular 
to ovate or oblong bracts (the 2 lowest protecting 
the inflorescence in bud and quickly caducous). 
Flowers a 5- or 6-merous, sessile. Hypan- 
thium + cylindrical, abaxially with conspicuous, 
unbranched, elongate hairs with or without glan- 
dular apices, intermixed with minute glandular or 
glandular-stellate hairs, adaxially glabrous or with 
a few hairs around apex. External calyx lobes 5 
or 6, conspicuous, elongate, terete. Internal calyx 
lobes 5 or 6, distinct (noncalyptrate), + 


or 6, * white, narrowly triangular with acute api- 


Volume 76, Number 2 Judd 489 
1989 Miconieae 


SE A n i 
We D /4 d E 1 mm 
FicuRE 5. Pentossaea coriacea. — A. Habit. —B. Leaf.—C. Flower. —D. Petal.— E. Stamen.— F. Berry.— 6. 
Calyx and inner surface of hypanthium; note absence of androecial fringe. 


490 


Annals of the 
Missouri Botanical Garden 


ces, 1 -veined, glabrous or occasionally with a single 
dorso-apical projection (hair), rarely with a few 
glandular hairs. Stamens 10-12, + isomorphic, 
glabrous, geniculate, inserted at apex of hypan- 
thium, androecial fringe lacking; filament narrowly 
ovate to oblong, flattened dorsally; anther narrowly 
ovate to oblong, sometimes slightly curved, the 2 
anther sacs dehiscing confluently by means of a 
single small apical pore, the connective sometimes 
slightly prolonged beyond the bases of the anther 
sacs. Ovary inferior or nearly so, 3—5-locular, with 
axile placentation and numerous ovules, the apex 
occasionally prolonged into several blunt append- 
ages surrounding the style. Style cylindrical, + 
straight; stigma capitate, papillose. Berries red to 
blue, with indumentum similar to hypanthium. Seeds 
small, rounded- to angular-obovate, the testa smooth 
to slightly roughened. 


Pentossaea angustifolia (DC.) W. Judd, comb. 
nov. Leandra angustifolia DC., Prodr. 3: 
154. 1828 

Pentossaea brachystachya (DC.) W. Judd, 
comb. nov. Clidemia brachystachya DC., 
Prodr. 3: 156. 1828. 

Pentossaea confertiflora (DC.) W. Judd, comb. 
nov. Clidemia confertiflora DC., Prodr. 3: 
156. 1828. 

Pentossaea congestiflora (Naudin) W. Judd, 
comb. nov. Clidemia congestiflora Naudin, 
Ann. Sci. Nat. Bot., Ser. 3, 17: 344. 1852. 

Pentossaea coriacea (Naudin) W. Judd, comb. 
nov. Clidemia confertiflora DC., Prodr. 3 
156. 1828 

Pentossaea heteronervis (Naudin) W. Judd, 
comb. nov. Sagraea heteronervis Naudin, 
Ann. Soc. Nat. Bot., Ser. 3, 18: 98. 1852. 

Pentossaea marginata (Desr.) W. Judd, comb. 
nov. Melastoma marginata Desr. in Lam., 


Encycl. 4: 32. 1797 


Additional new combinations will certainly be- 
come necessary following a detailed revision of the 
genus. 

The axillary-flowered 4-merous species of Os- 
saea and the very similar species traditionally placed 
in Clidemia sect. Sagraea (also axillary-flowered 
and 4-merous; see Cogniaux, 1891) are typically 
separated by the technical character of apically 
acute vs. blunt petals. However, I found this char- 
acter to vary continuously within the group, and 
several species placed in Clidemia sect. Sagraea 
actually have acuminate or acute petals (e.g., C. 
oligantha Urban, C. fuertesii Cogn., C. domin- 
gensis (DC.) Cogn., C. crossosepala Griseb.), while 
others treated in Ossaea may have blunt-tipped 
petals (e.g., O. barahonensis Urban & E. Ekman 


or O. setulosa Urban). Some taxa have been con- 
sidered members of both groups by various authors. 
In addition, phenetically very similar species have 
sometimes been placed in the two "genera," e.g., 
Clidemia crossosepala and Ossaea hirtella (Sw.) 
Triana (see Adams, 1972), and petal shape and 
apex occasionally vary infraspecifically, as in Cli- 
demia fuertesii. Thus, all 4-merous, axillary-flow- 
ered members of the ““druse-containing group” (Fig. 
2) with free internal calyx lobes, no androecial 
fringe, and a usually well-developed strigose in- 
dumentum are placed together in Sagraea (Fig. 
6; also see key) 

Necramium (Britton, 1924), 
4-merous axillary flowers having ovate, sharply 
acuminate petals and with an indumentum of long 
strigose hairs and irregularly branched-stellate hairs, 
clearly also belongs to this clade. A close association 
between Necramium and some species of Ossaea 
was recognized by Wurdack (1973: 721-722), 
who stated that Necramium was "realmente bien 
acomodado dentro de Ossaea DC. sensu Cogniaux 
pero la necesidad por una transferencia posterior 


a genus with 


cuando ese conjunto artificial se disperse se opone 
a la sinonimización inmediata." Wurdack (pers. 
comm.) suggested affinity of Necramium to Ossaea 
sect. Octopleura (Griseb.) Cogn. This is judged 
unlikely, however, since the latter group has ter- 
minal inflorescences (although frequently deflexed 
to a seemingly lateral position, see Judd, 19864), 
a glandular dorso-basal connective appendage, and 
strongly costate fruits. 

Necramium (Britton, 1924) is not the oldest 
name available for this clade, the names Ossaea 
and Sagraea (de Candolle, 1828) both having 
priority. Because the types of Sagraea (S. cap- 
illaris (Sw.) DC., here designated) and Ossaea (O. 
scalpta (Vent.) DC., here designated) fall within 
the group circumscribed above, and the two genera 
were described at the same time (de Candolle, 
1828), either name is available for this group. The 
name Sagraea is chosen here, since I wish to avoid 
the use of Ossaea, a name that has been used for 
a great many discordant species-groups (see Wur- 
dack, 1962, 1973, and Judd, 1986a). It is note- 
worthy that Triana (1871) considered these axil- 
lary-flowered species (along with a few discordant 
elements) to be generically separable (as Sagraea) 
from the terminal-flowered Clidemia. 

Sagraea may be most closely related to Mecra- 
nium, a more specialized genus also showing 
4-merous flowers, this a potential synapomorphy 
(15, Figs. 1, 2). Sagraea is considered to be meta- 
phyletic (Donoghue, 1985; Mishler, 1985; Mishler 
& Brandon, 1987) because its ca. 60 species show 
no autapomorphies in some of the generated clado- 


Judd 


Volume 76, Number 2 
1989 Miconieae 


A 

Zh 

LFH 

A E 
[L4 vy 


PEO ty, Sod 
e t E 


470 A TR , 
aa 


La - 
Ss, at 
stg faz MM CMM 4 fa 
TLE $ 


di ue 
LL’ 


LET hdl 


N 


KS 
SS 
4 Es ASS 


FIGURE 6.  Sagraea (represented by the species better known as Ossaea barahonensis). — A. Habit. —B. Leaf. — 
C. Flower — D. Longitudinal section of flower showing hypanthium; note lack of androecial fringe. — E. Elongate 
multicellular hair. —F. Stamen. — G. Petal. 
mined. Generic recognition of this phenetically 


grams. However, in other equally parsimonious 
rather variable group is considered appropriate and 


trees, the group appears monophyletic, being de- 
fined by a reversal to strigose hairs (5) or by a 
parallel evolution of 4-merous flowers (15). The 
group is possibly paraphyletic, but no character 
conclusively demonstrating this has been deter- 


preferable to leaving its species in several artifi- 
cially delimited genera. 

Several species of Sagraea have evolved for- 
micaria at the base of the petiole, e.g., Clidemia 


492 


Annals of the 
Missouri Botanical Garden 


C iEZII I 
Mz 


FIGURE 7. Mecranium multiflorum (Desr.) Triana. — A. Habit. —B, C. Leaves. — 
— F. Hypanthium, androecial fringe, and ovary apex. — G. Stamen. — H. Berry.— I. Se 


tococoidea (DC.) Gleason and C. crenulata Glea- 
son; these species have sometimes been confused 
with Maieta (see Gleason, 1931). The structure 
and position of these formicaria are quite different 
from those of the Maieta and likely due to con- 


D. Flower bud. — E. Flower. 
d. 


vergent evolution. Plants with a climbing habit and 
strongly anisophyllous leaves occur in Clidemia 
epiphytica (Triana) Cogn., a species that is also 
cytogenetically distinctive (Solt & Wurdack, 1980). 


As discussed above, anisophyllous leaves are char- 


Volume 76, Number 2 
1989 


Judd 493 


Miconieae 


acteristic of Maieta, and this feature undoubtedly 
has evolved in parallel in the two genera. 

It is not considered appropriate to make the 
many necessary new combinations in Sagraea until 
a thorough taxonomic revision of the genus has 
been completed. The species considered to belong 
in Sagraea and used in this cladistic investigation 
are: Clidemia aphanantha (Naudin) Sagot, C. 
capituliflora Cogn., C. ciliata D. Don, C. cren- 
ulata Gleason, C. crossosepala Griseb., C. cur- 
soris Wurd., C. discolor (Triana) Cogn., C. di- 
varicata (C. Wright ex Griseb.) Cogn., C. 
domingensis (DC.) Cogn., C. epiphytica (Triana) 

n., C. fendleri Cogn., C. fuertesit Cogn., C. 
graciliflora Huber, C. grisebachii Cogn., C. het- 
eroneura Cogn., C. insularis Domin (= Sagraea 
capillaris (Sw.) DC.), C. involucrata (Griseb.) C. 
Wright ex Sauv., C. minutiflora (Triana) Cogn., 
C. monantha L. Williams, C. obliqua Griseb., C. 
oligantha Urban, C. plumosa (Desr.) DC., C. pur- 
pureo-violacea Cogn., C. pusilliflora Cogn., C. 
d pod cs C. quadripetala Almeda, C. rub- 

a (Aublet) Mart., C. rubrinervis Cogn., C. sep- 
laa UM C. sericea D. Don, C. sessiliflora 
(Naudin) Cogn., C. silvicola Gleason, C. tococoi- 
dea (DC.) Gleason, C. trichotoma C. Wright, C. 
ulei Pilger, C. umbrosa Cogn., Ossaea curvipila 
Urban & E. Ekman, O. hirtella (Sw.) Triana, O. 
navasensis Britton & Wilson, O. nipensis Britton 
& Wilson, O. petiolaris (Naudin) Triana, O. ru- 
fescens (Griseb.) C. Wright, O. scabrosa (L.) DC., 
O. scalpta (Vent.) DC., O. setulosa Urban, and 
O. woodsii W. Judd & Skean. 

Mecranium (Fig. 7) is a distinctive monophyletic 
species group distinguished from other genera con- 


KEY TO AXILLARY-FLOWERED MICONIEAE 


taining druse crystals by its internal calyx lobes 
fused into a domelike, apiculate, and membranous 
cap that is ruptured irregularly by elongation of 
the style and emergence of the petals at anthesis 
(22), the external calyx lobes reduced in length 
and often inconspicuous (reversal of 19), androecial 
fringe (i.e., a ring of erose to ciliate appendages 
arising from apex of hypanthium [torus] internal 
to the stamens [32 )), and usually sparse and poorly 
developed indumentum (7); see Judd (19862) and 
Skean & Judd (1986). Because these apomorphic 
characters are found also in the monotypic Ek- 
maniocharis (E. crassinervis; Urban, 1921; Judd, 
1986a; Skean, in press), it is clear that this genus 
belongs in Mecranium. Likewise, Ossaea acumi- 
nata (Judd, 1986a) shows all these apomorphies 
except that of a domelike, apiculate cap. In this 
species the apices of the internal calyx lobes are 
distinct and imbricate, although the lobes are con- 
nate for most of their length, forming a structure 
that may be similar to the condition from which 
an apiculate cap was derived. It is obvious that O. 
acuminata is a basal member of the Mecranium 
clade. The nomenclatural transfers will be made 
by James D. Skean, Jr. in a monographic study of 
Mecranium currently near completion. A more 
detailed analysis of the phylogeny of this genus will 
be presented by Skean (in prep.) 

n summary, the phylogenetic relationships (and 
taxonomic diversity) among the axillary-flowered 
Miconieae are most clearly represented by a taxo- 
nomic system recognizing the following nine genera 
as delimited above: Henriettea, Huilaea, Killipia, 
Kirkbridea, Loreya, Maieta, Mecranium, Pen- 
tossaea, and Sagraea. 


la. Plants with large elongate ae (megastyloids) in leaf and stem tissues (easily observable with a hand 


ns); flower 


abaxially, or with a fe ada airs near 


ers usually clearly | petals densely to sparsely pubescent 
ex, occasionally 


2a. Flowers with radiately idend SDN AN expanded "amm pe with usually robust, fleshy anthers; 


f very slender to stout, elon 


ngate, unbranched, multicellular hairs 


. Loreya 
mentum of slen der 


e.g., stellate hairs, peltate scales, elongate hairs with minute ee near base, or elongate hairs 


with a suprabasal expanded region 
3a. Anther 


y + spherical crystals (druses) in le 


- 
c 
X 
£e 
25 
w we 
= 
> 
> 
2) 
Ca 


rue without dorso-basal projections; nord tad 
af a 


n and narrowed basal attachm 
s with 2 well-developed dorso-basal projections; as clearly pedunculate ..... Kirkbridea 


sually sessile and fasciculate .......... Henriettea 
stem tissues, Los megastyloids; flowers + 


sessile or pedicellate; petals usually glabrous or with a single apical h 
owers clearly pedicellate, i.e., pedicels ca. 1-1.5 cm long; rana pendulous and 3-flowered 


5 cm lon 


Huilaea 


with peduncle ca. 10-3 


4b. Flowers + sessile; inflorescences various but the peduncle shorter than ca. 6 cm. 
5 


lowers 5- or sometimes Ó-merous. 
6a. Plants with pouchlike formicaria (ant-domatia) at bases of leaf blades; leaves often clearly 
anisophyllous Maieta 
6b. Plants without formicaria; leaves + isophyllo 


yilo 
7a. Petals narrowly triangular, with acute ue plants with elongate, multicellular, un- 


494 


Annals of the 
Missouri Botanical Garden 


branched hairs (often strigose), with or without glandular tips; anthers lacking dorso- 


basal projections 


entossaea 


7b. Petals ovate to elliptic, with + 
anthers wit 
5b. Piden 4-merous 


blunt apices; plants lacking elongate, multicellular hairs; 
h 2 dorso-basal projections 


Killipia 


[e] 
> 
— 
Z 
- 
o 
m 
z 
> 
a, 
> 
& 
u 
m 
c 
2- 
> 
< 
qe 
a 
E 
E 
S 
E 
e 
5. 
d 
ju 
op 
= 5 
2 w 
d 
a 
Q1 
e 
h 
o 
S R.Z 
or 
=] 
oa 
e 
p 
EA 
= 
o 
i= 
ga 
> 
o 
e 
(e 
£e 
2 
o 
5 
Had 
<= 
E: 
i" gi 


conspicuously strigose, almost always with 


a 
elongate, branched or unbranched, multicellular hairs, often with glandular apices ......... Sagraea 


LITERATURE CITED 

Abams, C. D. 1972. Flowering Plants of Jamaica. Rob- 
ert MacLehose & Co., University Press, Glasgow. 

AuBLET, J. B. C. F. 1775. Histoire des Plantes de la 
Guiane oo Volume 1. Pierre-Frangois Didot 
i is 

Baas, P. 1981. A note on stomatal types and crystals 
in the ped of Melastomataceae. Blumea 27: 475 

7 

BaiLton, H. E. 1877. Mélastomacées. Pp. 1-65 in 

Historie des Plantes, Volume 7. L. Hachette et Cie, 


aris. 
BENSON, W. hi 985. Amazon ant- plants. Pp. 239- 
266 in E Prance & T. E. Lovejoy (editors), 


eius Pergamon Press, Oxford. 

BRITTON, N. L. 1924. Studies of West Indian plants — 
XII. 71. A new genus of Melastomaceae from Trin- 
idad. Bull. pop Bot. Club 51: 6-7. 

CANDOLLE, a Ti 1828. Melastomaceae. Prodromus 
3: 99 


COGNIAUX, Fa cor Ce of Clidemia sect. Mi- 
coniopsis. [n: C. F. P. von Martius (editor), Flora 

Brasiliensis 14(4): 496 
l Melastomaceae. Pp. in A. & 

i" Mà Candolle (editors), a a 

1 7. Masson, Paris 
& 


Diuienese R. R. T. THORNE, 1984. The order 
yrtales: circumscription, variation, and relation- 
nn. Missouri Bot. Gard. 71: 633-699. 


ships. A 
DoNocHuvE, M. J A critique of the biologi a 
Ln concept and recommendations for a phylo 
etic oe Bryologist 88: 172-181 
juod ICH, M. 1987. Common-is- epe: : piss 
on by tree a yst. Bot. 12: 217-237. 
GLEASON, H. A. 1925. Studies on the flora of sere 
South America— VIII. A 
lastomataceae. Bull. ae rw Clu = 
1931. The relationships of certain myrme- 
cophilous melastomes. Bull. Torrey Fin Club 5 
73-85. 


í 


l A synopsis of the Melastomataceae of 
British Ciana. Brittonia 1: 127-184. 

1958. Melastomataceae. In: R. E. Woodson, 
Jr. & R. W. Schery (editors), Flora of Panama. Ann. 


Missouri Bot. Gard. 45: 203-304 

Jounson, L. A. S. & B. G. BRIGGS Myrtales and 
Myrtaceae —a a Bal veis, Ann. Missouri 
Bot. Gard. s 756. 

Jupp, W.S. [o studies in the Miconieae 


Mo I. us 
sition. Brittonia 38: 15 
1986 eee placement of Calycogo- 


tion in inflorescence po 
l. 


nium p (Melastomataceae). Brittonia 38: 
238-2 


MACBRIDE, i F. 1941. Flora of Peru (Melastomata- 
ceae). Fieldiana, Bot. 13: 249-52]. 

Mabbison, W. R., M. J. DonocHue & D. R. Mappisow. 
19 Output ünalysis and parsimony. Syst. Zool. 
33: 03. 

MISHLER, B. D. . The i ah a develop- 
mental, and phylogenetic basis of species concepts 
in bryophytes. SUR 88: 207 bir 

. & R. N. DC 


1987. Iudbadüslibe plu- 
ralism, and the o im concept: Biology 
ir Pus phy 2 -4] 

Nero, C. & N. rad 1978. Estudo de 


Lr NP em algumas espécies de Boragi 
:hrysobalanaceae, Melastomataceae e Ru- 
bia . Acta Amazonica 8: 45-49. 

PITTIER, N 939. Description of Llewelynia. In: Notas 
Veri s de Venezuela IV. Bol. Soc. Venez. Ci. 

Du 307-309. 

RENNER, S. Systematic studies in the Melastomata- 
ceae: hela Loreya, and Macairea. Mem. New 
York Bot. Gard. (in pre 

SCHNELL, R. 1967. Camión a l'étude des genres 

einn amazoniens 7ococa Aubl. et Maieta Aubl. 


(Mélas cées) et de leurs poches foliaires. Adan- 
sonia, sér. D$, : 525-532 
SKEAN, J. D., Jr 


Rediscovery of Ekmaniocharis (Me- 
lastomataceae). Moscosoa 5 (in press 
& W. S. Ju 9 


UDD. An Mecranium 
Si lal from Hispaniola. dinum. 38: 
23 c 
SoLr M. L. J. Wukback 1980. 


bre pe agp 
i. in ds Melastomataceae. 47 
20 


Phytolog 
STEVENS, P. F. 1980. Evolutionary polarity M 
states. Annual. Rev. Ecol. S l 33 
1984. Metaphors and typology in 2: de 
opment on botanical systema atics 1690-1960, or the 
art of putting new wine in old bottles. Taxon 33: 
169- 2l l. 


1986. Evolutionary classification in PEU 
1960 0-1985. J. Arnold Arbor. 67: 313-330 
SWOFFORD, D. L. 1985. PAUP: Phylogenetic abs 
Jsing pupil ii 2.4. Illinois Natural His- 
tory Survey, Ur 


Priana, J. 1871 i Mélastomacios. Trans. Linn. Soc. 
London 28: 1-188. 
TROLL, W. Die Infloreszensen. Typologie und 


1964. 
on im Aufbau des Vegetationskorpers. Teil. 1 


oe T 1921. Description of Ekmaniocharis. In: 


Volume 76, Number 2 Judd 495 
1989 Miconieae 
Plantae Haitienses Novae vel Rariores a Cl. Er. L. 9 Wuirrin, T. Observations on some upper Am- 


Ekman 1917 Lectae. Ark. Bot. 17(7): 48-49. 
Vier, G. J. C. M. van & P. Baas. 1984. Wood 
anatomy and lacas of the Myrtales. Ann. Mis- 
souri Bot. Gard. 71: 783- 
, J. Kork-NOORMAN & B. J. ER WELLE. 
1981. Wood anatomy, [cnni and pbyogeny 
of the Melastomataceae. Blum 3-4 
R F. Poy ot inflorescences, J. 
n. Soc. Bot. 59: 
1988. The n s of inflorescences in 
the Myrtales. Ann. Missouri Bot. Gard. 75: 226- 
310. 


WELE, B. J. H. TER & J. KoEk-NoonMaAN. 1981. Wood 
anatomy of neotropical Melastomataceae. Blumea 
27: 335-394. 

. M. W. MENNEGA. 1977. On the presence 

of large styloids i in the secondary xylem of the genus 

Henriettea ra IAWA Bull. 1977/2: 

31-35. 


WHEELER, Q. D. 1981. The ins and outs of character 
analysis: a iade to Crisci and Stuessy. Syst. Bot. 


6: 297-306 


azonian TORREN Melastomataceae. Sida 5: 33- 
1 


WiLEv, E. O. 1981. Phylogenetics. Wiley, New York. 
Wurpack, J. J. 1957. Certamen Melastomataceis IV. 
Brittonia 9: 101-109. 
1962. Melastomataceae of Santa Catarina. 
Gellowia 14: 109-217 
1972. Comet Melastomataceis XVIII. 
Phytologia 22: 399-418 
1973. Melastomataceae. /n: T. Lasser (edi- 
in Flora of Venezuela. No. 8. Instituto Botanico, 
Ministerio de Agricultura y Cria, Caracas. 
1976. Endemic Melastomataceae of the Sierra 
Nevada de Santa Marta, Colombia. Brittonia 28: 
138-143. 


Melastomataceae. /n: G. Harling & E 
Sparre (editors), Flora of Ecuador. No. 13. Uni 
Góteborg & Riksmuseum, Stockholm 

1986. Atlas of hairs for neotropical Melas- 
ie taoeae. Smithsonian Contr. Bot. 63: 1-80. 


A SURVEY OF Susanne S. Renner? 
REPRODUCTIVE 

BIOLOGY IN NEOTROPICAL 
MELASTOMATACEAE AND 

MEMECYLACEAE! 


ABSTRACT 


In the Neotropics, the arena and de e comprise over 3,000 species in 106 genera. Pollination 
observations have been reporte 26 species in 35 genera of Melastomataceae and for four species in one genus 
of Memecylac ceae. Genetic llinc bo, has been found in 22 Melastomataceae species, self-compatibility in 
25 species. A s ingle m e e tested i is self- -compatible. Agamospermy is known in 19 New World and 

ace 


is known in a few species in one genus of Melastomataceae. The principal mode of promoting outcrossing in both 
families is spatial separation of pollen and stigma (herkogamy), achieved by the pollen being enclosed in poricidal 
anthers, which have to be manipulated to release pollen. New World Memecylaceae are pollinated in the same way 
and by some of the same bee species as most Melastomataceae. All bee species known to visit Melastomataceae and 
Meise :ylaceae are list ed; they comprise a ius seda of the a bee fauna. It is not yet c wouie 


in the ee mec ia beyond that ife en nhan cing the visual attractiveness of the eni and making ihe stamens 
easier to grasp. In most Melastomataceae, flowers offer only pollen; however, some 60 species in 8 gene r 
nectar as a reward for pollinators (probably, some species in at least one additional genus do, too). The nec dea 


species are pollinated by birds, bats, rodents, and bees. e nectar is rich in sucrose and, as a rule, its production 
is correlated with floral morphological changes relating to the fact that, except for bees, nectar consumers do not 
vibrate the stamens to collect pollen actively. I suggest that the capacity for developing nectaries is ic in 
WA SEN cid ut s seed in mos moden members The ae sea of Meinecylucénn = minute 


way a staman attractiveness. In a dispersal system m dn as occurred. with 40% of Es 
neotropical Melastomataceae having capsular an and wind-dispersed seeds and 60% having soft, juicy berries taker 


mainly by birds, but also by D m onke ats, other mammals, turtles, and other reptiles. All Meno le 
have berries, and their se ds are dispersed by bird ue) an h. I conclude that the east and west Gondwanian 
lineages of Melastomataceae and emecylaceae, i a coevolved he earliest bees in the lo id- 


Cretaceous. This may have been the time when the ancestors of these families switched from nectar-reward flowers 
to pollen-r reward flowers. In the further extensive radiation of jm families, special adaptations to pollinators were of 
limited importance; diversification in seed dispersal systems, vegetative characters, and edaphic adaptations were more 
important. 


The neotropical Melastomataceae and Meme- the field and the availability of several recent treat- 
cylaceae are particularly suitable for a study of ments in monographs and floras. As Gleason (1932) 
reproductive biology in view of the relative ease stated, “a dozen or more species . . . grow naturally 
with which members of these families are found in on every square mile of tropical America, unless 


'T thank the oo m for identifying bees: M. C. Almeida (Trigona), R. i ssler (Euglossini), C. 


Eickwort (Halictidae), V aaf (Oxaea), C. D. Michener (Exomalopsir A S. Laroca and J. R re Hakin (Halictidae), 
^ n ire e it opini, ono ‘a, Meliponini, D. Roubik (Melipona, Trigona), R. Snelling eae Ceratini), and 

cept for specimens retained by the specialists bees are deposited in the entomological collection 
" oí Instituto Nacional de Melee da Amazónia (INPA). It is a pleasure to dale the support I received 
at INPA, especially from Dr . Rodrigues, M. F. da Silva, and M. L. Absy, also from R. Bierregárd and J. Rankin- 
de-Merona, gels bes tae field directors, respectively, of the INPA-WWF “Minimal Critical Size of 
Ecosystems" project. Fieldwork in Brazil was supported by the Deutscher Akademischer Austauschdienst and the 


Deutsche Forschungsgemeinschaft. Fieldwork in Venezuela was funded by ale from the National Science Foundation 

and the Smithsonian Scholarly Studies Program. I am grateful to Klaus Kubitzki and John Wurdack for their help 

during different phases of this work and to T. Morley, C. Todzia, D. Roubik, H. Tobe, B. Stein, R. Lundin, and K. 
remer for comments on the manuscript. 


: Bot tanical Institute, University of Aarhus, Nordlandsvej 68, DK-8240 Risskov, Denmark. 
ANN. Missouni Bor. Garb. 76: 496-518. 1989. 


Volume 76, Number 2 
1989 


Renner 
Reproductive Biology in Melastomataceae 
| 


& Memecylaceae 


497 


TaBLE 1. New World Melastomataceae and Meme- 
cylaceae. Tribal arrangement following Cogniaux (1891), 
except for more recently described, transferred, or syn- 
onymized genera. Species numbers are those of the most 
recent treatment, or based on a herbarium survey (Ren- 
ner, 1986). indicates a recorded observation, citation 
in Table n — no observation presently know 


olli- 
nation 
Species Obser- 


Number vation 


MELASTOMATACEAE 
BERTOLONIEAE 
ertolonia 8 m 
Boyania l 
Diplarpea l = 
Macrocentrum 21 = 
Maguireanthus l = 
Monolena 15 = 
iari l + 
Salping 8 + 
CANC 1 — 
Triolena (incl. Diolena) 22 = 
Tryssophyton l = 
MERIANIEAE 
Acanthell 2 = 
Adelobotrys 25 + 
— 30 = 
Behuri 3 
Benev m 1 = 
Bisglaziova 1 — 
Centronia 15 + 
Dolichoura — 
Graffenrieda (incl. Calyptrella 44 + 
and anthus) 
Huberia 6 — 
Meriania 74 + 
Merianthera 3 — 
Neblinanthera 1 
Ochthephilus l = 
Phainantha 4 — 
Tes mou 3 — 
MICROLICIE 
ucque — 
sie T (incl. Pyramia) 21 = 
Castratella l = 
Centradeni 6 + 
Centradeniastrum l -= 
Chaetosto 12 = 
Eriocnema 1 
Lavoisiera 46 + 
Lithobium 1 = 
Microlicia 100 = 
Poteranthera 2 = 
Rhynchant 15 + 
Siphanthera final, Farringtonia) 16 + 
Stenodon 1 — 
Trembleya 11 + 
TIBOUCHINEAE 
Aciotis + 
Acisanthera 17 + 
Appendicularia -= 
Brachyotum 50 + 


TABLE l. Continued. 


Polli- 


ation 


Species Obser- 


Number vation 


Chaetolepis 10 
Comolia 22 
Comoliopsis l 
Desmoscelis l 
Ernestia 16 
ritzschic 1 
Heterocentron 6 
Loricalep 1 
cairea 22 
Mallophyton l 
Marcetia 23 
Microlepis 4 
Nepsera l 
Pilocos 3 
Pseudernestia l 
terogas 4 
Pterolepis 15 
andemania 1 
Cra 1 
Svitr 1 
Tibouchina (incl. Itatiaia and Pur- 243 
purella 
Tibouchinopsis 2 
RHEXIEAE 
Monochaetum 45 
Pachylo 6 
Rhexi 13 
MICONIEAE 
Anaectocalyx 3 
ellucia T 
Calycogonium 23 
Catacoryne 1 
Chalybea l 
Charianthus 11 
Clidemia 117 
Conostegi 43 
Ekmaniocharis 1 
enriettea 12 
Henriettell 51 
Heterotrichum 10 
] 4 
Killipia 4 
Kirkbridea 1 
Leandra (incl. Platycentrum) 175 
Llewellynia 1 
Loreya 13 
atet 2 
Mecranium 21 
Miconia ca. 1,000 (at least 958) 
Mommsenia 1 
Myriaspora 1 
Myrmid 2 
Necramium 1 
Ossaea 91 
Pachyanthus 16 
Pleiochiton 7 
Tetrazygia 21 
Tococa 54 


1+ | + | 


+ 


IE d cB ag i 


| ++ | 


c 


Td 14+ gd c ag d 


+ 


498 


Annals of the 
Missouri Botanical Garden 


TABLE l. Continued. 


Polli- 


nation 
Species Obser- 


Number vation 


CYPHOSTYLEAE 


lomaieta 1 x 
Alloneuron q — 
Cyphost yla l = 

BLAKEEAE 
Blakea 100 + 
Topobea 62 
MEMECYLACEAE 
Mouriri 81 d 
Votomita 8 — 


the land is under intense cultivation, and the var- 
ious genera extend from the coastal marshes at 
sea-level to the high páramos above the tree line." 
The Melastomataceae are a large family (v.i.) and 
diverse in terms of habit, reproductive and vege- 
tative architecture (Cremers, 1983), and indument 
(Wurdack, 1986). Memecylaceae comprise about 
89 species in the Neotropics and are much less 
diverse in these characters. 

The present survey is based on my observations 
of the reproductive biology of many Brazilian and 
some Venezuelan species and on a review of the 
literature. The survey is presented in five sections. 
First, an outline of the classification and repro- 
ductive morphological features of the Melastoma- 
taceae and Memecylaceae is given; second, breed- 
ing systems and cytology are discussed; third, the 
pollination spectra found in the two families are 
described; fourth, a brief survey is made of seed 
dispersal in the two families; fifth, the reproductive 
biological information is related to the fossil record 
and distributional data to contribute to a better 
understanding of the historical evolution of the two 
families. 

Flowering and fruiting phenology of neotropical 
melastomes and Memecylaceae are not dealt with 
here, but reference ma made to Snow (1965), 
Croat (1978), Opler et al. (1980), Hilty (1980), 
Lumer (1982), and Renner (1984 and in prep.). 
Suffice it to say that synchronous, episodic flow- 
ering with the episodes of short duration (a few 
days) characterizes some of the arborescent species 
of the largest genus, Miconia, whereas extended 
flowering periods (several weeks or months) are 
characteristic of many of the shrubs, herbs, epi- 
phytes, and some of the tree species. 


CLASSIFICATION AND REPRODUCTIVE 
MORPHOLOGICAL FEATURES 


Worldwide, the Melastomataceae A. L. Juss. 
(excluding the ca. ten species of Crypteronia, Ax- 
inandra, and Dactylocladus, i.e., the Crypteronia- 
ceae; Dahlgren & Thorne, 1984) and the Me- 
mecylaceae DC. consist of ca. 190 genera and 
approximately 4,800 species. In the Neotropics, 
the Melastomataceae comprise over 3,000 species 
in 107 genera (Table 1) and the Memecylaceae 
89 species in two genera. About half of these genera 
contain only one to four species, whereas one ge- 
nus, Miconia, has some 


lastomataceae as a subfamily, it has been pointed 
out (see Morley, 1953; Dahlgren & Thorne, 1984; 
Johnson & Briggs, 1984) that the differences be- 
tween these families are such that their union ob- 
scures rather than reflects our knowledge of the 
two groups. The Memecylaceae consist of the Asian 
genera Memecylon (ca. 300 spp.) and Lijndenia 
(4 spp.) (Bremer, 1982, 1983), the African Spa- 
thandra (6 spp.) and Warneckea (31 spp.), and 
the neotropical Votomita (8 spp.; Morley, 1963) 
and Mouriri (81 spp.). There is a consensus that 
the Melastomataceae and Memecylaceae belong to 
the core families of the order Myrtales (Dahlgren 
& Thorne, 1984; Johnson & Briggs, 1984). 

The classification of the Melastomataceae sensu 
stricto is essentially that of Triana (1867), who 
was closely followed by Cogniaux (1891). Wurdack 
(1980) has realigned the New World tribes in a 
sequence generally resembling that of A. P. de 
Candolle (1828). The main dichotomy in the Me- 
lastomataceae is between the capsular-fruited and 
the berry-fruited genera. The genera are arranged 
into tribes on the basis of fruit and seed characters 
and the presence and position of connective ap- 
pendages (D. Don, 1823; Triana, 1867; Cogniaux, 
1891; Gleason, 1929; comp. also Whiffin & Tomb, 
1972) 

Most Melastomataceae are characterized by 
poricidal anthers, the connectives of which are 
frequently prolonged (however, species of Miconia 
sects. Chaenopleura and Chaenanthera have an- 
thers that open by 1-2 slits). They usually have 
a pronounced foliar venation consisting of the mid- 
vein and one to four pairs of subparallel longitudinal 
veins. Memecylaceae, on the other hand, mostly 
have pinnate venation; their anthers open by short 
slits or pores. In a reproductive biological context, 
the presence of an elliptic, concave gland on the 
dorsal side of the connective is a unique feature 
characterizing most Memecylaceae. 


Volume 76, Number 2 
1989 


Renner 499 
Reproductive Biology in Melastomataceae 


& Memecylaceae 


The flowers in both families are actinomorphic 
or weakly to strongly zygomorphic as a result of 
movements of the stamens and style immediately 
after the expansion of the petals (Troll, 1922; 
Ziegler, 1925; pers. obs.). In Melastomataceae, 
the flowers are usually borne in well-developed 
cymose inflorescences, whereas in Memecylaceae 
the cymes are often condensed or reduced to a 
few flowers in fasciculate tufts. In both families, 
the mostly 4- or 5-merous flowers are perigynous 
or epigynous with a cup-shaped hypanthium that 
bears on its rim the calyx lobes, petals, and sta- 
mens. The antepetalous stamens are usually longer 
than the antesepalous ones, which may be stami- 
nodial; this dimorphism is more or less pronounced 
and has given occasion to much speculation con- 
cerning its role in pollination. In Melastomataceae 
with poricidal anthers, the anthers are one- or two- 
pored, a character that is consistent at the species 
level but varies within genera; there is also con- 
siderable variation in the size and position of the 
pores. Pollen grains are usually free, small, and 
rather smooth, although polyads and tetrads are 
known in species of Tococa and Miconia (Patel et 


al., 1984). 


BREEDING SYSTEMS 


The principal mode of promoting outcrossing in 
most Melastomataceae and Memecylaceae is the 
very effective spatial separation of pollen and stig- 
ma in the flowers (herkogamy). This is achieved 
by the pollen being enclosed in tubular anthers with 
apical or subapical pores or slits, which are gen- 
erally minute, and positioned well distant from the 
punctiform or capitate stigmas. Also, the stamens 
are usually shorter than the styles. In bud, the 
stamens are always folded over in such a way that 
the anther pores point downward and are maxi- 
mally distant from the stigma, which lies at the tip 
of the bud. Thus, chances for automatic selfing 
before and during anthesis are very low or nil. If, 
however, during anthesis, an anther or the style 
bends in such a way that the pore is brought into 
contact with the stigma, automatic selfing may 
occur. This has been reported for Monolena 
trichopoda (Melastomataceae; Warner, 1981) 
where, late in anthesis, the stamens fold over and 
the thecae are effectively stuck to the stigmatic 
exudate. 

No pollen escapes from the anthers unless me- 
diated by a vector, which at least potentially may 
arrive bearing conspecific pollen. Almeda (1977, 
1978) suggested that vigorous mechanical move- 


ment of the anthers by gusty winds or rains might 
release pollen in species of Monochaetum and Cen- 
tradenia (Melastomataceae). Additionally, he often 
observed pollen in water droplets connecting an- 
thers and stigma and suggested that this might be 
another possible route for selfing to occur. How- 
ever, bees, which regularly collect pollen from Me- 
lastomataceae, use vibrations of around 420 Hz or 
higher (this frequency was found to effect ample 
pollen emission from the large anthers of Bellucia; 
see below). Therefore, it seems unlikely that the 
low-frequency vibrations produced by wind could 
release pollen This 
impression is supported by the lack of fruit set in 
bagged flowers of nonagamospermic species ex- 
posed to strong wind (Renner, 1984). While autog- 
amy may be extremely rare, vector-mediated self- 
ing (i.e., geitonogamy), a function of pollinator 
behavior and number of conspecific flowers avail- 
able at a time, may be frequent in self-compatible 


rom melastome anthers. 


species. 

Temporal separation of male and female func- 
tion has been suggested for species of Monochae- 
tum (Almeda, 1978) in which *'the ill-defined stig- 
matic region . . . appears to be non-receptive during 
the first day." This may be a rare condition be- 
cause, as a rule, poricidal flowers that offer only 
pollen as a reward (subsequently also called “‘pol- 
len-only flowers”) receive foreign pollen at the same 
time a visitor harvests pollen from them; in female- 
phase flowers, there is nothing to reward visitors 
(only mimicry or deception might account for visits 
to female-phase flowers). Homogamy, i.e., the si- 
multaneous functioning of the male and female 
parts, is thus to be expected in pollen-only flowers. 
Temporal separation of the male and female func- 
tion in single flowers has been reported twice in 
nectariferous melastomes (Ule, 1896; Vogel, 1957); 
in both cases, the flowers were protandrous. 

The Melastomataceae are not entirely her- 
maphroditic. Wurdack (1964) was the first to de- 
scribe dioecism in Miconia dioica Wurdack; her- 
barium specimens of this species had female flowers 
with abortive stamens and male flowers with poorly 
developed stigmas. He also suspected dioecism in 
some other minute-flowered Andean species of Mi- 
conia (M. dielsii Markgraf, M. clathrantha Tr. 
ex Cogn., M. minuta Gleason; Wurdack, 1980: 
261-262) and in M. rubens (Sw.) Naudin from 
the West Indies and Venezuela (Wurdack, 1964). 
Field observations on the reproductive biology of 
these and some other suspected dioecious mela- 
stomes are lacking. 


At least one species of Lijndenia (Memecyla- 


500 


Annals of the 
Missouri Botanical Garden 


ceae) has hermaphrodite-flowered individuals and 
purely male-flowered ones (Bremen, 1982). 
GENETIC COMPATIBILITY 

As shown in Table Z, genetic self-incompatibility 
occurs in at least 22 species in the two families. 
As Darwin remarked (in a letter to J. D. Hooker 
on Nov. 28, 1871, in F. Darwin (ed.), 1903), 
melastomes *'are troublesome beasts to fertilise.” 
This is due to the closed anthers and mostly punc- 
tiform stigmas. In some large-flowered species, such 
as Bellucia (Melastomataceae), it is possible to 
collect large amounts of pollen for experimental 
purposes by holding a tuning fork to the stamens. 
Self-compatibility is found in 26 species (Table 2). 
Since no efforts have ever been made to distinguish 
pseudogamy from self-compatibility, several of the 
supposedly self-compatible species reported earlier 
(Darwin, 1876; Ziegler, 1925) may have set fruit 
as a result of agamospermy rather than of self- 
fertilization. Related species such as Tococa guia- 
nensis and T. bullifera or Centradenia grandi- 
flora and C. paradoxa (all Melastomataceae) differ 


O 


in their compatibility conditions (Table 2). Th 
humid stigmatic surface and the binucleate pollen 
of the Melastomataceae and Memecylaceae (Tobe 
& Raven, 1984) indicate that the incompatibility 
system is gametophytic (Nettancourt, 1977) 


AGAMOSPERM Y 


Experimental proof of agamospermy in mela- 
stomes is readily obtained by cutting off the style 
in freshly opened unvisited flowers. (To ensure that 
flowers are unvisited before the style is cut off, 
they must be bagged the evening before.) Apart 
from the 22 apomictic Melastomataceae listed in 

able 3, polyembryony, with additional embryos 
baina formed by nucellus or suspensor cells, has 
been reported from the Asiatic species Osbeckia 
hispidissima Wight, Sonerila wallichii Benn., and 
Melastoma malabathricum L. (Subramanyam, 
1942, 1944, 1948). Polyembryony is often cor- 
related with agamospermy (Gustafsson, 1946-47). 
Mouriri nervosa is the only member of the Me- 
mecylaceae so far tested for agamospermy, with 
negative results (Renner, unpubl.). In several of 
the neotropical species showing apomixis, for in- 
stance in Clidemia novemnervia and Macairea 
theresiae, the anthers contain very few pollen 
grains. In cultivated neotropical Leandra cordi- 
folia Cogn., Subramanyam (1942) also found that 
a low percentage of fertile pollen grains was com- 
bined with the frequent formation of two embryo 
sacs. In most neotropical species, agamospermy is 
correlated with low stainability of fresh pollen in 


cotton blue with lactophenol. This stain indicates 
presence of callose, a saccharide found in the intine 
of pollen. Lack of an intine could make a grain 
unviable and is usually accompanied by lack of 
cytoplasm. However, in Aciotis acuminifolia and 
Miconia argyrophylla, pollen grains stain heavily, 
yet these species are capable of producing seeds 
apomictically. In Clidemia and 
Maieta poeppigii, the stamens do not unflex com- 


novemnervia 


pletely upon anthesis but remain curved, and are 
thus not available for legitimate bee pollinators. 
Both species regularly set seeds apomictically. 

82) observed that Blakea gracilis, 
Blakea sp. (originally identified as B. grandiflora), 
and Topobea brenesii ripened a large number of 
seedless fruits from emasculated flowers; however, 
in T. pittieri, she found that of the bagged flowers, 
35% produced mature fruits containing seeds. Dar- 
win (1876) and Ziegler (1925) observed fruit set 
in cultivated plants of Monochaetum, Tibouchina, 
Centradenia, Calvoa, and Dissotis in the green- 
house. Assuming that there were no bee pollinators 


Lumer 


in the greenhouses, it is likely that such fruits 
resulted from agamospermy rather than from 
autogamy, which, as pointed out above, is me- 
chanically difficult in Melastomataceae. 

The number of species investigated is still too 
small to draw firm conclusions about correlations 
between habit, habitat, and breeding system. Of 
the agamospermous species, 13 are shrubs of var- 
ious types of savannas or of secondary vegetation; 
five are treelets, two are herbs, and one is a climber. 
The sample is clearly biased towards shrubby 
species, however, because of their accessibility. 
CYTOLOGY 

Some 300 of the over 3,000 species of neo- 
tropical Melastomataceae and only one species of 
Memecylaceae (Mouriri myrtilloides) are known 
cytologically (Solt & Wurdack, 1980). Only a few 
of these have been examined twice. Melastome 
chromosomes are very small (0.5-1 micron; Solt 
S Wurdack, 1980) and stain poorly. 
common base numbers are 9 and 17 (Wurdack & 
Kral, 1982). Based on Solt & Wurdack's results, 


the incidence of polyploidy in the Melastomataceae 


The most 


seems high. It cannot yet be answered whether 
polyploidy is more frequent in species with strong 
agamospermous reproduction than in those which 
lack apomixis, nor whether there is a correlation 
between polyploidy and growth habit. 


POLLINATION SYSTEMS 


In neotropical Melastomataceae and Meme- 
cylaceae, the most widespread pollination system 


Volume 76, Number 2 Renner 501 
1989 


Reproductive Biology in Melastomataceae 
& Memecylaceae 


TABLE 2. Compatibility systems in Melastomataceae. 


Sys- 
tem! Reference 
MELASTOMATACEAE 
MERIANIEAE 
Adelobotrys rachidotricha Brade SC Renner, 1984 
Graffenrieda latifolia (Naudin) Triana SC Sobrevila & Arroyo, 
1982 
BERTOLONIEAE 
Bertolonia marmorata Naudin SC Ziegler, 1925 
MICROLICIEAE 
Centradenia floribunda Planchon SC Darwin, 1876 
C. grandifolia (Schldl.) Endl. ssp. grandiflora SIC Almeda, 1977 
C. paradoxa (Kraenzlin) Almeda SC Almeda, 1977 
Rhynchanthera grandiflora (Aubl.) DC. SC Renner, 1984 
TIBOUCHINEAE 
Vepsera aquatica (Aubl.) Naudin SIC Renner, 1984 
Monochaetum amabile Almeda SC Almeda, 1978 
M. calcaratum (DC.) Triana i M. ensiferum Hook.) SC Darwin, 1876 
M. floribundum (Schldl.) Nau SC Almeda, 1978 
M. neglectum Almeda SC Almeda, 1978 
M. talamancense Almeda SC Almeda, 1978 
M. vulcanicum Cogn. SC Almeda, 1978 
RHEXIEAE 
Rhexia alifanus Walter SIC Kral & Bostick, 1969 
R. aristosa Britton SIC Kral € Bostick, 1969 
R. cubensis Griseb. SIC Kral & Bostick, 1969 
R. lutea Walter SC Kral & Bostick, 1969 
R. mariana var. interior (Pennell) Kral & Bostick SIC Kral & Bostick, 1969 
R. mariana L. var. mariana SIC Kral & Bostick, 1969 
R. mariana var. ventricosa (Fern. & Griscom) Kral € Bostick SIC Kral & Bostick, 1969 
R. nashii Small SIC Kral & Bostick, 1969 
R. nuttallii James SIC Kral & Bostick, 1969 
R. parviflora Chapman SIC Kral & Bostick, 1969 
R. petiolata Walter SC Kral & Bostick, 1969 
R. salicifolia Kral & Bostick SIC Kral € Bostick, 1969 
R. virginica L. SIC Kral € Bostick, 1969 
MICONIEAE 
Bellucia acutata Pilger SIC Renner, 1986/1987 
B. dichotoma Cogn. (as B. imperialis Saldanha & Cogn.) SIC Renner, 1984 
B. grossularioides (L.) Triana SIC Renner, 1986/1987 
Conostegia macrantha Triana SIC  Lumer, 1982 
Miconia araguensis Wurdack SC Sobrevila & Arroyo, 
1982 
M. dodecandra (Desr.) Cogn. SC Sobrevila & Arroyo, 
1982 
M. laevigata (L.) DC. SIC Sobrevila & Arroyo, 
1982 
M. macrodon (Naudin) Wurd. (as Heterotrichum macrodon (Naudin) 
Planchon) SIC Ziegler, 1925 
M. sp. 254 SIC Renner, 1984 
M. sylvatica (Schldl.) Naudin SC Sobrevila & Arroyo, 
1982 
M. tuberculata (Naudin) Triana SIC Arroyo & Cabrera, 
1977 
Tococa bullifera DC. SC Renner, 1984 
T. longisepala Cogn. SIC Renner, 1984 
BLAKEEAE 
Blakea anomala J. D. Smith SC Lumer, 1982 


B. chlorantha Almeda SC Lumer, 1980 


Annals of the 
Missouri Botanical Garden 


TABLE 2. Continued. 


tem! Reference 


B. pe Hemsley 


s B. grandiflora Hemsley) 
mith 


B. prea a D.S 

Topobea brenesii Standley 

T. durandiana Cogn. 
MEMECYLACEAE 


Mouriri nervosa Pilger 


SC Lumer, 1982 
SC Lumer, 1982 
SC Lumer, 1982 
SC Lumer, 1982 
SC Lumer, 1982 


SC Renner, 1984 


' SC = self-compatible, SIC = self-incompatible. 


involves female bees; it occurs in an estimated 98% 
of the species. Pollination by hummingbirds, bats, 


MELASTOMATACEAE 


POLLINATION BY POLLEN-COLLECTING BEES 


and rodents is found in an estimated 60 species of 


Meriania, Centronia, Brachyotum, Tibouchina, 
Miconia, Huilaea, Chalybea, and Blakea (Table 
4), and probably also in Axinaea (see below). The 
mode of pollination is determined by the type of 
reward offered: Melastomataceae and Memec yla- 
ceae with pollen-only flowers are, as a rule, polli- 
nated exclusively by bees because the morphology 
of the anthers excludes all other pollen-collecting 
vectors. In contrast, the nectar-offering species 


attract a wide variety of pollinators. 


a) Methods of pollen collection and kinds of bees 


Pollination by bees has been reported for some 
120 species in 31 genera of Melastomataceae (Ta- 
ble 4). In addition, a few species occurring at Finca 
La Selva, Costa Rica, and near Kourou, French 
Guiana, have been observed to be pollinated by 
small and medium-sized bees (Bawa et al. 1985; 
Roubik, 19792). Many unidentified melastomes are 
indicated as visited by bees in apidological studies 
(e.g., Sakagami et al., 1965, and Roubik, 1979b). 


TABLE 3.  Agamospermous Melastomataceae. 
Habit Habitat Reference 
Aciotis acuminifolia (DC.) herb forest understory Renner, 1984 
riana 
Clidemia capitellata (Bonpl.) D. — shrub savannas and disturbed vegetation Renner, 1984 
E d SR DC. climber ^ forest understory Renner, 1984 
C. fendleri Cogn shrub forest understory Sobrevila & Arroyo, 1982 
C. hirta (L.) D. Don shrub disturbed vegetation Renner, 1984 
C. novemnervia ed Triana shrub disturbed vegetation Renner, 1984 
C. rubra (Aubl. i. shrub savannas and disturbed vegetation Renner, 1984 
Macairea theresiae hs logn. shrub Amazonian white sand savannas Renner, 1984 
Maieta guianensis (Aubl.) DC shrub forest understory original 
Maieta poeppigii Cogn. shrub forest understory original 
Miconia alata (Aubl.) DC. shrub disturbed vegetation Renner, 1984 
M. albicans (Sw.) Triana shrub savannas (cerrado) Renner, 1984 
M. argyrophylla DC. treelet secondary forest Renner, 1984 
M. jn dió (Benth.) Tr. shrub disturbed vegetation Renner, 1984 
M. egensis Cogn. treelet secondary forest Renner, 1984 
M. nine iba ) DC. treelet secondary forest Renner, 1984 
M. prasina (Sw treelet secondary forest original 
M. regelii Cogn treelet secondary forest Renner, 1984 
M. spinulosa Naudin shrub forest understory la! : Hc 75 1982 
M. stenostachya DC. shrub vanna Renne 
M. tomentosa (L. C. Rich.) D. treelet disturbed vegetation uer Tee 
Don ex DC 
Rhexia mariana L. herb swamps, marshes Etheridge & Herr, 1968 


Volume 76, Number 2 
1989 


Renner 


503 


Reproductive Biology in Melastomataceae 
& Memecylaceae 


TABLE 4.  Pollinators of Melastomataceae and Memecylaceae. 
Important Pollinators' References 
MELASTOMATACEAE 
BERTOLONIEAE 


Opisthocentra clidemioides 
Hook. f. 
Salpinga secunda DC. 


MERIANIEAE 
Adelobotrys rachidotricha Brade 
Centronia phlomoides Triana 


Graffenrieda fruticosa Wurdack 


G. polymera Gleason subsp. neb- 
linensis 
G. reticulata Wurdack 


Meriania longifolia Cogn. 
M. tomentosa (Cogn.) Wurd. 
(7 Centronia excelsa (Bonpl.) 
DC.) 


MICROLICIEAE 
Centradenia grandifolia (Schldl.) 
Endl. 
Lavoisiera cordata Cogn. 
L. glandulifera Naudin 
Rhynchanthera grandiflora 
(Aubl.) DC. 


R. hispida Naudin 


R. sp. 


Siphanthera cordifolia (Benth.) 
Gleason 
Trembleya phlogiformis DC. 
TIBOUCHINEAE 


Aciotis acuminifolia (DC.) 
Triana 


A. annua (DC.) Triana 

A. circaeifolia (Bonpl.) Triana 

A. polystachya (Bonpl.) Triana 

Acisanthera uniflora (Vahl) 
Gleason 

Brachyotum ledifolium (Desr.) 
Triana 


Brachyotum ca. 40-45 species 


Desmoscelis villosa (Aubl.) Nau- 


Euglossa intersecta; Melipona ful- 


va 
Euglossa sp., Eulaema nigrita; Meli- 
pona sp. 


Ven E sp. 


Boris SPP -i A ai sp.; Melip- 
ona late 
Bombus spp.; nee lateralis 


Bombus spp.; Melipona lateralis; 
Dialictus sp.; Neocorynura sp. 
Xylocopa sp. 


hummingbirds 


Bombus sp.; Melipona sp. 


Bombus spp. 
Bombus s 


PP- 
Xylocopa frontalis, X. tegulata, X. 


muscaria; Centris nitens, C. flavi- 


frons, Eulaema nigrita; Monoe- 
ca sp. 

Xylocopa idein! Centris fus- 
cata; Ptiloglos 

Augochloropsis pa ani A. 
sparsilis, A. terrestris, Augo- 
chloropsis sp.; Pseudaugochlo- 
ropsis graminea 

Dialictus sp. 


Thygater analis, Bombus sp.; Xylo- 
copa sp. 


Megommation ogilvei; Paratetra- 
pedia duckei; Melipona fulva, 
Melipona sp.; Exomalopsini 

halictids; Paratetrapedia sp. 

lon sp. 

Paratetrapedia sp.; Augochloropsis 
callichroa, Augochloropsis sp.; 
Euglossa sp. 

halictids 


hummingbirds 
hummingbirds 
Dialictus sp.; Augochloropsis sp.; 


cf. tegulata; Augo- 
chlora sp.; Melipona sp. 


ylocopa 


Renner, 1984 

Renner, 1984 

Renner, 1984 

Vogel, 1988 

Renner, in press 

Renner, in press 

Renner, in press 

Vogel, 1978 

van der Werff, pers. comm.; Neill 
et al., label data 

Almeda, 1977 

Renner, 1984 

original 

Renner, 1984, and original 


Renner, 1984 
Laroca, 1970 
N. Ramirez, pers. comm., 1986 


Renner, 1984; Semir, pers. comm., 
1982 
Renner, 1984 


Renner, 1984 
Renner, 1984 
Renner, 1984, and original 


original 
Lagerheim, 1899 


Lagerheim, 1899; Wurdack, 1954; 
Fitzpatrick et al., 1979; B. Stein, 
pers. comm., 1985 

Roubik, 1979b; Renner, 1984, and 


original 


504 


Annals 


of the 
Missouri Botanical Garden 


TABLE 4. Continued. 


Important Pollinators' 


References 


Ernestia tenella (Bonpl.) DC. 
Macairea pachyphylla Benth. 
M. parv co Benth. 

M. rufescens DC. 


Monoc penat alpestre Naudin 
. amabile Almeda 


M. compactum Almeda 


. deppeanum (Schldl. & 
Cham.) Naudin 
. exaltatum Almeda 


M. 
M. 
M. cordatum Almeda 
M. 
M 


M. floribundum (Schldl.) Naudin 


= 


1. linearifolium Almeda 
M. macrantherum Gleason 


M. neglectum Almeda 


= 


talamancense Almeda 

M. tenellum Naudin 

M. trichophyllum Almeda 

M. vulcanicum Cogn. 

Nepsera aquatica (Aubl.) Naudin 

Pterogastra divaricata (Bonpl.) 
Naud. 

Purpurella see Tibouchina 


Sandemania hoehnei (Cogn.) 


Tibouchina arborea (Gardner) 
logn. 


8 
T. aspera Aubl. 


T. benthamiana (Gardner) Cogn. 
T. clavata (Pers.) Wurd. 


T. cleistoflora Ule (— T. 
din Ule, 1896, non 
nd. = Itatiaia ist 
inm (Ule) Ule, 1 
T. frigidula (DC.) a 


cleisto- 


T. gardneriana (Triana) Cogn. 
T. grandifolia Cogn 


Augochloropsis sp. 

Melipona sp. 

"uglossa sp. 

halictids 

Bombus sp.; Melipona sp.; Antho- 


phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus Es . Melipona sp.; Antho- 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 
phoridae 

Bombus sp.; Melipona sp.; Antho- 


phoridae 
Bombus sp.; Melipona sp.; Antho- 


P 
Megaloptidia contradicta; Megom- 


mation sp.; 


halictids 


Melipona sp. 


Augochloropsis 
roa, Augochloropsis sp.; 
Melipona fulva 


Bombus morio 


Megommation sp.; 
ic 


Aylocopa transitoria; Bombus sp.; 
Augochloropsis sp.; Euglo 
2 Melipona sp.; Ptiloglossa 


Pus hloropsis 


S Sp. 
Bombus morio; Pseudaugochlorop- 


s graminea 


Bom bus sp. 


Bombus sp.; Oxaea flavescens; 
Pseudaugochloropsis graminea 
Trigonopedia sp. 


1alictids 


original 

N. Ramirez, pers. comm. 1986 
N. Ramirez, pers. comm. 1986 
origina 

Almeda, 1978 

Almeda, 1978 

Almeda, 1978 

Almeda, 1978 
Almeda, 1978 
Almeda, 1978 


Almeda, 1978 


N 


Almeda, 1978 
Almeda, 1978 
Almeda, 1978 
Almeda, 1978 
Almeda, 1978 
Almeda, 1978 
Almeda, 1978 
Roubik, 1979b, Renner, 1984 


original 


Renner, 1984, and original 


F. Mueller, 1873 


Roubik, 1979b, Renner, 1984 


original 


Laroca, 1970 


Ule, 1896 


original 


original 
original 


Volume 76, Number 2 
1989 


Renner 
Reproductive Biology in Melastomataceae 


505 


Memecylaceae 


TABLE 4. Continued. 


Important Pollinators' 


References 


T. grossa (L. f.) Cogn. 


T. hospita (DC.) Cog 
T. longifolia (Vahl) Baillon 
T. moricandiana (DC.) Baillon 


T. mutabilis (Vell.) Cogn. 


T. ursina (Cham.) rn 


RHEXIEAE 
Rhexia virginica L. 
MICONIEAE 
Bellucia acutata m 
B. aequiloba Pilg 
B. dic coña eos (= 


B. im- 


B. grossularioides (L.) Tr. 


B. pentamera Naudin (— 
nanthera 

halybea corymbifera Naudin 
(7 Pachyanthus corymbifera 
(Naudin) Cogn.) 
Clidemia bullosa DC. 


C. capitellata (Bonpl.) D. Don 
C. hirta (L.) D. Don 


C. japurensis DC. 


C. rubra (Aubl.) Mart. 
Henriettea horridula Pilger 
Henriettella caudata Gleason 
Huilea macrocarpa Uribe 

H. penduliflora Wurd. 
Leandra micropetala Cogn. 
L. secundiflora (DC.) Cogn 
Leandra s 

Loreya riparia Renner 


— 


L. spruceana Triana 


B. axi- 


bats (Anoura geoffroyi); humming- 


Bombus morio 

Micrommation larocai 

Augochloropsis callichroa 

Bombus morio 

Bombus atratus; Xylocopa augusti; 

xomalopsis sp.; Augochloropsis 

a A. euphrosyne, A 
iris, A. leucotricha, A. scabri 
frons, A. sparsilis, A. terrestris 

“bees”; bumblebees 

Xylocopa frontalis; Centris lilacina 

Xylocopa spp.; Epicharis sp. 

Ptilotopus superbus; Centris lila- 
cina, Centris sp. nov.; Oxaea 

pe Epicharis conica, E. 
ustica, E. affinis 

pes X. tegulata, X. "similis 

Bombus spp.; Euglossa intersec- 

ta, E. cf. ignita; Eulaema me- 

riana, Eulaema nigrita, E. moc- 

zaryi 

m his Centris spp.; Epi- 
chari nica; Bombus sp.; Eu- 


glos ^ 
^on (m Centris spp.; 


picharis sp. 


hummingbirds 


Melipona eburnea; Augochloropsis 
s 


dis hloropsis sp 


Augochloropsis eee A. calli- 


pot halictids; as a 
M. lateralis, Melipon 


Euglossa sp. 
Pes dM sp.; Melipona ebur- 


nea, M. compressipes, Melipona 


halictids hoe 

Xyloc a Melipona sp. 

ERA sp.; Melipona sp. 

hummingbirds 

hummingbirds 

Augochloropsis sp.; Exomalopsini 

e du QE sp. 

halict 

Xy und frontalis 

Eulaema meriana; Xylocopa fron- 
talis 


Vogel, 1957 


Laroca, 1970 


Leggett, 1881; Eyde & Teeri, 1967 


Renner, 1984, 1986/1987 
Renner, 1986/1987 
Renner, 1984, 1986/1987 


Roubik, 1979b; Renner, 1984, 
9 


Renner, 1984, 1986/1987 


1986 


B. Stein, pers. comm., 


Renner, 1984, and original 


Renner, 1984 
Renner, 1984, and original 


Renner, 1984 


Renner, 1984 
Renner, 1984 
Renner, 1984 
Snow & Snow, 1980 
Snow & Snow, 1980 
Renner, 1984 
Renner, 1984 
Renner, 1984 


original 
Vogel, 1966; Renner, 1984, in 
press 


506 


Annals of the 


Missouri Botanical Garden 


TABLE 4. 


Continued. 


Important Pollinators' 


References 


Miconia alata (Aubl.) DC. 


= 


EX 


nia sp. 
Miconia sp. 1004 


pool 


alborufescens Naudin 
affinis DC. 

argentea (Sw.) DC. 
argyrophylla DC. 


. dispar Benth. 


egensis Cogn. 


. gratissima Benth. 
. ibaguensis (Bonpl.) Triana 


. minutiflora (Bonpl.) DC. 
(824) 


myriantha Benth 


. nervosa (Smith) Triana 


pileata DC. 


. poeppigii Trian 
. radulifolia ( (Benth. ) Naudin 


M. robinsoniana Cogn 


. rufescens (Aubl.) DC. 
. splendens (Sw.) Griseb. 
. tomentosa AL. C 


Rich.) D. 


Miconia es 


ora egensis DC. 


Myr 
Tococa 'nillifera DC. 


T. coronata Benth. 


T. guianensis Aubl. 


T. longisepala Cogn. 


T. pachystachya Wurd. 
T. subciliata (DC.) Triana 
T. tepuiensis Wurd. 


Melipona fulva; Lophothygater de- 
corata; Paratetrapedia sp.; Au- 
gochloropsis hebescens, A. cu- 
preola, A. callichora 
Pseudaugochloropsis graminea; 
Exomalopsis auropilosa; gen. 
nov. near Megommation 

Augochloropsis callichroa 

medium-sized bees 

"numerous spp. of bees" 

halictids 

Augochloropsis sp.; Exomalopsini; 
Melipona marginata; Xyloco 
similis 

Xylocopa ornata; Exomalopsis au- 
ropilosa; Paratetrapedia sp. 

Xylocopa do Melipona sp. 

Augochloropsis sp.; Xylocopa tegu- 
ata, X. muscaria Melipona sp.; 
Exomalopsin 

Augoc o callichroa; halic- 
tids; Melipona sp. 

Melipo 


sp 
uma dia sp.; Augochloropsis 


sp. 
Augochloropsis s 
Melipona fulva; Paratetrapedia sp. 
Paratetrapedia sp. 
Xylocopa darwinii 


To eburnea; Thygater sp. 
Melipo 
X Yiocopa ontalis, X. similis; Me- 


lipona spp. 
Augochloropsis sp. 
Melipon 
Pm pr Melipona fulva 
Eulaema cingulata 
as 


TES 
;uglossa intersecta, E. piliventris, 
E. chalybeata, E. ignita; Eulae- 
ma meriana; Melipona fulva; 
Paratetrapedia sp.; Megaloptid- 
ia Sp. 

Euglossa sp., E. intersecta; Augo- 
chloropsis s 

Melipona compressipes, Melipona 
sp.; Xylocopa sp.; Eulaema sp.; 
Augochloropsis sp. 

Eulaema mocsaryi, E 
Euglossa intersecta, Centris sp.; 
AXylocopa frontalis; Melipona 
fulva 

Bombus spp.; Xylocopa sp. 

Euglossa sp. 

Bombus spp. 


meriana; 


Renner, 1984 and original 


Renner, 1984, and original 


original 


Renner, 1984 
Renner, 1984, and original 


Renner, 1984; Ducke, 1902; Mori 
& Pipoly, 1984 


Renner, 1984 
Renner, 1984 


Renner, 1984 
original 
Renner, 19 
a et D. 1966; McMullen, 
Renner, 1984 

Renner, 1984 

Renner, 1984 


Renner, 1984 
original 

original 

Dodson, 1966 
Roubik, pers. comm., 
original 

Renner, 1984 


1987 


Renner, 1984 and original 


Roubik, 1979b; Renner, 1984 


Renner, 1984 and original 


Renner, in press 
Renner 84 
Hanes in press 


Volume 76, Number 2 
1989 


Renner 
Reproductive Biology in Melastomataceae 
Memecylaceae 


507 


TABLE 4. Continued. 
Important Pollinators' References 
Tococa sp. Melipona quinquefasciata; Laroca, 1970 
Exomalopsis fulvofasciata 
BLAKEEAE 


Blakea anomala J. D. Smith 


B. austin-smithii Standley 
B. chlorantha Almeda 
B. gracilis Hemsley 


B. penduliflora Almed 
B. sp. (a 

ley) 
B. tuberculata J. D. Smith 


Topobea brenesii Standley 


T. durandiana Cogn. 


T. praecox Gleason 


MEMECYLACEAE 
Mouriri guianensis Aubl. 


M. brevipes Hook. 


M. myrtilloides (Sw.) Poiret 


M. nervosa Pilger 


as B. Dose un Hems- 


Bombus ephippiatus, B. volucel- 
oides; Eulaema seabrai; Epi- 
charis sp.; Melipona fasciata, 
M. flavipennis; Neocorynura 
rupa 

rodents (Peromyscini, Oryzomini) 

rodents (Peromyscini, Oryzomini) 

Bombus epihippiatus, B. volucel 
loides; Eulaema seabrai, E. cin- 
gulata, E. P rptu Epicha- 
ris spp.; ona fasciata; 
Xylocopa aci halictids 

rodents (Peromyscini, Oryzomini) 

Bombus epihippiatus, B. volucel 
loides; Melipona fasciata; ha- 
lictids 

Bombus epihippiatus, B. volucel- 
lo 


ides; Eula aema seabrai; Xyloco- 


a fronta 
Bombus Ri qup B. volucel- 
loides; Eulaema seabrai, E. poly- 
chroma; Epicharis sp.; Melipona 
fasciata; Xylocopa frontalis; 
Crawfordopsis sp.; Pseudaugo- 


c us 2: Pseudaugochloropsis 
igerrim 


Eula aema “mexicana” (Surely 
another species, since E. mexi- 
cana does not occur in Panama; 
J. Ackerman, pers. comm., 


1985) 


Ceratina sp.; Euglossa spp 

Eulaema nigrita; Xylocopa fronta- 
lis 

Euglossa imperialis, E. tridentata, 
Euglossa sp.; Paratetrapedia 


catcarata 

Euglossa chalybeata, E. crassi- 
punctata, E. purpurata, E. viri- 
dis, Eulaema mocsar yi, 
riana; Xylocopa frontalis, X. 
similis, X. ornata; Melipona ful- 
va, M. marginata 


Lumer, 1982 


Lumer & Schoer, 1986 
Lumer, 1980, 1982 
Lumer, 1982 


Lumer & e 1986 


Lumer, 


Lumer, 1982 


Lumer, 1982 


Lumer, 1982 


Croat, 1978 


Renner, 1984, and original 
original 


Buchmann & Buchmann, 1981 


Renner, 1984, and original 


' “Important pollinators” 
Melastomataceae. Note that 
ascertained at present. 


excludes 
“halictids”” 


the numerous Trigona species, which are omnipresent pollen scavengers on 
stands for many different species, the correct names of which cannot be 


508 


Annals of the 
Missouri Botanical Garden 


Bees gathering pollen from Melastomataceae flow- 
ers must be able to shake it out of the anthers or 
else pull pollen grains out through the pores with 
their tongues. This latter mode of exploiting mela- 
stome flowers is used by a group of Trigona bees 
and does not usually result in pollination because 
these bees are too small to contact the stigmas. 
Another subgroup of Trigona bees, characterized 
by having toothed mandibles, cuts up melastome 
anthers and robs their pollen (Laroca, 1970; 

ner, . 
The method of pollen collecting used by all other 
bees to extract pollen from melastome anthers was 
first described correctly by van der Pijl (1954). 
When approaching a flower, in flight, the bees 
orient themselves with the group of stamens and 
the style and then straddle the stamens. They usu- 
ally grasp the filaments or the bases of the anthers, 
quite often using their mandibles. Therefore, after 
having been visited, flowers often show necrotic 
spots on the outside of the stamens where the bees 
held onto them. When the bees position themselves 
over the androecium, their potentially pollen-bear- 
ing surface contacts the stigma even before they 
start exploiting the anthers. Also, they strive to 
establish a firm contact with as many stamens as 

fT, 


their body size permits, going tc 
in order to bundle all 8, 10, or 12 stamens. Close 
contact between the bee and the stamens is nec- 
essary for pollen collection by means of thorax 
vibrations, which are transmitted to the stamens 
and cause the pollen to stream out of the anther 
pores. Electrostatic forces due to the net positive 
charge of the bees and the negative charge of pollen 
greatly aid pollen-load adhesion (Thorp, 1979). 
When a pollen load, taken with forceps from a 
freshly captured bee, is released near this bee, it 
jumps back onto it (pers. obs.). The very rapid 
distortions of the thorax-box (with the wings at rest) 
are made possible by the unique contractile prop- 
erties of the bees’ fibrillar indirect flight muscles. 
Pollen collecting by this method is always accom- 
panied by a characteristic buzzing sound and has 


Michener, 1962). 

The rapid vibrations used when harvesting mel- 
astome pollen presumably produce a high temper- 
ature of the flight muscles, 
this method of pollen collecting is particularly ap- 


been termed “‘buzzing method” (van der Pijl, 1954; 
) 


which may mean that 


propriate during the relatively cool early morning 
hours. Also, there are ecological constraints on the 
activity of dark-colored bees during the hottest 
hours. Buzz pollination continues with very high 
humidity of the air and even during slight rains. 
In most melastomes, pollen deposition and re- 


ception is on the lower abdomen (sternotribic) or 
lateral on the bees (pleurotribic); it is on the back 
(nototribic) in a few species with downwards-curved 
styles (e.g., species of Meriania, Adelobotrys, Sal- 
pinga) and in Desmoscelis villosa. In these cases, 
bees assume quite unexpected positions in order to 
vibrate the stamens; in Adelobotrys rachidotricha, 
for instance, they buzz with the head pointing out- 
ward and downward. Relatively large flowers, such 
as those of Blakea, Topobea, and Bellucia, are 

by a ua range of bees (Lumer, 1982; 
Renner, 1986/1 87). For example, stamens of 
Bellucia are us by 33-mm-long Centris and 
8-mm-long Augochloropsis bees. In the case of 
self-compatible species, visits by bees too small to 
contact the stigma and deposit foreign pollen may 
still result in self-pollen landing on the stigma when 
such small bees vibrate one or a few stamens. 

Bees visiting Melastomataceae exhibit consid- 
erable steadiness during individual foraging flights: 
pollen loads often consist of almost pure melastome 
pollen with very few grains from nectariferous flow- 
ers or with grains from Cassia mixed in (Renner, 

984). Cassia has buzz-pollinated flowers, and bees 
often visit several such flowers (see below). 

In Brazilian Amazonia, | have captured 80 
species of bees, including the Africanized honey 
bee, Apis mellifera scutellata Lepeletier, vibrating 
melastomes. (A list of all bees, including the species 
of Trigona, may be obtained upon request.) Ad- 
ditional species have been captured in extra-Am- 
azonian areas and at higher altitudes. To date, at 
least 100 species of bees have been found polli- 
nating melastome flowers (Table 4; this does not 
include the numerous species of pollen-scavenging 
Trigona bees). The relatively few groups of bees 
that have been successful in tropical forest envi- 
ronments, the euglossines, halictids, Xylocopa, and 
Melipona (Michener, 1979) 


tant pollinators of lowland melastomes. In savan- 


are the most impor- 


nas, Centridini, Oxaeidae, Exomalopsini, and Ptilo- 


a 
o 


glossa (Colletidae) become important, as 
bumblebees in southern Brazil and at higher alti- 
tudes. The bee fauna in neotropical forests and 
savannas is still incompletely known; however, based 
on bee inventories in Panama, Costa Rica, French 
Guiana, Surinam, and at the mouth of the Amazon 
(Heithaus, 1979; Roubik, 1979a, b; and ref. in 
Michener, 1979), it may be estimated that only 
about 300 species of bees occur in these forests, 
a number that includes non-flower-visiting parasitic 
and leaf-cutter bees. Thus, between 80 and 100 
species collecting pollen from Melastomataceae 
represents a high proportion of the total Amazonian 
bee fauna 


Volume 76, Number 2 
1989 


Renner 509 
Reproductive Biology in Melastomataceae 


& Memecylaceae 


b) Floral attractants 

Melastome pollen is an important pollen source 
for neotropical bees. This is clear from a survey 
of literature on bee foraging: it is collected by 
female euglossines (Moure, 1970; Janzen, 1971; 
Michener et al., 1978; Roubik, 1979a, b; Dressler, 
1982; Renner, 1984), meliponines and halictids 
(Ducke, 1901, 1902; Laroca, 1970; Roubik, 
1979a, b; Michener et al., 1978; Absy & Kerr, 
1977; Absy et al., 1980), mixed with Malpighi- 
aceae oil by anthophorine bees (Vogel, 1974; Ren- 
ner, 1984), and makes up 96-100% of the nest 
provision of the colletid bee Crawfordapis (Roubik 
& Michener, 1985). A chemical analysis of Bel- 
lucia dichotoma pollen revealed high nitrogen and 
total crude protein. (I am indebted to S. Buchmann 
for analyzing a pollen sample in 1983.) In this 
species, the pollen has a strong perfume resembling 
that of the flowers. 

Secondary attractants for bees include sweet or 
perfumelike floral odors produced by the petals, 
the connective, and the upper section of the style 
(as determined by neutral red staining; pers. obs.; 
method, Vogel, 1963), and the presence of a land- 
ing platform provided by the androecium. I have 
never seen bees land on the petals. Frequently, the 
filaments are adorned with glandular or eglandular 
hairs, which may make them easier for the bees 
to grasp. Also, direct alighting on the stamens may 
make visits to flowers with poricidal anthers par- 
ticularly rewarding compared with flowers requir- 
ing further crawling and searching after landing. 

Most Melastomataceae seem to be visited and 
pollinated early in the morning, sometimes 20 min- 
utes before sunrise (F. Mueller, 1873; Frankie, 
1976; Lumer, 1982; Renner, 1984), and many 
species close their petals by late morning. However, 
flowers of Monochaetum and Rhynchanthera open 
late, two to four hours after sunrise (Almeda, 1978; 
pers. obs.). Shedding of the flower parts usually 
starts at the end of the second day. Again, Mono- 
chaetum differs in having flowers that stay open 
for three days (Almeda, 1978), and I have seen 
Graffenrieda fruticosa flowers staying attractive 
for at least seven days. In the latter case, the 
adverse climatic conditions at 2,200 m altitude, 
where Graffenrieda fruticosa occurs, may place a 
high selective advantage on longer flowering pe- 
riods which presumably heighten the flowers' 
chances of being visited. 

The typical bee colors of most Melastomataceae 
flowers, white, purple, or pink, are due to total 
reflection and anthocyanins (Lowry, 1976; Bobbio 
et al., 1985). Yellow and orange flowers evolved 
repeatedly in isolated species throughout the fam- 


ily. Contrasting marks at the base of the petals and 
color changes during anthesis are frequent in 
Melastomataceae (F. Mueller, 1873; Ule, 1896; 
Wurdack, 1980; Renner, 1984). The colors change 
from white to red, from yellow to red, from white 
to purple, or from pink to white. The bright white 
of the small flowers of many herbaceous understory 
melastomes, e.g., species of Aciotis, Leandra, Cli- 
demia, Miconia, and Ossaea, may be of selective 
advantage under conditions of low light intensity. 
Some melastome flowers tested (using a Kodak 18 
Wratten filter) did not absorb ultraviolet light (Lu- 
mer, 1982), as was to be expected from the absence 
of flavonols from their petals. 

The androecia of many species in the capsular- 
fruited tribes Merianieae, Bertolonieae, Micro- 
licieae, Tibouchineae, and Rhexieae show a strong 
morphological differentiation into two sets of sta- 
mens. In addition, in such flowers the connective 
appendages may be large and stiff, forming 5, 10, 
or 15 usually yellow spurs, sometimes positioned 
like true anthers. Based on his brother's (Fritz 
Mueller) observations on a southern Brazilian species 
of ** Heeria," Hermann Mueller (1881, 1883) was 
the first to suggest that this dimorphism was the 
ultimate result of a **division of labor" between the 
two sets of stamens (his illustration shows that his 
brother was probably dealing with a species of 
Microlicia; Heeria is synonymous with Hetero- 
centron, which does not occur in southern Brazil). 
According to this idea, the conspicuous anthers 
attract the bees and offer “food pollen” as a re- 
ward, whereas the less conspicuous ones contain 
“functional pollen." The “functional pollen" is sup- 
posed to be placed on a part of the bee where it 
will not be groomed off and used as food for larvae. 
It will instead function in fertilization. Hermann 
Mueller’s idea caught on immediately (Forbes, 
1882; F. Mueller, 1883) and is usually mentioned 
as typical of the pollination strategy of Melasto- 
mataceae (e.g., Cammerloher, 1931; Percival, 
1965; Kugler, 1970; Proctor & Yeo, 1972; Ba- 
ker, 1978; Faegri & van der Pijl, 1979). 

A partitioning of the bee surface would be nec- 
essary for selection to favor the pollen grains from 
one set of stamens over those from the other and 
thus lead to pollen dimorphism. Pollination mech- 
anisms involving vibration of anthers, however, 
may not allow a sufficiently precise placement of 
pollen from two sets of stamens on bees. Forbes 
(1882), working with unnamed species of Mela- 
stoma, reported that “only the pollen from the long 
stamens seemed [sic] to be fertile"; yet he found 
pollen from both types of stamens on the stigmas. 
To date, with the exception of Forbes's report 


510 


Annals of the 
Missouri Botanical Garden 


(1882), no morphological or physiological differ- 
ences between pollen grains from dimorphic sta- 
mens have been found in any of the Melastoma- 
taceae studied (Darwin, 1876; Ziegler, 1925; 
Whiffin, 1973; Almeda, 1977; Renner, 1984). 
Further experimental fieldwork is needed to assess 
whether the staminal dimorphism and the stamen 
appendages have a function beyond that of en- 
hancing the visual attractiveness of the flowers and 
making the stamens easier to grasp for bees. In 
view of the importance of pollen aromas for guiding 
bee behavior (cf. Dobson, 1987), it is uncertain if 
the visual cues provided by the yellow connective 
appendages by themselves elicit pollen-collecting 
behavior 
POLLINATION OF NECTAR-PRODUCING MELASTOMATACEAE 
BY BIRDS, BATS, RODENTS, AND BEES 

Some 60 species of Meriania, Centronia, 
Brachyotum, Tibouchina, Miconia, Huilaea, 
Chalybea, and Blakea, all of them, except the 
species of Miconia, occurring at higher altitudes 
(1,600-3,400 m), offer nectar as a reward for 
their pollinators. On the basis of their character 
syndrome (see below), Meriania drakei (Cogn.) 
Wurd., M. furvanthera Wurd., 
and several other species of Meri- 
Centronia, and Tibouchina are 


Axinaea weber- 
baueri Cogn., 
ania, Áxinaea, 
expected to produce nectar, too. 
he first reports on nectar production in mel- 
astomes are by Ule (1895, 1896). In three species 
of Tibouchina (Table 4) growing on the Brazilian 
Itatiaia mountain range, he observed that nectar 
was produced by the staminal vascular bundle and 
secreted either directly from the filaments or some- 
times through a slit in the filaments. He suggested 
bumblebees as the probable pollinators of these 
species. At Ule's request, the chemist T. Peckolt 
analyzed the stamens of another species that did 
not secrete nectar (Ule gave the name as 7. glare- 
osa Cogn.; there is no such name, and he probably 
meant 7° granulosa Cogn.). Peckolt found that 
ugar was present “in great quantity," which sug- 
eed to Ule (1896) that this was likely true for 


other species of Tibouchina as well. In several 


Pry 


other nectariferous melastomes, slits in the upper 
ventral side of the filaments have been observed, 
e.g., in Tibouchina cleistoflora Ule (Ule, 1896; 
pers. obs.), T. grossa, Blakea chlorantha (Tobe 
et al., in prep.), Brachyotum ledifolium, B. an- 
dreanum Cogn., B. benthamianum Triana, and 
B. rugosum Wurd. (Lagerheim, 1899; pers. obs.). 
They have also been found in non-aectariferous 
species— such as Centradenia grandiflora, Het- 
erocentrum roseum, Triolena pustulata (as Ber- 
tolonia pubescens) —and in paleotropical species 


of Dissotis and Medinilla (Ziegler, 1925). These 
slits appear not to be consistently formed sutures, 
but rather spontaneous ruptures. The anatomy of 
the nectariferous tissue is being studied by Tobe 
et al. (in prep.). 

eports of nectar in Miconia minutiflora (Mori 
& Pipoly, 1984) are unclear, since its presence in 
the minute flowers (2.5 mm in diameter, style 4 
mm long) was only presumed. In central Amazonia, 
where I have observed this species, it did not pro- 
duce nectar (Renner, 1984). However, in another 
Miconia species in the Belém region at the mouth 
of the Amazon, Roubik measured floral nectar of 
ca. 20% mgar tad and collected Meli- 
, pers. comm. 198 


Recently, nectar N has also been con- 
firmed in the Antillean endemics Miconia sintenisii 
Cogn. and Mecranium amygdalinum (Desr.) 

/right (Dent, pers. comm.). Miconia robinson- 
iana, also with minute white flowers, was included 
under "nectar sources" 
records for Xylocopa darwinii by Linsley et al. 
1966). McMullen, who made further observations 
on the floral visits of this bee on Galápagos 
(McMullen, 1985), has never noticed any nectar 
in M. comm., 
1987) 

Except in Miconia, nectar production in Me- 
lastomataceae is correlated with floral morpholog- 
ical changes that relate to the fact that the nectar 
'Therefore, 


in a summary of flower 


~ 


robinsoniana (McMullen, pers. 


collectors do not vibrate the stamens. 
when nectar instead of pollen is the pollinator re- 
ward, the latter must be deposited on a visitor 
during nectar uptake. This is achieved by the an- 
thers having rather large pores and being stiffened, 
particularly at the base. Pressure on the base of 
the anthers exerted by the head of a bee, the bill 
of a bird, or the front paws of rodents clasping the 
flowers ejects pollen onto Hi visitor. Lagerheim 
1899) therefore speaks o ellow”” anthers; he 
found that a match the thickness of a hummingbird 
bill pushed into the flower of Brachyotum ledifoli- 
um so that it pressed the anther bases brought 
forth a pollen cloud 3 cm long. The flowers of the 
nectariferous species are inclined or pendent and, 


— 


in the case of the bird-pollinated ones, usually dark 
ed, deep 
Brachyotum ledifolium the petals are yellow, but 
the hypanthium is covered with bright red hairs. 
etals do not 


urple, or bluish to almost black. In 


In nectariferous melastomes, the 
spread at anthesis but remain convolute almost like 
in bud. Vogel (1957) reported a nocturnal cabbage 
odor in the predominantly bat-pollinated Tibouchi- 
na grossa. The other nectariferous melastomes are 
reported as scentless. 


Volume 76, Number 2 
1989 


Renner 511 
Reproductive Biology in Melastomataceae 
& Memecylaceae 


The bulk of nectariferous melastomes seem 
hummingbird-pollinated (Table 4). Most of them 
belong to the genus Brachyotum, which comprises 
50 species, the majority ornithophilous (Wurdack, 
1965, 1980; Lagerheim, 1899; Vogel, 1957; Fitz- 
patrick et al., 1979; label data). Other bird-polli- 
nated species are Meriania tomentosa (Cogn.) 
Wurd. (Centronia excelsa (Bonpl.) DC.; van der 
Werff, pers. comm.; D. Nei 
tariferous and hummingbird-pollinated species in 
the Miconieae are Huilea macrocarpa, H. pen- 
duliflora (Snow & Snow, 1980) and probably also 
Chalybea corymbifera (in which Stein observed 
nectar but no pollination events; pers. comm., 
1987). The Colombian Tibouchina grossa is pol- 
linated by bats and hummingbirds (Vogel, 1957; 
Stein, pers. comm., 1987) and certainly the same 
is true of T. Stricta Wurd. and T. reticulata Cogn. 

New World rats and mice of the tribes Pero- 
mycini and Oryzomini pollinate at least three nec- 
tariferous species of Blakea in Costa Rica (Lumer, 
1980, 1982; Lumer & Schoer, 1986). Lumer 
found that bees and hummingbirds rarely visited 
the flowers of these species during the day but that 
nectar secretion was nocturnal. The Blakea species 
involved are hemiepiphytic, and their flowers are 
positioned close to the stem where the rodents can 
reach them conveniently. 

The nectar of Blakea chlorantha contains ca. 
1195 sucrose (Lumer, 1982, and pers. comm.); 
that of Huilea macrocarpa has 12-16% with a 
mean of 13.495 (Snow & Snow, 1980); that of 
Chalybea corymbifera 16.5%; and that of 
Brachyotum ledifolium 20-22% (Tobe et al., in 
prep.). It would thus be classified as relatively rich 
in sucrose in the nectar class system of Baker & 
Baker (1983), who found that sucrose-rich or su- 
crose-dominated nectar prevails in hummingbird 
flowers. Thus, nectariferous melastomes offer a 
reward “intended” for pollinators other than bees. 

The evolutionary origin of pollination systems 
involving nectar in Melastomataceae is considered 


below 


ll, pers. comm.). N 


MEMECYLACEAE 


In their pollination mechanism, the neotropical 
Memecylaceae closely resemble the Melastoma- 
taceae, and the above description of bees’ methods 
of pollen collecting from melastome flowers also 
applies to them. In Mouriri, the anthers are stiff, 
with thickened endothecial walls, and are slightly 
prolonged just above the insertion of the filaments. 
The prolongations, termed "caudae" (Morley, 


1953), are used as footholds by the bees exploiting 


Mouriri anthers. Pollination in four species of Mou- 
riri has been observed (Table 4); the main polli- 
nators seem to be euglossines, followed by Xyloco- 
pa and Melipona species. They are legitimate 
visitors, i.e., large enough to contact the stigma 
regularly while collecting the reward. Memecyla- 
ceae anthers differ from those of Melastomataceae 
in bearing dorsal concave glands, 0.3-0.8 mm 
long. (For illustrations of the permutations in size 
and position of these glands in Mouriri and Vo- 
tomita, see Morley, 1953, 1976, and 1985.) None 
of the legitimate pollinators show any interest in 
the staminal glands (Buchmann & Buchmann, 
1981; Renner, 1984, and unpubl.). 

Earlier (1984), I reported that Melipona mar- 
ginata and M. fulva sometimes collected the se- 
cretion from the glands with the front legs, and 
that they mixed this substance with their pollen 
loads between pollen-collecting bouts on the wing. 
Further field observations, however, have con- 
vinced me that Melipona does not regularly touch 
the glandular surface when buzzing. The bees grip 
the stamens by the anther caudas just below the 
glands, and it is difficult to see what they do with 
their feet or their mouth parts when vibrating be- 
cause they curve tightly over the anthers to allow 
the pollen stream to settle on their bodies. Between 
buzzing bouts, Melipona bees (and the euglossines) 
rotate over the androecium so as to drum out the 
pollen from all ten anthers. Staining of their freshly 
collected corbicular loads with Fehling solution 
showed that the pollen had been wetted with sugar. 
With Sudan III (a lipid stain), Mouriri pollen grains 
stained red, but the secretion holding them together 
did not; this implies that the pollen contains lipids, 
whereas the secretion did not. 

Mouriri flowers are also visited by several species 
of Trigona. These small bees do not regularly con- 
tact the stigma but rob pollen from the anthers, 
which they are unable to buzz. Their role as pollen 
scavengers and robbers in Melastomataceae is well 
known (Renner, 1983). When cutting up anthers 
of Mouriri nervosa to reach pollen grains further 
down, Trigona williana very rarely touched the 
glands (Renner, 1984, and unpubl.). Most often, 
T. williana cut off the anthers above the glands, 
then stopped and started cutting and robbing the 
pollen of another anther. This shows that this pollen 
robber, like the legitimate pollinators, does not reg- 
ularly collect the glandular exudates. Buchmann 
& Buchmann (1981) have observed T. pallens 
manipulating the glands of M. myrtilloides with 
the front legs and mouth parts. 

The secretion of the staminal glands is yellow, 
viscous, stains red in Sudan III, and tests positive 


512 


Annals of the 
Missouri Botanical Garden 


for fatty substances. That of the relatively large 
(0.8 mm long) glands of M. lunatanthera forms 
myelin figures in water after a drop of sodium 
hydroxide is added (pers. obs.; method from Vogel, 
1974). The secretion in M. myrtilloides has been 
reported to be a lipoidal mixture of many com- 
pounds (e.g., phenolic Rud. small amounts 
3 fatty acids, and 
free fatty acids) and to be * een rich in non- 
(Buchmann & Buchmann, 


of glucose and saponin, at least 
lipid constituents" 
1981) 


Since the secretions from the floral glands con- 
tain lipids, the relationship between Mouriri and 
oil-collecting bees is of particular interest. Floral 
oils are collected by bees belonging to relatively 
few genera of Anthophoridae (e.g., Centris, Epi- 
charis, Monoeca, Paratetrapedia, Tapinotaspis, 
Tetrapedia; cf. Vogel, 1974; Ne Simpson, 

1981). Although many of these collect pollen from 
Melastomataceae (comp. Table 4), to date only a 
single anthophorid bee species, Paratetrapedia 
calcarata, has been observed on Mouriri (Buch- 
mann & Buchmann, 1981). This bee, due to its 
small size (7-8 mm long), rarely vibrated more 
than two or three of the ten anthers in each flower 
and only rarely touched the stigma (l.c.). Also, 
Buchmann € Buchmann (1981) reported that 
Paratetrapedia calcarata was an infrequent vis- 
itor of M. myrtilloides (16 times during 50 hours) 
and not a legitimate pollinator. 

The secretion of the staminal glands forms a 
shiny film on the gland cuticle and in this respect 
differs greatly from the lipid-filled blisters of other 
epithelial elaiophores (Vogel, 1974). Paratetra- 
pedia has scrapers on the fore-basitarsi (Neff & 
Simpson, 1981) and ruptures the cuticle of the 
elaiophores from which it collects oils. While scru- 
tinizing hundreds of stamens from Mouriri nervosa 
and M. guianensis under the stereoscope in order 
to collect their glandular secretions, I never found 
signs of the glands having been scraped. This was 
perhaps to be expected, since the secretion forms 
a film on the outside of the cuticle. 

A further problem in understanding the role of 
the staminal glands is that they are so small and 
positioned in such a way that the collection of any 
exudate either with the fore-basitarsi or the mouth 
parts by the medium-sized bees that pollinate Mou- 
riri is difficult to envisage. Visitors small enough 
to be attracted by the minute quantities of secretion 
offered (such as the Paratetrapedia species ob- 
served by Buchmann & Buchmann, 1981) are not 
large enough to touch the stigmatic surface when 
manipulating the glands, especially since the styles 
in Mouriri are always longer than the stamens and 
the stigmas are held well away from the anthers. 


A possible role of the staminal glands suggested 
by Morley (1976) is odor production. Mouriri 
species often have a strong perfume scent (Buch- 
mann & Buchmann, 1981; Renner, 1984). Per- 
haps a volatile oil would persist longer when mixed 
with the fatty oils and other compounds. Buchmann 
& Buchmann (l.c.) suggested that the glands, which 
absorb ultraviolet light, might visually guide the 
bees to the pollen. Extending this idea, one might 
ascribe a pseudonectary role to the staminal glands. 
It is interesting that in the Bonnetiaceae, at least 
23 species of Caraipa and Mahurea have very 
similar minute urceolate glands positioned subapi- 
cally on the connective (Kubitzki, 1978). The role 
of these glands, as of those of Mouriri, is obscure. 


SEED DisPERSAL IN MELASTOMATACEAE 


Of the neotropical Melastomataceae, 40% have 
capsular fruits and wind-dispersed seeds (Renner, 
1986). The remaining 60% have soft, juicy berries 
and endozoochorous seeds. Animal-dispersed Me- 
lastomataceae are most diverse in lowlands and 
montane forests, whereas the capsular-fruited tribes 
are richest in species in various types of savannas. 
Melastome capsules are often sturdy and persist 
on the plants for many months, during which the 
seeds are gradually shaken out. There are no data 
on dispersal distances in the wind-dispersed species; 
however, the many striking disjunctions observed 
1987b, 
press) probably attest to their good dispersability. 


in savanna melastomes (Renner, and in 
Small herbaceous understory species with broadly 


conical capsules ee O Bertolo- 


nia) may also have thei 
by raindrops. 


Melastome berries range in size from five to ca. 


d ballistically 


35 mm in diameter. The smaller ones are of out- 
standing importance for small frugivorous birds 
throughout the Neotropics (Huber, 1910; Snow, 
1965, 1971, 1981; Ricklefs, 1977; Skutch, 1980; 
Lumer, 1982; Wheelwright et al., 1984; R. Bier- 
regård, pers. comm., and many additional refer- 
ences). Epiphytic Melastomataceae are mostly bird- 
dispersed (Renner, 1986), as are many of the most 
successful pioneer species, for example, Clidemia 
hirta, which has become a noxious weed in Africa, 
Asia, and Hawaii. 

Fruit attributes, such as position, size, aroma, 
and color, indicate that different animal classes act 
as the principal dispersers in different species groups. 
For example, black spider monkeys are the pre- 
dominant consumers and likely dispersers of the 
green, slightly astringent and relatively large ber- 
ries of the tree Bellucia grossularioides in Suri- 
nam (Roosmalen, pers. comm. ). In French Guiana, 


Volume 76, Number 2 
1989 


Renner 513 
Reproductive Biology in Melastomataceae 
& Memecylaceae 


this species is a favorite with the PAilander opos- 
sum (Atramentowicz, 1982), which also eats Hen- 
riettea and Loreya fruits. However, a search of 
recent literature on feeding records (Renner, 19872) 
showed that Melastomataceae are not among the 
preferred foods of either monkeys or bats (for bats, 
Fleming, 1986; but compare Foresta et al., 1984, 
and Fleming, 1988). Perhaps because the majority 
of Melastomataceae are shrubs or understory trees 
with terminally positioned infructescences, they are 
unsuited for foraging monkeys and bats. Also, the 
taste of the mostly sweet, watery melastome berries 
may not be to the liking of monkeys, which ap- 

parently prefer slightly acidic fruits (Sourd & Gau- 

tier-Hion, 1 ). 

Fruits that have fallen to the ground are often 
eaten by turtles (Moskowitz, pers. comm.; Roos- 
malen, pers. comm.), lizards (W. Magnusson, pers. 
comm.), rodents (Magnusson & Sanaiotti, 1987), 
tapirs (Huber, 1910), and ants (pers. obs.). 


SEED DISPERSAL IN MEMECYLACEAE 


All Memecylaceae have berries, ranging in size 
from 0.5 to 10 cm long; color at maturity is most 
commonly yellow to red, less often it is purple- 
black to black (Morley, 1976). The smaller-fruited 
species are surely bird dispersed; some larger-fruit- 
ed ones are monkey-dispersed (Roosmalen, 1985). 
Fish eat the fruits of at least one riverine Mouriri 
species (Goulding, 1980), which is curious, since 
Morley (1976) reported that two other species are 
used for poisoning fish. 


REPRODUCTIVE BIOLOGY AND THE EVOLUTION OF 
MELASTOMATACEAE AND MEMECYLACEAE 


Floral morphology in Melastomataceae and Me- 
mecylaceae can only be understood as the result 
of coevolution between the earliest ancestors of 
these groups and pollen-collecting bees. Flowers of 
the Old World members of both families have the 
same basic construction as those of their New World 
relatives and, although little information is available 
on their floral biology, van der Pijl's (1939) detailed 
description of the pollination mechanism in some 
species of Osbeckia, Medinilla, and Melastoma 
makes it likely that they are pollinated in the same 
manner as the New World ones. The pollen offered 
in the poricidal stamens of both families is acces- 
sible exclusively to bees, but not to many of the 
most common ones, such as the numerous species 
of Trigona and the paleotropical Apis, which ap- 
parently do not vibrate stamens. Most Melasto- 
mataceae and Memecylaceae have “modern” pol- 
len flowers in the sense of Vogel (1978), in which 
the visitor is forced into a fixed position while 


harvesting the reward and in which the position of 
the stigma is of relatively little consequence be- 
cause of the deposition of pollen on a large part 
of the visitor's surface 

There appears to be little opportunity for par- 
titioning the bee body. Sharing of pollinators, noted 
rather frequently among sympatric buzz flowers 
(Linsley € Cazier, 1963; Macior, 1970, 1971; 
Thorp & Estes, 1975; Lumer, 1982; Renner, 
1984), results in hybridization in the absence of 
genetic barriers. This is consistent with the fact 
that systematists working with Melastomataceae 
and Memecylaceae often notice hybrids (Kral & 
Bostick, 1969; Whiffin, 1973; Morley, 1976; Al- 
meda, 1978; Solt & Wurdack, 1980; Renner, in 
press). The data on Melastomataceae breeding sys- 
tems corroborate the suggestion that apomixis may 
be more widespread in the tropics than formerly 
thought (Kaur et al., 1978; Ashton, pers. comm., 
1987). Apomictic seed production would make such 
species independent of pollinators, while retaining 
the advantages of producing seeds, i.e., the poten- 
tial for dispersal and dormancy. Most notably, it 
would give them the capacity to build up a new 
population from a single propagule. 

Statements that “details of the structure of the 
anthers doubtless relate to mechanisms of polli- 
nation" (Cronquist, 1981) or “in a family, such 
as the Melastomataceae, where floral evolution has 
been extensive, one may logically conclude that 
the diverse floral types represent adaptations to 
different pollination systems” (Almeda, 1977) are 
not corroborated by field data. Rather, Melasto- 
mataceae seem to confirm Macior's (1971) sug- 
gestion that buzz pollination is so successful a sys- 
tem that, once it is established, hardly any other 
pollination mechanism evolves, even in species-rich 
groups. Compared with other families, such as 
Lecythidaceae, Gesneriaceae, and Bignoniaceae, 
for which the overall pollination spectrum is fairly 
well known, Melastomataceae and Memecylaceae 
show little diversification in floral morphology and 
pollination strategies. In this they resemble Mal- 
pighiaceae (Anderson, 1979) and Solanaceae (Sy- 
mon, 1979), which also have relatively specialized 
pollination systems and faithful bee pollinators. Such 
ack of diversification could be interpreted as the 
result of being stuck on an adaptive peak. 

Melastomataceae that offer nectar are pollinated 
by a broader range of pollinator classes, namely 
by hummingbirds, bats, rodents, and bees; this 
statement applies to the individual species as well, 
several of which have more than one important 
pollinator type, for instance birds and bats in 
Tibouchina grossa and bees and rodents in Blakea 


chlorantha. Cruden (1972) suggested, as a possible 


514 


Annals of the 
Missouri Botanical Garden 


ecological significance for nectar production at 
higher altitudes, that under adverse climatic con- 
ditions the homoiothermic birds and mammals may 
be more reliable pollinators than bees. If valid, this 
explanation would apply to all now known nectar- 
producing Melastomataceae, except the species of 
Miconia. 

Whether the thousands of pollen-only Melas- 
tomataceae derived from ancestors with nectar 
flowers or whether the proto-Melastomataceae nev- 
er possessed nectaries (as suggested by Vogel, 1978 
is a difficult question. Phylogenetically, the extant 
nectariferous species belong to groups not closely 
related: Meriania and Centronia are in the cap- 
sular-fruited Merianieae (as is Axinaea, which 
probably also has some nectariferous species, see 
above) Brachyotum and Tibouchina are in the 
likewise capsular-fruited Tibouchineae; Chalybea, 
Huilea, and Miconia are in the berry-fruited Mi- 
conieae; and Blakea is in the also berry-fruited 
Blakeeae. Thus nectar production occurs in at least 
five evolutionary lines: the first leading to Meriania 
and Centronia (which are not closely related to 
each other) the second to Brachyotum and 
Tibouchina (the first genus is a segregate of the 
second; Wurdack, 1954), the third to Chalybea 
and Huilea (two intimately related genera), the 
fourth to Miconia (with 1,000 D only two 
of which to date are securely known to produc 
nectar), and the fifth to Blakea. dila could 
have n ida in all these lineages. 

oblem is confounded by the slits observed 


— 


in the hea of some non-nectariferous Mela- 
stomataceae exactly in the same position as in some 
of the nectariferous species, where nectar is se- 
creted through them. They occur in species of 
Heterocentron (Tibouchineae), Bertolonia (Ber- 
tolonieae), and Centradenia (Microlicieae) and in 
the paleotropical genera Medinilla and Dissotis 
(compare above under nectariferous melastomes) 
and may be rudimentary, nonfunctional nectaries. 
Thus, to date, only in two of the neotropical tribes— 
the small Rhexieae with 64 species and the Cy- 
phostyleae with nine species—are nectar produc- 
tion or presumed rudimentary nectaries absent. In 
view of the presence of nectaries in at least five 
neotropical lineages and the indications of their 
former presence in two further ones, their parallel 
independent origin becomes hard to accept. In- 
stead, I suggest that the capacity for developing 
nectaries is basic in the Melastomataceae but sup- 
pressed in most modern members. 

To understand the origin of the memecyloid 
connective gland, whatever the chemical compo- 
sition of its secretion, the following observations 
seem relevant. Functioning nectaries have been 


observed on the hypanthia of Memecylon edule 
Roxb. (= M. ramiflorum Desr.; Burck, 1 )an 
M. floribundum Blume (Zimmermann, 1932). Also, 
I have frequently seen ants on the hypanthia of 
Mouriri nervosa, which may be an indication that 
this neotropical Memecylaceae also secretes e 
albeit in minute quantities. Evolutionarily, the c 
pacity to secrete minute quantities of dece 
directly through the hypanthium epidermis, with- 
out conspicuous morphological structures, may be 
correlated with the development of the staminal 
glands. However, data on the pollinators and the 
chemical composition of the exudates of paleo- 
tropical Memecylaceae are needed before the sig- 
nificance of the staminal glands in the family as a 
whole can be assessed. The Memecylaceae have 
a. 340 Old World and 89 New World species, 
and it should be noted that, in the opinion of Morley 
(1953), the New World Memecylaceae are more 
basal than the Old World ones. 

There is a great amount of parallelism between 
the Old World and the New World tribes in the 
Melastomataceae: the Oxysporeae, Sonerileae, Dis- 
sochaeteae, and Osbeckieae resemble more or less 
closely the Merianieae, Bertolonieae, Miconieae, 
and Tibouchineae, respectively, of the New World 
(Gleason, 1932). Based on wood anatomy, the dif- 
ferentiation into at least some of these tribes and 
into Melastomataceae and Memecylaceae occurred 
before the breakup of Pangaea (Vliet et al., 1981). 
However, the coevolutionary relationships between 
bees and proto-Melastomataceae, which are a con- 


O 


ditio sine qua non for the evolution of the mela- 
stome androecium, can only have begun during 
the mid-Cretaceous when bees arose (Michener, 
1979). The mid-Cretaceous may thus have been 
the time when the ancestors of these families 
switched from nectar-reward flowers to pollen-re- 
ward flowers (with the pollen presented in poricidal 
anthers and available only to bees). Likely, the 
Melastomataceae and Memecylaceae originated 
during Aptian times (lower mid-Cretaceous, 11 
million years before present), when the gap between 
Brazil and Ghana was still very narrow; their east 
and west Gondwanian lineages, probably coevolved 
in parallel with the earliest bees. The oldest fossil 
melastome leaves are known from the early Eocene 
(Hickey, 1977), about 53 million years before pres- 
ent, and Eocene and Miocene leaves occur widely 
scattered throughout the Northern Hemisphere 
(Wurdack & Kral, 1982). Therefore, members of 
the family (and some of their pollinators?) at one 
time may have dispersed via Europe between North 
America and Africa. 

The Melastomataceae, with two-thirds of their 
species in the Neotropics, have much of their di- 


Volume 76, Number 2 
1989 


Renner 515 
Reproductive Biology in Melastomataceae 
& Memecylaceae 


versity centered in areas of varied topography — 
the Andes, Guayana, and south-central Brazil. The 
neotropical Memecylaceae, on the other hand, are 
centered in the Amazonian lowlands, where they 
appear adapted to different soil types (pers. obs.). 
The very early shift from anemochory to zoochory 
in the Melastomataceae was probably one of the 
reasons for their success in the understory of closed 
forests, where animal dispersal is more advanta- 
geous than wind dispersal. (The capsular-fruited 
tribes are much more diverse in open habitats and 
have relatively few species in forests.) The crucial 
points in the radiation of large groups within the 
Melastomataceae, besides their different seed-dis- 
persal strategies, have been the acquisition of par- 
ticular plant architectural, indument, and leaf ve- 
nation characters. Because adaptations to different 
pollinator types played such a limited role in the 
radiation within the two families, the floral biolog- 
ical data cannot be used to support either of the 
two suggested phylogenies for the Melastomataceae 
(Wurdack in Welle & Koek-Noorman, 1981; Vliet 
et al., 1981). 
LITERATURE CITED 
Assy, M. L. & W. E. Kerr. 1977. Algumas plantas 
visitadas para obtengao de n por operárias de 
Melipona Vae ie merrillae em Manaus. Acta 
Amazonica 7: 309-3 
. BEZERRA & W. E. Kerr. 1980. Plantas 
nectariferas utilizadas por duas espécies de Melipona 
da Amazónia. Acta Amazonica 10: 271-281. 
ALMEDA, F. 1977. Systematics of the neotropical genus 
Centradenia (Melastomataceae). J. Arnold Arbor. 


85: 73-108 


Systematics of the genus Monochae- 
tum om in nn and Central Amer- 
ica. Univ. Calif. . Bot. -134. 
.R 197 lord bec in neo- 

tropical Malpighiaceae. Md ay 11: 219-223. 
ARROYO, M. T. K. & E. CaB 1977. Preliminary 
selí: Hur tag test i some tropical cloud forest 
species in Venezuela. Incomp. Newsletter 8: 72-76. 
ATRAMENTOWICZ, M. 19 Influence du milieu sur 
l'activité locomotrice et la reproduction de Caluro- 


Ms philander (L.). Rev. Ecol. (Terre Vie) 36: 373- 


TN H. 1978. Chemical aspects of the pollination 

biology of woody Po in the tropics. Pp. 57-82 
. B. Tomlinson & M. H. Zimmermann (editors), 

Tropical Trees as Living Systems. Cambridge Univ. 
Press, d London, New York, Melbourne. 

AKER. 1983. Floral nectar sugar con- 
stituents in relation to pollinator type. Pp. 117-141 
in C. E. Jones & R. J. Little (editors), Handbook of 
Experimental Pollination Biology. S & AE Scientific 
and Academic Editions, New res 

Bawa, K. S., S. H. BULLOCK, D. R. Perry, R. F. COVILLE 
& . H. Grayum. 1985. Rode biology of 
tropical ~ rain forest trees. II. Pollination 
systems. Amer. J. Bot. 72:346-356. 

BonBio, F. O., P. 5 BOBBIO & E H. DEGASPARI. 


1985. 


up iade from Tibouchina grandiflora. Food 
Chem. 18: 153-160. 

BREMER, ko 1982. Lijndenia, a re-established iid 
tropical am of the orar ^n ceae — Mem 

laceae. Nord. J. Bot. 2: 121- 

198 . Taxonomy of M iis (Melasto- 
aage) i in Borneo. Opera Bot. 69: 1-47. 
BucHMANN, S. L. & M. D. BUCHMANN. 1981. Anthe- 

cology of Mouriri myrtilloides (Melastomataceae: 
Memecyleae), an oil flower in Panama. Biotropica 


: Beitraege zur Kenntnis bn myr- 
mekophilen Pflanzen und der Bedeutung der 
extranuptialen Nektarien. Ann. Jard. Bot. eg 
10: 75-144. 
CAMMERLOHER, H. 
Vlg., Berlin. 

pue A. P. DE. 28. Melastomataceae. Pro 
omus Systematis Naturalis Regni Vegetabilis 3: 99- 

202. 


1931. 


Bluetenbiologie I. Borntraeger 


COGNIAUX, A. . Melastomaceae. /n: A. & C. de 
Candolle, Revue ue Phanerogamarum 7: 1-1256. 
m G. Architecture Végétative et Struc 
> Inforesentel de Quelque Melastomataceae 
. Univ. Strasbourg, Fra 
. 1978. Flora of Barro Colorado Island. 
“Stanford Univ. Press, Stanford, Californ 
j a A. 1981. An Integrated dise: ‘of Classi- 


fication of Flowering Plants. Columbia Univ. Press, 
New ‘Yor 
CRUDEN, R. W. 1972. Pollinators in high-elevation eco- 


systems: ap Kor cM of birds and bees. Sci- 

ence 176: 14 440 

DAHLGREN, R. & R. F. Coe 1984. The order Myr 
tales: circumscription, variation, and futon 
Ann. Missouri Bot. Gard. 71: 633-699. 

Darwin, C. 1876. The Effects of Cross- and Self-Fer- 
sg in the Vegetable Kingdom. John Murray, 


1903. More Letters of Charles 
, New 


x ue 3 (editor). 
, Volume II. D. Appleton & Comp. 


d H. E. M. 1987. Role of flower and pollen 


aromas in host-plant recognition by solitary bees. 
23. 


Ethology of some bees of the 
tribe Euglossini. J. Kansas Entomol. Soc. 39: 607- 


629. 

Don, D. 1823. An 21 of the natural family of 
plants ir Melastomaceae. Mem. Wernerian Nat. 
Hist. Soc. 4: 276- 329. 

DRESSLER, R. E 1982. Biology of the orchid bees (Eu- 
glossini). m Rev. Ecol. Syst. 13: 373-394 
cn A. 190 «s cios de ueber Bluetenbesuch, 

ds etc. der Para vorkommenden 


Bienen. Z. Syst. Hymenop. Dipterol. l: 25-32, 49- 


1902. Beobachtungen ueber i cda d 
Erscheinungszeit etc. der bei Para vorkommende 
Bienen. Allg. Z. Entomol. 7: 321-326, 360- 368, 
400-405, 417-422. 


ETHERIDGE, A. L. & J. M. HERR. 1968. _ The devel- 
ent of th in Rhexia 


mariana. Canad. J. Bot. 46: 133-139. 
Eype, R. H. & J. A. TEERI. Floral antomy of 
Rhexia virginica (Melastomataceae). Rhodora 69: 
63-178 


FaEcnI, K. & L. VAN DER PL. 1979. The Principles 


516 


Annals of the 
Missouri Botanical Garden 


of Pollination Ecology, 3rd revised editi 
Press, O 

FITZPATRICK, J. W J. W. TEnBORGH. 
|« 


A new y species of Meses m from Peru. 
77-186. 


Pergamon 


Wilson Bull. 91: 
FLEMING, T. H. 1986. Opportunism versus specializa- 
tion: n Soo of feeding in frugivorous bats. Pp. 
l in A. Estrada & T. H. Flen Ps (editors), 
Fruor and Seed Dispersal. Junk, e echt. 
1988. The short-tailed fruit i Chi- 
cago Press, Chicago and London. 
Forges, H. O. 1882. Two kinds of stamens with dif- 
ferent functions i in the same flower. Nature 26: 386. 
Foresta, H. DE 
PREVOST. 
de la régénération naturelle après Sp en forêt 
guyanaise. Rev. Ecol. (Terre Vie) 39: 369-40 
FRANKIE, G. W. 1976. Pollination of widely dispersed 
trees by animals in Central Ame n with emphasis 
on bee pollination systems. Pp. 151-159 ín J. Burley 
B. T. Styles (editors), Tropical Trees. Variation, 
Breeding, and Conservation. Academic Press, Lon- 
don. 
GLEASON, H. A. 1929. Studies on the flora of northern 
South America — XII. Cyphostyleae, a new tribe in 
ora a Bull. Torrey Bot. Club 56: 97- 
112 


a. 


Iniv. 


A synopsis of the Melastomataceae of 
British Cui Brittonia 1: 127-184. 
GOULDING, M. 1980. The fishes and the forest. 
California Press, Berkeley, California. 
Gus1 hann. A. 1946-47. a in higher plants. 
cta Univ. Lund. 42/43: 1-370. 
ae E. R. 1979. Cano structure of neo- 
tropical flower ie bees and wasps: diversity and 
phenology. Ecology 6 : 190-202 


Univ. 


Hickey, L. J. 1977. St TOATA and Laser d s 
the Golden V rid Formation (early Te oe 4 we 
ern North Dakota. Geol. Soc. Amer. no. 

Hu ty, » I "Flowering and fruiting pero 
in a premontane rain forest in Pacific Columbia 

ropica 12: 292-300. 

HUBER, a 1909[1910]. Mattas amazónicas 

Bol. Mus. Paraense Hist. Nat. 6: 91-216. 


Janzen, D. H. 1971. Englossion n as long-distance 

pollinators of Base a = ience 171: 203-205. 

JouNsoN, L. A. S B. G. Bric 1984. Myrtales and 

Myrtac eae—a phylogenetic nn. Ann. Missouri 
-756. 


QA. >. Ha JoNc, . SANDS, H. T. CHAN, 
E. oS & le > Aso. 1978. A omixis 


st. ets 271: 440 

.& The genus Rhexia 
Mitad. Sida 3: 387- -440 

KuBITZKI, K. 1978. Caraipa and Mahurea er 

ae). Mem. New York Bot. Gard. 29: 38. 

. Einfuehrung in die ce E 
Fischer Vlg., Stuttgart. 

LAGERHEIM, C. Ueber die Bestaeubungs- u 
Aussa iii. aaa von Brachytoum ledi. 
folium (Desr.) C 1899: 105-122. 

Laroca, S. 1970. ‘Contribuição para o conhecimento 
das relações entre abelhas e flores: coleta de pólen 
das anteras tubulares de certas Melastomataceae. 
Revista Floresta 2: 69-74 

Leccer, W. H. : Fertilization " due virginica 
L. Bull. Torrey Bot. Club 8: 04. 


NEFF, J. 


LINSLEY, E. G. & M. A. CAZIER. 1963. Further obser- 
vations on bees which take pollen from pus of the 

genus Solanum. Pan-Pacific Entomol. 39: 1-18. 
. M. Rick & S. G. STEPHENS. d Ob- 


more on nee floral relationships of the ar tain 


carpente . Pan-Pacific Entomol. 42: 1- 
Lowry, : n. Anthocyanins of the Msn: 
tace 


e, Myrtaceae and some allied families. Phyto- 
ue 15: 513-516 
Lumer, C. 1980. Rodent pollination of Blakea (Me- 
lastomataceae) in a Costa Rican cloud forest. Brit- 
tonia 32: 512-517. 
1982. The 


e pollination ecology, iis A" 
tems and phenology o of Blakea and Topobec 
lastomataceae) in Monteverde, ed Rica. Thesis. 
City University, New York w York. 

& R. D. SCHOER. 1986. Pollination of d 
austin- smith and B. penduliflora (Melastomata- 
ceae) by small rodents in Costa Rica. Biotropica 18: 

363-3064. 

MCMULLEN, C. K. 


, 


1985. Observations on insect visitors 


to flowering plants of Isla Santa Cruz. Part I. The 
endemic C id bee. Not. Galapagos 42: 24-25 
Macior, L. 970. Pollination ecology of Do- 


re ee is stinum (Primulaceae). Bull. Tor- 
rey Bot. Club 97: 150-153 


971. Co-evolution of plants and animals- 
— insights from plant-insect interactions. 
Taxon 20: 17-28. 
MacNUSSON, W. E. € T. M. SanarlorrI. 1987. Dispersal 
of Miconia > by the rat Bolomys lasiurus. J. 
Trop. Ecol. 7-278. 
MICHENER, C. D. aer An interesting method of pollen 
collecting by bees from flowers with tubular anthers. 
Revista Biol. Trop. 10: 167-175. 
1979. Biogeography 4 the bees. Ann. Mis- 
souri Bot. Gard. 66: 277-34 
, M. L. Winston & R. Duos 1978. Pollen 
manipulation and related activities and structures in 
1 m Apidae. Univ. Kansas Sci. Bull. 


51: 515 
Moni, S. A. & T. 1 PiproLY. 1984. Observations on the 
big bang flowering of Miconia minutiflora (Mela- 


stomataceae). Brittonia 36: 337-341. 
MonrEv, T. 1953. The genus Mouriri (Melastomata: 
ceae). Univ. California Publ. Bot. 26: 223-312. 
. Votomitia Aublet (Melastomatconik 
Bull. Er Bot. Club 90: 1-16. 
976. M regis (Melastomataceae). Fl. 
Np 15: 1-295. 
Five new taxa of New World Memecy- 
TNR uu i Ann. Missouri Bot. Gard. 
(255 557 
Moure, J. S. 1970. The species of euglossine bees of 
Central America belonging to the subgenus Eugloss- 
ella (Hymenoptera, Apidae). Anais Acad. Brasil. Ci. 
2: 147-157 


(letters). /n: A. Moeller (editor), 
. Werke, E e und Leben. Volumes 
I-II. Fischer Vlg., Je 
1883. Two kinds x stamens with different 
m— in the same flowe ature 27: 364-365 
MUELLER, H. 81. Two kinds. al stamens with different 
functions in the same flower. Nature 24: 307 
. 1883. Pe pee bei Staubgefaessen von 
Pollenblumen. (Von Hermann Mueller mii n 
tungen von Fritz Mueller. ) aps 13: 241-25 
L. € B. B. Simpson. 1981. 


Oil-collec it 


Volume 76, Number 2 
1989 


Renner 517 
Reproductive Biology in Melastomataceae 
| 


& Memecylaceae 


structures in the oe bong qu 
morphology, function, and use in systematics. J. K 
sas Entomol. Soc. 54: 95-123 

NETTANCOURT, D. DE. 1977. Incompatibility in Angio- 
digg og Berlin. 

OPLER, P. AKER & C. W. FRANKIE. 1980. 
ae crate characteristics during secondary 

ssion in yis ar lowland forest ecosystems. 

epa 12 (suppl.): 40-46. 


PATEL, V. . J. SkvarLa & P. H. Raven. 1984. 
Pollen characters in relation to the delimination of 
Myrtales Missouri Bot. Gard. 71: 858-969. 

PrERcIVAL, M. oe 1965. Floral Biology. Pergamon Press, 


xford. 
"un TOR, M. & P. Yeo. 1972. The Pollination of Flow- 
rs. ee Publ. Comp., New Yor 
RENNER, S. S. 1983. The widespread occurrence of 
anther Pus by Trigona bees in Melastoma- 
taceae. Biotropica 15: 257-267. 

1984. Phaenologie, Bluetenbiologie und Re- 
kombinationssyteme einiger zentralamazonischer 
Melastomataceen. Doctoral Dissertation (printed 
copy), Univ. Hamburg, 19 

. 1986. The neotropical epiphytic Melasto- 
mataceae: an fruges fruit types, and 
floral ME Selbyana 4-11 
à b neis bo of aa 


cia Ma Acta Amazonica 16/17: 
208. 


1987a. 


; Seed dispersal. Progr. Botany 49: 
413-432. 


rs hoehnei (Tibouchineae: 

Melastomataceae): taxonomy, distribution, and biol- 
ogy. Brittonia 39: 441- 

Reproduction and evolution in some genera of 

neotropical Melastomataceae. In: G. Gottsberger & 
G. T. Prance er eu Vip of Tropical Woody 
Angiosperms t. Gard. (in press.) 

—— —— —. Floral biological observation on id is Fs 
tatei (Sarraceniaceae) and other plants from Cerr 
de la Neblina in oo Pl. Syst. Evol. (In press. ) 

RickLEFS, R. E. 1977. A discriminant function analysis 
of assemblages of fruit-eating birds in Central Amer- 
ica. Condor 79: 228-231. 

RoosMALEN, M. G N. 1985. Fruits of the Guianan 
Flora. Inst. Syst. Bot., Utrecht Univ., Netherlands. 

Rousik, D. W. 1979a. Competition studies of colonizing 
| honeybees and native pollinators in Sout 
America. T as, Lawrence, Kansa 


79b. Africanized honey bees, stingless bees, 
in de Secunia of tropical dpa pollinator com- 

nities. Proc. 4th International Symposium on Pol- 
estan. Maryland Agric. Exp. i Misc. Publ. 1: 
403-417. 


& C. D. MICHENER. 1985. Nesting biology of 
Crawfordapis So Colletidae). J. Kansas 
ntomol. Soc. 57: 662-671. 
SAKAGAMI, S. F., poem & W. KERR. 
6 Behavier studies of the stingless m with 
special reference to the oviposition process. J. Fac. 


Sci. Hokkaido Imp. Univ., Ser. 4, Zool. 15: 578- 
607. 


pios H, ^. F. 1980. Arils as food for tropical American 
qoi 82: 31-42. 
a : D. W. 1965. A possible selective factor in the 
evolution of fruiting seasons in tropical forest. Oikos 
15: 274-281. 


1971. diera aspects of fruit-eating by 
Ibis 113: 202. 
19 Tropical frugivorous birds and their 
food plants: a world survey. Biotropica 13: 1-14. 
& B. K. Snow. 1980. Re D between 
hummingbirds and flowers in the Andes of Colombia. 
Bull. Brit. Mus. (Nat. Hist.) Zool. 38: 105-139 
SOBREVILa, C. & M. T A 2. Breeding 
systems in a montane cloud forest in Venezuela. Pl. 
q 


bids: 


J. J. Wurpack. 1980. Chromosome 
brun in the Melastomataceae. Phytologia 47: 
Sourb, C. Ps A. GAUTIER- ped 1986. Fruit selection 
y a forest guenon. J. Anim. Ecol. 55: 235-244. 
SUBRAMANYAM, K. 1942. ode and embry- 
ogeny in a few members of the Melastomataceae. J. 
Indian Bot. Soc. 21: 
19 A akai to the life-history of 
Indian Acad. Sci 


Sonerila wallachii Benn. Proc. 
-120 


19: 


jt An d aes study of Melastoma 
ralabathricum L. J. Indian Bot. Soc. 27: 11-19. 
Bun. D. E. 79. Sex Poeta in Solanum (Solanaceae) 
and the role of pollen a insects. Pp. 385- 
397 in J. G. Hawkes, R. r & A. D. Skelding 
(editors), bie Biology p Taxonomy of the Solanace- 
ae. Academic Press, 
Thorp, R. W. 19 St al behavioral, and phys- 
iological adaptations of bees o z collecting 
po Ann. Missouri Bot. Gard. 66: 788-812. 
Estes. 1975. 
bees on flowers of Cassia ara 


du ecd of 


J. Kansas 


The number of cells 


es). Bot. 


H. Raven. 1984. 
in the pollen of er (Myrtal 
Mag. (Tokyo) 97: 

TRIANA, J. 1867. M aceae. Pp. 725-773 in G. 
Bentham & J. D. Hooker, a paca Reeve 

Co., London. 
Trott, W. 1922. Ueber Staubblatt- und Griffelbewe- 
1 und ihre teleologische Deutung. Flora 15: 
VI. 


Ta die Bluetheneinrichtungen von 

Purpurella agn einer neuen Melastomatcee. 

Ber. Deutsch. Bot. Ges. 13: 415-420 
1896. Weiteres zur blustheaeticichtunp von 

Purpurella ML Pee Er Verwandten. Ber 

Deutsch. Bot. Ges. 14: 169-178. 

irse Bot. n Syst. 42: 234- 


1909. 


1939. Over de meeldraden van enkele 
Melastomataceae. Trop. Natuur 28: 169-172. 
195 Xylocopa and fl in the tropics. 
d. Akad. Wetensch., Ser. C 57: 


35. 
VAN DER Pur, L. 


I-III. Proc. Kon. 
413-423, 541-562. 

VuiEr, G. J. €. M. ben J. ipe vadis Us B. J. H. 
TER WELLE. 198 Wood anatomy, classification 
and phylogeny of ils Ms Blumea 

-473. 


VocEL, S. 195 Fledermausblumen in Suedamerika. 
Oesterr. Bot. Z. 10 530. 
962 [1963]. Duftdruesen im Dienste der 


Bestaeubung. Ueber Bau und Funktion der irr 
ys ps Akad. Wiss. Abh. Math. iaa iss. Kl. 
163. 


1966. Parfuemsammelnde Bienen als Be- 


518 


Annals of the 
Missouri Botanical Garden 


staeuber von 2 und Gloxinia. Oesterr. 
Bot. Z. 113: -361. 
1974. eas und oelsammelnde Bienen. 
Pp. 284-547 in W. Rauh (editor), Tropische und 
subtropische Pflanzenwelt. 7. Steiner, Wiesbaden. 
1978. Evolutionary shifts from reward to de- 
ception in pollen flowers. Pp. 89-96 in A. J. Richards 
(editor), The Pollination of Flowers by Insects. Ac- 
ademic Press, London, New York. 

1988. Neu er kannte bzw. new dokumentierte 
Fledermausblumen aus drei Kontinentem. Tagun ngs- 
berichte Deutsche Bot. Ges.) Giessen 1988: 188. 

WARNER, R. Systematics of Central im 


Monolena (Malsstomatucpae): Thesis, Univ. Min- 
nesota, St. Paul, Minnesota 
WELLE, B. J. H. TER & J. KOEK-NOORMAN. 1981. Wood 


anatomy of the neotropical Melastomataceae. Blu- 
mea 27: LUN 
WHEELWRIGHT, N. T., W. A. Haber, K. G. MURRAY & 
C. GUINDON. on Tropical. fruit -eating birds and 
their food plants: a survey of a Costa M lower 
3-19 


73. Analysis of a hybrid swarm between 
Heterocentron elegans and H. D ae um (Me 
lastomataceae). Taxon 22: 413 

& A. S. TomB. 1972. The M signifi- 


cance of seed morphology in the neotropical capsular- 
fruited Melastomataceae. Amer. J. Bot. 59: 411- 


Wurpack, J. J. A revision of the genus Brach- 
yotum ee Mem. Ne 
York Bot. Gard. 8: 343-407. 

. Md us Pp. -186 in B. 
Maguire, J. J. Wurdack, and Con The Bot- 
any of the Db. Highland — Part V. Mem. New 

York cee Gard. 10 


Coclamibh Melastomataceis. XI. Phy- 

n 11: 377-400. 

———. 1980. Melastomataceae. In G. Harling & B. 

Sparre (editors), Flora of Ecuador. 13: 1-406. Swed- 
ish Natural Science Research Council, Stockholm. 

l Atlas of hairs for neotropical Melas- 
tomataceae. Smithsonian Contr. Bot. 63: 1-80. 

& R. KRAL. 1982. The genera of Melasto- 
mataceae in the Southeastern United States. J. Ar- 
nold Arbor. 63: 429-439. 

1925. Beitraege zur Kenntnis des An- 
und der pig ee einiger Me- 
astomataceen. Bot. Arch. 9: 398-467. 
Ape .G. 1932. Ueber die extrafloralen Nek- 
tarien der Angiospermen. Beih. Bot. Centralbl. 49: 
99-196. 


> 


FLORAL NECTARIES IN 
MELASTOMATACEAE AND 
THEIR SYSTEMATIC 

AND EVOLUTIONARY 
IMPLICATIONS! 


Bruce A. Stein? and Hiroshi Tobe’ 


ABSTRACT 


e vast majority of the 4,500-5,000 species in the primarily tropical m rne Risus im do not produce 


floral nectar, instead relying on pollen as the pollinator reward. To deter 


.0 
genera of Memecylaceae, a family traditionally placed within Melastomataceae . We 
. No struct 
r, most nectar- -secreting species appear to produce the nectar 


to clarify the site of nectar secretion in several genera 


mine = an 
pence 
mical ies os six species in three 
also o co a field observations 

a) 


ura differentiated parenc yma 


from a thickened staminal vascular bundle. Within the order Myrtales, this type of nonstructural androecial nectary 
is limited to Melastomataceae and is apparently very rare among angiosperms as a whole. Two additional methods 
eve 


explanation for the diversity of nectary types within the family and for the scattered phylogenetic positions of nectar 


producing taxa 


The family Melastomataceae, with ca. 200 gen- 
era and 4,500-5,000 species, is the largest in the 
order Myrtales. This primarily tropical family is 
distributed worldwide but particularly well repre- 
sented in tropical and subtropical areas of the New 
World. Most members of the family produce no 
floral nectar, relying instead on pollen as the prin- 
cipal pollinator reward (for review see Renner, this 
volume). Species in at least two genera, Tibouchina 
(= Purpurella) (Ule, 1896) and Brachyotum 
(Lagerheim, 1899), however, have long been known 
to produce nectar. Additional observations of nec- 
tar production, or presumed production, have been 


made more recently on these and several other 
melastome genera, including Blakea (Lumer, 1980; 
Lumer & Schoer, 1986), Huilaea (Snow & Snow, 
1980), and Miconia (Mori & Pipoly, 1984). Most 
reports that note the location of nectar secretion 
within the flower generally found the nectar to 
emanate from the stamens. This strongly contrasts 
with the situation in most myrtalean families in 
which floral nectaries are located either on the floral 
tube, the gynoecium, or at the floral tube-gynoe- 
cium junction. 

oral nectaries of Melastomataceae have been 
little studied, and nothing has been reported pre- 


' Our special thanks to Peter H. Raven for his continued help and sg a gre in this study. For contributing 


fixed flower material for anatomical observations, we express gratitude to Frank Alm 
gs, H. Churchill, Enrique Forero, Michael 
ed E 
V. Pancho, Ching-I ds Barry Prigge, pesas Sal, | Solomon, ia cud Sytsma, and Elsa Zardini. We 


Breedlove, Barbara C. Brig 
Blas F. Hernaez, Peter C. Boch, A. H. M. Jayasuriya 


thank William H. Haber 


ecile Lumer, Mic 


o 
urdack, nie Frank A 


eC St 
3 adis dE Yoshida College, Kyoto uie. jo 


ANN. 


and Susanne R 
A Wurdack for identification at field. collected plants. 
Almeda for comments on earlier drafts of this paper. This study 
85- R M 


eda, Paul E nis 
Solin William A. Haber, Barry Hammel, 
Maxwell, ‘Gordón McPherson, Bruce elson, Juan 


n and National Science 


9 
ngton, Virginia 22209, U. S. A. 
606, Japan 


Missouni Bor. GARD. 76: 519-531. 1989. 


520 


Annals of the 
Missouri Botanical Garden 


viously about their anatomical structure. The pri- 
mary purpose of this study is to elucidate the struc- 
ture of Melastomataceae nectaries through 
anatomical examination and to confirm the unusual 
location of nectar secretion through field obser- 
vations. Our investigation of floral nectary struc- 
ture extends to representative genera of Me- 
a family traditionally included in 
Melastomataceae as a tribe or subfamily. The an- 


mec ylaceae, 


ther connective of Memecylaceae contains a dis- 
tinctive depressed gland structure that had previ- 
ously been described as a nectary (Burck, 1891; 
Subramanyam, 1949). In Mouriri, however, these 
glands have recently been found to secrete a lipid- 
rich substance, suggesting that they are elaiophores 
(Buchmann & Buchmann, 1981 


MATERIALS AND METHODS 


We examined anatomically flowers of 40 species 
in 17 genera of Melastomataceae, representing 
known nectar producers and genera and species 
not reported to secrete nectar. In addition, six 
species of Memecylaceae in three 
investigated. FAA-fixed collections were made by 
the authors and by numerous collaborators world- 
wide (Table 

After dehruration through a t-butyl alcohol se- 
ries and embedding in Paraplast with MP 57- 


58 C?, flowers (and sometimes stamens alone) were 


enera were 


sectioned transversely and longitudinally at 8-10 
um thickness; the sections were stained with he- 
matoxylin, safranin, and fastgreen FCF and were 
mounted with Entellan. Dark staining by hema- 
toxylin represents a concentration of cytoplasm in 
cells, indicating high metabolic activity such as seen 
in active nectaries. In other families of the Myr- 
tales, this staining method has fully demonstrated 
the location and size of nectariferous tissue (Tobe, 
pers. obs.); if present, nectaries are thus easily 
distinguished from other tissue by their dense stain- 
ing. 

We noticed that certain species, particularly 
those known to secrete nectar, have a conspicu- 
ously thickened staminal vascular bundle. Thus, 
for comparative purposes, the thickness of the 
staminal vascular bundle is presented as a relative 
value (Table 1). The “relative thickness" of stamen 
vascular bundles is calculated as: radial thickness 
of the vascular bundle (t)/radial thickness of the 
filament (T). Calculations were made on the basis 
of two or three stamens. Relative thickness values 
are presented as a range, since the thickness of 
the filament and the vascular bundle can differ at 
varying points along the same filament. 


Field observations of nectar secretion in 
Brachyotum, Chalybea, and Tibouchina were 
conducted while field collecting floral material for 
this study. Fresh flowers were examined for evi- 
dence of nectar, and the location of nectar secretion 
within the flower was noted. Nectar quantity and 
sugar concentration were measured using micro- 
capillary tubes and a Reichert temperature-com- 
pensated hand refractometer (model 10431). Nec- 
tar sugar concentration measurements (% brix) 
were subsequently converted to sucrose-equiva- 
lents (weight by total weight). 


RESULTS 


We did not detect densely staining structural 
nectaries in the flowers of any of the species ex- 
amined of Melastomataceae and Memecylaceae. 
Three significant features were noted, however. 
First, stamens of some melastomes, particularly 
those known to secrete nectar, have a markedly 
thickened vascular bundle in relation to the fila- 
ment or anther connective. Relative thickness of 
staminal vascular bundles of nectar-producing and 
non-nectar-producing species are presented in Ta- 
ble 1. The relative thickness values of staminal 
vascular bundles in nectar-producers normally ex- 
ceed 0.3 (mean 0.34), whereas 
stomes examined range between 0.1 and 0.3 (mean 
0.23), a statistically significant difference (P < 
0.001). Second, although no densely staining struc- 


most other mela- 


— 


tural nectaries were found, many of these thickened 
vascular bundles showed relatively dark-staining 
phloem cells, indicating that they are cytoplasm 
rich and may be fulfilling a nectary function. Third, 
many of the nectar-secreting species have external 
slits that develop on or near the geniculum of the 
filament and that may represent a nectar emission 
pathway. 

Information on the site of nectar secretion and 
on known or putative pollination vectors for nectar- 
producing Melastomataceae is summarized in Table 
2. Examined genera that are known or strongly 
suspected to produce nectar are discussed individ- 
ually below. Members of several other genera may 
also secrete nectar but were not examined. These 
include two additional Andean genera that hum- 
mingbirds have recently been observed visiting, 
Centronia excelsa (Bonpl.) DC. (Neill, pers. comm., 
1987) and Meriania tomentosa (Cogn.) Wurdack 
(van der Werff, pers. comm., 1987) 


MELASTOMATACEAE 


Blakea. 


anatomically, with B. chlorantha Almeda known 


Three species of Blakea were examined 


Volume 76, Number 2 Stein & Tobe 521 
1989 Floral Nectaries in Melastomataceae 


TABLE 1. 


Species examined anatomically. Relative thickness of staminal vascular bundles given at right. Tribal 
positions of genera essentially follow Cogniaux (1891) and van Vliet et al. (1981). 


Relative Thickness 
of Staminal Bundle, 


Species 


Collection Voucher 


t 
(see Figs. 5 and 10) 


Melastomataceae 


Tribe Microlicieae 


Rhynchanthera BOUM (J. D. Costa Rica, Grayum et al. 6009 (MO) 0.24-0.26 
Smith) Gleaso 
Tribe Tibouchinea 
Brachyotum penu (Bonpl.) Ecuador, Stein & D'Alessandro 2709 0.32-0.40 
Triana 
B. ledifolium (Desr.) Triana Ecuador, Stein et al. 2701 (MO) 0.39-0.45 
B. ledifolium (Desr.) Triana Ecuador, Stein 2892 (MO) 0.32-0.37 
B. lindenii Cogn Ecuador, Stein 2890 (MO) 0.33-0.34 
B. microdon (Naud.) Triana Bolivia, Solomon 13089 (MO) 0.33-0.36 
B. sanguinolentum (Naud.) Triana Bolivia, Solomon 13859 (MO) 0.32-0.33 
Heterocentron elegans (Schlecht.) Cultivated, Royal Botanic Gardens, Sydney, 0.16-0.23 
Kuntze Griggs 7145 (NSW) 
Tibouchina bicolor (Naud.) Cogn. Bolivia, Solomon 13862 (MO) 0.30-0.36 
T. clavata (Pers.) Wurdack Cultivated, Mathias Botanical Garden, 0.29-0.31 
Prigge 6525 (UCLA) 
T. grossa (L.f.) Cogn. Colombia, Stein & McDade 3147 (MO) 0.38-0.41 
T. heteromalla Cogn. Cultivated, Mathias Botanical Garden, 0.19-0.24 
Prigge 6445 (UCLA) 
T. laxa Cogn. Cultivated, Mathias Botanical Garden, 0.32-0.33 
Prigge 6211 (UCLA) 
T. multiflora Cogn. Cultivated, Mathias Botanical Garden, 0.24-0.28 
Prigge 6524 (UCLA) 
T. semidecandra (DC.) Cogn. Cultivated, Missouri Botanical Graden, Tobe 0.34-0.37 
s.n. (no voucher 
T. stenocarpa (DC.) Cogn. var. boli- Bolivia, Solomon 13231 (MO) 0.32-0.35 
viensis Cogn 
T. urvilleana (DC. ) Cogn. Cultivated, Mathias Botanical Garden, 0.32-0.34 
Prigge 6444 (UCLA) 
Tribe Rhexiea 
Pr D floribundum (Schlecht.) Mexico, Breedlove & Almeda 567 16 0.27-0.33 
Naud. (CAS) 
Tribe Sonerileae 
Sonerila Pius Korth. Thailand, Maxwell s.n. (PDA) 0.23-0.27 
Sonerila Sri Lanka, Jayasuriya 3009 (PDA) 0.22-0.25 
Sonsrila a sp Sri Lanka, Jayasuriya 3030 (PDA) 0.26-0.27 
Sonerila sp Sri Lanka, Jayasuriya 3094 (PDA) 0.24-0.25 
Tribe Bertolonieae 
Bertolonia maculata DC. Cultivated, Missouri Botanical Garden, Zar- 0.23-0.24 
ini s.n. (no voucher) 
Monolena multiflora Warner ined. Pune Churchill 4155 (MO) 0.17-0.23 
Triolena hirsuta (Benth.) Triana Costa Rica, Schatz 999 (WIS) 0.17-0.20 
Tribe Dissochaeteae 
Medinilla fuchsioides Gardn. Sri Lanka, Jayasuriya 3351 (PDA) 0.23-0.24 
M. myriantha Merr. Cultivated, Mathias Botanical Garden #78- 0.10-0.13 
069 (no voucher) 
Tribe Miconieae 
Bellucia pentamera deos Costa Rica, Hammel et al. 14149 (MO) 0.08-0.10 
Chalybea corymbifera d. Colombia, Stein et al. 3610 (MO) 0.26-0.33 
Miconia dodecandra DE ) Cogn. 0.14-0.22 


Mexico, Breedlove & Almeda 57466 
(CAS) 


522 Annals of the 
Missouri Botanical Garden 
TABLE 1. Continued. 
Relative Thickness 
of Staminal Bundle, 
t 
Species Collection Voucher (see Figs. 5 and 10) 
M. melanotricha a Gleason anama, McPherson 8060 (MO) 0.13-0.22 
M. minutiflora (Bonpl.) D Venezuela, Berry 4417 (VEN) 0.26-0.29 
M. reducens Triana Panama, Sytsma 4032 (MO) 0.18-0.19 
Tribe Blakee 
Blakea hae Almeda Costa Rica, Haber 1197 (MO) 0.21-0.27 
B. foliacea Gleason Panama, Churchill 5463 (MO) 0.21-0.25 
B. sp. nov. Costa Rica, Grayum et al. 3553 (MO) 0.18-0.23 
ed ned Cogn. Panama, Churchill 5477 (MO) 0.11-0.13 
Tribe Astr 
yiii candolleana Cogn. Philippines, Hernaez 3829 (CAHP) 0.19-0.23 
A. ferruginea Elmer Taiwan, Peng 5247, 5503 (HAST) 0.21-0.29 
A. meyeri Merr. Philippines, "n 3632 (CAHP) 0.17-0.20 
Memecylaceae 
Tribe Memecyleae 
Memecylon cantleyi Ridley Cultivated, Singapore Botanical Garden, 0.23-0.25 
Maxwell s.n. (no voucher 
M. caeruleum Jack. Cultivated, Singapore Botanical Garden, 0.15-0.23 
Maxwell s.n. (no voucher 
Mouriri oo (Sw.) Poir. Panama, Hamilton 2883 (MO) 0.20-0.25 
rvosa Pilge Brazil, Renner 249 (US) 0.16-0.18 
Tribe Pternandreae 
Pternandra caerulescens Jack. Cultivated, Singapore Botanical Garden, 0.23-0.24 
Maxwell s.n. (no voucher 
P. echinata Jack. 0.17-0.24 


Cultivated, Singapore Botanical Garden, 
her) 


Maxwell s.n. (no voucher 


to produce nectar. Two additional nectar-producing 
species have recently been reported, B. austin- 
smithii Standl. and B. penduliflora Almeda (Lu- 
mer & Schoer, 1986). There is some disagreement 
concerning the site from which nectar is secreted 
in this genus. Lumer (1980, and pers. comm., 
1985) stated that in B. chlorantha nectar appears 
to be secreted in the area at the base of the stamens, 
either at the junction of the filament and floral tube 
or at the base of the filaments. According to Haber 
(pers. comm., 1985), however, nectar is secreted 
on the abaxial surface of the filaments (the side 
facing the corolla wall), with the nectar forming 
thick droplets between the filament and the corolla. 

Anatomical sections of Blakea chlorantha flow- 
ers do not show a densely staining nectary any- 
where on the floral tube wall (Fig. 1), nor do sta- 
mens have an externally or internally differentiated 
nectariferous structure (Figs. 2, 3). Stamens are 
folded inward in bud, with a geniculum present just 
above the midpoint. Slitlike structures are present 
on the adaxial surface of the geniculum (arrows in 
Figs. 1-3). The relative thickness value of the 
vascular bundles is fairly low (0.21-0.27) com- 


pared with other nectar producers (Fig. 5). These 
vascular bundles, however, are relatively densely 
staining, particularly in the upper part of the fil- 
ament and at the connective (Figs. 3-5). A major 
part of the staminal vascular bundle appears to 
consist of small, 


cular bundle are not specialized (Figs. 4, 5), sug- 
gesting that they are not nectariferous. 

Stamens of Blakea sp. nov. (fide F. Almeda) 
and B. foliacea are folded inward in bud at the 
anther—filament junction, and filament slits are not 
present. Although, like B/akea chlorantha, they 
have a thin more or less densely staining vascular 
bundle, they differ in that the staminal vascular 
bundle is surrounded by a thick-walled tanniferous 
epidermis and by some underlying tanniferous cell 
layers. Blakea sp. nov. and B. foliacea stamens 
are unlikely to secrete nectar through such tan- 
niferous tissue, and nectar production has never 
been reported in either of these two species. 


Brachyotum. All Brachyotum species investi- 
gated in the field were found to produce nectar, 


Volume 76, Number 2 
1989 


Stein 


& Tobe 523 
Floral Nectaries in Melastomataceae 


TABLE 2. Summary of data on nectar secretion and pollinators in nectar-producing Melastomataceae. 
Species Nectar Secretion Site Secretion Pollinators References 
Blakea austin-smithii Base of filaments Nocturnal Rodents Lumer & Schoer, 
tandl. 1986 
B. chlorantha Almeda Abaxial surface or base of Nocturnal Rodents Lumer, 1980 
aments 
B. penduliflora Almeda Base of filaments Nocturnal Rodents Lumer & Schoer, 
1986 
m ledifolium Adaxial surface of filaments Diurnal Hummingbirds Lagerheim, 1899; 
sr.) Triana and oth- Stein, pers. obs. 
- Br rachyotum species 
C halybea corymbifera Base of stamens? (see text) Diurnal Hummingbirds? Stein, pers. obs. 
Naud 
Huilaea macrocarpa Not known Diurnal Hummingbirds Snow & Snow, 1980 
Uribe 
Medinilla magnifica Tips of petals Diurnal Not known Tobe et al., in prep. 
Lindl. 
Miconia minutiflora Not known (stigma?) Morning Bees Mori & Pipoly, 1984 
(Bonpl.) DC 
Tibouchina cleistoflora Adaxial surface of filaments Not known Bees Ule, 1896; Renner, 
e this volume 
T. grossa (L.f.) Cogn. Adaxial surface of filaments Nocturnal/ Bats and hum- Vogel, 1957; Stein, 
Diurnal mingbirds pers. obs. 
T. hospita (DC.) Cogn. Adaxial surface of filaments Not known Bees Ule, 1896; Renner, 
this volume 
T. itatiaiae (Wawra) Adaxial surface of filaments Not known Not known Ule, 1896 


ogn. 


confirming previous reports (Lagerheim, 1899; Vo- 
gel, 1957; Wurdack, 1965), and it is likely that 
this entire Andean genus is nectariferous. From 
field observations, the nectar appears to be secreted 
by the filament at the slitlike structure located on 
or just below the geniculum on the adaxial filament 
face (Fig. 6) and forms large droplets suspended 
between adjacent pairs of filaments. Nectar con- 
centration ranged from a high of 20-22% sucrose- 
equivalents in B. ledifolium (Desr.) Triana to a 
low of 15-16.5% in B. campanulare (Bonpl.) 
Triana, values consistent with most hummingbird- 
pollinated flowers (Baker, 1975). 

natomical sections of stamens do not show any 
differentiated nectariferous structure in the fila- 
ments. Stamens of Brachyotum ledifolium have a 
thick vascular bundle running the length of the 
filament (Fig. 7) to the connective (Fig. 8). The 
staminal vascular bundle is densely staining and 
contrasts with the surrounding nonspecialized 
parenchyma cells. All other examined species of 
Brachyotum contain a similar thick, densely stain- 
ing staminal vascular bundle: B. campanulare (Fig. 
10), B. lindenii Cogn., B. microdon (Naud.) Triana, 
and B. sanguinolentum (Naud.) Triana (Fig. 9). 
This similarity in staminal anatomy suggests that 
all these species exude nectar from the filament. 


Chalybea. | Chalybea is a monotypic genus of 
the Colombian Andes closely related to, if not con- 
generic with, the nectar-producing genus Huilaea. 
Poorly known and consisting only of C. corymbif- 
era Naud. (= Pachyanthus corymbiferus [ Naud. | 
Cogn.), Chalybea recently has been resurrected 
on the basis of material newly collected for this 
study (Wurdack, 1988). We have observed nectar 
secretion from C. corymbifera and measured the 
nectar sugar concentration at 16.5% sucrose- 
equivalents. Pinpointing the site of secretion is 
problematic. Nectar was collected from the summit 
of the inferior ovary; however, the nectar was 
observed and measured several hours after the 
normally pendulous flowers were collected and could 
have migrated to this location. Stamens are bent 
inward in bud and have neither a pronounced ge- 
niculum nor filament slits (Figs. 11, 12). Anatom- 
ical sections do not show a differentiated nectarifer- 
ous structure at the ovary summit (Fig. 11). While 
stamens do not have either externally differentiated 
structures or densely staining nectariferous tissue 
(Figs. 11, 12), they do contain a thick vascular 
bundle running through the filament (Fig. 14) to 
the connective (Fig. 13), suggesting staminal se- 
cretion. Tanniferous parenchyma cells surrounding 
the staminal vascular bundle appear unfavorable 


524 


Annals of the 
Missouri Botanical Garden 


FIGURES 1-5. Blakea chlorantha. - 
ansverse sections (TS) of he and of filame 
e d at the inner side of the bend. Abbreviation: sta, 


br nd va n (LS) of flower. 


—2. Stam S of stamen. 


3. 
3 indicate a eee ion or break 


. Arrows 1 


stamen. t in Figure 5 are thickness of filament and 


of vascular bundle respectively, which are measured for ii ulation of teh thickness of staminal vascular bundles 
5. 


(see text for explanation). Scale bars — 1 mm in Figures 


for secretion of nectar. These tanniferous cells, 
however, are much less conspicuous in the lower 
part of the filament, and nectar thus may exude 
from the filament in this location, perhaps ac- 
counting for its observed presence on the ovary 


summit. 


Huilaea. Snow & Snow (1980) reported nectar 
production by Huilaea macrocarpa Uribe and 
measured the nectar sugar concentration to range 
from 12 to 16% with a mean of 13.4%. No in- 
formation was provided as to the precise location 
of nectar secretion in these pendulous flowers. 

Tibouchina. 
examined, including the Andean T. grossa, which 
is known to produce nectar. Brazilian Tibouchina 


Nine species of Tibouchina were 


species reported to secrete nectar include 7. cleis- 


toflora Ule, T. hospita (DC.) Cogn., and T. ita- 


3; 0.2 mm in Figures 4, 


tiaiae (Wawra) Cogn. (Ule, 1896). Our observa- 
tions of nectar secretion in T. grossa differ somewhat 
from those of Vogel (1957), who considered the 
nectar to exude from the inner wall of the floral 
tube below the insertion of the stamens. 7ibouchina 
grossa flowers are open-campanulate and oriented 
horizontally with all stamens arranged in a half- 
circle in the lower side of the corolla. We observed 
nectar droplets forming on the adaxial surfaces of 
the filaments near the geniculum. The drops are 
at first suspended between adjacent filaments and 
occasionally between the filaments and style. The 
rather copious nectar then flows down the filaments 
to collect either in the floral tube or on the convex 
lower petals. We measured sugar concentration of 
T. grossa nectar at 14.5% sucrose-equivalents. 
No densely staining differentiated structure was 
found in the filaments of Tibouchina grossa (Fig. 


Volume 76, Number 2 Stein & Tobe 525 
1989 Floral Nectaries in Melastomataceae 


FIGURES 6-10. Brachyotum. —6. Stamen of Brachyotum ledifolium. —7, 8. T ections (TS) of filament 
and of anther of Brachyotum ledifolium. —9. TS of anther of "d hare Mer ea cal —10. TS of anther 
(above) and of filament (below) (the adaxial sides of which are opposite) of achyotum campanulare. Note thick 


gia hr bundles in all species. oan (T and t) in Figure 10 plan iius to those in Figure 5. Scale 
bar in Figure 6; 0.2 mm in Figures 7-10. 


19) 
eu KFE 
Mae os > 
o yd 


FIGURES 11-14. Chalybea con mbifera. — 11. Longitudinal section of flower. — 1 2. Stamen. — 13, 14. Transverse 
sections of anther and of filament. Abbreviations: sta, stamen; vs, vascular bundle. Scale bars — 1 mm in Figures 
11, 12; 0.2 mm in Figures D p 


526 


Annals of the 
Missouri Botanical Garden 


_FIGURES 15-21. 
. Transverse sections (TS) of anther and of filament 
Longitudinal section (LS) of flower bud of Miconia minutiflora. — 19. 
—20. Upper half of ast of flower bud of Mic 
i sta, stamen; sty, style. Scale 
05 mm in n 19. 


18, nad 21; 


5). Instead, stamens of T. grossa have a thick, 
densely staining vascular bundle surrounded by 
nonspecialized parenchyma cells (Figs. 16, 17). 
Relatively thick staminal vascular bundles are ob- 
served in all Tibouchina species investigated (see 
Table 1), although nectar production has so far 
not been reported in those other species. 


Medinilla. The Old World genus Medinilla in- 
cludes certain species that morphologically appear 
to be good candidates for bird pollination. We ex- 
amined anatomically one such species, Medinilla 
fuchsioides Gardn., but found the staminal vas- 
culature to be unexceptional. Subsequent to this, 


Tibouchina and Miconia. —15. Stamen of Tibouchina grossa, which is bent at a hinge. — 16, 
of Tibouchina grossa, showing thick vascular bundle. T 


TS of young anther of Miconia minutiflora. 


onia melanotricha. — 21. TS of filament of Miconia melanotricha. 
bars = 1 mm in Figures 15, 20; 0.2 mm in Figures 16, 17 


however, we found cultivated material of M. mag- 
nifica Lindl. at the Berlin Botanical Garden to 
secrete nectar from the petal tips. Anatomical in- 
vestigations of this very unique nectar secretion 
method are currently under way (Tobe et al., in 
prep). 
Miconia. |n discussing a mass flowering episode 
of Miconia minutiflora (Bonpl.) DC, Mori & Pipoly 
(1984) reported the presence of nectar as a pol- 
linator reward in this species. In other localities, 
however, this lowland species has not been found 
to secrete nectar (Renner, 1983). 

Only young buds of Miconia minutiflora were 


Volume 76, Number 2 
1989 


Stein & Tobe 527 


Floral Nectaries in Melastomataceae 


available for the present study. Anatomical sections 
of this material indicate no structure likely to pro- 
duce nectar in either the stamens or around the 
floral tube-gynoecium junction (Fig. 18). Flowers 
are very small, with the filaments supplied by a 
relatively thin vascular bundle (Fig. 19). Anatom- 
ical sections show, however, that M. minutiflora 
has a remarkably conspicuous stigma supplied by 
ample, densely staining vascular tissue (Fig. 18). 

Hummingbirds have been observed visiting Mi- 
conia melanotricha (Triana) Gleason (Almeda, pers. 
comm., 1986), and this species may produce nec- 
tar. The flowers appear morphologically well suited 
for hummingbird pollination, given their large size 
(10-15 mm in diameter an -25 mm long) and 
red petals. This species also has longitudinally de- 
hiscent anthers rather than the apically poricidal 
anthers typical of bee-pollinated melastomes. An- 
atomical sections do not show densely staining 
structures or tissue, and the stamens have a thin 
vascular bundle (Fig. 21). As in M. minutiflora, 
however, the stigma and supporting vascular tissue 
are unusually densely staining (Fig. 

iconia robinsoniana Cogn. was reported to 

produce nectar in a survey of the foraging ecology 
of the Galápagos Islands carpenter bee, Xylocopa 
darwinii (Linsley et al., 1966). Subsequent obser- 
vations of this species have not reconfirmed the 
presence of nectar (McMullen, pers. comm. in Ren- 
ner, this volume). During our own field investiga- 
tions of this species, no suitable material could 
located to resolve this question. An additional, as 
yet unidentified, Miconia species from lowland 
Amazonia has recently been reported to produce 
nectar (Roubik, pers. comm. in Renner, this vol- 
ume). 


MEMECYLACEAE 


Memecylon and Mouriri have a peculiar de- 
pressed gland on the backside of the anther con- 
nective (Figs. 22, ). This structure was early 

escribed as a nectary (Burck, 1891; Subraman- 
yam, 1949; Venkatesh, 1955) but recently has 
been hypothesized to be an elaiophore, or oil-pro- 
ducing gland (Buchmann & Buchmann, 1981). 
Anatomical sections reveal this gland to be com- 
posed of more or less radially enlarged epidermal 
cells, which are clearly distinguishable from other 
cells of the connective (Figs. 23, 25). 

Pternandra is variously included in or excluded 
from the Memecylaceae. Its uncertain status is 
based largely on the absence of the anther con- 
nective glands found in other members of Me- 
mecylaceae. The two species examined in this study, 
P. caerulescens Jack. and P. echinata Jack., lack 


these gland structures on the surface (Figs. 26, 
2'7) and in section (Fig. 28) 


DISCUSSION 


POSITION AND STRUCTURE OF FLORAL 
NECTARIES IN MELASTOMATACEAE 


The present study shows that, in contrast to 
nearly all other myrtalean families, Melastomata- 
ceae lack histologically differentiated nectariferous 
tissue either on the inner surface of the floral tube, 
on the gynoecium, or at the floral tube-gynoecium 
junction. While nectar production in Melastoma- 
taceae is rare, a diversity of unusual nectary types 
are implicated, including secretion from the sta- 
mens, petals, and perhaps stigmas. Most docu- 
mented nectar-producing melastomes secrete nec- 
tar from the stamens, although we did not find any 
differentiated nectariferous structures or tissue in 
that organ. On the basis of observations and an- 
atomical structure, we thus conclude that in most 
nectariferous Melastomataceae nectar is produced 
by a nonstructural nectary consisting solely of the 
staminal vascular bundles. 

Compared with melastome species generally, 
stamens of nectar-producers have a much thicker 
vascular bundle containing many small, densely 
staining cytoplasm-rich phloem cells. An ultra- 
structural survey of these cells could offer addi- 
tional evidence of their nectar-secreting capability, 
since there is a clear connection between cytolog- 
ical constituents and processes of nectar secretion 
(for review of ultrastructural features see Fahn, 
1979). 

The nonlocalized nature of these androecial nec- 
taries may help resolve the g reports about 
the specific sites of nectar secretion (Table 2). If 
nectar is produced by the staminal vascular bundle, 
it could emanate from almost anywhere along the 
stamen. Additionally, if the direction of the nectar 
stream within a stamen is affected by gravity, the 

d 


secretion site could differ between stamens an 
among individual flowers. 

Several observers, including one of us (B.A.S.), 
have noticed that in certain genera (e.g., Brachy- 
otum and Tibouchina) the location of nectar se- 
cretion appears to be closely associated with a small 
slit in the adaxial surface of the filament (see arrows 
in Figs. 1-3, 6). The development and significance 
of these slits is somewhat controversial. We found 
no anatomical differentiation of this structure. Since 
stamens in Melastomataceae are folded or bent 
inward in bud, becoming more or less straight at 
anthesis, the observed slitlike structures may form 
spontaneously and be unrelated to nectar secretion. 


528 


Annals of the 
Missouri Botanical Garden 


IGURES 22-28. Me 
anthers of Meme ylon suns foes 


r 


lourir 


and P. « 


al 
Purnandns echinata 


section of stamen of nal caerulescens. Abbreviation: el, anther elaiophore. Scale bars 
28; 0.2 mm in Figures 22, 23; 0.05 mm in Figure 25. 


24, 26, 27, and: 


Such slits are, in fact, reported in some species not 
known to secrete nectar (Renner, this volume). 
However, an apparent association of these slits with 
nectar-producing species, their absence in related 
non-nectariferous species (e.g., Blakea chlorantha 
vs. B. sp. nov. and B. foliacea), and field obser- 
vations of nectar droplets formed precisely at these 
indentations strongly argue for a role in nectar 
secretion ese structures may form through a 
deterlonition of a small amount of tissue on the 
inner side of the geniculum, providing an exit path- 
way for nectar produced by the staminal vascular 
bundle. Clearly, a systematic survey of the occur- 
rence of these slits throughout the Melastomata- 
ceae would be useful in clarifying their role. 
Although nectar production has been reported 
in Miconia minutiflora (Mori & Pipoly, 1984; 
Mori, 1986), Renner (this volume) 


pers. comm., 


ds de d a Pternandra. 
d M 


nervosa, show 
sections of anther glands S Memec ylon caeruleum and Mouriri nervosa. 
aerulescens, showing absence of glands on backside Ben connective. — 28. Longitudinal 


— 22, 24. Scanning electron micrographs (SEM) of 
ing anther glands on backs 5 pi Lo nective. — 23, 25. 
SEMs of anthers of 


= 0.5 mm in Figures 


has questioned whether the fluid observed was truly 
Mico- 


nia minutiflora does not appear capable of se- 


nectar. From our anatomical examination, 


creting nectar from the stamens in the manner of 
the other confirmed nectar-producers we investi- 
gated. If nectar production in this species is re- 
confirmed, the conspicuous stigma, which is sub- 
tended by cytoplasm-rich vascular tissue, should 
be closely examined for a possible secretory role. 
Stigmatic nectar secretion is rare within the an- 
giosperms but occurs in Asclepiadaceae and Ara- 


ceae (Fahn, 1979). 


EVOLUTION OF FLORAL 
MELASTOMATACEAE 


NECTARIES IN 


Nectaries generally consist of an epidermis sub- 
tended by specialized parenchyma cells (Esau, 1977; 


Volume 76, Number 2 
1989 


Stein & Tobe 
Floral Nectaries in Melastomataceae 


529 


Fahn, 1979). The phloem-based nectaries we have 
found in Melastomataceae, which have no struc- 
turally differentiated parenchyma, are a rather ex- 
treme deviation from this typical structure. The 
nonstructural androecial nectaries found in Melas- 
tomataceae are particularly anomalous in Myrtales, 
where, of the 14 families currently included in the 
order (Dahlgren & Thorne, 1984), only Melasto- 
mataceae, Memecylaceae, and Rhynchocalyca- 
ceae lack densely staining structural nectaries. Eyde 
(1981) has discussed evolutionary shifts in the lo- 
cation of floral nectaries in flowers of various myr- 
talean families and concluded that the ancestral 
site of nectar secretion for Myrtales, as for many 
other dicots, is the junction between the gynoecium 
and the floral tube or, in tubeless flowers, the junc- 
tion with the androecium. From this ancestral po- 
sition, nectaries evolutionarily have migrated onto 
the floral tube, or even onto the androecium in 
some tubeless species of Lopezia and Epilobium 
of the Onagraceae (Eyde & Morgan, 1973; Eyde, 
1981). Androecial nectaries are also found in other 
groups, including virtually all members of the Cary- 
ophyllales (Zandonella, 1977 
droecial nectaries in these other families are more 
typical differentiated structures and thus not com- 
parable to the nonstructural androecial nectaries 
of Melastomataceae, which are certainly unique 
within the Myrtales and perhaps among angio- 
sperms as a whole. 

Given the ancestral myrtalean nectary type and 
position proposed by Eyde (1981), it seems clear 


that structural nectaries were lost in the evolu- 


. However, the an- 


tionary lineage ancestral to Melastomataceae and 
Memecylaceae. Such nectaries appear to have been 
lost independently in the lineage leading to Rhyn- 
chocalycaceae (a monotypic family consisting only 
of the South African Rhynchocalyx lawsonioides 
Oliver), since on many other grounds this family 
is more closely related to other members of the 
Myrtales than to Melastomataceae (Tobe & Raven, 
1984). 

Pollinator interactions appear to have played a 
key role in both the ancestral loss and subsequent 
re-evolution of nectaries within the Melastomata- 
ceae. Melastomataceae are, on the whole, a famil 
uniquely suited for bee pollination. The distinctive 
poricidally dehiscent anthers are directly related 
to vibratile extraction of pollen by bees, a phe- 
nomenon known as buzz pollination. Bees foraging 
for pollen in this manner rarely forage simulta- 
neously for nectar, and then only to mix it with 
the pollen to improve the pollen's handling prop- 
erties (Buchmann, 1983). Although poricidal an- 


thers are found in numerous angiosperms (for re- 
view see Buchmann, 1983), few plant families are 
so completely characterized by this feature as Me- 
lastomataceae. Given the widespread distribution 
of poricidal anthers within all lineages of Melas- 
tomataceae, this feature appears to be the ancestral 
condition for the family. Thus, if buzz pollination 
became established in the lineage ancestral to Me- 
lastomataceae, there would be little or no selective 
advantage in maintaining energy-expensive nec- 
taries. By attracting undesirable and potentially 
destructive floral visitors, nectaries may even have 
been selected against in protomelastomes. 
Nectaries appear to have re-evolved indepen- 
dently in several distinct lineages within the family, 
most of which share certain ecological and repro- 
ductive traits. Cruden (1972) has pointed out the 
potential reproductive advantage ensuing from a 
shift from bee to bird pollination in wet-tropical 
montane ecosystems. Similarly, Heinrich & Raven 
1972) have noted the increased energy reward 
necessary to attract pollinators operating at higher 
elevations and to support vertebrate rather than 
invertebrate pollinators. Nectar, composed prin- 
cipally of carbohydrates, is a much richer energy 
source than pollen and is almost invariably the 
floral reward used to attract vertebrate pollinators. 
Not surprisingly, most confirmed nectar-producing 
Melastomataceae (species of Blakea, Brachy- 
otum, Chalybea, Huilaea, and Tibouchina) occur 
at high or relatively high elevations, and are known 
or presumed to be pollinated by vertebrates (Table 


— 


According to traditional and recent classification 
systems for the family (Cogniaux, 1891; van Vliet 
et al., 1981; Wurdack, 1980), confirmed nectar- 
producing Melastomataceae are found in at least 
four different tribes: Blakeae, Dissochaeteae, Mi- 
conieae, and Tibouchineae (see Table 1). The phy- 


monophyletic lineage within the family. In addition, 
the majority of genera in these four tribes are not 
known to produce nectar. Except for the lowland 
species of Miconia discussed above, most of the 
nectar-producing taxa can be viewed as —— 
derived within their respective tribes in s of 
pollination systems and ecological/ aia! pubs 
erences. With regard to the latter, the Central and 
northwestern South American montane terrains that 
are inhabited by many of these species (and, if the 
above interpretation is correct, may have provided 
the ecological impetus for the development of nec- 
tar rewards) are generally more recent (Zeil, 1979) 


530 


Annals of the 
Missouri Botanical Garden 


than the origin of the Melastomataceae, fossils of 
which are known from as early as Eocene times 
(Hickey, 1977). 

wo hypotheses can be advanced to account for 
the unusual systematic distribution of nectaries 
within the family. The first is that such nectaries 
are the basal condition in Melastomataceae and 
have been lost in the vast majority of species. Civen 
the structural nectaries present in other families 
of Myrtales and the rather specialized morpholog- 
ical state of most of the nectariferous species, how- 
ever, such an interpretation is untenable. Renner 
(this volume) advocates a modified version of this 
hypothesis, suggesting that “the capacity for de- 
veloping nectaries is basic in the Melastomataceae 
but suppressed in most modern members.” This 
supposition seems equally difficult to accept. Rath- 
er, we propose a second hypothesis, which is that 
nectaries have evolved independently a number of 
times within the family and developmentally have 
followed the easiest course available. T 
evidence point to this. First, three different nectary 
types have developed within the family, each in- 


e. Two lines of 


volving different floral organs (stamens, pistil, pet- 
als). Although rare within angiosperms, the most 
common melastome nectaries, nonstructural an- 
droecial nectaries, involve very little modification 
of existing structures and thus appear to be an 
ipd developed solution to producing nectar. It is 

cult, however, to reconcile this view of the ease 
of developing such nectaries with the relative rarity 
of nonstructural androecial nectaries in the angio- 
sperms as a whole 


SYSTEMATIC IMPLICATIONS OF NECTARY 
ANATOMY FOR MEMECYLACEAE 


Memecylaceae traditionally have been included 
in Melastomataceae as a tribe or subfamily (e.g., 
Cogniaux, 1891; Morley, 1976). Based on floral 
nectary characters, Memecylaceae agree with Me- 
lastomataceae in completely lacking densely stain- 
ing structural nectaries. The term “disc” has some- 
times been used to describe certain structures within 
the flowers of Memecylon (see Bremer, 1983). 
Given the complete lack of nectariferous tissue, 
this term should, however, be avoided. As previ- 
ously discussed, loss of structural nectaries seems 
to be a feature ancestral for Melastomataceae, and 
this is probably also the case for Memecylaceae. 
We regard loss of structural nectaries as a syn- 
apomorphy uniting these two families. 

Memecylaceae have generally been circum- 


scribed to include the Old World Memecylon and 


the New World Mouriri and Votomita, all con- 
taining anther connective glands. Based on obser- 
vations of Mouriri myrtilloides (Sw.) Poir, Buch- 
mann & Buchmann (1981) suggested that the 
connective glands of Memecylaceae are elaio- 
phores that secrete lipids for nutritional use b 
pollinating bees, primarily in larval o 
Although the secretion is lipid rich, its chemic 
complexity and the presence of components seem- 
ingly toxic to insects have led others to be cautious 
about this interpretation (Renner, this volume; 
Simpson & l 

Morley (1976) considered these distinctive but 
little-understood connective glands to be one of the 
most characteristic features of the "Memecyleae' 
(7 Memecylaceae), even though they are occa- 
sionally lost, as in Mouriri exadenia Morley. 
Whatever its taxonomic rank, the group is now 
generally considered to include additional genera, 
among them Pternandra, which lacks connective 
glands (Dahlgren & Thorne, 1984). The present 
study reconfirms this observation: all examined 
species of Memecylon and Mouriri have anther 
glands on the backside of the connective, whereas 
all examined species of Pternandra lack them. 
Therefore, even if Pternandra is aligned with Me- 
mecylon, Mouriri, and Votomita at the familial 
level, evolution of the anther glands appears re- 
stricted to the lineage including the three last- 
mentioned genera. Available evidence from stam- 
inal anatomy does not resolve the correct familial 
placement of Pternandra but does suggest that if 
included in Memecylaceae, it should be regarded 
as the sister group to the rest of the family. This 
is generally the approach taken by van Vliet et al. 
(1981) in placing Pternandra in its own tribe Pter- 
nandreae of their “Memecyloideae” (= Memecy- 
laceae). 


LITERATURE CITED 


Baker, H. G. 1975. Sugar concentrations in nectars 
from hummingbird flowers. Biotropica 7: 37-41. 
E K. Taxonomy of Memecylon aad 


mataceae) in Borneo. Opera Bot. 69: 
Bum HMANN, S. L. 1983. Buzz pollination in ae 
In: C. E. Jones & R. J. Little (editors), Handbook 
of Experimental Pollination Biology. Van Nostrand, 


BUCHMANN. 1981. sang rad of 
Mouriri myrtilloides (Melastomataceae: Memec 
1648), an oil flower in Panama. Barone 13 (suppl.): 


Buick, W. 91. Beitráge zur Kenntnis der myr- 
mekophilen Pflanzen und der Bedeutung der extra- 
nuptialen Nectarien. Ann. Jard. Bot. Buitenzorg 10: 
75-144. 


Volume 76, Number 2 
1989 


Stein & Tobe 531 


Floral Nectaries in Melastomataceae 


COGNIAUX, A. Mélastomatacées. In: A. de Can- 
dolle (editor), Mongraphiae Phanerogamerum 7: 1- 
1256. 


CRUDEN, R. W. 1972. Pollinators in high-elevation eco- 
systems: relative effectiveness of birds and bees. Sci- 
ence 176: 1439-1440. 

DAHLGREN, R. & R. F. THORNE. 1984. The order Myr- 
tales: circumscription, variation, and relationships. 

Ann. Missouri Bot. Gard. 71: 633-699. 

. 1977. Anatomy of ves Plants, 2nd edition. 
Joh Wiley & Sons, New 

EyDE, R. H. 1981. O structures and ev 
lution in Ludwigia (Onagraceae). III. cubana 
nectaries, conclusions. Ann. Missouri Bot. Gard. 68 
379-412 

. T. MORGAN. 1973. Floral structure and 
evolution in Lopezieae (Onagraceae). Amer. J. Bot 

7711-787. 

1979. Secretory Tissues in Plants. Academic 

Press, London. 

A 'B. € P. H. Raven. 1972. Energetics and 
pollination ecology. Science 176: 597-602 

Hickey, L. J. 1977. Stratigraphy and paleobotany of 
the Golden Valley formation vel ML of west- 
ern North Dakota. Geol. Soc. 

pace a 1 Über die o 

—pinnchtungen von B dm teal 

m (De iid m rae Bot. iu 1899: 105-12 

Leslie, E. G., C. M. Ric . G. aan 1966. 
Tibi on the foral relationships of the Galá- 
pagos carpenter bee. Pan Pacific Entomol. 42: 1- 


60: 
FAHN, i 


LUMER, C. 
lastomataceae) in a 
tonia 32: 512-51 

——— & R. 


1980. Rodent pollination of B/akea (Me- 
a Rican cloud forest. Brit- 


kea 
austin-smithii and a (Melastomata- 
ceae) by small rodents in Costa Rica. E ud 18: 


1984. di e on the 
ra (Melas- 


J. PIPOLY. 


Memecyleae iMaka Fl. 
Neotrop. 15: 1-295. 


RENNER, S. S. 1983. The widespread occurrence of 
anther destruction by Trigona bees in Melastoma- 
taceae. Biotropica 15: 257-267. 

1989. A survey of reproductive ET in 
neotropical Melastomataceae and Memecylaceae. 
Ann. Missouri Bot. Gard. (this volume). 

SiMPSON, B. B. & J. L. NEFF. 1981. Floral rewards: 
alternatives to pollen and nectar. Ann. Missouri Bot. 
Gard. 68: 301-322. 

SNow, W. B. K. SNow. 1980. Relationships 
between hummingbirds and flowers in the Andes of 
Colombia. Bull. Brit. Mus. (Nat. Hist.), Zool. 38: 


-139. 
deo ue K. On the nectary in the oo 
mecylon heyneanum Benth. Curr. Sci. 11: 

41 5-4 19. 

ToBE, H. & P. H. Raven. 1984. The embryology and 
relationships of Rhynchocalyx Oliv. (Rhynchocaly- 
caceae). Ann. Missouri Bot. Gard. 71: 836-843. 

Une, E. 1896. Weiteres zur Blütheneinrichtung von 
Purpurella ins ape und Verwandten. Ber. 


Deutsch. Bot. v : 169-178. 
Van VLIET, C. J. C d KorEK-NoonMaN & G. J. H. 
TER WELLE jon Wood anatomy and classifi- 


cation of the Melastomataceae. Blumea 27: 463- 
4 


1955. The structure and dehiscence 
and Mouriri. Phyto- 


VENKATESH, C. S. 
of the anther in Memecylon 
morphology 5: 435-440. 

VocEL, S. 1957. Fledermausblumen in Südamerika. 

. Bot. Z. 104: 491-530. 

Certamen Melastomataceis. Phy- 

400 


Melastomataceae. In: G. Harling & B. 

Sparre (editors), Flora of Ecuador 13: 1-405. 

Certamen Melastomataceis 38. Phy- 
tologia 64: 293- 301. 

ZANDONELLA, P. 1977. „Apports de l'étude comparée 
des nectaires floraux à la conception cie d 
de dw Be E URSI Ber. Deutsch. Bot 
Ges. 90: 25. 

ZEIL, W. 1979. The Andes: A pei ae Review. Ge- 


briider Borntraeger, Berlin 


A SYSTEMATIC REVISION OF Paul E. Berry’ 
FUCHSIA SECT. QUELUSIA 
(ONAGRACEAE) 


ABSTRACT 


Fuchsia sect. Quelusia consists of nine closely related species, eight from southeastern Brazil and one from southern 


m F. regia and 


It is one of onl s. Unique characters of the section include the nina 


violet, convolut 


Southern Hemisphere. With the Old World sect. Skinnera as the sister group of all the American species of Fuchsia, 
sect. Quelusia represents an early offshoot of the ancestral South American fuchsias, distinct from the tropical Andean 
and Central American sections in its polyploidy and derived, 3-porate pollen. The nearest relative and sister species 
of sect. Quelusia i is i lycioides, a specialized xerophyte and the sole member of sect. Kierschlegeria, which occurs 
in central Chile just north of the range of F. magellanica. The Andean and Brazilian populations of sect. Q 


probably became cd in 
to the east of the 


subspecies are i described, F. regia sub 
ccur sympatrically with alber members of the section, and it forms natural en 
The only naturally occurring octoploid populations in the genus occur in F. 


the only species now found to 
with at least five different os 
and appear to be recently derived. 


the late Tertiary, when the austral temperate forests were broken 
des, as these were strongly uplifted. While F. magellanica has not differentiated erac tuam 
throughout its range in the southern Andes, sect. Quelusia has radiated extensively in the subtropical m 


up by inc 


Fuchsia regia, on 


Fuchsia regia is 


Fuchsia is a distinctive genus of Onagraceae 
with close to 105 species belonging to ten sections 
(Berry, 1982; Berry et al., 1988). The genus con- 
sists mostly of mesic shrubs confined to cool, moist 
habitats, with nearly 34 of the species concentrated 
in the tropical Andes. The remaining species occur 
in Mexico and Central America, Hispaniola, south- 
eastern Brazil, the southern Andes, New Zealand, 
and Tahiti. 


In recent years, Fuchsia has been the subject 
of a series of studies designed to refine the tax- 
onomy and to improve our understanding of the 
evolutionary relationships within the group. A com- 
prehensive analysis of leaf flavonoids (Averett et 
al., 1986) confirmed the distinctiveness of the New 
Zealand species of the widely disjunct sect. Skin- 
nera from their American congeners. Studies of 
pollen morphology and ultrastructure in a broad 


! This study was made possible by financial support provided by the American Philosophical Society, the Consejo 


Nacional de Investigaciones Cientificas y Tecnológicas (CONICIT, Venezuela), the C 


e study. Haroldo Cavalca 


sucre field guides in Brazil. José Badini, José Fernando Baumgratz, Lois Bako. Rejan 
oberto Klein, Teresinha Silva Oliveira, Solange Vasconcellos de Pessoa, and 
arlier version of this 


assistance. Valuable 


an e 
J. O. Wright. The staff of the Jardim Botânico a 


Bolívar, in (o enezuela. I thank 
B, BAA, BAB, BH, BHCB, MH, BM, : 
ICN, K, LE, LG, LIL, L , MA, IC 


? Dep partamento de Biologia de Organis 
Current address: Missouri Botanica! Garden 


, P.O. Box 


ANN. 


L, LP, LY, BM, 
A, S, SGO, SI, SP, uk TCD, TEX, U, UC, UEC, US, VALD, W, 
ismos, Universidad Simón Bolivar, A calle o 89000, Caracas 1081, Venezuela. 
299, St. Louis, Missouri 63166-0299, U.S.A. 


ional de 


Guedes, Lui 
aulino Reitz also ede field 


R, OXF, P, PACA, PEL, POM, R, 


Missouni Bor. Garb. 76: 532-584. 1989. 


Volume 76, Number 2 
1989 


Berry 533 
Fuchsia sect. Quelusia 


array of species have helped to define sectional 
limits by detecting consistent differences in viscin 
types, aperture number, and grain size 
(Skvarla et al., 1976, 1978; Nowicke et al., 1984; 
Praglowski et al., 1983). Systematic revisions for 
seven of the ten sections in the genus have been 
completed, based on broad cytological surveys, a 
thorough review of the major collections of her- 
barium specimens, and extensive field examination 
of natural populations (Breedlove, 1969; Berry, 
1982, 1985; Breedlove et al., 1982; Berry et al., 
1988). The present study uses the same method- 
ology to produce a revision of the species of sect. 
Quelusia, a group centered in southeastern Brazil 
and forming the largest section outside of the trop- 
ical Andes. 

A better understanding of the systematics of 
sect. Quelusia is particularly desirable, as the species 
form a closely related, yet unspecialized group that 
may have retained many of the primitive char- 
acters of the genus. Biogeographically, the section 
is confined to ancient forest areas in the southern 
Andes and in southeastern Brazil that show clear 
austral-Antarctic affinities. Ultimately, this may 
have been the source area from which other sec- 
tions differentiated and spread both to Australasia, 
when more or less direct overland connections were 
possible across Antarctica until the mid-Tertiary, 
and to Central America, where several small sec- 


thread 


tions presently occur 

orticulturally, most of the hundreds of hybrid 
Fuchsia cultivars have been derived primarily from 
members of sect. Quelusia (Reiter, 1944), yet there 
has been much confusion as to which taxa were 
involved in the early crosses. Accurate information 
on the different native species of sect. Quelusia 
can also be expected to stimulate interest in their 
preservation, cultivation, and use in hybridization 
for new ornamental cultivars. 

In the only recent monograph of Fuchsia, Munz 
(1943) recognized five species in sect. Quelusia, 
together with Fuchsia hybrida Hort. ex. Vilm., a 
name used to encompass most of the common 
garden hybrids. Fuchsia magellanica and F. regia 
included three and four varieties, respectively; these 
species have the broadest geographical distributions 
in the genus and also show complex patterns of 
morphological variation. In this study, Fuchsia 
magellanica is treated as a single, cytologically 
uniform, polymorphic species with no clear geo- 
graphical pattern of differentiation. Fuchsia regia, 
on the other hand, has tetraploid and octoploid 
populations, with geographically differentiated 
variants that are treated here as comprising three 
intergrading subspecies. Both F. alpestris and F. 


glazioviana are reinstated as distinct species, and 
two previously unrecognized species are described, 
F. brevilobis and F. hatschbachii. As herein de- 
limited, sect. Quelusia consists of 11 taxa, with 
nine different species. 


HABITAT AND DISTRIBUTION 


Fuchsia sect. Quelusia is an entirely Southern 
Hemisphere group of species, bs. a latitudinal 
range extending from 18*S to 

outh America in two large vi widely disjunct 
areas: the southern, temperate Andes of Chile and 
Argentina, and the coastal mountains and high 
plains of southeastern Brazil (Fig. 1). 

The only species occurring in the southern An- 
des is F. magellanica, which is characteristic of 
the wet, temperate, Valdivian forests that are cen- 
tered in the Región de Los Lagos of Chile. These 
forests are considered to be relicts of an ancient 
vegetation type that includes many taxa of Aus- 
tralasian affinity, such as Nothofagus, Drimys, 
Podocarpus, and Araucaria (Rambo, 1951; 
Hueck, 1972; Lötschert, 1981). Fuchsia magel- 
lanica also extends farther north into drier areas 
in Chile, where it occurs in moist microhabitats as 


. It occurs in 


far north as Santiago and Valparaiso, and then 
southward to the subantarctic scrub forests of Tier- 
ra del Fuego. Throughout its range, F. magellan- 
ica occurs as low as sea level, but only at its 
northernmost range does it reach its upper alti- 
tudinal limit of 1,750 m. Seasonality increases 
southwards, so that in Tierra del Fuego, flowering 
is restricted to the summer months of December 
to March, and plants are strongly deciduous in the 
freezing winter months. 

e other sie species of the section all occur 
in the highlands of southeastern Brazil. A single 
species, F. regia, is widespread and covers the 
entire range of the section in Brazil, from 18?20'S 
to 29930'S. The remaining species have much more 
restricted distributions, several occurring on a sin- 
gle mountain range. 

Fuchsia regia consists of three largely allopatric 


subspecies Subspe ec ies 


mainly in dense 
dawa forests at the crests and on the seaward- 
facing escarpments of the coastal mountains from 
Rio Grande do Sul to northern São Paulo state, at 
elevations of 500-1,450 m. In Santa Catarina, 
however, it is also found in enclaves of montane 
vegetation on the summits of the isolated *morros" 
that emerge just 100 to 300 m above the Atlantic 
lowland forest in the Itajai Valley (Klein, 1978). 
It has also been found at sea level in restinga forests 
near the Santa Catarina- Rio Grande do Sul border, 


534 


Annals of the 
Missouri Botanical Garden 


where Reitz (1961) found a number of other plants 
typical of the high inland serras. 

Fuchsia regia subsp. reitzii is restricted to 
thickets and low forest on the drier planalto, or 
southern highlands, that occupies much of the in- 
terior of Paraná, Santa Catarina, and Rio Grande 
do He It occurs at altitudes between 900 and 

m in areas subject to strong winter frosts 
ed 1984). The range of subsp. reitzii in this 
area coincides quite closely with Araucaria an- 
gustifolia, the dominant element of the planalto 
forests (Hueck, 1953). In Rio Grande do Sul and 
southern Santa Catarina, the joris ends abruptly 
to the east at the **Aparados” or ** ' sheer 
cliffs that drop down beh of meters to the 
Atlantic coastal plains. At these sites, subsp. reitzii 
reaches its eastern limit, growing sympatrically 
along a narrow band with the westernmost popu- 


aimbes,' 


lations of subsp. serrae. 

Fuchsia regia subsp. regia occurs further north 
in a series of separate mountain systems, from 
northern Sào Paulo to Rio de Janeiro, Minas Gerais, 
and Espirito Santo. It grows in cloud forests be- 
ginning at 1,000 m and reaches open campos 
above treeline as high as 2,400 m on Mt. Itatiaia. 

Several species are restricted to high altitudes 
and are known from just one area. Fuchsia bra- 
celinae grows only between 2,300 and 2,850 m, 
above treeline in the “campos de altitude" of a 
few peaks of the Serra do Caparaó. This is the 
highest mountain range of southern Brazil, situated 
along the Minas Gerais- Espirito Santo border. Sim- 
ilarly, F. campos-portoi occurs only in campos 
around Mt. Itatiaia, in the Serra da Mantiqueira, 
between 2,100 and 2,550 m. Fuchsia coccinea 
occurs naturally only at the summit of four of the 
highest mountains in Minas Gerais, between 1,400 
and 2,000 m 

Two species are endemic to several of the high, 
granitic *morros" characteristic of the Serra do 
Mar in Rio de Janeiro state. Fuchsia alpestris 
grows among bushes at the base of granitic outcrops 
in the Serra dos Órgãos and around Santa Maria 
Madalena. Fuchsia glazioviana is found as high 

as 2,100 m in open campo vegetation on morros 
around Nova Friburgo and Santa Maria Madalena. 
It also descends into e forest as low as 1,500 
m on the same mountai 

Two newly described pcm from Paraná and 
southern Sáo Paulo states have broader ranges. 
Fuchsia brevilobis occurs in forests of low, inland 
mountain ranges at 600—900 m, but on the eastern 
slopes of the Serra do Mar in Paraná, it is found 
from 400 m to sea level. Fuchsia hatschbachii 
occurs further inland in Paraná, in drier forests 


than the coastal mountains, and is apparently re- 
stricted to limestone and sandstone outcrops be- 


tween 950 and 1,150 m. 


REPRODUCTIVE BIOLOGY 


The members of sect. Quelusia all conform to 
a very similar pattern of floral biology. The flowers 
are hermaphroditic and self-compatible, as well as 
protogynous and herkogamous. Consequently, the 

enus has been characterized as 
crossing" (Raven, 
pendent, with the stigma exserted well below the 
anthers, a prime example of “approach herkoga- 
my” (Webb €: Lloyd, 1986), where the stigma is 
contacted as the visitor enters the flower prior to 
picking up pollen. Protogyny is incomplete, how- 
ever, because the anthers begin to dehisce within 
a day after anthesis, while the stigmas remain re- 
ceptive from the onset of anthesis until flower se- 
nescence several days later. Despite the presence 
of these outcrossing mechanisms, geitonogamous 
pollen transfer may still lead to effective selfing. 
Automatic selfing can also occur, since pollen usu- 
ally hangs down in clumps joined by viscin threads, 
which are able to contact the stigmatic surface 
before the flower senesces. 

The flowers of sect. Quelusia are highly spe- 
cialized for pollination by hummingbirds. Each 
species has red, odorless, hanging, tubular flowers 
with copious nectar secreted at the base of the 
floral tube, all standard features in hummingbird- 
pollinated plants (Faegri & van der Pijl, 1979). 
Unlike other bird-pollinated fuchsias, however, the 
petals are much larger and strongly convolute; they 
thus act functionally as a prolongation of the floral 
tube, further restricting the access of insects to the 
nectary. eep blue-violet color of the petals 
of sect. Quelusia is unique in the genus, althoug 
its significance in pollination is not clearly under- 
stood. The abundance of viscin threads in Fuchsia 
is among the most copious in the family and has 
been considered a probable adaptation to bird pol- 
lination (Nowicke et al., 1984) 

Hummingbirds are commonly observed visiting 
the flowers of sect. Quelusia throughout its native 
range, but few published accounts have identified 
the species of the hummingbird visitors. Curtis 
(1836) cited an early account by J. Anderson, the 
botanist on Philip King's voyage to the Magellan 
Straits, who observed numerous hummingbirds vis- 
iting F. magellanica around Port Famine, even 
after days of constant rain and sleet at near-freez- 
ing temperatures. Hauman-Merck (1912) reported 
frequent visits of Eustephanus galeritus (Mol.) on 


Volume 76, Number 2 
1989 


Berry 535 
Fuchsia sect. Quelusia 


F. magellanica near Corral, in Valdivia, Chile. In 
Brazil, Ruschi (1982) reported Colibri serrirostris 
(Vieillot) visiting flowers of F. regia. A Plovercrest 
hummingbird, Stephanoxis lalandi (Vieillot), was 
observed visiting Fuchsia in the open campos area 
of Mt. Itatiaia, Rio de Janeiro state by Mitchell 
(1957), as well as by Ruschi (1982) in Teresopolis 
at 900 m. Three hummingbird species, Colibri 
serrirostris, j euco- 
chloris albicollis, were common around F. bra- 
celinae in the Serra do Caparaó, at 2,400 m, 
during a visit in February 1985 (L. Gonzaga, pers. 
comm.). It is unlikely that any strong pollinator 
discrimination exists between the different species 
) flowers are very 


Stephanoxis lalandi, and 


of Fuchsia in Brazil, since the 
similar in size and overall floral pattern. On both 
F. campos-portoi and F. brevilobis, I have seen 
two different hummingbird species visiting flowers 
of the same bush in rapid succession, and I also 
watched an individual hummingbird visit flowers of 
F. brevilobis right after leaving flowers of a neigh- 
boring plant of F. regia subsp. serrae. 

Besides hummingbirds, several insect species 
have also been observed visiting flowers of sect. 
Quelusia in their native habitat. Apis mellifera 
was a common visitor to flowers of F. coccinea at 
Serra da Piedade, Minas Gerais, during our visit 
in February 1985; it was also observed on 
campos-portoi at Mt. Itatiaia in January 1985. 
Hauman-Merck (1912) described the visits of 

bus chilensis to F. magellanica in southern 
Chile, and thought it to be a more important visitor 
there than hummingbirds. Both of these bee species 
hang from the anthers to collect pollen and ma 
attempt to reach the nectary at the base of the 
tube; Bombus, especially, is able to make contact 
with the stigma and acts as an effective pollinator. 
I also saw a butterfly visiting F. regia subsp. serrae 
near Mogi das Cruzes, Sào Paulo, Brazil; it grasped 
the stamens attempting to enter the floral tube for 
nectar and made contact with both the anthers and 
the stigma in the process. When species of sect. 
Quelusia are cultivated outside their native range 
in areas where hummingbirds do not occur, such 
as England, they are frequently visited by honey 
bees and bumblebees (J. Wright, pers. comm.). 

Seed dispersal is probably effected by frugivo- 
rous birds, although no direct observations or pub- 
lished reports have been made. 


MORPHOLOGY 


Habit and stems. Plants of sect. Quelusia are 
all shrubs or lianas. Stems are usually flexuous and 
require the support of neighboring vegetation, but 


low, densely branched shrubs are common in such 
species as F. coccinea and F. bracelinae. Fuchsia 
regia can become a large, forest liana, with stems 
to 20 cm in diameter climbing as high as 15 m 
into the tree canopy, but it can also be found 
growing as a compact shrub in open situations. On 
large, climbing plants of F. regia and F. alpestris, 
it is common to find hanging branches several 
meters long. Short, divaricating side shoots are 
typical of young plants of F. regia subsp. serrae. 

Vegetative reproduction occurs in most taxa 
through stem-layering, especially when long, flex- 
uous stems lay over ground litter. Fuchsia bra- 
celinae and F. coccinea reproduce locally by un- 
derground, stoloniferous stems, and some 
populations of F. regia subsp. serrae ] 
roots on shoots well above the ground, as described 
by Lindley (1841) for F. radicans. 


aerial 


Leaves. Phyllotaxis in sect. Quelusia is op- 
posite or whorled. The most corumon condition in 
the section is ternate leaves, but individual plants 
often combine leaves in pairs, threes, and fours on 
different shoots. Exceptions are F. alpestris, almost 
always with opposite leaves, and F. regia subsp. 
reitzii, with leaves all in whorls of 3, 4, or 5. 
Fuchsia glazioviana has unusually short inter- 
nodes, just 3-12 mm long, with the leaves densely 
packed on the stems. 

The texture of the leaves of most species is 
membranous, but F. regia subsp. serrae has co- 
riaceous leaves, and those of F. regia subsp. regia 
are mostly subcoriaceous. Leaf surface is a useful 
character in some taxa; it is smooth and lustrous 
in F. regia subsp. serrae, with a dark green color 
above and a paler tone below. Leaf surfaces of 
Fuchsia glazioviana and F. hatschbachii are sim- 
ilar, but the surface is less brilliant. Impressed leaf 
veins are especially characteristic of F. bracelinae, 
and to a lesser M dii in F. brevilobis. Secondary 
venation in leaves of F. campos-portoi can be 
almost ds while in F. regia subsp. re- 
itzii a conspicuous tertiary venation is usually ev- 
ident. 

Leaf size and shape vary considerably within 
taxa, but some differences can be readily distin- 
guished. The leaves of Fuchsia glazioviana, F. 
bracelinae, and F. campos-portoi are small, most- 
y less than 4 cm long, and very narrow in F. 
campos-portoi. 'The largest leaves occur in F. al- 
pestris and F. regia subsp. regia and are mostly 
6-14 cm long. Ovate leaves with cordate bases 
are diagnostic of F. coc 

eaf margins are caló gland-den- 


ticulate, but in F. campos-portoi and F. regia 


Missouri Botanical Garden 


Annals of the 


536 


Ue E y \ y HE 
ON Pel 4 3 : 
|^. > ‘N ANM o | 

: EN AT *}3 


100 — 200 2300 
y 


o 
| A cal 
| p 


‘Fuchsia sect, Quelusia 


Distribution of 


SOUTH AMERICA 


Las ca E 


d aw ry? LES 
NO P Len ATES Y ES 
AAA ZN Ke 


e, DEN Wr 


Native distribution of Fuchsia sect. Quelusia. 


FIGURE 1. 


s 
por aad 
_”T 
+ | 
| 
| 
—T » 

ines E = - 
o o o 
^ ^ * ^ 


Volume 76, Number 2 
1989 


Berry 537 
Fuchsia sect. Quelusia 


subsp. reitzii they are conspicuously gland-serru- 
late, with the teeth angled towards the leaf apex. 
Fuchsia magellanica has mostly serrulate mar- 
gins, but they are sometimes crenate, with shallow 
lobes between the teeth. 

Petiole length is useful in discriminating taxa in 
sect. Quelusia. Fuchsia coccinea and F. brace- 
linae have nearly sessile leaves, with petioles 1—3 
mm long, while they exceed 10 mm in F. regia 
subsp. regia and F. regia subsp. serrae. Often the 
proportion of the petiole to the blade length is very 
informative: high (> 1:4) in F. regia and F. mag- 
ellanica, and low (1:8 «) in F. hatschbachii and 
F. alpestris. Petioles are stout and over 1 mm 
thick in F. alpestris, F. coccinea, F. regia subsp. 
serrae, and F. regia subsp. regia; they are slender 
in F. magellanica. 

Stipules in all but one taxon are small, lanceo- 
late, and deciduous. Fuchsia regia subsp. serrae 
is unique, with prominent, persistent stipules that 
are thick and recurved, usually fused in pairs be- 
tween adjacent leaves, and give a characteristic 
nodose appearance to the stems. 


Flowers. 
in pairs in the axils of young leaves near the branch 


Flowers are all borne singly or rarely 


tips. Rarely more than two or three whorls of leaves 
bear flowers at the same time on a single stem. 
The pedicels are pendulous, except in F. coccinea, 
where they are divergent from the stem before 
hanging down in the distal half. 

Section Quelusia is the only group in the genus 
where the sepals are much longer than the floral 
tube. In addition, the sepals are always connate to 
varying degrees at their base, which in effect 
lengthens the tubular portion of the flower. The 
petals are strongly convolute and form a cone 
around the stamens and stigma. The nectary con- 
sists of a smooth to slightly lobed band ca. 1 mm 
thick lining the inner, basal portion of the floral 
tube. A flower from sect. Quelusia is illustrated 
diagrammatically in Figure 2. 

The floral tubes of most species are cylindrical- 
fusiform, but F. campos-portoi is distinct with its 
short, subrhombic tubes. Sepal connation is only 

-6 mm in most taxa. Fuchsia brevilobis is ex- 
ceptional, with sepals connate for 10-16 mm, al- 
most completely concealing the corolla and longer 
than either the free sepal lobes or the true floral 
tube. Fuchsia regia subsp. serrae also has sepals 
joined for over 4 their length; the buds are no- 
ticeably four-ridged, and the free lobes sometimes 


recurve after anthesis. 
Fruits. Berries in sect. Quelusia are generally 
oblong-ellipsoid and dark purple to black when ripe. 


n F. regia subsp. reitzii, the berries are mostly 
globose, while in F. magellanica they are much 
more elongate than in the other species. Ovules 
are biseriate in each locule and number between 
60 and 120 per fruit. 


CYTOLOGY AND PALYNOLOGY 


Cytology. Concurrently with this revision, 
Hoshino & Berry (1989) made an extensive cy- 
tological survey of all 11 taxa of sect. Quelusia, 
examining 103 individuals from natural popula- 
tions and 11 cultivated plants. Their results showed 
the section to be entirely polyploid, predominantly 
tetraploid (n — 2x — ; i 
= 4x = 44) found in two species. All other sections 
in the genus, except for the tetraploid, monotypic 
sect. Kierschlegeria, are entirely or mostly diploid 
(n=x= 11). 

Tetraploidy was found in all specimens examined 


ut with octoploidy (n 


of six Brazilian species and in the more than 40 
populations that were studied from the widespread 
F. magellanica of the southern Andes (see Hoshino 
& Berry, 1989, Table 1). Normal bivalent for- 
mation was observed in all of these collections. 

Octoploidy was limited to the three subspecies 
of F. regia and a single cultivated individual of F. 
alpestris. Fuchsia regia subsp. reitzii and F. regia 
subsp. serrae were consistently octoploid, but tet- 
raploid individuals were also found in F. alpestris 
and F. regia subsp. regia. In all octoploids ex- 
amined, varying numbers of quadrivalents were 
present at meiotic Metaphase I and Anaphase I. 
Although tetraploid and octoploid plants occurred 
sympatrically in several localities, no hexaploid in- 
dividuals were detected. The chromosome numbers 
of the individuals examined by Hoshino & Berry 
are indicated in this paper under the specimen 
citations of the respective taxa. 

Palynology. Pollen grains of Fuchsia have 
distinctive, onagraceous features such as viscin 


threads, paracrystalin e-spongy ektexine, protrud- 
ing apertures, and a solid endexine. The genus is 
unique, however, in possessing predominantly two- 
aperturate, bilaterally symmetrical pollen grains. 
Radially symmetrical, three-aperturate grains also 
occur in the genus but are confined to polyploid 
species and are clearly derived from two-aperturate 
ancestors, a result of the increased cell volume of 
polyploid cells. Most sections of the genus can be 
distinguished by a combination of aperture number, 
viscin thread ultrastructure, and t 
sculpture elements. Detailed pollen descriptions of 
the different sections are provided in Praglowski 


et al. (1983) and Nowicke et al. (1984) 


e of exine 


538 


Annals of the 
Missouri Botanical Garden 


The members of sect. Quelusia have radially 
symmetrical and three-aperturate pollen grains, a 
condition consistent with the polyploidy found in 
all species of the section. Viscin threads are seg- 
mented-beaded, and the exine has globular sculp- 
ture elements. Praglowski et al. (1983) reported a 
size range of 33-60 um for P (polar axis length) 
and 59-85 um for E (equatorial axis) for the seven 
taxa they examined. Rare four-aperturate grains 
have been found in F. coccinea and F. regia; 
occasional two-aperturate grains were also found 
in F. magellanica and were considered to be re- 
versions to the ancestral condition of the genus 
(Praglowski et al., 1983; Nowicke et al., 1984). 
Octoploid individuals of F. regia cannot be distin- 
guished from tetraploids by differences in pollen 
morphology, although octoploid pollen grains may 
be significantly larger (Hoshino, pers. comm.). 

he only other section of Fuchsia characterized 
by three-aperturate pollen is the monotypic sect. 
Kierschlegeria (F. lycioides). Its pollen can be 
distinguished from that of sect. Quelusia, however, 
by the smooth viscin threads with elongated sculp- 
ture elements (Nowicke et al., 1984). 


FLAVONOID COMPOUNDS 


Because of the ornamental appeal of Fuchsia, 
several studies have examined the role of antho- 
cyanins in the different flower colors in the genus 
(Harborne, 1963; Yazaki & Hayashi, 1967; Noz- 
zolillo, 1970; Yazaki, 1976; Crowden et al., 1977). 
Flower colors are determined almost entirely by 
the 3-glucosides and 3,5-diglucosides of all six com- 
mon anthocyanins. Red coloration in the sepals 
and floral tube is due mostly to cyanidin and peon- 
idin pigments, whereas the characteristic blue-vi- 
olet color of the petals of all members of sect. 
Quelusia is due to the dominance of malvidin de- 
rivatives, with peonidin in much lesser concentra- 
tions (Nozzolillo, 1970; Yazaki, 1976; Crowden et 
al., 1977). Yazaki (1976) indicated that the intense 
blue-violet color of some cultivars is due to the 
simultaneous presence of other co-pigments in the 
cell, particularly quercetin glycosides. In some cul- 
tivars, the petals change color from blue-violet to 
purple-red with age; Yazaki (1976) has shown this 
to be due to the combined effect of the flavonoid 
co-pigments, together with a decrease in the pH 
of the cell sap brought about by the accumulation 
of organic acids. 

Averett et al. (1986) carried out a comprehen- 
sive survey of the foliar flavonoids in 80 taxa of 
Fuchsia and reported the flavonoid profiles of four 
taxa belonging to sect. Quelusia. Current taxo- 
nomic changes in the section, however, have mod- 


nectary | 
floral tube 


insertion of 


sepal 
connation 


FIGURE 2. Schematic longitudinal section of a flower 
from Fuchsia sect. nee with terminology used in 
the species description 


ified these results, and a revision is presented in 
Table 1. Based on the same samples examined by 
Averett et al. (1986), two additional taxa are now 
included, F. brevilobis and F. regia subsp. serrae. 
From these changes, it is apparent that it is F. 
brevilobis, not F. regia, that has a distinctive fla- 
vonol diglucoside. Fuchsia magellanica remains 
the only species in the section known to possess 
flavones (compounds 12 and 15), which could be 
used as excellent markers for detection of the par- 
entage of this species in cultivars of uncertain or- 
igin, although flavones might not be expressed in 
all F. magellanica hybrids. Otherwise, the re- 
maining flavonol monoglucosides are all of common 
occurrence in other taxa of the genus. 


NATURAL HYBRIDIZATION 


Because of the restricted distributions of most 
of the Brazilian species of Fuchsia, there is no 
range overlap among them, except with the wide- 
spread F. regia. Fuchsia regia occurs sympatri- 
cally with five other species, and in eac 
morphologically intermediate, putatively hybrid in- 
dividuals have been detected (Table 2). Populations 
of F. regia subsp. serrae also occur together with 
each of the other subspecies of F. regia, forming 
extensive hybrid swarms in areas of sympatry. 


case, 


Volume 76, Number 2 


Berry 539 
Fuchsia sect. Quelusia 


1989 
TABLE l. Foliar flavonoids in species of Fuchsia sect. Quelusia.' 
Compounds? 

Taxa Examined l 2 3 5 6 7 8 9 12 15 
F. brevilobis F m + + ax 
F. picis -portoi * + + + + + 
F. coc + + + + + 
F. ei MS + + + + + + + + 
F. regia subsp. regia + + + + E F + 
F. regia subsp. serrae + E + +F T P + 


' Based on the same samples as reported in Averett et al. (1986); changes in Per des are as follows: Davidse 


et al. 10904, Hatschbach 


Ramamoorthy 42976, Ramamoorthy & Reitz 1163, 


& Ramamoorthy 42974 (F. d Conrad & Diet 
and 


ich 2055, Hatschbach & 
am ud & Vital 685 (F. regia subsp. serrae); 


imamoormy & Vital 673, 674, 676, Lai (F. regia o 


* + = present in all o os E 
codes as in Averett et al. (19 = =K. 3-O-glu; 2 
Q-3-O-gluc; 7 = Q-3-O-gal; 8 = 9-3:0xham; 9 = Q-3- 


x bain Q = quercetin, A = apigenin, L = ed. pee — glucose, p — galactose, rham — 


arabino 


The ee orae dta intermediacy of several char- 
acters was the main criterion used in detecting 
hybrids iind different taxa; in most cases, both 
putative parents were growing sympatrically close 
by. Details of the different hybrid combinations 
found are provided in the Appendix. 

Natural hybridization among sympatric taxa of 
sect. Quelusia appears to occur readily, as there 


e only 
indication that reduced fertility might occur in hy- 
brids was from one artificial cross between a tet- 
raploid individual of F. regia subsp. regia and an 
octoploid individual of F. regia subsp. serrae, where 
the hexaploid progeny showed low pollen fertility 
(Hoshino & Berry, 1989). More experimental work 
is needed on the chromosomal behavior of crosses 
between tetraploid and octoploid taxa in sect. Que- 
lusia. 

In most cases, the area where two species over- 
lap is small, and the frequency or abundance of 
hybrids is low compared with the parental popu- 
lations. Fuchsia campos-portoi, F. coccinea, and 
F. glazioviana are all high-altitude species that 


TABLE 2. 


ene 
= present in some cols but at | in others. Same numerical 
= hu. 3-K 


-O-rham-glu; 6 = 
15 = L-7-0- v Allesviations K = 


rhamnose, ara = 


-gal; 


12 — A-7-O-glu; 


grow in the open **campos de altitude" at or above 
treeline. Hybrids between these species and F. re- 
gia occur where they descend into the upper forest 
zone, or when F. regia extends beyond its normal 
limit into the open campos. Fuchsia brevilobis has 
the inverse of this pattern, generally occurring at 
lower altitudes than F. regia subsp. serrae, with 
probable hybrids in limited areas where their ranges 
overlap. Fuchsia bracelinae may have grown sym- 
patrically with F. regia until the past century, when 
extensive cutting and burning eliminated the upper 
forest belt of most of the Serra do Caparaó. 

The situation among the subspecies of F. regia 
is notably different. These taxa are much more 
widespread and overlap more extensively than any 
of the other species in the section. Fuchsia regia 
subsp. reitzii is typical of the planalto region in 
Rio Grande do Sul and southern Santa Catarina 
states, but it contacts populations of the more mesic 
F. regia subsp. serrae along the easternmost edge 
of the planalto near the crests of the Serra Geral, 
producing large, widely variable, intermediate pop- 
ulations. Extensive intergradation between F. regia 
subsp. regia and F. regia subsp. serrae occurs 


Sympatric occurrence and natural hybridization among the native taxa of Fuchsia sect. Quelusia in 


Brazil. An X indicates that both sympatry and natural hybridization occur. Abbreviations are of the first three letters 
of the specific (capitals) or subspecific (lower case) epithets. 


Subspecies of 
F. regia 
subsp. regia X == = X X = 
subsp. reitzii — = € = = = = 
subsp. serrae = mE X = — = X X 
ALP BRA BRE CAM COC GLA HAT reg rei 


540 


Annals of the 
Missouri Botanical Garden 


along the eastern edge of the Serra da Bocaina 
and the Serra do Mar in northern Sào Paulo state. 
Although F. regia is the only species presently 
found in widespread contact with other species of 
sect. Quelusia, the numerous hybrid combinations 
detected indicate that interspecific genetic recom- 
bination may have been an important mechanism 
in providing a rich source of novel, adaptive ge- 
notypes in the fuchsias of southern Brazil. 


EVOLUTION OF SECTION QUELUSIA 


Various lines of evidence, including chloroplast 
DNA restriction fragment analysis (Sytsma & Smith, 
1988) and leaf flavonoid analyses (Averett et al., 
1986), have suggested the Old World sect. Skin- 
nera to be the earliest offshoot of the genus and 
the sister group of all the American sections of 
Fuchsia. The oldest fossil pollen grains of the genus 
have been found in New Zealand and Australia 
from Oligocene and Miocene deposits (Pocknall & 
Mildenhall, 1984; Berry et al., in prep.). Despite 
the early divergence of this group, the members 
of sect. Skinnera exhibit several specialized fea- 
tures for the genus, including the derived condition 
of male sterility, reduced petals, presence of flavone 
sulphates, and a variety of unusual life forms, such 
as large trees and procumbent creepers. 

The most likely area of origin of Fuchsia is 
South America, where the greatest concentration 
of species as well as the sections with the most 
generalized features in the genus occur (Raven, 
1979a; Berry, 1982). Whereas the large Andean 
sections appear to have arisen and radiated during 
the Neogene uplift of the tropical Andes (Berry, 
1982, upied by sect. 
Quelusia have changed little since the mid-Ter- 
tiary, when temperate forests were much more 
widespread in southern South America (Simpson, 
1973). With the current Australasian fossil record, 
it is now apparent that Fuchsia formed part of the 


5) the areas now occ 


Nothofagus and podocarp-dominated temperate 
forests that occurred more or less continuously 
across southern South America, Antarctica, and 
Australia until at least the late Eocene (Zinmeister, 
1987; Truswell, in press). The wet, temperate for- 
ests of the southern Andes and the montane forests 
of southeastern Brazil are now the only remnants 
of these austral temperate forests in South Amer- 
ica, which, like Fuchsia, share a large number of 
disjunct plant groups that also have close relatives 
in Australasia (Raven, 1979b) 

Although there are no fossil records to indicate 
an early presence of sect. Quelusia in South Amer- 


ica, the section follows a pattern consistent with 
other relicts of the southern temperate forests. It 
is ecologically conservative, with most individuals 
restricted to moist sites in cool, forest habitats. 
Morphologically, there is little divergence in the 

section, with species differentiated by relatively 
minor variations on a common pattern of floral and 
vegetative characters. Many of the features of sect. 
Quelusia are unspecialized in the genus, including 
the axillary, bisexual flowers, well-developed petals, 
numerous seeds, 
threads. It is possible that some of the primitive 
characters in sect. Quelusia were preserved throug 


and segmented-beaded viscin 


the buffering effects on evolutionary divergence 
caused by paleopolyploidy in the group (Stebbins, 
1985; Hoshino & Berry, 1989) 

The ancestor of the modern-day species of sect. 
Quelusia was probably tetraploid, with derived, 
three-porate pollen (Hoshino & Berry, 1989). As 
such, it could not have given rise directly to the 
other, mostly diploid, sections of the genus, but 
rather represents an early offshoot of the ancestral 
stock of Fuchsia in South America. Early on, sect. 
Quelusia developed its own floral syndrome for 
hummingbird pollination: the relatively short, nec- 
tar-holding floral tube; large, attractive sepals; and 
petal cone and sepal tube that further extend the 
tubular portion of the blossom. 

Section Kierschlegeria, containing only F. ly- 
cioides restricted to the dry coastal scrub of central 
Chile, is the only other section of Fuchsia that is 
entirely polyploid and has three-porate pollen grains. 
Because of this similarity and its geographic prox- 
imity to F. magellanica, Berry (1982) and Raven 
1988) postulated that it is related to sect. Que- 
lusia. Using chloroplast DNA restriction fragment 
analysis, Sytsma & Smith (1988) demonstrated 
that sects. Quelusia and Kierschlegeria form a 
distinct lineage from the rest of the genus, defined 
by three synapomorphies. Furthermore, within this 


— 


lineage F. lycioides could not be distinguished from 
the two species of sect. Quelusia studied, F. ma- 
gellanica and F. regia. This evidence of a close 
relationship is remarkable, since F. lycioides has 
a high number of specialized features, such as 
functional dioecy, spinose leaf bases, low seed num- 
ber, smooth viscin threads, small flowers, and sum- 
mer deciduousness. Evidently, the evolutionary shift 
of sect. Kierschlegeria to a xeric habitat in the 
southern Andes entailed a large number of mor- 
phological changes corresponding to a much small- 
er degree of genotypic divergence. 

Within sect. Quelusia, we are faced with the 
question of why there is just one species in Chile 
and Argentina, while in Brazil there are eight species 


Volume 76, Number 2 
1989 


Berry 
Fuchsia sect. Quelusia 


in an area of comparable size. Fuchsia magellan- 
ica is a classic example of the lack of differentiation 
that is characteristic of the /Vothofagus forests of 
Chile. These habitats presumably have been stable 
and continuous for long periods of time, providing 
little opportunity for speciation to occur. Most taxa 
inhabiting these areas have retained primitive char- 
acters for their respective groups (Simpson, 1973). 
Although the current distribution of F. magellan- 
ica extends beyond the Nothofagus forests, vir- 
tually all of the area now occupied by this species 
south of 44°S latitude was covered by ice during 
the Pleistocene (Vuilleumier, 1971) 

In contrast to the temperate latitudes of Chile 
and Argentina, the Brazilian members of sect. Que- 
lusia are centered in the subtropics. There, Fuch- 
sia is limited to the cool, moist habitats of the 

oastal mountains and the interior planaltos. Most 
of the cold-adapted taxa of Brazilian fuchsias are 
vicariant species that are now restricted to just a 
few high mountain peaks. Allopatric speciation in 
this group of species was likely enhanced by the 
joining and subsequent separation of populations 
caused by the cycles of Pleistocene glacial and 
interglacial events (Simpson, 1979; Klein, 1984). 

Two of the Brazilian species have adapted to 
warmer, lower-altitude habitats that are more con- 
tinuous over much of the southeastern part of the 
country. Fuchsia regia extends over the entire 
range of the section in Brazil and forms a broad 
taxonomic complex treated here as comprising three 
subspecies, each adapted to a slightly different set 
of environmental conditions. This species shows an 
intermediate stage in the morphological differen- 
tiation of populations between the more uniform 
F. magellanica in the southern Andes and the 
distinct, high-altitude species of Brazil. Fuchsia 
regia overlaps geographically with most of the 
other species in the section and hybridizes with 
them whenever they come into contact; these hy- 
brids may constitute an important source of genetic 
recombination for new adaptive genotypes in the 
section. It is also the only species in the genus with 
naturally occurring octoploid populations, which 
may lead to partial reproductive isolation with the 
other tetraploid taxa in the section. 

Two newly described species from Paraná, Brazil 
both consist of tetraploid populations that occur 
within the range of the octoploid subspecies of F. 
regia. The different ploidy levels may afford suf- 
ficient reproductive isolation for these species to 
remain distinct. One of the new species, Fuchsia 
hatschbachii, 1s the only species in the section that 
appears to be edaphically specialized, occurring on 
limestone and sandstone substrates on the eastern 


edge of the southern planalto. Populations of F. 
regia subsp. serrae occur within m of F. 
hatschbachii near Curitiba, Paraná, but the two 
have never been observed to grow together under 
natural conditions. The other new species, Fuchsia 
brevilobis, is specialized in its small flowers with 
elongate sepal tubes, and it has become adapted 
to lower altitudes than most neighboring popula- 
tions of F. regia. 

Overall, the evolutionary scenario of sect. Que- 
lusia differs significantly from the two, more spe- 
ciose, Andean sects. Fuchsia and Hemsleyella. 
Those species radiated recently into a broad ex- 

anse of new, cool montane habitats that were 
created with the recent uplift of the tropical Andes 
(Berry, 1982, 1985). Section Quelusia, on the 
other hand, has occupied a more ancient and stable 
vegetation type in which disruptive selection may 
have played less of a role in the divergence of 
populations. 


CLADISTIC ANALYSIS 


This section examines the relationships of the 
taxa of sect. Quelusia and the closely related F. 
lycioides by means of a formal cladistic analysis. 
The advantages of this approach are that it provides 
clearly stated hypotheses about evolutionary re- 
lationships based on shared, derived character 
states, and it is based on an objective, repeatable 
methodology. 

e data were analyzed on an ATT 6300 mi- 
crocomputer by the Wagner parsimony method of 
the PAUP software package (version 2.4.1., Swof- 
ford, 1985), using the branch and bound algorithm 
(Hendy & Penny, 1982). Character polarity was 
determined by outgroup comparison (Hennig, 1966; 
Watrous & Wheeler, 1981), using sect. Skinnera 
as the outgroup 
character states in sect. Skinnera are almost cer- 


. In two characters, however, the 


tainly apomorphic in the genus; so in these cases 
the other genera of Onagraceae were used as the 
outgroup. 

Twenty-two characters were used in the phy- 
logenetic analysis, with the character states listed 
in the data matrix (Table 

Fifteen equally parsimonious trees were pro- 
duced, each with 29 steps and a consistency index 
of 0.931. Topologically all the trees are identical, 
as they represent arbitrary resolutions of a single 
tetrachotomy that appears in the trees. The unique 
topology of these trees is presented in Figure 3. 
According to these results, sects. Kierschlegeria 
and Quelusia form a monophyletic group defined 
by five synapomorphies, but within this clade F. 


542 


Annals of the 
Missouri Botanical Garden 


Data matrix for characters of taxa used in the cladistic analysis of Fuchsia sects. Quelusia and 
6, 


Kierschlegeria. Sect. Skinnera is used as the outgroup. Characters 9, 1 


7 are unordered. A zero represents 


the plesiomorphic states, and higher numbers represent the apomorphic states. In those characters where no zero 
appears, the plesiomorphic conditions are uncertain, and the character states were coded as unordered. 


Characters 
Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 
sect. Skinnera 0 0 0 0 0 0 0 0 1 01 0 0 1 01 1 0 0 ? 0 ? 
F. lycioides 1 1 1000002 101 21 O 1 2 2 0 0 0 0 ? 
F. alpestris 1 ? 0 ? 1) 1 1 1 3 0 0 0 0 0 O 3 3 0 0 1 21 O0 
F. bracelinae 1] 1 O ? 1 1 1 1 3 0 O 0 0 0 0 3 3 0 1 O 1 O0 
F. brevilobis I 1 1 2 1 1 1 3 0 O D @ 0 0 3 3 0 D L 1 9 
F. campos-portoi 1 1 O 1 1) 1 1 1 30000003 30000 1 
F. coccinea 1l 1 O 1 1 1 11 1 3 0 O O 09 Q0 O 3 3 0 1 O |) 90 
F. glazioviana 1 1 O ? 1 li 1 l 3 0 0.0 0 0 O 3 3 0 0 O0 0 1 
F. hatschbachii l1 1 O 1 1 1 tt 3 0 O 0O O 0O O 3 3 0 0 0 0 Ọ 
F. magellanica 1 1 0 O 1 1 I 1 3 0 0 0 0 0 0 3 3 0 0 0 0 0 
subsp. regia | ? O i 1 1 l 1 3 0 0-0 O O 0 3 3 0 O 1 0 0 
subsp. serrae l| 2 0. bh 2 1 1 1 3 0 D D O 0 O 3 3 1 B l1 0 B 
subsp. reitzii L 2 O ? 1 1 1 1 3 0 0 0 3 3 0 0 0 
Pollen grains: 0 — 2-porate, 1 — 3- 


Viscin threads: 0 — segmented- um ]- 
Flavones: O — present, 1 = abse 


Leaves: 0 = ae l = opposite/ whorled 


d WA ad 


Sepals: 0 = unfused, 1 = fused < < P of length, 2 — fused > 


, Tor 
Ploidy level: 0- diploid, 1 — a 2- - ee 
oth 


V4 of length 


th band around ome part of floral tube, 2 — thick lobes at base of tube around style, 3 — 


cain ce by dominant gene 


Stamens: 0 = included within sepals, 1 = exserted beyond sepals 
Floral tubes en length: O — (sub)equal, 1 — sepals noticeably longer 
Nectar: — smoot 
man a from lower part of floral tube 
10. Stigma: O = subentire, 1 = 4-lobed (on functional female flowers) 
11. Pollen color: 0 = cream, 1 = blue 
12. Seed number per fruit: 0 = > 50,1 
13. Petiole bases: 0 = deci l = persistent and spin 
14. Sexual system: 0 = aioe l = male 
15. Sexual system: 0 = hermaphrodite, 1 = male sterility present and determination unknown 
16. Petal color: | = ink or pale purple, 3 — blue-violet 


17. Petal imbrication: 1 — separate, 2 = slightly sis ng, 3 — strongly convolute 
stent 


18. Stipules: 0 = small and deciduous, 1 = thick- 
19. Petioles: 0 — 
20. Habit (branch type): 0 = shrubb 


> 3 mm long, 1 = < 3 mm lon 
y, 1 = lianoid 


22. Internodes: 0 = 


lycioides is the sister group to all of sect. Quelusia. 
Section Quelusia is well delimited by the presence 
of four synapomorphies, and F. lycioides possesses 
eight unique, derived character states (autapo- 
morphies). Within sect. Quelusia, F. magellanica 
appears as the sister group of all the Brazilian taxa, 
which are grouped by a single character, loss of 
flavones. The Brazilian taxa of Fuchsia are poorly 
differentiated by the cladistic analysis, forming a 
polychotomy with four different lineages. Fuchsia 
hatschbachii is an isolated line in Brazil, differing 
from F. magellanica only by the loss of flavones. 
The three subspecies of F. regia form a separate, 
mainly octoploid lineage, supporting the monophy- 


crassate and persisten 


nt glabrous or sparsely pubsecent, 1 = parts of plant densely pilose 
mm, l = m 


ly of F. regia sensu lato. Fuchsia glazioviana and 

. campos-portoi are sister species within the third 
lineage. The fourth lineage includes four species, 
which are divided into two species pairs, the high- 
altitude F. bracelinae and F. coccinea, and F. 
alpestris and F. brevilobis. Two homoplasies (par- 
allelisms) occur in the cladogram: long-connate se- 
pals in F. brevilobis and F. regia subsp. serrae, 
and lianoid habit in two species pairs. 

The paucity of characters that discriminate be- 
tween the taxa of sect. Quelusia makes the results 
of the cladistic analysis within the section quite 
tentative, but it is also a reflection of the very close 
relationship of the species. The proposed sister- 


Volume 76, Number 2 
1989 


Berry 543 
Fuchsia sect. Quelusia 


- 
g fog 
e «o E 5 °? $ o o 2 
© 2 . $952 2 & 8B 
€ . E - o $ $ & r ” = 
E gpg 5 8 $3 
x o = > F1 Q 5 8 — x 
o > E 8 83 o o o 0 8 
5-2 5-2 
18 
20 19 20 
T 
2-2 +22 21 
+3 +4 
+9-b 
FIO 
+12 
13 5-1 
+15 6 
r 16-b 7 
+ 17-b 8 
FII i} 
+ 14 2-1 
9.c 
16 -c 
17-c 


Fic . Cladogram of Fi oo sects. ag 2 
Kierschlegeria (F. peri . Skinnera is used a 
the outgroup. Double crossbars indicate la. See 


Table 3 


group relationship of F. magellanica to the Bra- 
zilian members of the section is consistent with the 
biogeography of the group and the evolutionary 
hypotheses discussed previously, specifically the 
southern origin of the genus in South America and 
the secondary radiation of the section in the high- 
lands of southeastern Brazil. According to this hy- 
pothesis, the Brazilian taxa differentiated, after their 
separation from the Andean F. magellanica, from 
an ancestor that had lost flavones. The occurrence 
of two parallelisms between F. brevilobis and F. 
regia subsp. serrae is remarkable and may be the 
result of past hybridization between the two species. 


HisTORY OF CULTIVATION 
Members of sect. Quelusia were by far the most 
important species used in the early stages of fuchsia 


breeding. For the most part, however, fuchsia 
breeders in the past century kept few records about 


the parentage of their introductions. As a result, 
it is impossible to determine with certainty which 
species have been involved in the parentage of 
most of the modern cultivars (Reiter, 1944). An- 
cestry of sect. Quelusia can be assumed in cultivars 
that share the following exclusive **quelusioid" traits: 
fairly large, axillary flowers with sepals equal to or 
longer than the floral tube, and large, violet, round- 
ed, and strongly convolute petals. 

Following Reiter (1944), one can distinguish 
three stages in the history of fuchsia breeding dur- 
ing the past century. First, a variety of species and 
natural varieties were accumulated in cultivation. 


different introductions from sect. Quelusia, and 
then in a more important step, these were crossed 
with Mexican and Central American species of sect. 
Ellobium. Some of the resulting strains were fur- 
ther crossed with species of other sections, such 
as F. boliviana (sect. Fuchsia) and F. lycioides 
(sect. Kierschlegeria). Although some other cross- 


than the quelusioid crosses. The final stage involved 
the exploitation and selection of mutations, which 
no longer depended upon the introduction of native 
material. In the remainder of this section, I will 
review the role of sect. Quelusia in the first two 
stages of fuchsia breeding, then summarize the 
present-day situation of sect. Quelusia species in 
cultivation. 


Species introductions. With the possible ex- 
ception of a short-lived introduction of F. triphylla 
in the early 18th century (Wright, 1975), F. coc- 
cinea was the first species of Fuchsia introduced 
into cultivation in Europe. Plants of this species 
were growing at Kew Gardens in London in 1788 
(Aiton, 1789), and an illustration of it was published 
by Sims (1789). Hooker (1869) suggested that 
Sims’s figure might have represented F. magel- 
lanica rather than F. coccinea, because of the 
prominent petioles on the lower leaves. This is 
clearly not the case, however, as the figure ac- 
curately represents the unique arching pedicels and 
ovate, subsessile upper leaves of F. coccinea, and 
Sims's own specimen deposited at Kew is F. c 
cinea (pers. obs.). Although Dryander stated med 
the plants he described were introduced from Chile 
(Aiton, 1789), the species is native to just a few 
mountains in Minas Gerais, Brazil. 

The next introduction from sect. Quelusia was 
F. magellanica. An array of different variants of 
this species appeared in England in the 1820s 


544 


Annals of the 
Missouri Botanical Garden 


under several names, starting with F. gracilis, 
which was reported from the Edinburgh Botanic 
Garden in 1822 (Lindley, 1824). Fuchsia conica 
and F. gracilis var. multiflora were both intro- 
duced into England from Chile in 1824 (Lindley, 
1827a, b). Fuchsia discolor was introduced from 
Port Famine in the Magellan Straits around 1830 
(Lindley, 1835). I have seen no other evidence, 
such as old herbarium specimens from cultivated 
individuals, to indicate that F. magellanica was 
present in Europe prior to 1822. Soon after F. 
magellanica was introduced, however, it was con- 
fused with F. 
coccinea (see de Candolle, 
1869 

Fuchsia regia was first grown in England be- 
tween 1838 and 1840, under the name F. radi- 
cans (Lindley, 1841), and Fuchsia alpestris was 
cultivated in the Glasgow Botanic Garden as early 


as 1842 (Gardner, 1843). 


coccinea and was often cited as F. 


1828, and Hooker, 


Interspecific crosses. Interspecific hybridiza- 
tion among members of sect. Quelusia must have 
begun in the mid-1820s, but the first evidence for 
this was the appearance of F. globosa in a Hor- 
ticultural Society exposition in London in 1832 
(Lindley, 1833). Although the origin and parentage 
of this plant were not known to Lindley, the illus- 
tration and the description strongly suggest that it 
was a cross between F. magellanica and F. coc- 
cinea. A great proliferation of names of cultivars 
that were either variants of F. magellanica or 
crosses with F. coccinea followed in the next de- 
cades, including F. riccartonii, F. tenella, F. re- 
The history of these 
plants is unfortunately too poorly documented to 


curvata, and F. corallina. 
determine their precise parentage, and names such 
as riccartonii have been applied to quite different 
cultivated strains (J. Wright, pers. comm. 

A major step in the search for novel, interesting 
traits in Fuchsia cultivars was the use of F. fulgens, 
a diploid species of the Central American section 
Ellobium, in crosses with selections from sect. Que- 
lusia. One of the first such hybrids produced was 
exhibited in London in 1839, a cross of F. fulgens 
with F. globosa called “Standish's Fuchsia” (Lind- 
ley, 1840), which must have been either triploid 
or a spontaneously doubled hexaploid. Harrison 
(1841) also made a large number of crosses of F. 
fulgens with sect. Quelusia hybrids, publishing fig- 
ures of the flowers of several of the different hybrids 
he selected. A hybrid between F. cordifolia (— F 
splendens, sect. Ellobium) and F. globosa was 
grown in 1842, under the name F. exoniensis 
(Paxton, 1843). Fuchsia regia was also used in a 


series of crosses starting in 1840, leading to the 
development of darker petal color and improve- 
ment of flower shape (Wright, 1979). Fuchsia 
boliviana, a member of sect. Fuchsia from South 
America, may also have been used in the early 
crosses with selections of sect. Quelusia (J. Wright, 
pers. comm. ). 

he 19th-century crosses between members of 
sect. Quelusia and sect. Ellobium can be consid- 
ered the "mainstream" fuchsias later treated col- 
lectively under the names F. hybrida Hort. (Siebert 
& Voss, 1896; Munz, 1943) and F. speciosa Hort. 
(Bailey, 1900). In the strictest sense, these names 
have been attributed to crosses between F. mag- 
ellanica and F. fulgens. At the other extreme, 
they have been used to include almost all garden 
fuchsias. Reiter (1944) suggested that F. hybrida 
might best be treated as a horticultural complex 
including all cultivars of sects. Quelusia x Ellob- 
ium ancestry, including many of complicated ori- 


Chaudhuri (1956) made cytological observations 
that suggest how crosses between F. magellanica 
and F. fulgens might have facilitated the rapid 
advances in fuchsia breeding during the mid-1 800s. 
In his crossing experiments, the tetraploid (n — 2x 
— 22) F. magellanica and the diploid (n — x — 
11) F. fulgens yielded triploid progeny in which a 
high percentage of unreduced gametes were pro- 
duced. As a result, the F, crosses were partially 
fertile, and hexaploids (6x) were produced in the 
F, generation, as well as tetraploid (4x) and pen- 
taploid (5x) individuals in the respective backcross- 
es. All of these crosses retained some degree of 
fertility. Crosses between F. magellanica and other 
diploids, such as F. splendens and F. denticulata, 
did not have the same propensity for unreduced 
gametes as F. fulgens, and their hybrids were large- 
ly sterile. Thus, F. fulgens may have been a critical 
species that helped prevent total sterility in crosses 
with the polyploid members of sect. Quelusia and 
with later cultivars of even higher ploidy levels. 
O. Wright in England has 


confirmed Chaudhuri's findings, producing a com- 


More recently, J. 


plete set of 3x to 9x progeny, entirely by selfing 
and backcrossing the original F. magellanica X 
F. fulgens hybrids (J. Wright, pers. comm.). Wright 
further reports that none of these hybrids showed 
any obvious similarity to F. fulgens. 

Fuchsia magellanica is the only species in sect. 
Quelusia able to withstand winter conditions with 
long periods of freezing temperatures, so that all 
of the hedge plants of fuchsia grown outdoors in 
England and Ireland are variants of this species or 
crosses with a strong F. magellanica parentage. 


Volume 76, Number 2 
89 


Berry 545 
Fuchsia sect. Quelusia 


Many of the recent introductions from Brazil, how- 
ever, such as F. campos-portoi and F. coccinea, 
are proving to be frost-resistant as well (J. Wright, 


pers. comm. ). 


Native species in cultivation today. Many of 
the early species introductions have been main- 
tained for long periods of time in cultivation without 
crossing. Thus, Hooker (1869) was able to trace 
plants of F. coccinea found in the Oxford Botanic 
Garden in 1867 back to the original introduction 
of that species at Kew in 1788. Although this 
species may have been later lost from cultivation 
in England and Europe, it was successfully estab- 
lished on several of the Atlantic islands off the 
north African coast, such as Madeira, St. Helena, 
and the Canary Islands, in the early 1800s, per- 
sisting to present day (Hansen, 1969; A. Calero, 
pers. comm.). Plants of F. alpestris grown today 
in England and continental Europe have particu- 
larly small leaves similar to Gardner's type speci- 
men and may well date back to his original intro- 
duction in 1842. Wild strains of F. magellanica 
are apparently still present in England, and the 
species has become widely naturalized in parts of 
Ireland, New Zealand, Hawaii, South America, and 
eastern and southern Africa. 

As a result of the author's collections of native 
fuchsias in Brazil between 1985 and 1987, and 
seeds of different strains of F. magellanica ob- 
tained from Chile and Argentina, many new species 
and strains from sect. Quelusia have been recently 
introduced into cultivation. All the taxa recognized 
in this treatment, except F. bracelinae and F. 
glazioviana, are now in cultivation, with the most 
complete collections grown at the University of 
California Botanical Gardens at Berkeley and in 
the private garden of Mrs. Drude Reiman-Dietiker 
in the Netherlands. Other important collections 
include those of Mr. J. O. Wright in England, 
members of the different amateur fuchsia societies 
on the west coast of the U.S., and the Nederlandse 
Kring van Fuchsia-Vrienden in the Netherlands. 


SYSTEMATIC TREATMENT 


Taxonomic history. The earliest known spec- 
imens of Fuchsia from South America were col- 
lected in the Magellan Straits around 1690 by 
George Handisyd, who was surgeon aboard the 
British ship Modena (Middleton, 1909). These col- 
lections correspond to F. magellanica and are 
deposited in the eighth volume of Hans Sloane's 
herbarium at BM. Louis Feuillée published the first 
description and illustration of this species under 
the name Thilco (Feuillée, 1725), but it was not 


taken up by Linnaeus, who described Fuchsia in 
1753 in Species Plantarum. More specimens of 
Fuchsia were collected in the Magellan Straits in 
January of 1768 by Philibert Commerson, natu- 
ralist on the French survey expedition led by Louis 
de Bougainville. Commerson's specimens were ex- 
amined by Lamarck, who designated them the type 
collection for F. magellanica, which he described 
in 1788. Another early collection of this species 
was made by Ruiz and Pavón between 1778 and 
1788, near Concepción, Chile; they later described 
it as F. macrostema (Ruiz & Pavón, 1 ] 

Domingo Vandelli published the genus Quelusia 
in 1788, based on Brazilian plants. He did not 
name a species, and his description and illustration 
dealt mostly with the flower, making it difficult to 
distinguish which of the Brazilian species he was 
treating. José M. da C. Vellozo, a contemporary 
of Vandelli, described Quelusia regia from Brazil, 
providing locality information and an illustration 
that included leaves and flowers. Due to political 
problems, however, Vellozo's name for the species 
was not published until 1829 (Borgmeier, 1937; 
Carauta, 1973). 

Jonas Dryander described Fuchsia coccinea in 
1789 based on cultivated plants that had been 
introduced in England the previous year. The source 
of his plants remains unclear, however, since Aiton 
(1789) attributed their introduction from Chile by 
a Captain Firth, while Salisbury (1791) stated that 
they were brought from Brazil by Vandelli around 
1787. In reprinting Vandelli's publication of Que- 
lusia, Roemer (1796) added a footnote wherein 
he stated that Vandelli’s genus was the same as 
Dryander's F. coccinea. Meanwhile, Salisbury 
(1791) had already synonymized Quelusia under 
F. coccinea. 

In the first conspectus of the genus, de Candolle 
(1828) designated Quelusia as a section of Fuch- 
sia; by that time, the identity of F. magellanica 
and F. coccinea were already badly confused, and 
de Candolle considered them to be synonymous. 

Collections made in Brazil by Auguste de Saint- 
Hilaire between 1816 and 1822 resulted in the 
description of four new species by Jacques Cam- 
bessédes in 1830. None of these species have been 
maintained in the present treatment, but the name 
F. integrifolia Camb. was long used for F. regia. 

During the 1820s and 1830s, there was a sud- 
den surge of collections of Fuchsia from Brazil and 
Chile, and many plants were shipped for cultivation 
in Europe. Several new taxa, most of them now 
included under F. regia and F. magellanica, were 
described by John Lindley, George Gardner, and 
Karel Presl. During the second half of the 19th 


546 


Annals of the 
Missouri Botanical Garden 


century, Auguste Glaziou made extensive collec- 
tions in eastern Brazil, when this area was off limits 
to most foreign botanists. The rare F. glazioviana 
was described from one of his finds 

During the first half of this century, Curt Brade 
and Ynes Mexia collected fuchsias from the two 
highest peaks of southern Brazil, Pico da Bandeira 
and Itatiaia. Their collections led to the description 
of two local endemics, F. bracelinae and F. cam- 
pos-portoi. 

The leading figure in Fuchsia taxonomy was 
Philip Munz, who published a comprehensive re- 
vision of the genus in 1943. He refined the sectional 
concepts and limited sect. Quelusia to a closely 
knit group of species from Brazil and one from the 
southern, temperate Andes. He recognized five 
species in the section, as well as F. hybrida, which 
covered most of the cultivated hybrids. Two of his 
species were polytypic: F. magellanica had three 
varieties, and F. regia four. 

Since Munz, many new collections have accu- 
mulated, especially from the southern states of 
Brazil, where Balduino Rambo, Raulino Reitz, Rob- 
erto Klein, and Gert Hatschbach have made sig- 
nificant contributions. Their collections have led 
me to recognize four new taxa of Fuchsia from 
the south of Brazil in this treatment. During two 
field trips in 1985 and 1987, I was able to collect 
and study field populations of all the Brazilian taxa. 


In distinguish- 
ing species within the section, I attempted to rec- 
ognize basic, morphologically definable units that 
are also correlated with geographical, ecological, 
or cytological discontinuities between populations. 
It is assumed that these units also reflect the evo- 
lutionary divergences that have taken place in the 
group, but such hypotheses must be subjected to 
much more rigorous testing methods before they 


Criteria for taxonomic rank. 


can be substantiated. 

Subspecific rank was used when a wide-ranging 
species exhibited geographically or ecologically dis- 
tinct morphological variants, but these were linked 
by intermediate populations along localized areas 
of sympatr 

Although Jiri varieties have been recog- 
nized in the past for some species of Fuchsia, there 
is little biological basis for recognizing varieties. 
Similar pale color variants, for example, have ap- 
peared independently in a series of geographically 
distinct populations of F. magellanica, and they 
are probably the result of a single, recessive gene 
mutation (Chaudhuri, 1956). I do not consider this 
kind of variation worthy of formal taxonomic rec- 
ognition in a biosystematic revision such as this, 


although I do recognize the utility of assigning 
names to such plants for horticultural purposes. 

In widespread species, such as F. regia and F. 
magellanica, it has not been feasible to distinguish 
the genetic basis for the many minor variants that 
exist from environmentally induced variations or 
inherent plasticity in the plants. Most likely, ge- 
netically determined variations will best be rec- 
ognized through careful observation of plants cul- 
tivated under uniform growing conditions. In this 
way, amateur and professional nurserymen can 
make valuable contributions to our taxonomic un- 
derstanding of sect. Quelusia in the future, espe- 
cially if they maintain well-documented records of 
as porch of their stock and the constancy 
patterns of the different characters. 

E view of the uncertain phylogeny of the species 
of sect. Quelusia, they are listed as follows in 
alphabetical order. 


Fuchsia section Quelusia (Vandelli) de Candolle, 
Prodr. 3: 36. 1828. Quelusia Vandelli, Fl. 
Lusit. Brasil., 23, fig. 10, 1788; pod 
Script. Pl. Hisp. Lusit. Brasil., 101, tab. 7, 
fig. 10. 1796. LECTOTYPE: Fuchsia coccinea 
Dryander. Quelusia was published without the 
protologue naming a species, but Vandelli's 
original text and engraving were reproduced 
by Roemer, who stated in a footnote that the 
description corresponded to F. coccinea. Salis- 
bury (1791) and de Candolle (1828) treated 
F. coccinea as the type of Quelusia. There- 
fore, Munz's (1943) choice of F. magellanica 
as the lectotype of Quelusia and Berry's (1982) 
designation of F. regia as the lectotype cannot 
be upheld. 


Erect, scandent or climbing shrubs, subshrubs, 
or lianas. Leaves opposite or whorled. Flowers her- 
maphroditic, pendulous and axillary near the branch 
tips. Sepals normally reddish, longer than the floral 
tube and connate at the base. Petals purple, strong- 
ly convolute, erect, shorter than the sepals. Sta- 
mens erect, exserted beyond the sepals, the anti- 
sepalous filaments longer than the antipetalous ones. 
Pollen grains triporate or occasionally 4-porate, 
with segmented-beaded viscin threads. Nectary ad- 
nate to the base of the floral tube, smooth or ridged. 
Stigma clavate to ei ded obscurely 4-cleft at 
the apex. Berries with ca. seeds. Gametic 
chromosome number n — 22, 


Distribution (Fig. 1). Southeastern Brazil, from 
Minas Gerais to Rio Grande do Sul, and the Andes 


Volume 76, Number 2 
1989 


Berry 547 
Fuchsia sect. Quelusia 


of southern Chile and Argentina from 33% to 55°S 
latitude 


Etymology. The name Quelusia was given by 


Vandelli after Queluz, the summer palace of Por- 


KEY TO THE SPECIES OF FUCHSIA SECTION QUELUSIA 


tuguese royalty located outside of Lisbon (Don, 


Common names. Chile: chilco, chilca, palo 
blanco, fucsia, tilco. Brazil: brinco de princesa. 


: Leaf measurements are m on mature, fully expanded leaves. Dimensions are intended to cover the full 


iude ol dry and fresh specimen 


ce pilos 


— 


a. Petioles mostly 1-3 m 


2a. Leaves elliptic, with acute to aed base, pilose on all veins below; pedicels hanging, 12-20 m 
l 


on 


mm 
bracelinae 


g : 
2b. Leaves ovate, with subcordate base, pilose er s on lower % of midvein below; pedicels divergent 
18-50 mm lon 9. Ece 

m long, glabrous to pilose 

Sepals connate at the base (from the point of petal insertion) for 2 8 mm and 


from stems and pendent only in distal V5, 
Petioles mostly more than 3 mm 
3a. 


T 


lengt 


coccinea 


> V4 of their total 


4a. Petioles Ed 4-8 mm long, leaves 10-25 mm wide, upper surface lightly sulcate-nerved; stipules 


. brevilobis 


nconspicuo 
4b. Petioles mostly 10-25 mm long, leaves 25-45 mm wide, upper surface smooth; stipules crassate 
9. 


and pers 


a subsp. serrae 


w 
c 


. Sepals ae at nm base for 2-7 mm and - %4 of their total length 


5a. Leaves 2-6 mm wide, narrowly lanceolate; floral tube subrhombic, 4-6 mm long „uuu 


campos-portoi 
5 


Fic 
5b. Leaves > 8 mm wide, less than 5 times as long as wide; floral tube cylindrical- la, 5-3 


m ion 


6a. bee 15-30 mm long, 8-15 mm wide, mack grouped on branches with internodes 3- 


12 mm long; floral tube 5-7 mm long and glabro 
Leaves longer or wider than above, internodes > 12 mm long; or floral tubes pubescent or 


6b. 


c 


> 7 mm long. 


. glazioviana 


7a. Petioles < 6 mm long, leaf blade > 8 times as long as the petiole, leaves mostly opposite. 


8a. Plants glabrous, leaves 8-15(-30) mm wide; floral tube 10-15 mm lon 


7. F. hatschbachü 


8b. Plants densely pilose, leaves 20-70 mm wide; floral tube 5-9 mm long ..................... 


-J 
c 


eaves ternate or whor 


. Petioles (3-)5-35 mm long, leaf blade < 6 times as long as the petioles, some or 
ed. 


. F. alpestris 
all 


9a. Mostly erect shrubs; leaves membranous, usually glabrous, 15-60 mm long; margin 


serrulate and often shallowly lobed between 
mm a berry narrowly oblong, 4-7 mm thick when ripe; 


tube mostly 2-3.5 
southern Chile and Argentin 


Ko) 
¡al 


pilose, 20-140 m 


ripe; Brazil 


teeth; petioles 0.3-1 mm thick; floral 


. magellanica 


. Mostly scandent shrubs or eem leaves membranous to coriaceous, oo to 

mm long; margin entire to gland-s 

floral tube 3-7 mm wide; berry oblong, ellipsoid, or globose, 9-13 mm thick when 
9. F 


m thick; 


serrate; petioles 


. regia 


l. Fuchsia alpestris Gardner, Bot. Mag. 69: 
tab. 3999. 1843. TYPE: Brazil. Rio de Janeiro: 
Organ Mountains, “in moist, bushy, rocky 
places," 1,570 m, Mar. 1841, George Gard- 
ner 5706 (holotype, OXF; isotypes, BM, 
CGE—-2 sheets, G, GH, K—2 sheets, NY, 
P, UC, W). Figure 4. 


Fuchsia mollis Krause, Engl. Jahrb. 37: 600. 1906. 
TYPE: Brazil. Rio de Janeiro: in forest near Nova 
Friburgo, 900 m, Oct. 1896, E. Ule 4418 (holotype, 
B, destroyed in World War Il, photograph at F; 
isotypes, CORD, R— #41706, RB). 

Scandent shrubs 1-5 m tall, branches generally 
long and hanging, up to 5 m long. Branchlets 
reddish tan, densely pilose with erect, whitish hairs 


ca. ] mm long. Leaves opposite, occasionally ter- 
nate, membranous, oblong, broadly ovate or nar- 
rowly elliptic-ovate, (55-)60-140 mm long, 20- 
70 mm wide, apex acute to acuminate, base 
rounded to subcordate or subauriculate, moderate- 
ly pilose above, densely pilose below, sometimes 
purple-flushed when young, often yellowing before 
dehiscing; margin entire to subdenticulate, second- 
ary veins 5-9(-10) per side. Petioles densely pu- 
bescent, 3-6 mm long, stout and 2-4 mm thick 
on older leaves, canaliculate above. Stipules nar- 
rowly triangular, 1 -2 mm long, 0.8-1.5 mm wide, 
adjacent ones sometimes fused, generally covered 
by hairs. Flowers few and solitary in upper leaf 
axils; pedicels pendulous, 25-40 mm long. Ovary 


548 Annals of the 
Missouri Botanical Garden 


ii ea ef Aly 


E 4. Fuchsia alpestris. —a. Flowering branch. —b. Details of entire and longitudinally split flower. — c. 
Pair of large leaves from basal portion of stem.— d. Detail E young stem with stipules and petioles. —e. Tip of 
flowering shoot with nearly sessile leaves. From Berry et al. 4418. 


Volume 76, Number 2 
1989 


Berry 
Fuchsia sect. Quelusia 


@ F. coccinea 
BB F. bracelinae 

(9 F. alpestris 

(€ F. glazioviana 

A r. campos - portoi 
Ak F. brevilobis 

O F. hatschbochii 


45* 40* 
r1 L 


FIGUR 


5. Geographical distributions of the Brazilian species of Fuchsia sect. Quelusia, except for F. regia. 


GURE 5. 
Contours higher than 800 m are indicated by the stippled areas. Capital letters are state abbreviations. MG = Minas 


Gerais, ES = 
Grande do Sol. 


subquadrangular, pilose, 6-8 mm long, 2-3.5 mm 
wide. Floral tube cylindrical-fusiform, (5-)6-8(- 
10) mm long, 3-4 mm wide, pubescent outside, 
pilose inside; nectary 4-6 mm high, 1 mm thick, 
irregularly ridged. Sepals 18-26(-28) mm long, 
connate for 4-6 mm at the base, pilose outside; 
free lobes 3.5-4.5 mm wide at base, spreading at 
anthesis. Tube and sepals red to dull pink. Petals 
violet, broadly obovate, 10-14 mm long, 8-9 mm 
wide. Filaments red, 26-35 mm and 24-32 mm 
long; anthers red, oblong, 2.5-3.5 mm long, 1.2- 


Espirito Santo, RJ — Rio de Janeiro, SP — Sáo Paulo, PR — Paraná, SC = Santa Catarina, RS — Rio 


1.6 mm wide. Style red, pilose in basal V5; stigma 
red, clavate, 2.5-3.5 mm long, 1.1-1.6 mm wide. 
Berry oblong-ellipsoid, 14-16 mm long, 8-10 mm 
wide, purple when ripe; seeds oblong, 1.8-2.2 mm 
long, 1-1.5 mm wide. Gametic chromosome num- 
ber n — 22, 44. 
Distribution (Fig. 5). In thickets and low for- 
est near granitic outcrops of several morros in 
north-central Rio de Janeiro state, Brazil: near 
Nova Friburgo, Santa Maria Madalena, and in the 


550 


Annals of the 
Missouri Botanical Garden 


Serra dos Órgãos between Teresópolis and Petró- 
polis, (1,100-)1,400-1,600 m. Flowers mainly 
from November to March. 

Additional specimens examined. BRAZIL. RIO DE 
JANEIRO: Teresópolis, Bailey & Bailey 1296 (BH), von 
Bayern 19539 (F, NY), von Bayern in Sep. 1888 (M); 

26 km SE of Itaipava, road from Teresópolis to Petrópolis, 
Berry et al. 4418 (MO, RB; n = 22); Fazenda Portugal, 
adiós dos Ór rgàos, Brade 19467 (RB); nis t cy Su- 

are II, Teresópolis to Nova Friburgo, Braga 1535 (RB); 


me .A. CALIFORNIA: tl 
plant, A 2575 (RSA). T 

S: "Hollands che Rading, Berry & Bra o 001- 
86 (MO; n = 44). Also seen cultivated in England and 
France. 

Munz (1943) treated Fuchsia alpestris as a 
variety of F. regia, but the former can be readily 
distinguished from F. regia by having large, mostly 
opposite, short-petioled leaves. The older leaves 
have a characteristic subcordate base, and the en- 
tire plant has characteristic dense, pale pubes- 
cence. The flowers are considerably smaller and 
often much paler than those of the subspecies of 
F. regia that occurs in the same area. 

This species is rare in nature and is apparently 
confined to shrubby patches bordering exposed rock 
on the flanks of large, granitic domes. Fuchsia 
regia subsp. regia always occurs nearby, and some 
intergradation between the two species has been 
found at Pedra Dubois, near Santa Maria Madalena 
(see Appendix). 


2. Fuchsia bracelinae Munz, Proc. Calif. Acad. 
Sci., Ser. 4, 25: 7, pl. 1 2. 1943. TYPE: 
Brazil. Espírito Santo: Serra do Caparaó, rocky 
open campo, 2,650 m, 25 Nov. 1929, Ynes 
Mexia 4013 (holotype, GH, photograph at 
BH; isotypes, A, BM, S, G—DEL, GH, 
MO, NY, P, S, UC, US, Z). Figure 6. 


Few- to many-branched subshrubs 10-60 cm 
tall, or occasionally scandent shrubs to 2 m high 
in dense shrubbery. Branchlets terete, generally 
reddish purple, densely pilose with erect, whitish 
hairs ca. 1 mm long; older branches with freely 
exfoliating bark. Leaves 3-5-verticillate, membra- 
nous, narrowly elliptic-lanceolate, 20-50 mm long, 
7-17 mm wide, apex narrowly acute, base round- 
ed, green and slightly strigose above with conspic- 
uously impressed veins, paler and usually purple- 
flushed below and densely pilose on the sulcate 
veins and margins; margin subentire to gland-den- 
ticulate, secondary veins 4-7 per side. Petioles 


green to reddish, pilose, 1-3 mm long. Stipules 
lanceolate, with a thick base, 1.5-2 mm long, 
adjacent ones often fused basally, subpersistent. 
Flowers solitary in upper leaf axils; pedicels slender, 
pilose, pendulous, 12-20 mm long. Ovary oblong, 
strigillose to pilose, 4.5-8 mm long, 2-3 mm wide. 

loral tube cylindrical-fusiform, 3.5-7 mm long, 

-4 mm wide, lightly strigillose to pilose outside, 
glabrous inside; nectary 2.5-3.5 mm high, ca. 1 
mm thick and slightly ridged. Sepals 19-26 mm 
long, narrowly lanceolate to elliptic-ovate, connate 
at the base for 3-5(-6) mm; free lobes 5-8 mm 
wide at the base, apex acuminate, spreading at 
anthesis. Tube and sepals light red to pink. Petals 
deep violet, broadly obovate, 10-15 mm long, 6.5- 
9 mm wide, apex rounded. Filaments reddish pur- 
ple, 24-30 mm and 17-21 mm long; anthers 
elliptic, 2.5-3 mm long, 1.2-1.6 mm wide. Style 
light red, glabrous; stigma reddish, clavate to sub- 
globose, 1.5-3 mm long, 1-1.5 mm wide, exserted 
5-14 mm beyond the anthers. Young fruits oblong, 
mature ones not seen. Gametic chromosome num- 
ber n — 


Distribution (Fig. 5). Endemic to the highest 
peaks of the Serra do Caparaó (Pico da Bandeira, 
Pico do Cristal, Pico do Cruzeiro, and Pico do 
Calçado) along the border of the states of Minas 
Gerais and Espirito Santo. Grows in open campo 
above treeline, between 2,280 and 2,850 m, most 
common at 2,700-2,800 m. Flowers in the sum- 
mer, principally from November until March. 


Additional specimens examined. BRAZIL. ESPÍRITO 
SANTO: Pico da Bandeira, Berry 4524 (MO, Bh n= 22), 
4530, 4532, 4533, 4534, 4535 (MO, RJ; n = 22); Irwin 
2745 (F, MICH, NY, R, TEX, UC). MINAS GERAIS: Pico 
da Bandeira, Berry 4525 (MO, RJ), 4531 (MO, RJ; n 
= 22), Campos de Caparaó, Ule in 1890 (R 
H41703); Schwacke in 8 Feb. 1890 (R #41702); Serra 
do Caparaó, Torgo 6 (HB, MBM). 


Fuchsia bracelinae is distinguished from other 
members of the section by having dense, stiff pu- 
bescence and small, narrow, verticillate leaves. It 
inhabits only the rocky, open campos of the highest 
peaks of southern Brazil, in the Serra do Caparao. 
Individuals are found along rocks, in meadows, or 
in protected shrub and bamboo patches. This area 
is subject to frequent and severe winter frosts, and 
F. bracelinae is found to within 60 m of the 2,897- 
m-high summit of Pico da Bandeira. 
winter dieback of the above-ground stems probably 
accounts for the small stature of most plants, but 
underground stems lead to extensive vegetative 
reproduction in areas with good soil accumulation. 

Fuchsia regia occurs lower down on the same 
mountain range, but there is no sympatry at pres- 


Extensive 


Volume 76, Number 2 551 
1989 


Berry 
Fuchsia sect. Quelusia 


FIGURE 6. Fuchsia bracelinae. —a. Detail of flower in longitudinal section. —b. Petal, spread open.— c. Floral 
bud just prior to anthesis.—d. Flowering branch.—e. Leaf whorl and detail of stipules and petioles. From Berry et 
l. 4524. 


ent; extensive forest areas at intermediate eleva- m) has narrow, moderately dentate leaves that re- 
tions existed until the past century but have been semble F. campos-portoi, but the Irwin collection 
eliminated by cutting and fires. The collection /rwin is probably just a stunted individual of F. brace- 
2808 (US #2324661; Serra do Caparaó, 2,600  linae. 


552 


Annals of the 
Missouri Botanical Garden 


RE 7. Fuchsia brevilobis. —a. Flowering branch. —b. Leaf Er = Ripe fruit.—d. Detail of entire and 
longitudinally split flower. —e. Petal, spread open. From Berry et al. 4 


3. Fuchsia brevilobis P. Berry, sp. nov. TYPE: 
Brazil. Sao Paulo: 38 km NE of the Sao Paulo- 
Paraná border near the 267 km post on High- 
way 116, bamboo forest on marshy ground, 
680 m, 8 Mar. 1976, Gerrit Davidse, T. P. 
Ramamoorthy & D. M. Vital 10902 (holo- 
type, MO #2931564; isotype, SP (under Da- 


vidse & D'Arcy 10902); n — 
Figure 7. 


22, 2n = 44. 


Frutex scandens, ramulis foliisque junioribus puberu- 
lentibus vel pilosis. Folia membranacea, lanceolata vel 
elliptico-ovata, basi rotundata, apice acuta, 2-7.5 cm 
longa, 1-3 cm lata, nervis secundariis 3-6; petiolis 
(2-)4-8(-14) mm longis. Tubi florales fusiones 7-10 


Volume 76, Number 2 
1989 


Berry 553 
Fuchsia sect. Quelusia 


mm longi. Sepala 17-25 mm "is basi (8-)11-16 mm 

connata, lobis liberis 5-11(-13) mm longis. Petala spa- 

thulata 12-18 mm longa, 5-7 mm lata, a tubo sepalorum 

fere occulta. Numerus gameticus chromosomatum n — 
22. 


Scandent shrubs 2-6 m tall, or occasionally 
lianas in trees to 8 m high, with basal stems 1-6 
cm in diameter; branches scandent or drooping to 
3 m long. Young growth densely pilose to puber- 
ulent, older stems exfoliating. Leaves opposite, ter- 
nate or occasionally in whorls of 4, firmly mem- 
branous, lanceolate to narrowly ovate, 20-75 mm 
long, 10-25(-30) mm wide, acute to occasionally 
acuminate at the apex, rounded at the base, subgla- 
brous to sparsely pubescent above with impressed 
veins, sparsely puberulent to villous below, espe- 
cially along nerves; margin entire to remotely den- 
ticulate, secondary veins 3-6 per side. Petioles 
usually dull purple, (2-)4-8(- 1 4) mm long, subgla- 
brous to densely pilose. Stipules 0.5-1.1 mm long, 
0.7-1.2 mm wide, purplish, broadly triangular, 
thick-stubby and divergent, deciduous. Flowers sol- 
itary in upper leaf axils; pedicels pendulous, 17- 
28(-42) mm long. Ovary oblong, 5-10 mm long, 
2-4 mm wide, loosely pilose, green. Floral tube 
subcylindric to fusiform, 7-10 mm long, 2-3 mm 
wide at the base, 3.5-4.5 mm wide in the middle, 
often constricted to ca. 3 mm wide at the point of 
insertion of the stamens beneath the widening of 
the sepal tube, glabrous to pilose outside, pilose 
inside; nectary green, slightly ridged, 1-1.2 mm 
thick and 2.5-3 mm high. Sepals 17-25 mm long, 
connate for (8—)1 1-16 mm at the base and forming 
a tube, this enlarging to 8-10 mm wide before the 
separation of the sepals; the free lobes narrowly 
triangular, 5-11(-13) mm long, 3-5 mm wide at 
the base, acute to acuminate at the apex, spreading 
to slightly recurved at anthesis. Tube and sepals 
red to red-pink. Petals almost entirely enclosed by 
the sepal tube, light to dark purple, spathulate, 
12-18 mm long, (5-)6-7 mm wide, rounded at 
the apex, the base slender and adnate to the sepal 
tube in the basal 3-4 mm above the insertion of 
the filaments. Filaments reddish pink, 22-38 mm 
and 19-35 mm long; anthers red-purple, oblong, 
2.5-3 mm long, 1-1.5 mm wide. Style reddish, 
pilose in lower half; am clavate, purple, 1.2- 
1.6 mm long, ca. wide, exserted 5-10 mm 
beyond the anthers. Berry e ellipsoid, 15- 

22(-26) mm long and - mm wide at 
maturity, black when ripe; seeds a 
2-2.6 mm long, 1-1.6 mm broad, ca. 0.5 mm 
thick. Gametic chromosome number n — 22. 


Distribution (Fig. 5). In montane forest of the 
Serra de Paranapiacaba in southern Sào Paulo 


state; in Paraná state in the Serra da Virgem Maria 
and Rio Capivari drainage, and then on the low- 
ermost slopes of the Serra do Mar and in coastal 
restingas, | to 900 m. Flowers 
throughout the year, principally from November 


from sea leve 


Additional specimens examined. BRAZIL. SÀO PAULO: 
Miguel Arcanjo & 


ot 
Sete Barras, Prance et al. 6860 (MBM, SP, UEC); Re- 
erva Capào Bonito, m S of Itapetininga, Sakane 
543 (SP); 50 km S of Itapetininga, Gibb 3279 


Apiai-Sao Paulo, km 294, Vianna in 5-5-64 (ICN #2820). 
PARANA: Prainhas, Mun. Morretes, Hatschbach et al. 
13412 (MBM, U); Cordeiro et al. 215 (MBM; n = 22); 
below Eng. Lange train station to Prainhas, Cordeiro et 
al. 218 (MBM; n = 22); Porto da Cima, Mun. Morretes, 
Berry et al. 4495 (MBM, MO; n = 22); Dusén 11964 
(S), 14133 (G— DEL, K, S, US), 14340 (A, Sy; Hatsch- 
bach 43303 (MBM); Sesmaria, 2- à km W of Rio Ca- 
pivari, Mun. Bocaiúva do Sul, Berry & Juarez 4441 
(MBM, MO; n = 22), 4442, 4443, 4444, 4445 (MBM, 
MO); Hatschbach 20936 (HB, MBM, UC); Hatschbach 
& Ramamoorthy 42973 (MBM, MO), 42974 (MBM); 
10 km E of Curitiba, Lindeman & Haas 2539 (MBM, 
U); Estrada da Graciosa, Serra do Mar, 850 m, Lindeman 
& Haas 5879 (U); road Rio Taquary-Rio Divisa, Mun. 
Campina Grande do Sul, Hatschbach 7009 (MBM); Serra 


Hatsehtach 7056 (MB); Rio Sáo Joàozinho, Mun 
ranaguá, Berry et al. 4496 (MBM, MO); Falkenberg 
2234 (ICN); Sapitanduva (cultivated), Berry et al. 4494 
MBM, MO); Falkenberg 2222 (FLOR); Mun. DE 
Cacatu, Hatschbach et al. 50788 (MBM; n = 22); Mur 
Campina Grande do Sul, oda BR-2, Ribeirào do Ced. 
ro, Hatschbach 8946 (MBM). 


~ 


Fuchsia brevilobis is distinguished by its re- 
markably long sepal tube, which almost entirely 
encloses the petals, and by the membranous, pu- 
bescent, narrowly ovate leaves. There are two dis- 
tinct areas where this species occurs, with notable 
morphological differences in their populations. In 
the low ramifications of the Serra do Mar in north- 


bescent, with especially long sepal tubes. Another 
group of populations is found on the lower, eastern 
flanks of the Serra do Mar in central Paraná, from 
400 m to sea level. Most plants from these pop- 
ulations have longer petioles than those of higher 
altitude, and they vary considerably from popu- 
lation to population in color (dull pink to bright 
red), degree of sepal connation, and leaf shape and 
texture. 

Several populations of F. brevilobis occur in the 
restinga or swamp forest close to sea level, a very 


554 


Annals of the 
Missouri Botanical Garden 


FIGURE 8. 
stipules, and a longitudinally divided flower. From Berry et al. 4435. 


unusual habitat for Brazilian fuchsias. These plants 
grow as lianas and are found along rivers or streams, 
perhaps enabling them to withstand the heat and 
high evapotranspiration of the Atlantic lowlands. 
The population from Cacatu has unusually narrow, 
lanceolate leaves with acuminate tips. 
Throughout most of its range, this species is 
sympatric with F. regia subsp. serrae. The pres- 
ence of plants morphologically intermediate be- 
tween the two taxa indicates that hybridization has 
occurred between them (see Appendix). The pop- 
ulations of F. regia subsp. serrae in northern Pa- 
raná have exceptionally long sepal tubes, which 
may be the result of introgression with F. brevi- 


lobis. 


4. Fuchsia campos-portoi Pilger & Schulze, 
Notizbl. Bot. Gart. Berlin-Dahlem 12: 470. 
1935; Rodriguesia 2: 94, figs. 1-5. 1935. 
TYPE: Brazil. Rio de Janeiro: Itatiaia, Serra da 
Mantiqueira, 2,200 m, 27 Dec. 1934, Robert 
Pilger & Curt Brade 31 (holotype, B, de- 
stroyed in World War II; lectotype, RB, here 
designated; isotype, BH). Figure 8. 


Fuchsia campos-portoi. Flowering shoots with details of an individual leaf underside, a leaf node with 


Subshrubs to woody scandent bushes 0.3-2 
(-3) m tall, often growing among rocks. Young 
growth subglabrous to finely canescent or pilose; 
older stems 1-4 cm thick, with flaky, coppery 
brown bark. Leaves mostly ternate, occasionally 
opposite or 4—5-whorled, firmly membranous, nar- 
rowly elliptic-lanceolate, 12-40 mm long, 2-6(-8) 
mm wide, narrowly acute at the apex, acute to 
cuneate at the base, dark green and subglabrous 
above, paler and mostly glabrous below or with 
villous hairs along veins and margins; margin gland- 
serrulate, with teeth angled towards the apex, sec- 
ondary veins 3-7 per side, at times inconspicuous; 
internodes short, mostly 10-16 mm long. Petioles 
(2-)3-8 mm long. Stipules 0.5-1 mm long, nar- 
rowly lanceolate or filiform, deciduous. Flowers 
solitary in upper leaf axils; pedicels slender, pen- 
dulous, (4-)8-20 mm long. Ovary oblong, 4-5 
mm long, ca. 2 mm wide, strigose to strigillose. 
Floral tube subrhombic, 4-6 mm ong, 3.5 mm 
wide in the middle and ca. 2.5 mm wide at the 
base and summit, glabrous to strigose outside, gla- 
brous inside; nectary 2.5-3 mm high, shallowly 
8-lobed. Sepals 12-20 mm long, lance-elliptic, con- 


Volume 76, Number 2 
1989 


Berry 555 
Fuchsia sect. Quelusia 


nate for 3-5 mm at the base; free lobes 4-7 mm 
wide at base, spreading at anthesis. Tube and sepals 
red to dark pink. Petals violet, broadly obovate, 
9-12 mm long, 7-8 mm wide, rounded to slightly 
retuse at the apex. Filaments red, 15-28 and 13- 
25 mm long; anthers purple-red, elliptic-oblong, 
1.5-2 mm long, 1-1.5 mm wide. Style glabrous, 
exserted 5-10 mm beyond the anthers; stigma 
clavate, 1.2-2 mm long, 0.7-0.9 mm wide. Berry 
cylindrical-oblong, 14-16 mm long, 7-8 mm thick; 
seeds 1.3-1.6 mm long, 0.7-1.1 mm wide. Ga- 
metic chromosome number n — 22. 

Distribution (Fig. 5). 
campos of the Itatiaia mountain massif, in the Serra 
da Mantiqueira, Rio de Janeiro and Minas Gerais 
border, from 2,100 to 2,550 m. Flowers mainly 
during the summer, from November to March. 


Endemic to the open 


Additional iip ie examined. BRAZIL. RIO DE 
JANEIRO: Ayuruoca, Glaziou 15948 (K, P); Itatiaia (Plan- 
alto), Andrade E Emerich 626 (R) 
mundo 826 (MO, R 
et al. 4435 (MO, RB; n = v 4436, 4438 (MO, RB), 
Brade 15160 m RB), 15676 (RB, POM, photos at NY, 
R, UC), 18008 (R, RB, ru 0335 (R); Campos Porto 
2705 (R, RB), yaks 826 (US); Dusén 178(R); Gounelle 
in Jan. 1899 (G—BOIS); Glaziou 4801 (P), 6522 (K 


bates 


n N v. 1958 
Sampaio in n Feb. 1954 (R); east 
25591 (M); t A. (HB, K, M, MBM); P 
7037 (HB, EL) Pereira & Pereira 7578 (B, 
HB, R, i Ramamoorhy 104 (MO), 814 (MBM, MO), 

s.n. (MO; 22); Santos 5169 (DS); Sucre 4645 (MO, 
RB); Ule 118 (R): Vianna 771 (R). 


Fuchsia campos-portoi is readily distinguished 
from other fuchsias by its small and very narrow, 
gland-serrulate leaves. It also has unusual, small 
flowers; the floral tube is nearly rhombic and very 
short, whereas the sepals are up to four times as 
long and are noticeably inflated in bud. 

This species inhabits the high-elevation campos 
of the Itatiaia massif, where it grows in shrubby 
patches or in open, rocky sites. Winter frosts are 
common in this area. Populations of F. regia subsp. 
regia grow together with F. campos-portoi, and 
several intermediate plants have been found that 
indicate the occurrence of local hybrids (see Ap- 
pendix). 


9. Fuchsia coccinea Dryander in Aiton, Hort. 
Kew., 1st edition, 2: 8. 1789. Sims, Bot. Mag. 
tab. 97. 1789. Mordant de Launay, Herb. 
Gen. Amat. 2: 89, fig. 1817. Hook., Bot. Mag. 
94: tab. 5740. 1868. Nahusia coccinea 


(Dryander) Schneevoogt, Icon. Pl. Rar. 1: 21. 
1792. Fuchsia elegans Salisbury, Ic. Stirp. 
Rar., 13, tab. 7. 1791, nom. illeg. (Salisbury 
thought he would avoid confusion in renaming 
F. coccinea, since all species known at the 
time had red flowers.) Fuchsia pendula Salis- 
bury, Prodr., 279. 1796, nom. illeg. (based 
on F. elegans Salisbury). TYPE: From a plant 
cultivated in 1788 at Kew Gardens, London, 
England, **Oct. 1788, Hort. Kew" (lectotype, 
LINN #670.3, Smith Herbarium, here des- 
ignated: see Fig. 9; isolectotype, BM). Figures 


Fuchsia montana Cambessédes in x pri a l. Brasil. 
Merid. 2(17): 275, tab. 135. TYPE: Brazil. 
Minas Gerais: near the summit a oe Sen a da Ca- 
raca, Dec. 1816-Mar. 1818, Auguste St. Hilaire 
B1 449 (holotype, P; isotype, P). 


Erect shrubs 0.5-1.5 m tall, or scandent to 7 
m high, often with extensive underground stems. 
Young growth puberulent to densely pilose, older 
stems 5-20 mm thick, with exfoliating bark. Leaves 
mostly ternate, sometimes opposite or in 4s, mem- 
branous or rarely subcoriaceous, narrowly ovate 
to ovate, 15-55 mm long, 7-25 mm wide, acute 
to subacuminate at the apex, subcordate or some 
times rounded at the base, generally dull, light 
green, paler below, sometimes strongly purple- 
flushed, subglabrous to puberulent on both sides, 
but generally densely pilose along the basal V$ of 
the midvein below; margins serrulate or occasion- 
ally subentire, secondary veins 4—6 per side. Pet- 
ioles short and stout, ca. 1 mm thick, 1-3 mm 
long, occasionally longer on basal leaves, usually 
densely pilose or hirsute. Stipules narrowly lan- 
ceolate, 0.8-1.4 mm long, deciduous. Flowers sol- 
itary in the upper leaf axils; pedicels slender, pu- 
bescent, 18-42(-50) mm long, divergent or arching 
from the stems and pendent only in the distal third. 
Ovary oblong-ellipsoid, 5-9 mm long, 2.5-3 mm 
thick. Floral tube fusiform, 5-10 mm long, 2.5- 
3 mm wide at the base, 3.5-4.5 mm wide in the 
middle and 3-4 mm wide at the top, subglabrous 
to puberulent outside, puberulent inside; nectary 
2-3 mm high. Sepals 15-24 mm long, 
oblong-lanceolate, lightly pubescent, connate at the 
base for 4-7 mm; free lobes 3.5-6 mm wide, acute 
at the apex, spreading at anthesis. Tube and sepals 
red to dark pink. Petals violet, obovate, 7-10 mm 
long, 6-8(-9) mm wide. Filaments red-purple, 18- 
38 mm and 13-30 mm long; anthers purplish, 2— 
2.5 mm long, ca. 1.5 mm wide. Style puberulent, 
red; stigma clavate, 1.5-2 mm long, exserted 5- 
15 mm beyond the anthers. Mature berry ellipsoid, 


smooth, 


556 


Annals of the 
Missouri Botanical Garden 


14-17 mm long, ca. 12 mm thick, purple; seeds 
oblong-triangular, 1.4-1.9 mm long, 0.8-1.2 m 


wide. Gametic chromosome number n = 22. 


Distribution (Fig. 5). 
campo near the summits of several of the highest 
mountains in Minas Gerais: Serra da Piedade, Serra 
da Caraga, Serra do Itacolomi, and Pico do Itambé, 
from 1,400 to 2,000 m. Flowers throughout the 
year, mainly from November to March. 


Occurs in rocky, open 


Rura Me de examined. BRAZIL. MINAS 
AIS: E slopes of Pico do Itambé, Serra do Espinhaço, 
Ande 'rson et p 35832 (HB, MICH, MO, NY, U, W); 


Serra do Gaviao, Pico do Itambe, Windisch & Gillamy 
183 (HB); ascent E Nun :0 do Itambé, Mun. San António 
do Itambé, 18°24’ UW, Furlan et al. 3058 (SPF); 
Serra da Piedade, p e n aete, rey Herb. #21373 
21374 (OUPR); vd et al. 4553 (MO, RB; n = = 29), 
4554, 4555, 4556, 7 (MO, SEA 4558 (MO, RB; n 
— 22); Glaziou in e pos 1893 (Py; Hoehne 6278, 
6281 (POM), 6279, 6280 (R); Irwin et al. 28777 (K, 
den. MO, NY, RB, US), 30326 (MO, NY, RB, U, 
US); Landrum 4275 (MBM); Magalháes Gomes 1451 
(R); Mello Barreto 7158, 7162, 8803 (F, BHMH), 7159 
(BHMH, R), 7160 (F), 7161 (BHMH); vicus & Pabst 
2662 (MO); St. Hilaire 2276 (P); Warming in 1907 
(P) collector unknown EO in Non 1839 
unknown in Sep. 
Pico Itacolomi, Badint 
boza 1056 (R); Bello 249 (R): iet et al. 4547, 4548 
(MO, RB); E io ed G — BOIS, HB, RB); Falken- 
berg 3381, 33 3385, 3386 (FLOR); Glaziou 
14703 (K, LE); a js 19001 (POM Magalháes Gomes 
3027, in 28 June 1896, in 20 Nov. 1893 (OUPR); 
Mello Barreto 9168 (BHMH. F); Morros de Vila Rica 
(Ouro Preto), dar s.n. b^ — 2 sheets); Serra da Ca- 
Martinelli & Tavora 2742 


M, 

, RB), Viger in July 1896 (R). 

No locality: Clauss 44 in Oct. 1839 (G), 88 in 1840 

(BM) Hausmann 267E (RB); Longsdorfin Sept. 1829, 

s.n. (LE); Netto in 1862 Schwacke 13665 (MO, 

RB); Vasconcellos in 25 Jun. 1901 (POM); Vellozo? 
E). 


Cultivated specimens. AZORES: Flores, Bena Mao, 
Pryor 79 (BM); Flores, Ponta Delgada, Gonçalves 4256 
(BM); Pico, Landroal, Goncalves 4436 (BM); Faial, Horta 
Alagoa, Goncalves 2278 (BM); Sao Jorge, Toledo, Gon- 
calves 3630 (BM). CANARY es Tenerife, Orotaoa, 
Burchard 177 (G— DEL, Z). Jav ] 
5824 (US); near Cinchona, ilr 91. 30 (BM), / 
in 16 July 1903 (NY). MabEIRa: Funchal, Bornmüller 

, Z); above São J ge, Pickering 40 PS 
n da Serra, Pond in July 1865- -6 (G, G— BOI 
— DEL). SuMaTRA: Karo Highlands, E o "his 
e & Toroes 6 624 O). U.S.A. KENTUCKY: Short in 
1864 (PH). 


= 


The type sheets of F. coccinea are the earliest 
known specimens of this species, and both were 
taken from the plant at Kew Gardens upon which 
Dryander based his description (Hooker, 1869). 
The source of the plants is unclear; both Sims and 


FIGURE 9. Photograph of the lectotype of Fuchsia 
NN. 


coccinea from the Smith Herbarium at L 


Dryander described the species as a native of Chile, 
introduced to Kew by a Captain Firth. Salisbury 
(1791), on the other hand, stated that Vandelli 
introduced the species from Brazil around 1787. 

Fuchsia coccinea is a rare species known from 
just four of the highest mountain ranges in Minas 
Gerais. It typically grows in rocky, open campo or 
occasionally in forest patches. It has long been 
confused in the literature with F. magellanica, but 
F. coccinea is easily distinguished by its ovate, 
subsessile leaves with a conspicuous tuft of long 
hairs at the base of the lower surface. Its divergent 
or arching pedicels also differ markedly from the 
fully pendent flowers of the other species in the 
section. 

Fuchsia regia subsp. regia is known to occur 
sympatrically with F. coccinea at the Pico do Itambé 
and near the summit of the Serra do Itacolomi, 


Volume 76, Number 2 Berry 557 
1989 Fuchsia sect. Quelusia 
> 
FIGURE 10. Fuchsia coccinea. —a. Flowering branch; note the characteristic arching pedicels. —b. Flower in 


longitudinal section. —c. Ripe fruit, same scale 


stem, show 
4558 


which has been badly disturbed by fires and grazing. 
A series of intermediate variants between the two 
taxa indicates that hybridization has taken place 
in these areas (see Appendix). 


6. Fuchsia glazioviana Taubert, Engl. Bot. 
Jahrb. 15, beibl. 34: 10. 1892. TYPE: Brazil. 
Rio de Janeiro: Alto Macahé de Nova Fribur- 
go, 14 Aug. 1888 (fide specimen at R), 4. 
Glaziou 17614 (holotype, B, destroyed in 
World War II, photograph at F; lectotype, F, 


as (a).—d. Pet 
wing the cordate leaf base and rial tuft of hairs at the base of the abaxial midvein. From Berry et al. 


al, spread open. — air of leaves from the lower 


here designated; isotypes, G, LE, LY, P, R 
#10152). Figure 11 


Fuchsia santos-limae Brade, Arq. Jard. Bot. Rio de Ja- 
neiro 15: 10, fig. 2. 1957. TYPE: Brazil. Rio de 
Janeiro: Santa Maria Madalena, Rochela, 30 May 
1938, J. Santos Lima 405 (holotype, RB #37704; 
isotype, HB #25471, with no collector number but 
same date of 30-V-38). 


Shrubs 0.5-4 m high, often scandent in trees 
or low brush and with decumbent branches to 6 
m long. Branchlets subdivaricate and generally 


558 


Annals of the 
Missouri Botanical Garden 


FIGURE 11. 


densely packed, Honey pea dull purplish, 
readily exfoliating. Leaves opposite or ternate, firmly 
membranous to subcoriaceous, elliptic or lanceo- 
late, 15-30(-40) mm long, 8-15 mm wide, acute 
at the base and apex, dark sublustrous green and 
glabrous above, paler and subglabrous below or 
sometimes pilose at the base of the midvein; mar- 


Fuchsia glazioviana. —a. Flowering branch. 
open. — c. Detail of anther. — d. Detail of stigma. — e. Detail of stipules. —f. Leaf underside. From Berry et al. 4422. 


—b. Flower in longitudinal section and petal spread 


gins remotely gland-denticulate, with 4—5(—6) sec- 
ondary veins per side. Petioles 3-6 mm long, pur- 
plish and sparsely puberulent; internodes short, 3— 
12 mm long. Stipules broadly triangular, 0.6-1.2 
mm long and wide, thick at the base, purplish, 
deciduous. Flowers solitary in the upper leaf axils; 
pedicels slender, sparsely puberulent, pendulous, 


Volume 76, Number 2 
1989 


Berry 559 
Fuchsia sect. Quelusia 


12-26 mm long. Ovary oblong, 4-5 mm long, 2- 
2.5 mm wide. Floral tube cylindrical, 5-7 mm 
long, 2.5-4 mm wide, subglabrous outside, gla- 
brous inside; nectary smooth, 3-4 mm high. Sepals 
17-22 mm long, narrowly lanceolate, connate at 
the base for 4-5 mm; free lobes 3-4 mm wide, 
acuminate at the apex, spreading at anthesis. Tube 
and sepals red or pink. Petals purple, obovate, 9- 
12 mm long, 6-9 mm wide, rounded at the apex. 
Filaments red-purple, 22-32 and 16-28 mm long; 
anthers oblong, 2.5-3.5 mm long, 1.1-1.6 mm 
wide. Style red, glabrous to puberulent; stigma 
clavate, 2-3 mm long and 1-1.4 mm wide, exsert- 
ed 5-20 mm beyond the anthers. Berry shiny dark 
purple, subcylindrical, 10-16 mm long, 5-8 mm 
wide; seeds oblong, 2-4 mm long, 1-1.5 mm wide. 
Gametic chromosome number n — 22. 


Distribution (Fig. 5). 
cloud forest on a few high mountains around the 
towns of Nova Friburgo and Santa Maria Mada- 
lena, in Rio de Janeiro state, from 1,500 to 2,100 
m. Flowers throughout the year, mainly from No- 
vember to March. 


In open campo and in 


Additional specimens examined. BRAZIL. RIO DE 
JANEIRO: Nova Friburgo, Morro da Nova Caledónia, Berry 
et al. 4420 (MO, RB), 4422, 4423 (MO, RB; n = 22), 
4426, 4428, 4431, 4434 (MO, RB); de Lima 530 (RB); 
Martinelli et al. 2540 (RB); Pabst 8134 (HB); Nova 
2 au ces de la Bocaina do Conejo, Glaziou 12667 
(F, K, Y, P) Nova Friburgo, Morro da torre de 
TV, Ai et dl 789 (MO); Santa Maria Magdalena 
Pedra do Desengano, Farney & Caruso 1190 (RB); Gin. 
tos Lima & Brade 13286 (RB, HB #25466, fragment). 


Fuchsia glazioviana is distinguished by its small, 
tightly grouped, shiny, subcoriaceous leaves with 
short petioles. It is a local endemic known from 
only two mountains in the Serra do Mar of Rio de 
Janeiro, where it extends from upper cloud forest 
into open campo above treeline. On the Morro da 
Nova Caledónia, it is sympatric with F. regia subsp. 
regia in cloud forest habitats, and the two species 
hybridize locally (see Appendix). 


el 


. Fuchsia hatschbachii P. Berry, sp. nov. 
TYPE: Brazil. Paraná: Serra Sáo Luiz do Pu- 
runa, BR-277, Mun. Campo Largo, 25 Jan. 
1985, P. E. Berry, T. Plowman & F. Juarez 
4458 (holotype, MO #3504627; isotype, 
MBM; n = 22). Figure 12. 


Frutex erectus vel scandens 1-5 m sa e ple- 
rumque opposita, anguste lanceolato-ovata m lon 
a, 0.8-3 cm lata, apice attenuata vel a a basi 
Vines ann vel anguste — ene nne v^ 7) 
m longis, e. asim laminae; stipulis par- 
membran "x 7 mm longis, 0.6-1.2 mm latis. 

Tubi florales ‘cylindric 10- 15 mm longi, 3-5 mm lati. 


Sepala 18-26 mm longa, 3.5-5 mm lata, basi connata 
2-6 mm. Petala late obovata 12-17 mm longa, 8-12 
mm lata. Numerus gameticus chromosomatum n — 22. 


Erect to scandent shrubs 1—3 m tall or climbers 
in trees to 5 m above ground. Branchlets glabrous, 
dull purple, older branches ascending on sma 
shrubs or decumbent in large or climbing plants. 
Leaves mostly opposite, occasionally in whorls of 
3-4, firmly membranous to subcoriaceous, nar- 
rowly lance-ovate, generally 3 times longer than 
wide, 3.0-7.5(-11.0) cm long, 0.8-2.5(-3.0) cm 
side, apex attenuate to acuminate, base rounded 
to narrowly subcordate, sublustrous dark green 
above, pale below and glabrous on both sides except 
for small tufts of hairs 0.8-1 mm long at the base 
of the lower midvein in some plants; margin re- 
motely gland-denticulate, secondary veins 5-6(-7) 
per side. Petioles wine purple, (2-)3-6(-7) mm 
long, 1-2 mm thick, canaliculate above, at times 
making a marked angle at the junction with the 
midvein. Stipules narrowly triangular, 1-1.7 mm 
ong, 0.6-1.2 mm wide, adjacent ones occasionally 
fused, semideciduous. Flowers glabrous and uod 
in upper leaf axils; pedicels pendulous, (10-)20- 
30 mm long. Ovary cylindrical, 5-9 mm long, 
2.5-3 mm wide. Floral tube cylindrical, 10-15 
mm long, 3-5 mm wide; nectary 3-5 mm high. 
Sepals 18-26 mm long, connate at the base for 
2-6 mm; free lobes 3.5-5 mm wide at the base, 
spreading at anthesis. Tube and sepals red. Petals 
violet, broadly obovate, 12-17 mm long, 8-12 
mm wide. Filaments red, 40-45 mm and 32-40 
mm long; anthers red-purple, 3-3.5 mm long, 1- 
1.5 mm wide. Style red, loosely pilose in lower 
half; stigma 2-3 mm long, 1-1.5 mm wide, exsert- 

5-20 mm beyond the anthers. Berry ed 
lindrical, + 4-angled, 13-18 mm long, 7-10 m 
thick, anis when ripe; seeds oblong- da 
wide. Gametic 


— 


-1.8 mm long, 0.8-1.2 mm 
chromosome number n = 22. 
Distribution (Fig. 5). In low forests on lime- 
stone or sandstone outcrops, in the planalto north 
and west of Curitiba, Paraná, between 950 and 
1,150 m. Flowers mainly from November to March. 


Additional specimens examined. BRAZIL. PARANÁ: 
Serra Sáo Luiz do Puruna, BR-277, Mun. Campo Largo, 
Berry et al. 4459 (MBM, MO; n = 22), 4460, (MBM, 
MO), 4461 (MBM, MO; n = 22), 4462, 4463, 4464, 
4465 (MBM, MO); Brade 19494 (RB); Frenzel in 28 
Feb. 1951 (MBM #6372, MBM #75197); Hatschbach 
7541 (MBM); Hatschbach & Oliveira 42962 (MBM); 
Hatschbach et al. 42960, 42964 (MBM); Kummrow 
1512, 2217 (MBM); Pereira 5193 (F, HB, MB, MO 
RB); Sprea, Mun. Balsa Nova, Hatschbach 42966 (MBM). 

ampinhos, Mun. Bocaiúva do Sul, Hatschbach 1817 
(MBM, PACA); Campina dos Tavares, Mun. Bocaiüva do 


560 Annals of the 
Missouri Botanical Garden 


FIGURE 12. Fuchsia hatschbachii. —a. Flowering branch.—b. Details of leaves (upper surface).— c. Flowers. 
From Berry et al. 4458. 


Volume 76, Number 2 
1989 


Berry 561 
Fuchsia sect. Quelusia 


Sul, jii dad 7776 (MBM): Bacaetava, near cave, 
Mun. Boc ul, Kummrow 1022, 1418 (MBM), 
Ea a al 2998 8 (MBM; n = 22); Bateias, Mun. 
Campo Largo, Hatschbach 40746 (MBM); Tigre, Mun. 
Cerro Azul, Hatschbach 43632 (MBM); Colombo, Fal- 
kenberg 2215 (FLOR); Manancial da Serra, Braga 550 
(RB, SP); Acungui, Mattos & Labouriau in 1 Mar. 1948 
(RB) Paredáo, Mun. Jaguariaiva, Hatschbach 31130 


eira, 

Oliveira 299 (MBM); Curiola, Mun. Rio Branco do Sul, 
Hatschbach 16155 (MBM); Madre, Mun. Rio Branco do 
Sul, Hatschbach 7774 (MBM); Serra do Caete, Hatsch- 
bach 40698 (MBM); Mun. Palmeira, BR-277, Rio Pa- 
pagaio, Hatschbach 50824 (MBM). 

Fuchsia hatschbachii is most likely to be con- 
fused with F. regia but differs by having shorter 
petioles, narrow leaves with somewhat curved pet- 
ioles, shortly connate sepals, and small stipules. It 
is tetraploid, whereas all examined populations of 
F. regia from Parana are octoploid. Fuchsia 
hatschbachii grows within 20 km of populations 
of F. regia subsp. serrae and F. brevilobis but 
occurs in drier, more interior areas of the planalto 
and is unusual in its close association with limestone 
or sometimes sandstone outcrops. The species is 
dedicated to Gert Hatschbach, Director of the 
seu Botánico Municipal de Curitiba, who has done 
more than any previous botanist to collect and 
study the flora of Paraná. 


8. Fuchsia magellanica Lamarck, Encycl. 2: 
565. 1788; Tabl. encycl. 1, Volume 2(1): pl. 
282. 1792. Fuchsia magellanica var. typica 


Munz, Proc. Calif. Acad. Sci. IV. 25: 9. 1943. 
Fuchsia coccinea Dryander var. robustior 
Hooker, Bot. Antarc. Voy.: 269. 1847, pro 


parte. TYPE: Chile. Straits of Magellan, Jan. 
1768, Philibert Commerson (holotype, P— 
Lamarck Herbarium, also on microfiche; iso- 


types, BM, P—Jussieu Herb., LINN). Figure 
13 


Fuchsia macrostema Ruiz & Pavon, Fl. Peruv. Chil. 3: 
88, pl. 324, fig. b. 1802. Fuchsia macrostema Ruiz 
& Pavón var. grandiflora Hooker, Bot. Misc. 3: 
308. 1833. Fuchsia gracilis var. macrostema (Ruiz 
& Pavón) Lindley, Bot. Reg. 13: pl. 1052, 1827. 
der var. macrostema (Ruiz 


1847. 


TYPE: Chile: VIII Región ato Concepción, Que- 
brada de Carcamo, 1778-1788, Hipólito Ruiz & 
José Pavón (lectotype, MA, here designate 

Thilcum tinctori olina, Saggio Chili, md edition: 
146, 286. 1810. LECTOTYPE: plate Thilco 
Feuillée in J. Observ. Phys. Math. Bot. (Hit. Plantes 


Medicinales). 1725 here — ia (Feuillée de- 
scribed Thilco with five-merous flowers, which Mo 
ina thought worthy of pie recognition) 

Fuchsia sin Lindley, Bot. Reg. 10: pl. 847. 1824. 

magellanica m micis (Lindley) Bailey, 

Gyclop. Amer. Hort., O. TYPE: cultivated 

in Walton, England, Jo d Lindley in 1824 (holo- 

type, CGE). First illustrated in Bot. Mag. 2507. 

is where it was erron neously identified by Graham 
Fuchsia decussata Ruiz & Pavón. 

Fuchsia conica Lindley, Bot. Reg., t. 1062. 1827. Fuch- 
sia macrostemma var. conica (Lindley) Sweet, ful 
Fl. Gard. 6: pl. 216. 1833. Fuchsia magellan 
var. conica (Lindley) Bailey, Cyclop. Amer. Hort. 
614. 1900. TYPE: cultivated in London, England in 
1826, from seeds sent from Chile in 1824, John 
Lindley (not seen). 

Fuchsia gracilis var. multiflora Lindley, Bot. Reg. 13: 

. 1052. 1827. TYPE: cultivated at the Es 
Society, London, England, 1826, from Chilean seeds 
d in 1824, John Lindley s.n. (holotype, 


CGE; isotype, K). 
Fuchsia ped Lindley, Bot. Reg., pl. 1805. 1835. 
uchsia magellanica var. discolor (Lindley) Bailey, 
TUM . Amer. Hort., O. TYPE: cultivated 
at the Harticulurd Society, London, England, from 


h G 
Fuchsia chonotica R. A. Philippi, Dina 28: 687. 1856. 
uchsia coccinea Solander var. chonotica (R. A. 
Philippi) Reiche, Anales Univ. Chile 98: 485. 1897. 
TYPE: Chile: X Región (Los p Chiloé, Archi- 
piélago de Bod. 6 Mar. 1857, F. Fonk 153a 
B s SGO #53136, here VER. The lec- 
otype is mounted on the same sheet with Fonk 
1595, collected on 8-III-1857 from Puerto Low, 
hiloé 


C 
Bele, p Wer F. Philippi in R. A. Philippi. Bot. Zeit. 
,t. 9, fig. 6. 1876. TYPE: Chile: IX ias 
VR . Cautin, Toltén, Feb. 1875, 
Philippi s.n. holotype SGO #53139). 
a magellanica var. molinae Espinosa, Bol. Mus 
c. Chile 12: 102. 1929. TYPE: Chile, X Región 
(Los Lagos), ar quebrada de La Chacra, near 
Castro, 31 Jan. 1929, PAET n. (holotype, SGO 
#53152; isotype, SGO #5312 
Fuchsia magellanica var. ^a ; T Ans. Inst. 
Pat. (Chile) 10: 155. 1979. TYPE: Chile: XII Región 
(Magallanes), Isla Englefield, Seno Otway, 5 S 
71%51' eb. 1979, Edmundo Pisan 
Cárdenas 4947 (holotype, HIP #6892; oe 
MO, RNG, SGO #95030). 


Erect to semiscandent shrubs 0.5-3(-5) m tall. 
Branchlets tan to reddish, glabrous or occasionally 
sparsely strigillose with appressed hairs 0.3-0.4 
mm long; older stems with tan, flaky bark, up to 
25 cm diam. at the base. Leaves opposite, ternate 
or occasionally quaternate, membranous, (narrow- 
ly) elliptic-ovate, 15-55(-70) mm long, 5-25(-40) 
mm wide, acute to acuminate at the apex, mostly 
obtuse at the base, glabrous to strigillose along veins 
on both sides, paler below and sometimes with 
reddish veins; margin serrulate or crenate-dentate, 


562 Annals of the 
Missouri Botanical Garden 


FiGURE 13. Fuchsia magellanica. Flowering branch, longitudinally split flower, individual petal spread open, and 
detail of stem with stipules and petioles. From Solomon & Solomon 4599. 


veins; margin serrulate or crenate-dentate, often 1.3 mm long, 0.5-0.8 mm wide, deciduous. Flow- 
shallowly lobed between the teeth, secondary veins ers solitary or rarely in pairs in upper leaf axils; 
4-7 per side. Petioles slender, 0.3-1 mm thick, pedicels filiform, pendulous, (13-)20-55 mm long, 


3-20 mm long. Stipules (narrowly) lanceolate, 0.7- ^ glabrous to strigillose. Ovary (narrowly) oblong, 


Volume 76, Number 2 
1989 


Berry 563 
Fuchsia sect. Quelusia 


( 
30*—— — A 30° 
—. L4 M 
f 
» 
C 
` 
40*—— — 
50 
o 200 
80° 


FiGURE 14. Native distribution of Fuchsia magel- 
lanica. Stars indicate collections made in the early 19th 
century from populations that have since been eliminated. 


(5-)7-11(-14) mm long, 2-3 mm wide. Floral 
tube cylindrical to narrowly obconic, 7-15 mm 
long, 2-3.5 mm wide, glabrous to strigillose out- 
side, pilulose inside; nectary 1.8-3 mm high, ir- 
regularly lobed. Sepals (15-)17-25(-30) mm long, 
narrowly lanceolate, connate at the base for 2-5 
mm; free lobes (3-)4.5- 7(-8) mm wide at base, 
acuminate at the apex, spreading at anthesis. Tube 
and sepals generally crimson, rarely pale whitish 
pink. Petals purple or rarely pale pink, narrowly 
obovate, (8-)11-20 mm long, 5- mm wide, 
apex rounded. Filaments red-purple, (18-)22-35 
mm and (11-)19-31 mm long; anthers purplish, 


oblong, 2.5-3.5 mm long, 1-2 mm wide. Style 
purple-red, pubescent at the base; stigma clavate, 
purple-red, 2-3 mm long, 1-2 mm wide. Berry 
narrowly oblong, (10-)15-22 mm long, 4-7 mm 
thick; seeds irregularly oblong-triangular, 1.0-1.4 
mm long, 0.7-0.9 mm wide. Gametic chromosome 
number n = 22. 

In moist scrub and along 


Distribution (Fig. 14). 


Valparaiso, Chile (32*50'S) and Neuquén, Argen- 
tina (39%30'S) to southern Tierra del Fuego (55°S). 
From sea level (throughout the range) to 1,750 m 
in central Chile near Santiago. Flowering princi- 
pally from December to March, occasionally as 
early as October and as late as April. Widely cul- 
tivated throughout the world and naturalized in 
parts of South America, east Africa, New Zealand, 
Ireland, and Hawaii. 


resentative specimens examined. CHILE. v 
REGION (VALPARAÍSO). VALPARAÍSO: Viña del Mar, road to 
Concón, 23 Apr. 1939, Behn s.n. (NY); Concón, 14 Jan. 

1950, Boelcke 3920 (BAA, MO, SI); Poeppig 124 (BM, 
MO, P); Valparaiso, 7 Dec. 1930, Behn s.n. (MO, UC); 
Oct. 1829, Bertero 1008(F, G, GH, MO); 1832, Bridges 
198 (LE, W), s.n. (G, K, W); McRae in Feb. 1825 (LE); 
Las Docas, 26 Dec. 1949, Boelcke 3838 (BAA, SI); 
Cerro Campana, Cajón Grande, Pozo de Coipo, 8 Oct. 
1955, Garaventa 6606 (CONC); Limache, Cajón Grande, 
27 Dec. 1944, Boelcke 427 (BAA); Curauma, 31 Jan. 
1932, Garaventa s.n. ( ); Fundo le Valpa- 
raiso, 10 Oct. 1955, Skottsberg s. n.(S; Ns 
Verde, 15 km 
( 


San An Mar. 1982, Landrum 4338 (NY). REGION 
METROPOLITANA (SANTIAGO): SANTIAGO: irri ar de 
Piedra, 1,500 m, 2 Jan. 1927, Looser 148-H (SI); Que- 
ig Peñalolen, ay iat toad 33°29'S, 70°32’ W. 1,600 
, 22 Feb. 1928, RM 2299 (POM), 28 Feb. 1928, 
2300 (POM), 30 1929, Gar uper 828 ( y 
Santiago, “ad riv f ap ale " Gay 1234 
vain 22 Sep. 1918, Claude- Josep 


R o 280 (F, , NY, Py San 
Vicente, 1925, Pennell. 12866 cun s COLCHAGUA : Tal- 
caregue, 34?36'S, 70*53'W, 1870, without ieri (SGO 
#41427); Ciruelos, Dep. de San Fernando, 15 Nov 

1943, Aravena 38 (SGO); Hacienda Millahue 2 pu 
34°36'S, 71?16'W, 16 Feb. 1946, Kausel 1821 (SGO). 
VII REGION (MAULE). CURICO: road Curicó to Argentina, 
beyond Los Quenes, a. Arroyo Puerta, Andean foothills 
at 1,240 m, 13 Feb. 1936, Mexia 7863 (BH, BM, F, 


Quenes, 9 Mas 
(BAB, CONC); Cordillera Peteroa-Planchón, Canon Rio 
Claro, 950 m, 18 Feb. 1936, Mexia 4406 (GH, MO, 
UC) Húmeda Puntilla, aa Llico, 2 Feb. 1969, Vil- 
lagrán & Tapia s.n. (SGO), Fundo La Morita, 20 Dec. 
1968, Mahu 2787 (BAB), Aquelame, near Laguna Vi- 


564 


Annals of the 
Missouri Botanical Garden 


chuquén, 16 Sep. 1963, Barra s.n. (CONC #28414), 
Vichuquén, Dec. 1861, without collector (SGO #53132). 
TALCA: Constitución, Feb. 1891, Reiche s.n. (SGO 
#60893). LINARES: La ~~ ipa to Melado, 17 Dec. 
1953, Ricardi 2793 (CON Perge een 35°47'S, 
71%26'W, 1912, E '840 (NA). VIII REGION 
(BÍO-BÍO). NUBLE: Termas de Chillán, 24 Dav. 1907, Elwe 
s.n. (K); Banos de Chillan, 1,400-1,500 m, 29-31 rin 
1925, Pennell 12473 (F, PH, SGO, 
Las Trancas, i to Chillán, 28 Feb. 1979, Rodríguez 
1107 (CONC), 2 a above Recinto, road to Termas de 
Chillan, 1,500 m, 17 Jan. 1979, oe 4389 (MO); 
Guarilihue, ya 72°41'W, 21 Feb. A saree 
203 ed ungay, Fundo Baquedano . Luc 
9 Fe 957, Artigas 19587 (CONC). CONCEPCIÓN: von 
dps Le a Lala s.n. (MO); Rio Chivilingo, 4 km 
S of Coronel, 5 Jan. 1966, Meyer 9777 (K, MO, NA, 
Valk er 240 ? (cH, m 2s La Araucana at 
O'S, 72%55' 


54 C. 

| Nau 11359 NO 17 Oct. 1953, Sparre 

Ta (CONC); 5 km SE of n along Rio Bio- 
g 1033 (POM, US); 
1 . 


que 
11178 rei 7 Jan. 1976, Marticorena el el 18 
s 1969, Carrasco 200 ( 


«Tus 
e 5934 (CONC): 1 km 
W -facing coastal cliffs, W of Talca uano, Punta Hualpén 
Peninsula, 31 Dec. 1951, pra 226 (K, UC, US); 
Talcahuano and Pe Née? MA 11183584); 
. 1942, fans 6879 (CONC); 
Quebrada Honda, Puentes Mello, 21 Apr. 1976, Mar- 
tic orena et al. 1075 (CONC); San Vicente, above beach, 
1925, Pennell 12866 (GH, PH); Fundo Trini- 
tarias, S 29 Mi Ls Pfister 1232 (CONC); Cerro Car- 
acol, 14 Nov 7, Pfister 6477 (CONC); Concepción, 
Predio RE 26 Oct. 1950, Ricardi s.n. (CONC); 
Barrio M e bar aid 13 Mar. 1980, Riquelme s.n. (MO; 
44); Las C 


2n = anoas, road Concepción to Chalmévida, 
a ae 1980, Riquelme s.n. AME 2n= 44); vp 
, Germain s.n. (BM, F, 6); 1826-1830 ds 


a s.n. (BM); Oct. 1825, Macrae s.n. (BR, 

) Née s.n. (MA #183583). Bío-Bío: 4 km above i 
on road to Laguna La Laja, 930 m, 23 Jan. 1979, 
Solomon 4440 (MO); Antuco, V s.n. (GH); Cor- 
dillera Antuco, La Cueva, Feb. 1887, Rahmer s.n. (SGO); 
Laja, Salto del Trubunleo, 23 ri Ces Boelcke 6528 
(BAA), 90359 (BAB); 23 Jan. 1969, Ricardi & Mar- 
ticorena 5802 (CONC); road from Bio-Bio to Copahue, 

km E of Pitrillon, 13 Mar. 1976, Marticorena et al. 


976, Marticorena et al. 889 (CONC); road ion 
Bio- Bío to Copahue, 3 km past Bio-Bio, 13 Mar. 1976, 
Marticorena et al. 920 (CONC); road Bio-Bio to Santa 
Bárbara, 14 Mar. 1976, Marticorena et al. 1010 (CONC). 
ARAUCO: Cordillera de Nahuelbuta, Contulmo tunnel, La 
Huina, 19 Feb. 1955, Aravena 29 (BR, SI, UC); El 
Diamante, Cordillera de Nahuelbuta, 14 Mar. 1978, Que- 
zada 230 (CONC); Estación Lanalhue, 20 Feb. 1955, 
isa vena 35 (UC); Laraquete, 37?10'S, 73°15'W, 20 m, 

l Jan. 1979, Ser 4413 (MO); Quebrada del Rio 
ch ongol, 13 Dec. 1974, Rodríguez 634 (CONC); road 
Cost ebus to ER 11 Jan. 1972, Quezada et al. 


18(CONC); Trongol Bajo, 3 Mar. 1978, Cuevas 3 (CONC); 
1959, Behn 25364 (CONC); Eights 
o Sta. m ia de a Ricardi 9173 
i bul, 20 Jan. 1979, 
). IX REGION irte ANIA). MALLECO: 
Fundo Solano, Los Alpes, Cordillera de Nahue uta, 590 
m, 12 Jan. 1958, Eyerdam 1014 
Angol, 15 Nov. 193 i 
1892, Kuntze s.n. (F) , 22 
Mar. 1973, Rodríguez & Torres 155 (CONC); Ercilla, 
Feb. 1892, Kuntze s.n. (F, NY); between Manzanar 
Malacahuello, 7 Jan. 1977, Marticorena et al. 1226 
ONC); cuesta La Esperanza, 200 m, 3 Mar. 1980, 
Marticorena & Quez a 1669 (CONC); Ee Dy ces 
, Ricardi et al. 1969 (BAB, 


1980, Marticorena & Quezada 1670 (CONC); 
Termas de Tolhuaca, 1,140-1,180 m, Morrison & Wag- 
enknecht 17499 (BH, F, G, GH, MO, NA, S, UC); Volcán 
de Tolguaca, 1,300-1,400 m, 24 Feb. 1925, Pennell 
12787 (PH) Reserva Forestal Malleco, Los Guindos, 
38°03'S, 71°47'W, 7 Apr. 1978, Rodriguez aha eerie 
Rio Blanco, 27 Dec. 1952. Schwabe 13733 (CONC); 3 
km below Vegas Blancas, 37?45'S, 72%58'W, 24 Jan. 
1979, Solomon 4469 (MO); Termas del Rio Blanco, 2 
Jan. 1948, Pfister 7895 (CONC); Traiguén, Quechere- 
gua, 600 m, 1 Dec. 1947, dis 3327 (S); Pailahueque, 
Dec. . 1929, Pirión 201 dl ad Renaico Valley, 15 Jan 

, Elwes s.n. (K). « : Pucón, near Villaries. 
25 o m, Jan. 1929, Bue. aen s.n. (MO); 

aei 1979 (CONC); Villarrica, Fundo Flor del Lago, l 
Jan. 1946, Behn 26906 (CONC); Volcán Villarrica, 23 
Apr. 1952, Frodin 819 (BM); 3 km E of Puente Cor- 
rentoso, 17 km E of Villarrica, 39%16'S, 72%00'W, 30 
Jan. 1979, Solomon 4535 (MO); Laguna Conguillio, Feb. 
1963, Gleisner 194 (CONC); 3 km from Laguna Captrén, 
Temuco, 20 Jan. 1976, Marticorena et al. 830 (CONC); 
between Laguna Quililo and Melipeuco, 12 Jan. 1977, 
Marticorena et al. 1475 (CONC); Lastarria, 39?13'S, 
72°41'W, 3 Mar. 1980, Marticorena & Quezada 1712 
(MO; 2n — 44); S of Lautaro, banks of Rio Cautin, 7 
Mar. 1980, Marticorena & Quezada 1713 (MO; 2n = 
44); Rio Cautin, Nov. 1919, Hollermeyer s.n. (G); Depto. 
Victoria, 12 Mar. 1939, Morrison x Wagenknecht 17499 
(K); Lago Caburgua, 6 Feb. 1969, Navas 3419 (B); 
Toltén, Feb. 1875, Philippi s.n. (SGO #53139, SGO 
#41419); Termas de Palguin, 11 Jan. 1953, Ricardi 
2399 (CONC); Volcán Caine. Parque Nac. Los Paraguas, 
29 Jan. 1979, Solomon 4511 (MO); entrance to Parque 
Nac. Villarrica, 30 Jan. 1979, Solomon 4523 (MO); road 


1914, Calunn s.n. esa 
X REGION (LOS LAGOS). VALDIVIA: Calafquén, 21 Jan. 1927, 
Comber 1031 (K); Depto. dk ie beu 39?34' o 72°04'W, 
15 Jan. 1976, Marticorena et al. 439 (CC a Pan- 
guipulli, Mar. 1927, Hollermayer 1364 (A, BM, CAS, 
, K, MO, NY, S, UC, US, Z); ou Pirhua, 
Panguipulli, 17 Jan. 1976, Marticorena et al. 609 
(CONC); Purulón, 39°28'S, 72°41'W, 12 Jan. 1976 
(CONC); Lago Rinihue, ribera Norte, Llascahue Chico, 
Marticorena et al. 253 (CONC); desague del Lago Pd 
nihue, 13 Jan. 1976, Marticorena et al. 288 (CONC 
Paso Carirriñe, Panguipulli, 18 Jan. 1976, Marticorena 


Volume 76, Number 2 
1989 


Berry 565 
Fuchsia sect. Quelusia 


et al. ves (CONC); road to Choshuenco, Mois id 16 
Jan. 1976, Marticorena et al. 536 (CONC); pics 
Liquiñe e 16 km E of Panguipulli road, 31 Jan. 1978, 
Solomon 4544 (MO); road to camino del los i os Blanco 
and BL. 39?46'S, 71%45'W, 18 Jan. 1976, Mar 
661 (CONC); W slopes of Cordillera Pe- 
or on La Unión-Punta Hueicolla 


Eyerdam 10669 (F, N nión, Cerro 
Esmeralda, Feb. 1958, Schlegel 1525 (CONC). Calle- 
Calle, 30 Oct. 1896, Buchtien s.n. (BREM, CGE, F, G, 
H, , Llanaca acura, 22- 


j 3 (Sy; Niebla, 2 Nov. 195 
(VALD); estero Puente Negro, 39°58'S, 73°05'W, Mar- 
ticorena & Quezada 1671 (MO; 2n — 44); 2 km from 
Enco, 17 Jan. 1976, Marticorena et al. 585 (CONC); 
S of Valdivia, 2 km past road to Los Guindos, 39?40'S, 
73°02'W, Marticorena & Quezada 1672 (MO; 2n = 
44); Rio Cruces, S of Lanco, 7 Mar. 1980, Marticorena 
& Quezada 1711 (MO; 2n = 44); 1 km SE of Puente 
Piedra Blanca, 8 km S of Valdivia, 29 Jan. 1979, Solomon 
4550 (MO); Trumao, Valdivia, Feb. 1865, Philippi 633a 
(SGO); Amargo, Valdivia, 18 Nov. 1956, Schmitz 108 
(VALD); Valdivia, 3 Nov. 1850, Lechler 537 (CGE, G, 
O); Feb. ra a 35 (BM, CGE, G, O, P, S, TCD); 
Corral, Mar. , Pérez Moreau s.n. (RSA), 12 Oct. 
1929, dn F), 3 Oct. 1930; Corral-La Aguada, 


1948, Senn 4632 (GH, MO, RSA, US); Portezuelo De- 
solación, Volcán Osorno, airis ap 4713 (VALD); La 
Picada, Volcán Osorno, Sparre s.n. (S); Golgol, 40°34'S, 
71%56'"W, 6 Mar. 1948, Rudaiph. 472 6 (VALD); Pampa 
Alegre, sea shore, 14 Jan. 1948, P di: 3967 (S); Tres 
Esteros, 40941'S, 72*53'W, 15 1940, Rudolph 
4724 (VALD); Hueyusca, 40°55’S, 338 W, 16 Feb. 
1950, Rudolph 4711 (VALD); San Juan de la Costa, 
40°31'S, 73?24'W, eb. 1950, Rudolph 4710 
(VALD); Osorno, 15 Jan. 1885, Philipp & Borchers s.n. 
(BM); Osorno, S of Puente Rahue, 40?41'S, 73*08'W, 4 
Mar. rie d & Quezada 1676 (MO; 2n = 
44); 10 mi. E of ocean, road to Termas Pucatrihue and 
Osorno, » 29 Jan 195 8, a 10532 (F, NY, UC, 
US). LLANQUIHUE: Lago Llanquihue, 6 Mar. 1897, Dusén 
s.n. y Jan. 1909, Hicken s.n. (G), Llan nouus, Isla 
Loreley, 1913, Hosseus 45 (CORD); Frutillar Bajo, shore 
of po Eon Mar. 1980, Marticorena & Que- 

zada 7 (MO; 2n = 44); between Frutillar Bajo and 
Frias Ao, : Mar 1980, Marticorena & Quezada 
1678 (MO; 44); Ensenada, Lago Llanquihue, 8 
is. 1939, Everdon & Beetle 24621 (K); 2 km N of 
vog 4 Mar. 1 
I. RS Los R 


1707 (CONC; 2n — 44); Puerto Varas, Hieken 87 (SI); 


m 6 Feb. 1972, Cantino 7 (GH); Parque Na- 

nal Pérez Rosales, 41?09'S, 72°18’W, 8 Dec. 1971, 
Marticorena et al. 1926 (CONC); Saltos del Río uu 
hue, P 


aras, Pu 

Weldt 1248 (CONC); Puerto Varas, 4 Feb. 1945, Hos- 
seus 192 (CORD); Piedra Azul, 11 Feb. 1967, Ricardi 
5287 (CONC); foot of Cerro Hornohuinco, side o 
Chamiza—Lago road, 20 Jan. 1946, Pfister 6047 (CONC); 
Centinela, Jan. 1948, Sparre 4398 (S); Lago Chapo, 235 
m, 41%26'S, 72*35'W, 6 Mar. 1980, Marticorena & 
Quezada 1704 (CONC; 2n — 44); between Correntoso 
and Lago Chapo, 6 Mar. 1980, Marticorena & Quezada 
1703 (MO; 2n = 44); Chamiza, 41?29'S, 72°51'W, 6 
Mar. 1980, Marticorena & Quezada 1699 (MO; 2n — 
44); between Puerto Montt and Pargua, El Avellanal, 
41°45'S, 73°24’W, 4 Mar. 1980, Marticorena & 


Plowman 2604 (GH, 
(G), 14 Feb. 1925, Pennell 12675 (F, 
Volcán Osorno, m from Ensenada- 
road to ski slopes, 6 Feb. 1979, Selaman. 4599 (MO); 
Ensenada, Volcán Osorno, 8 Feb. 1939, Eyerdam & 
Beetle 24621 (GH, NA, UC); Petrohue, shores of Lago 
Todos Los Santos, 6 Mar. 1980, Marticorena & Quezada 
1709 (MO; 2n — 44); Lago Todos Los Santos, 7 Feb. 
1939, Eyerdam et al. 24591 (BH, G, GH, K, MO, NA, 
UC); Casa Panque, between Nahuel Huapi and Laguna 
Todos Los Santos, Dec. 1926, Edwards s.n. (BM), 3 Jan. 
1939, Hunnewell 16069 (GH); Nahuel Huapi, 29 Dec. 
1929, Stuckert s.n. (CORD). CHILOE: 1.5 km fro m Chacao 


al. . 
(CONC); near Ancud, 24-28 Jan. 1932, Macmillan & 
Erlanson 36 (US); Isla Jig Chaques, Golfo de Ancud, 
42?17'S, 73°10'W, 17 . 1968, McFarland & Bar- 
rett 35-14 (NA); Isla Chaullin, Golfo de Ancud, 43?03'S, 
73°27'W, 18 Dec. 1965, Schallenberger & Barrett 36-9 
(NA); between Paranqui a 


M 
CONO; 1 d to Dale, 42%20'S, 
ar eiae & Quezada 1696 
= - 44). QQ. Estero Mechaico and Rio San 
Ant oui. Ancud- Castro da 5 "e 1980, Marticorena 
& Quezada 1698 (MO; 2n 


(CONC); Caatro- Chonchi road, 12 Feb. 1967, Ricardi 
5290 (CONC); near Castro, 15 Mar. 1936, Mexia 8009 
(BM, BM, F, 6, K, MO, NY, S, UC, US); 4 km S of 
Mar. 1980, Marticorena & Quezada 
per (MO; 2n = - 44); Rio Mollueco, o to Quellón, 
5 Mar. 1980, Marticorena & Quezad 2 (MO; 2n 
= 44); 3 km past pm Alto, Castro to Quellón, 43%01'S, 
73°38'W, 5 Mar. 


corena & Quezada 1689 (MO; 2n = 44); 11 Jan. 1975, 
Marticorena et al. 55 (CONC); Quellón, Punta lapa en 
Nadi, 12 Jan. 1975, Marticorena & Quezada 78 (CONC); 
"eid 5 Mar. 198 
(MO; 2n = 44); 
(GH); is Quilán, 1 Jan. 1932, Junge 1483 (CONC); 
Valle del Rio Manao, 41?52'S, 73?32'W, 16 Mar. 1947, 


566 


Annals of the 
Missouri Botanical Garden 


Rudolph 47 14 (VALD); Piruquina-Ovejería, 6 Feb. 1 
Junge 1777 (CONC); Isla Chiloé, 1871, Reed 85 dx 
K), 1826-1830, King (Andersson) s.n. (BM, K, TCD); 
San Carlos de Chiloé, Née s.n. (MA). e Fabia Tic- 
toc, Golfo Corcovado, 43?36'S, 72°54’W, 19 Nov. 1968, 
Barrett 37-17 (NA); Rolecha, 41%55'S, 72*50'W, 5 Jan. 
1951, Pfister s.n. (CONC); Mt. OW Rio Palena, Nov. 
1923, Werdermann 61 (BM, CAS, F, G, GH, MO, SI, 
); Puerto Low, 8 Mar. 1857, Fonch 153G SR 
XI REGION (AISEN). AISEN: Islas Guaytecas, Melinka eb. 
1957, Peri & Marticorena 4104 (CONC); John- 
Ww A 


Tres Montes, 46?49'S, 75°16'W, 22 Nov. 
rris & Barrett 39-17 (NA); Golfo San Estéban, 
Fiordo Kelley, 46%55'S, 74*05'W, 28 Nov. 1972, Hoppe 
4 (RNG); Peninsula Taitao, s in 17 Dec. 
ee (GOT). COTHAIQUE: Fenchik, 45?33'S, 
7 , 13 Feb. 1959, Sc iced 2246 (CONC); Parque 
Nacional Río Simpson, Mar. 1968, Sanguiesa s.n. (S 
sE JARRERA: eón, Pues Lago Buenos Aires, 
11 Feb. 1939, von Rentzell 6223 (GH, MO); Lago 
General Carrera, Mina Silva vA E 72°20'W, 18 Dec. 
1939, Heim s. a 1k Valle del Rio Em slopes of Cerro 
Pina, 47%00'S, 73?15'W, 22 Dec 71, Davison F-40 


ci Barbara, 48°04 


); Puerto delia Pro- 
montorio Exmouth, 49°25'S, 74°19'W, . 1879, 
Ibar s.n. (SGO); Ring Dove Inlet, 49*44'S, 74?10'W, 24 
Jan. 1879, without ride (SGO #53127); Ultima Es- 
peranza, Gusinde 295 (GOT); Portland Bay, Midpoint 
Channel, 50%15'S, [ein a 1956, Tillman 55 pori 
sla Drummond Hay o Molyneux °l 
74°55'W, 11 Dec 11977, Roig 144 (HIP); Bahia Med 
Papudo, 50?22' S. 75°19'W, 25 Nov. 1968, Barrett et 
al. 42-7 (NA); Cordillera de Paine, 50° 54'S, 73°00'W, 
Jan. 1966, Tsujii 784 Naples Andean foothills, ca. 
51?00'S, 72%50'W, 1961, Anderson 18 ( ); Cerro 
Sa Tr Toro, La Peninsular 51°19'S, 72°46'W, 15 
Dec , Boelcke e 5 (HIP, RNG); Se 
a E Rio M n 19'S, 73°07'W, 2 


Ed Cost 


Mar. 1945, Biese 1486 (SGO); Puerto Bellavista, 51°31 'S, 
73°16’W, 11 Jan. 1977, Moore & Pisano 1628 (CONC, 
HIP, RNG); near Ultima Esperanza, 51?30'S, 73%00'W, 
Borge 339 (GH, K, NY, S, US); Seno Ultima Esperanza, 
Hicken 87 (SI) Isla Virtudes, Canal Elías, Puerto Vir- 
tudes, 51?33'S, 74%54'W, 9 Feb. 1976, Dollenz et al. 
1444 (HIP, RNG); re Roca, Seno Resi, 51?51'S, 
73%02'W, 24 Jan. 1978, Pisano 2877 7 (HIP, RNG); Seno 
Unión, Ancón, 52%09'S, 73%02'W, 12 Jan. 1976, Dollenz 
et al. 984 (HIP, RNG); Isla Long, 52*18'S, 73°38'W, 
| Feb. 1879, Guero s.n. (SGO); Smith Channel, Long 
Island, 1921, Asplund 126 (S); Cueva del Milodón, 13 
Jan. 1952, Pfister & Ricardi 12065 (CONC); Eberhardt, 
13 Apr. 1899, Borge 344 (GH, MO, NY, S, US), Donat 
384 (BM, CAS, F, GH, K, MO, NY, Z). MAGALLANES: 
Isla Riesco, Mina Elena, 53%02'S, 72%45'W, 30 Apr. 


1940, Santesson 7 751 (K); Estancia Cerro Pinto, N side 
kyring, 1 and 2 Feb. l 
(SGO #53128); Seno Otway, Punta H Seibert 
(M); Seno Otway, Punto Otway, 9 Feb. , Lee s.n. 
(US); Rio Caleta, Seno Otway, 53?09'S, un 32' W. 5 Jan. 
1979, Cárdenas 17 (HIP, MO); Isla Engelfield, Seno 
Otway, 53°05’S, 71*51'W, 24 Feb. 1979, Pisano & 
Cárdenas 4926 (HIP, MO, RNG, SGO); Seno Otway to 

unta Arenas, 1 Mar. 1945, Biese 1250 (F, GH); Fiordo 
Silva Palma, Angostura Titus, 53%26'S, 71°44’W, 9 Jan. 
1973, Pisano 3825 (HIP, MO, RNG); Borja Bay, Straits 
of Magellan, 53°32'S, 72°29'W, Feb. 1882, Coppinger 
s.n. (K); Ballena E E La of Isla Carlos III, 53?35'S, 
72%22'W, 18 973, Goodall 5 (NA); ees 
Brunswick, Bahía m Aguila, 53?47'S, 71°01'W, 

1975, Samsing s.n. (HIP); Bahia del Indio, Lote San 
Isidro, Río abel. 53%48'S, 71%00'W, 1 Mar. 1973, 
Pisano 4024 (HIP, MO, RNG); Puerto San Antonio, 
53%53'S, 70954'W, 1826-1830, Anderson et al. s.n. 
(W), King s.n. (TCD); Fuerte Bulnes, 53%35'S, 70°56' W, 
13 Mar. 1970, Pisano 2501 (CONC, HIP, MO, NA), 
25 Apr. 1951, Pisano s.n. (MO; 2n = 44); junction roads 
to Puerto Hambre and Fuerte Bulnes, 53?40'S, 70955'W, 
9 Mar. 1979, Solomon & Solomon 4691 (HIP, MO); 
Puerto Edén, Anliot 6143 (SGO); Port Gallant, 1838- 
1840, Hombron s.n. (P; 1838, d'Urville s.n. (TCD), 11 
Feb. 1877, Savatier 134 (K, P, RSA); Port Famine, 
1826-1830, King 75 (BM, CGE, K, TCD), 1834, s.n. 
(U, W); 1838-1840, le Guillou 41 (P, TCD); Shipton 
Glacier, Bay of Mountains, 53%55'S, 72?01'W, 13 Feb. 
1976, Barclay 6 (US); Isla Meer 54?00'S, 70°45'W, 
12 Mar. 1896, Dusén s.n. (P), 12 Mar. 1896, Norden- 
skjold 657 (0); Isla Dawson, Harris Bay, 25 Feb. 1908, 
Skottsberg 174 (S); Islote Offing, Isla Dawson, Pastore 
in 7 Feb. 1921 (SI #4842); Isla Capitán Aracena, Bahia 
Morris, 54?14'S, 71%01'W, 8-9 Dec. 1971, Moore 2745 
(BAB, NA, RNG); Magellan Straits, 1852, Andersson 
330 (MO), Handisyd in July 1690 (BM, Sloane Herb.), 
1826-1830, King s.n. (BM, F). TIERRA DEL FUEGO: Río 
Santa Maria, 60 km by Camino Sur, 53?43'S, 69*58'W, 
10 Feb. 1972, Pisano 3517 (HIP, MO); Canal Whiteside, 
uerto Yartou, 53%50'S, 70?18'W, 12 Nov. 1941, 
Santesson 1697 (K); S side of Bahía Inútil, 2 km E of 
uerto ie 53%52'S, 70°07'W, 13 Nov. 1971, Moore 
2439(K, RNG); Bahia Inútil, 1905, Crawshay s.n. (BM); 
Isle Grawe, gra 195 (W); N. Esperanza, Goteborg 
298 (LE , W) ; 


y 


A 
Hunziker 6909 E 
Los Andes Ms Dec. 
cerros N of San Mar ae de 
Dawson p; (BA, E Lag Log 12 Mar. 1926, 
Comber 813 (K), 2 Feb Lago Espejo, 7 Feb. 
1940, Cabrera 5976 (NY); Eu Fre 17 Feb. 1968, 
Cabrera & Crisci 19206 (CONC); Puerto Correntoso, à 
Feb. 1934, aer 11679 (BAA); road Bariloche- Co 
rentoso, 20 Mar. 1939, Cabrera 5044 (F, GH, NY); Isla 
Victoria, Parque s Nahuel Huapi, 9 Jan. 1946, Boelcke 
1781 (BAA) Isla Victoria, Zoological Station, 31 Mar. 
1946, de Barba 1122 (LIL); Lago Epilaufquén, 4 Feb. 
1948, 2 & Schwabe 2392 (BAA); 
5 Feb. X - 
ied 14 pl 1951, Diem 1774 (BAA, BAB, SI); Villa 
a Angostura, 30 Mar. 1980, Diem 3615 (MO; 2n = 
44); Lago Tromén, 19 Feb. 1941, Pérez Moreau s.n. 


Bo 
p 
n 
[e] 
p 
a 
I] 
= 
È. 
g- 
Es 
un 
m 
3 
= 
p 
mi 
æg 
3 
o 


Volume 76, Number 2 
1989 


Berry 
Fuchsia sect. Quelusia 


(RSA); Laguna Currilafquén, Feb. 1942, Pérez Moreau 

: n, Feb. 1942, d Moreau 
r. 1941, Pérez 
Feb. 1980, den & aa. s.n. (MO); 


in 


= 
N 
= T 
p 
A 


Isla Victoria, Puerto Pampa, 800 m 
BAB, MO). río NEGRO: dene Blest to 
S, MO, 


Raulies, 20 


n 9 
Blest, 8 Mar. 1914, Hosseus 1071 (CORD): Puerto Blest, 
25 Jan. 1930, Kies s.n. (BAA); Cerro ier Or, 
1,200 m, 21 Feb. 1979, Blumen & Solomon 4639 
(MO); road to i. d ig (LP); jose Mas- 
cardi, 8 Feb. 1940, Pérez Moreau s.n. (RSA); between 
Llao-Llao and Bahia López, 23 Mar. 055, Fabris 1134 
(F); Llao-Llao, Crespo 161 (SI); Feb. i Bahía López, 


(UC); Lago e E Jan 


Península Los Luizo 


ad to Cerro V. Lóp 19 Feb. 9, mon 
Solomon 4610 (MO); Puerto c Lago Nahuel Huapi, 
29 Mar. 1934, Ljungner ); Cascada de Lo 
Alerces, 13 Feb. 1965, Mn 202 (GH); El Bolsón 
Feb. 1945, Corvetto 324 y; near Bariloche, 
1965, Walter & Walter 479 (By; Bariloche, Coloma 
Suiza, 12 Mar A, N 


o ahue 

K). CHUBUT: Epuyén, F 
TER Futaleufü, 7 Dec. 1967, 
(BAA); Los Rá el Río Futaleufú, 21 Jan. 1945, Cas- 
tellanos s.n. (F, LIL); Colonia 16 de Octubre, Futaleufú, 
Jan. 1948, a s.n. (LIL); Parque Nacional de Los 
Alerces, Lago Futalaufquén, 6 Feb. 1955, Burkart 19851 
(MO); Lago Cisne, 10 Feb. 1944, Pérez Moreau s.n. 
(RSA); Valle de las Plumas, Lago General Paz, 2 Mar. 


E 


USE 
ea] 
O 
c 
— 
No) 
tx 
o 
Y 
a 
` 
a 
N 
[^ 
= 
` 
a 
a 
RE 
a 
"3 
P amm 
o] 
un 
> 
> 
> 
e 
c 
ad 
o 
"y 
E 
O 
c 


A, o 
15 Feb. 1961, Lourteig & Buchinger 194 (P); Cordillera, 
Jan. 1901, Burmeister s.n. (BAB); cerros N of La Comisa- 
ría, Feb. 1942, Pérez Moreau s.n. (RSA); Los Alerces, 
9 Jan. 1948, Soriano 2885 (BAA). SANTA CRUZ: Glaciar 
Spegazzini, 50%15'S, 73°15'W, 19 Mar. 1968, Anliot 
6016 (SGO), 19 Jan. 1967, Boelcke et al. 12612 (BAA, 
BAB, CONC, P, SD); Santa Cruz, 14 Nov. 1975, Boelcke 
16403 (BAA); S shore of Lago Argentino, 1907-1908, 


ue 98 (GH, NY, US); Cerro Buenos Aires, near 
Moreno Glacier, 5 Jan. 1959, James 669 (BM, DS); Valle 


-1901, pus d s.n 


Rio Grande, Cordini s.n 
64?00' W, 54%45'S, Webster s.n. (G), Estancia d 
cultivated from bush s s to have been broug m the 
western Beagle Channel by Thomas Bridges. 25 Mar. 
1968, Goodall 1677 (NA, ip UC, US), 8 Nov. 1968, 
1836 (NA, RNG, UC, US; H urberton Ed (culti- 
s = Jan. 1966, Goodall 298 (LP, RNG). 
sentatite a specimens. ARGENTINA: 
San a Villa Nou , Tucumán, 12 June 1945, 
Ortiz s.n. (NY); road E Lucía M T de figs 20 
1944, Torrent s.n. (NY). BOL a Paz 
Apr. 1946, Llanos 239 (BAB) is Ar eee 
June 1966, Cárdenas 6279 (LIL), Apr. 1961, 3969 
(LIL); Incachaca, Apr. 1938, Cárdenas 688 (LIL), Mar. 
1941, 2248 (LIL); Toralapa, 18 Dec. 1973, Cárdenas 
3753 (LIL); Ayopaya, Dec. 1935, Cárdenas 3369 (LIL); 
Carrasco, Sehuenka, 1964, Cárdenas 6153 (LIL). 
CoLoMBIA: Macizo de Bogotá, 10 Apr. 1946, Schultes 
7275a (COL). Kenya: Mt. Kenya forest, 5 Jan. 1967, 
Perdue & Kibuwa 8416 (Ky Nyeri, 16 Apr. 1971, 
Wandumbi School 757 (K). MEXICO: Orizaba, Veracruz, 
July 1855, Botteri 1047 (K). MALAYA: ere 
lands, 1 Apr. 1976, T. & P. 612 (K). RwWAnpa: 
5 Aug. 1977, Runyinga 724 (LG). PAGAN Jaegstal, 
; o s 


T 


Upper Waiakea Forest Preserve, 580 
Wagner et al. 4847 (BISH), Hawaii National Park, Fa- 


a 
Crater Rim Drive, pr. 1 , Fosberg 57479 (BISH); 
near Kilauea Crater, Degener 1588 (BISH); near Volcano 

House, 3,900 ft., 1 Sep. 1937, Neal 946 pron Kauai, 


Kokee State Park, 3. 500 ft., 
1075 (BISH). 


24 Apr. 1968, Herbst 


Fuchsia magellanica is separated from the Bra- 
zilian members of section Quelusia by 1,000-1,500 


(usually over 1⁄4 the length of the 
dentate leaf margins, which are sometimes shal- 
lowly lobed between the teeth. Fuchsia magellan- 
ica lacks the long, drooping branches of some of 
the lianoid species from Brazil, and it generally 
grows as an upright shrub. In cultivation it can 
also reach treelike proportions and is considered 
the hardiest (most frost-resistant) of all Fuchsia 
species. 

A number of different species and varieties have 
been recognized by others for what is considered 
here to be a monotypic species. Two different color 
varieties have been named for plants that lack the 
typical red coloration of the floral tube, sepals, leaf 


568 


Annals of the 
Missouri Botanical Garden 


veins, and petioles. One, F. magellanica var. mo- 
linae, comes from the island of Chiloe; the other, 
F. magellanica var. eburnea, is from the southern 
Magallanes Region of Chile. Similarly pale-colored 
plants are also known from Corral, Prov. Valdivia, 
Chile (Gunkel 2133, GH). All of these plants occur 
sympatrically with red-flowered populations of F. 
magellanica and probably result from minor re- 
curring mutations on pigmentation. 

Munz (1943: 9) thought that plants from the 
southern part of the range, which he called F. 
magellanica var. typica, had consistently smaller 
leaves and flowers than those from the north, but 
this is clearly not the case. Large-leaved plants 
occur throughout the range of the species. James 
Solomon (pers. comm.) examined many local pop- 
ulations in Chile and Argentina and found that 
individuals growing in the sun usually have smaller 
leaves and flowers than those nearby in shadier 
sites. Similarly, clines in leaf size and texture occur 
along climatic gradients, such as that along the 
shores of Lago Nahuel Haupi in Argentina. The 
vegetation in that area becomes increasingly xeric 
from west to east, and plants of F. magellanica 
respond to this gradient by developing lower, more 
compact plants with smaller, thicker leaves. 

Populations of plants with small, densely packed 
leaves are common on the island of Chiloé, where 
high levels of humidity predominate throughout the 
year. Larger-leaved individuals also occur together 
with these variants. Some of the phenotypic vari- 
ants of F. magellanica may also be genetically 
determined, as they are known to maintain their 
distinctive characteristics under uniform growing 
conditions. In horticultural circles, there has been 
a strong tendency to assign varietal names to some 
of these strains. 

In this treatment, I am not recognizing any 
varietal or subspecific taxa under F. magellanica, 
because it is apparent that a great deal of local 
variation exists over the broad geographical range 
of the species, without any clear geographical cor- 
relations. The species is cytologically uniform, with 
over 40 populations counted with 2n — 44. Since 
most of the present range of the species, south of 
44°S latitude, was completely glaciated during the 
major glacial advances of the Pleistocene (Vuilleu- 
mier, 1971), there has probably been very little 
time for populations to diverge in the southern part 
of the range. 

In his Flora of Tierra del Fuego, Moore (1983) 
cited localities of F. magellanica from as far south 
as the Beagle Channel at 55°S, and west to Staten 
Island, at 64°W. Collections from these areas were 
made in the past century, however, and it is un- 


likely that the species occurs there naturally today. 
According to D. Moore (pers. comm.), the intro- 
duction of goats on Staten Island early in the last 
century and the activities of Europeans in the vi- 
cinity of the Beagle Channel during the past 100 
years have badly deteriorated the woody vegetation 
of these areas. 

On the drier, Argentinian side of the Andean 
Cordillera, F. magellanica occurs natively as far 
north as the Lago Quillen, at 39*30'S. Northwards, 
the mountains become increasingly arid. On the 
Pacific side of the Andes, native populations are 
found as far north as Concón (32°50'S), just north 
of Valparaiso. These populations grow close to the 
coast, along streams in wooded ravines. Looser 
(1928) found native populations of F. magellanica 
east of Santiago (33?30'S), in the Quebrada de 
Penalolén, between 1,450 and 1,750 m, and Navas 
(1976) reported the species from the nearby Que- 
brada de Macul. 

The northernmost extension of Fuchsia magel- 
lanica barely overlaps with the southernmost pop- 
ulations of the xeromorphic F. lycioides (sect. 
Kierschlegeria) in the vicinity of Concón, north 
of Valparaiso, Chile. A number of specimens col- 
lected in this area during the last century are 
morphologically intermediate between the two 
species and may represent interspecific hybrids (see 


Appendix). 


9. Fuchsia regia (Vellozo) Munz, Proc. Calif. 
Acad. Sci. IV. 25: 13, pl. 1, fig. 5, 1943. 
Quelusia regia Vellozo, Fl. Flum. 149. 1829; 
Icon. 4: tab. 6. 1831. LECTOTYPE: The illus- 
tration in Tab. 6, Volume 4 of the /cones of 
Flora Fluminensis. Vellozo stated for the type 
locality: “in the Pharmacopolitan Alps on the 
way to the town of Cunha," which is located 
between Paraty and Cunha, in the Serra da 
Bocaina, near the Sáo Paulo- Rio de Janeiro 
border, Brazil. According to Borgmeier (1937), 
Vellozo collected in this area between 1779 
and 1787, and the original illustrations were 
made in the field from live plants. 


Erect to generally scandent or climbing shrubs 
0.5-5 m tall, or lianas up to 15 m high in trees. 
Branchlets wine purple, glabrous to puberulent, 
rarely densely pilose; main branches ascending in 
small plants, decumbent to drooping and 1-6 m 
long in large ones. Leaves opposite, or in whorls 
of 3, 4, or rarely 5, firmly membranous to cori- 
aceous, (narrowly) elliptic-ovate to ovate, 20-140 
mm long, 8-70 mm wide, apex acute to acuminate, 
base acute to rounded, glabrous to pilose on both 


Volume 76, Number 2 
1989 


Berry 569 
Fuchsia sect. Quelusia 


Aa 


Fuchsia regia 


@ subsp. regia 


@ subsp. serrae 


O subsp. reitzii 


45* 40* 
5 


E 15. Native distributions of the three 


subspecies of Fuchsia regia. Contours higher than 800 m are 
5.) 


Ficu : 
dicated by the stippled areas. Capital letters are Brazilian state abbreviations. (See caption of Fig. 


surfaces; margin entire to gland-serrate, secondary 
veins 4—10 per side. Petioles 1-3 mm thick, 3- 
35 mm long. Stipules triangular, membranous to 
thick-stubby, 0.8-3 mm long, 0.4-3 mm wide, 
adjacent ones sometimes fused, usually recurved 
with age, ephemeral to persistent. Flowers solitary 
or rarely in pairs in upper leaf axils; pedicels pen- 
dulous, 10-55 mm long, usually wine red. Ovary 

oblong, 5-12 mm long, 2-4 mm wide. Floral id 
cylindrical-fusiform, 5-16 mm long, 3-7 mm wide, 
glabrous to puberulent or pilose outside; nectary 


smooth or lightly ridged, 2.5-8 mm high. Sepals 


15-45 mm long, connate at the base for 3-14 
mm; free lobes 3-9 mm wide at base, spreadin 

to recurved or reflexed at anthesis. Tube and sepals 
red or rose. Petals purple, obovate to spathulate, 
10-25 mm long, 8-16 mm wide. Filaments red- 
purple, 25-45 mm and 20-38 mm long; anthers 
oblong, purplish, 2-5 mm long, 1-2 mm wide. 
Style glabrous to pubescent in lower 14; stigma 
clavate, 2-6 mm long, 1-2 mm wide, exserted 8- 
25 mm beyond the anthers. Berry oblong, ellipsoid 
or globose, 10-27 mm long, 9-13 mm thick, dark 
purple when ripe; seeds tan, oblong to narrowly 


Annals of the 
Missouri Botanical Garden 


triangular, laterally compressed, 1.6-2.5 mm long, 
1-1.6 mm wide. Gametic chromosome numbers n 


— 22, 44. 
Distribution (Fig. 15). This is the most widely 


distributed species of Fuchsia in Brazil, occurring 
in the states of Minas Gerais, Espirito Santo, Rio 
de Janeiro, Sáo Paulo, Paraná, Santa Catarina, 
and Rio Grande do Sul from 18?20'S to 29°30'S 
Most common between 700 and 2,000 m, on the 
upper, wettest slopes of ranges such as the Serra 
do Mar, Serra Geral, Serra da Mantiqueira, and 
Serra do Espinhaço. Fuchsia regia subsp. reitzii 
is associated with the Araucaria forests of the 
southern planalto from Paraná to Rio Grande do 
Sul. Plants occasionally occur as high as 2,400 m 
on Mt. Itatiaia and at sea level in coastal restingas 
of Santa Catarina and Rio Grande do Sul. Flowers 
mainly from October to April. 


This is the most variable of the Brazilian species 
of sect. Quelusia. Three distinct and largely allo- 
patric series of populations can be recognized based 
on differences in several floral and vegetative char- 
acters. They intergrade in areas where their ranges 
overlap, and the presence of octoploid populations 
in some or all of the populations sampled in each 
series is unique in the genus. For these reasons, 
the different series are treated here as subspecies 


of F. regia. 


KEY TO THE SUBSPECIES OF FUCHSIA REGIA 


la. Sepals strongly connate above the point a in- 
np of petals and stamens, joined for 
than of wm er len ngth, the free yel 
era ally recurved; stipules crassate and per- 
sistent, e a nodose appearance to young 
mer serrae 
: Sepals slightly connate at the base, Joined fe 
less than %4 of their total length, the free labes 
spreading; stipules membranous or thickened 
only at the base and young stems not apparently 
nodose. 
2a. Leaves in whorls of 3-5, membranous, ter- 
tiary venation conspicuous; margins gland- 
serrulate and petioles mostly shorter than 
10 mm; floral tubes 5-10 mm long, sepals 
17-28 mm long ......................... 9c. subsp. reitzii 
. Leaves mostly opposite or ternate, subcor- 


- 
c 


N 
c 


O mm; fl 
10-16 mm long, sepals 24-45 mm 
long 9a. subsp. regia 


9a. Fuchsia regia (Vellozo) Munz subspecies re- 
gia. Fuchsia regia (Vandelli) Munz var. typ- 
ica Munz, Proc. Calif. Acad. Sci. IV. 25: 13. 
1943. Vellozo's type illustration shows a plant 


with narrowly ovate-lanceolate leaves in pairs 
and threes, and flowers with very shortly con- 
nate, long sepals. These characters correspond 
to the northernmost subspecies, which reaches 
as far south as the type locality in northern 
Sào Paulo state. Figure 16 


xis A Cambessedes in eu Hilaire, H. Bras. 
d. 2(17): 273. 1830. Ty 


photograph at F; isotypes, P — again 
a two different branches, the one to the left num- 
red C' 80, F #1008268 “ex P," Fie 
Fuchsia pyrifolia K. Presl, Symbol. Bot. 2: 19, tab. 65. 
34. TYPE: Brazil, “ad Rio Dae > Presl? (ho- 
vid PRC— 445/a). 


Shrubs 0.5-5 m tall, occasionally erect but usu- 
ally scandent, or lianas to | igh in trees. 
Branchlets generally glaucous and wine purple, 
occasionally loosely to densely pilose, older plants 
with long hanging branches to ong. Leaves 
opposite, or in whorls of 3 and 4, subcoriaceous, 
elliptic to lance-elliptic or ovate, 40-140 mm long, 
10-50(-70) mm wide, apex acute to acuminate, 
base acute to cuneate or rounded, sublustrous green 
and glabrous to pubescent above, veins often red- 
dish-tinged, paler green and glabrous to pilose be- 
low, sometimes with tufts of pilose hairs at the base 
of the lower midvein; margin subentire to remotely 
gland-denticulate, secondary veins 6-9(-10) per 
side. Petioles stout, 2-3 mm thick, canaliculate 
above, (6-)10-28 mm long, wine purple. Stipules 
triangular, membranous or thickened at the base, 
+ recurved, 0.9-2(-3) mm long, 0.5-1.8 mm 
wide, adjacent ones sometimes fused, the base usu- 
ally persistent but the tip deciduous. Flowers sol- 
itary in upper leaf axils; pedicels 22-55 mm long, 
pendulous. Ovary cylindrical, 6-11 mm long, 3- 

mm thick. Floral tube narrowly to broadly cy- 
lindrical-fusiform, 10—16 mm long, 4-6 mm wide, 
glabrous to subpilose outside; nectary smooth to 
ridged, 5-8 mm high. Sepals 24-45 mm long, 
narrowly lanceolate, connate at the base for 3-7 
mm; free lobes 4-9 mm wide at base, spreading 
at anthesis. Tube and sepals red to rose. Petals 
violet, obovate, 13-22 mm long, 10-16 mm wide. 
Filaments red-purple, 25-44 mm and 23-38 mm 
long; anthers oblong, 3-5 mm long, 1.5-2 mm 
wide. Style glabrous i pilose in lower half; stigma 
red, narrowly clavate, 3.5-6 mm long, 1.2-1.6 
mm thick, exserted 10-27 mm beyond the anthers 
when fully extended. Berry oblong-ellipsoid, purple 
when ripe, 20-25 mm long, 9-12 mm thick; seeds 


Volume 76, Number 2 
1989 


Berry 
Fuchsia sect. Quelusia 


1.5 mm 


meo: 
-a 


Y 


N y 15mm 


Fuchsia regia subsp. regia. —a. Flowering branch. —b. Longitudinally split flower and individual 


FIGURE 
petal.— c. Slightly immature berry. —d. Seed from (c) in two lateral views. From 


1.8-2.8 mm long, 1.2-1.8 mm wide. Gametic 
chromosome numbers n = 22, 


Distribution (Fig. 15). In montane cloud for- 
ests of the Serra do Mar, Serra do Caparaó, Serra 
do Espinhaco, Serra da Caraqa, Serra da Manti- 
queira, Serra da Bocaina, and other subsidiary 
mountain ranges in Espirito Santo, Minas Gerais, 
Rio de Janeiro, and northern Sáo Paulo states, 


amamoorthy 683-2. 


1,000-2,400 m. Flowering mainly from October 
to April. 


Additional specimens examined. BRAZIL. ESPÍRITO 
SANTO: Pico de Forno Grande, Mun. Castelo, Brade 19275 


GERAIS: Carangola, Serra da Gramma, Chase 9579 (US); 
Kuhlmann 137 (RB); Mexia 4278 (A, BM, F, C, GH, 
MO, NY, UC, US); Serra da Gramma, trail to Araponga, 


572 


Annals of the 
Missouri Botanical Garden 


Mexia 4326 (A, BH, BM, F, G, K, MICH, MO, NY, P, 
PG, S, U, UC, US, Z), Araponga, Kuhlmann 33934 
(RB) Alto Caparaó, Tronquieras to Vale Verde, Souza 
& Marcia 85 (HBR, RB, US); — Alto Caparao, 
Berry et al. 4543 (MO, RB; n = ; Vale Verde, 
Caparaó, Hatschbach & Ahumada Ey (MBM); Pico 
da Bandeira, close to Caparaó, 1,500-1,600 m, Shep- 
herd et al. 5769 (MBM); ine to Fazendinha, Serra 
aparaó, Lanna 32 bos a ra de Macieiras, Caparaó, 
Moreira 47 (R); Rancho das Trés Barras, Serra Caparaó, 
Santos & Campo in 26 June 1950 (RB); Caparaó Velho, 
Torgo 30 (HB); Serra bs Arp a, Mello Barreto 7157 
d irri in Feb. 9 (Py Caraga, Tanquinhos, 
di x Tales 165 (BHCB) Ouro Preto, Berry et al. 
2 (MO, RB); pg 29191 (R, US); Claus- 
se i PE Macedo 3078 (MO, US); Morro do Baú, 
DUE od Badini pid cud Campo Grande, Mun. 
Ouro Preto, Badini 20113 (OUPR); Morro da Queimada, 
Ouro Preto, de Lima et al. 1369 (MO); Camarinhas, 
Godoy H 106 Esc. Pharmacia (OUPR); Fonte dos Amores, 
Mun. Poços de Caldas, Mello Barreto 11078 (BHCB); 
Pocos de Caldas, Barros 667 (SP); Hoehne 2703 (SP); 
Pocos de Caldas, Caixa d'Agua, Santos 5734 (DS); road 
Botelho- Pocos de piod Pabst 9500 (HB, MBM); Cal- 
das, Araujo s.n. (R 1685); Lindberg 346, 347 (S) 


Mosén 472 (MO, P, E 473, 4509 (S); Regnell vulp 
Regnell in 1862 (BR, K, M, NY), in 186 865 (LE, 
O, R), in 1 (P, W), in 1867, in 1871 (Sy bs 


Ribeiráo dos Bugres, "n 474 (S); Serra do Cipó, du 

2 US); Serra do Cipó, road t 
err Mello A 8855 (NY, R), 9204 (BHCB), 
à (MO. 


, i ico do slopes, 
35830 m A RB, US; Serra RE Itabira, Sello s.n. 
(LE, S); Rio , Duarte 926 (HB); 


Oiro Banco Weddell 1553 (P); Morros do Lobo, no coll. 
647 (M); Passa Quatro, lta uaré, Brade & Araujo 1981 
(MO); E 2120 (R); Fazenda dos Campos, Passa Qua 
tro, Vidal 1789 (R); Cidade Azul, Serra da Mantiqueira, 
Sapucai-Mirim, Kuhlmann 2915 (SP); Delfim Moreira, 
7 km from Isópolis, Mattos 15397 (SP); Sao Francisco 
dos Campos, Alto do Morro do Cabrito, divisoria SP-MG, 
Delfim Moreira, Kuhlmann 2466 (SP); Aguas de 
km NW of Cascata, road to Sao Roque da Fatura, 
s & Mattos 14192 (SP); Aiuruoca, Serra do Pa- 
ae. Silveira in Nov. 1897 (R #102490); Serra do 
Pict, Schwacke in 10 Dec. 1886 (R), Fazenda Santa 
Alda, Teixeira Soares, Sampaio in 1944 (R #41684); 
Fazenda Sao Benito, Serra da Mantiqueira, Vidal in Apr. 
1949 (R #41684); Hermilio Alves, Cacholina, Duarte 
2384 (MO); Serra do Itacolomi, Mun. Ouro Preto, Badini 
23131, 25273 (OUPR); poa 1058 (R); Berry et al. 
4544 (MO, RB; n = 22), 4545 (MO, RB); Claussen 7 
(F, LE, P, RSA); de Lima un (MO, RB); Macedo 2774 
(MO, NY, S); Magalhaes Gomes 1744 (OUPR); no lo- 
cality, Ackermann Ex (BR); Claussen in June 1840 
OXF); Miers 2433 (US); Langs- 
ds n. (LE); Pus in 1842 (P, U); Riedel 62 (P); 
St. Hilaire B' 200 (P) Velho 4555 (BM, K); 
Widgren in 1845 (BR, US). RIO DE JANEIRO: Monte Es- 
trela, Mieire 553, s.n. (LE); Serra da Estrela, Petropolis, 
Goés 75 (RB); Serra da Estrela, near Cachoeira de Véu 
da Noiva, Correias, Fazenda do Sampaio, Pabst in 5 Jan. 
1964 (HB); Petrópolis, Correias, Serra dos Orgáos, Fa- 
zenda Bom Fim, Carauta 693 (MO); Serra de Petropolis, 
Duarte 1163 (RB); Duarte & Pereira 4669 (RB); Vale 
das Videiras, Mun. Petrópolis, Martinelli & Gurken 508 


(RB); Martinelli et al. 169 (MO); Vale das Videiras, 
Morro da Cuca, Jouvin 3 (RB); Petrópolis, road to Fa 
zenda Inglesa, Rocha & Silva 14 (R, US); Itaipava, Vale 
de Cuiaba, Mun. Petrópolis, Martinelli et al. 1641 (RB); 
26 km from Itaipava, road to Teresópolis, Berry et al. 
4419 (MO, RB; n = 44); Petrópolis- Teresópolis, Lutz 
1734 (R); Petrópolis-Quitandinha, Lutz 1356, 1577 (R); 
Petrópolis, Araras, Morro Bolo de Milho, Sucre & Braga 
2808 (RB), Petrópolis-Itamaraty, Glaziou 8676 (R); Pati 
do Alferes, Mun. Petrópolis, Plowman & Martinelli 10125 
(F); Petrópolis, oo 8676 (P) Alto do UAE Fan 
Petrópolis, ia. ton s.n. (K); Serra dos Órgáo Mun. 
Petrópolis, Martinelli . et al. 1749 (RB); ais "a 
Teresópolis, Gentry & Peixoto 907 (M 
rque Nacional Ser d dos Órgãos, A 
RB); Fromm et al. 
3 (MO); Lem a R Mar. 1964 (NY); 
Lott 12 (MO, RB k Rodrigues ; Sampaio 1622 
(R); Santos et al. 202 (R), OCB (PEL, R); Vidal 11.147, 
11.654, 11.6489 Bh Wawia see ); Campo dos An- 
tas, Serra dos Orga 22 May 1948 (RB); 
Emygdio et al. in Mar. 2 (R); al 1917 (RB), 
25 July 1949 (HB, M Santo & Pessoa in 25 Feb. 
(R); Serra dos Orgaos, Araujo 
‘a Badini meee 22263 ! (OUPR Brade 


GH 225 tee NY, XF, P, TCD, W); 
Glaziou "is Simart 3733 (P); ergy Bee (LE); Miers 
3473 (TCD), 4453 (US), in = 8 (BM — mixed 
collection); Peckolt s.n. (Wy, Wilkes s.n. S US); Serra 
os Orgaos, Castelo, ers 10945 BA X Ad un Serra 
dos Órgãos, Vianna 654 (R), Brade 9918 (R); Teresó- 
polis, Bailey & Bailey. 1296 (BH); Teresopolis to Pedra 
da Baleia, Cardoso 145 f 21 d a Caledonia, 
Nova Friburgo, Berry et 21, 44 25 (n = oa 4432, 
4433 (n = 22) (all MO, aus ee de Frade, 
Pons 15876 (B, RB); Sgt et al. 6 621( M Conia, 
e & Aparicio 18703 (RB); Alto Soberbo, Gen 
& "Peixoto 937 (MO, RB); Itatiaia, Serra da Mantiqueira, 
including Maceiras and Planalto, Altamira ter 
/ 1 (M, RB); uei in 11 Oct. 1945 rip kar 
& Edmundo 829 (RB); Barth 1136 (US); Brade 17515 
PACA, RB); Campos deis 277 : (B, NY, dg RB); 
Carauta & 528 (MO); Dus n 81 (GH, MO, R, S, W), 650 
), in 25 May 1902 2 (S Eiten & Eiten 6662, 6674 
(SP, US); Emmerich 53 (R); Mb & Briolando 184 
(RB); Fromm 17 (R); Fromm et al. 164 (R); Ginzberger 
142, 143 (F, W); Hunt 6418 M Fd did 26436 
(RB); Lanna 54 NO Lanstyak 253 (MO); Lindeman 
& Haas 5203 (Uy Lutz in Jan. 1957 ir Markgraf & 
Brade 36: 5 1 (RB); Occhioni 16483 (RB 
3, 116, 815-823 (MO); 
var 78(R); Ule 148 (R); Abrigo Rebougas, Itatiaia, Berry 
et al. 4437, 4439 (MO, RB; n = 44); Oliveira 413 (RB); 
Pereira & Pabst 8222 (HB), 5178 (GH, RB); Santos 
5034 (US); Smith 1476 (GH); Sucre & Plowman 5178 
(GH, RB); Walter 71 (RB); no locality, St. Hilaire D 22 
(F, P), 800 (P); Luschnatte s.n. (OXF). SAO PAULO: Serra 
da Mantiqueira, between Itajuba and a a, Cas- 
art 23371 (MO); Pico Itapeva, Campos do Jordào, 
al & Ramamoorthy 4860 (MBM); Parque Estadual, 
Campos do Jordào, Mattos 15956-a (SP); Umuarama, 
Campos do Jordao, Kuhlmann 32492 (SP); Campos n 
Jordao, Bailey & Bailey 811 (BH); Campos Porto 3 
(B, RB), 2991, 2992 (RB), 3233 (MO); Davis et ji 
b (RB, SP, UEC); de Lima 1131 (MO; Emmerich 


~ 
S 


Volume 76, Number 2 
1989 


Berry 573 
Fuchsia sect. Quelusia 


15 (HB); Goncalves & Kuhlmann 2169 (SP); Friedrichs 
in Jan. 1944 (PACA); Hashimoto 313 (SP); dies 
33105 (B, RB); Leite in 1945 (A); Lutz & Lutz 1 

(Ry Mattos & Mattos 14467 (SP); Pabst 4212 (F, HE 
MBM, RB); Ramamoorthy & Vital 6 
674 (MBM, MO), 675 (MO), 676 (n = 44), 677, 680 
(n = 44), 682, 684 (MO), 817 (MB), Sakane 206, 211 
(SP); Serra da Bocaina, Gibbs et al. 4573 (F); Duarte 
7689 (HB, MO); Handro 808 (SP), Ramamoorthy $ 
de Lima 1172, 1174 (MO); Vianna 2949 (R, US); Vian- 
na et al. 286 (R, US); Serra da Bocaina, Mun. Sào José 
do Barreiro, km 16 near Silveiras, Shepherd & Shepherd 
12858 (SP, UEC); Serra da Bocaina, road to Mambucaba, 
Lutz in Apr. 1951 (R #116696); Serra da Bocaina, 
Sertào da Bocaina, margins Rio Pintado, Pabst 4744 
(HB, MO); Bananal, Sertao do Rio Vermelho, Serra da 
Bocaina, Brade 15268 (RB); Brade & Duarte 20129 


Botánico do Estado, Teixeira 241 (SP); Ypiranga, Jardim 
Museu Paulista, Luederwaldt s.n. (SP); Chácara dos Mor- 
rinhos, Pickel 4447 (SP). 


Fuchsia regia subsp. regia is the most wide- 
spread of the Brazilian fuchsias, occurring in four 
different states, centered in Rio de Janeiro and 
Minas Gerais. As a result of its fragmented distri- 
bution, with populations in a number of discrete 
mountain ranges, considerable morphological dif- 
ferentiation has taken place throughout the range 
of this subspecies. It is also cytologically hetero- 
geneous, with tetraploid and octoploid populations. 
All counts made from Minas Gerais and northern 
Rio de Janeiro have been tetraploid, whereas oc- 
toploid and tetraploid individuals occur together in 
northern Sào Paulo and south-central Rio de Ja- 
neiro states, in areas such as Itatiaia, Serra da 
Bocaina, and Serra dos Órgáos (see fig. 9 in Ho- 
shino & Berry, 1989). The different cytotypes 
cannot be distinguished morphologically, however, 
and their relationships are not well understood. 

Fuchsia regia subsp. regia can be distinguished 
from other taxa of sect. Quelusia by having large 
flowers with spreading sepals and only slight sepal 
connation, large leaves with petioles generally long- 
er than 10 mm, and mostly glabrous stems and 
foliage. 

This subspecies has a particularly broad ecolog- 
ical tolerance, extending from tall cloud forest at 
1,000 m to open campos as high as 2,400 m. 
When it occurs inside the forest, F. regia subsp. 
regia grows as a liana, flowering when its branches 
reach sufficient light in the tree canopies. As the 
vegetation decreases in height with greater ele- 
vation, it grows as a scandent shrub, and as a short, 
erect shrub when found on the edges of the open 
campos. This gradual transition in the habit of 
subsp. regia has been observed in several localities, 
such as the Serra dos Órgãos and the Morro da 


Nova Caledônia in Rio de Janeiro, and the Serra 
do Itacolomi and Serra do Itambé in Minas Gerais. 

As a result of its broad altitudinal and geograph- 
ical distribution, F. regia subsp. regia occurs sym- 
patrically with four other species from sect. Que- 
lusia. In each case, morphologically intermediate 
individuals have been found that indicate the oc- 
currence of interspecific hybridization (see Appen- 
dix). It also occurs sympatrically with F. regia 
subsp. serrae in northern Sào Paulo along the 
eastern slopes of the Serra da Bocaina, where it 
forms hybrid populations, with further intergra- 
dation southwards along the Serra do Mar as far 
as Caraguatatuba. These populations are discussed 
in the Appendix. 

Several collections of F. regia subsp. regia from 
widely distant localities are unusually pubescent 
yet do not appear to be the result of hybridization 
or introgression with pubescent species growing 
nearby, such as F. alpestris or F. coccinea. The 
specimen Campos Porto 3421 (RB, from Campos 
do Jordáo, Sào Paulo, 1,600 m) is similar in all 
morphological features to nearby glabrous popu- 
lations of F. regia, but the entire plant is covered 
by a dense, pilose indumentum. Three collections 
from Minas Gerais are similarly pubescent: Gard- 
ner 4555 (K), Claussen 88 (G), and Brade 16953 
RB, US, from the Serra do Caparaó). Martinelli 
et al. 169 (RB, Vale das Videiras, Petrópolis, Rio 
de Janeiro) is covered by short, fine pubescence, 
which is unusual among the populations of F. regia 
from Rio de Janeiro. These pubescent variants 
are apparently quite rare, and none were found 


~ 


during my fieldwork in Brazil. 


9b. Fuchsia regia subsp. serrae P. Berry, subsp. 
nov. TYPE: Brazil. Paraná: Mun. Morretes, 
Recanto Eng? Lacerda, Estrada da Graciosa, 
850 m, 28 Jan. 1985, P. E. Berry, T. Plow- 
man & F. Juarez 4493 (holotype, MO; iso- 
type, MBM; n = 44). Figure 17. 

ddr A pin iue ies edes, ie Bras. Merid. 2(17): 274. 

Par 


E: Brazil. à: “in forest near Ferraria, 


ra bs p Curitiba > May 
contains two different 


in and labeled as the type collection of F. 


Piehot jen Miers i in Lindley, Bot. . Reg. 27: pl. 66. 
841 


BM; 
different plants, but the holotype has only one spec 
imen on its sheet, with a pasted blue slip stating 


574 


Annals of th 
Missouri Hood Garden 


FicunE 17. 
leaf. 


Fuchsia regia ibig sp- 
ary e. Details of crassate, a teas From Berry et al. 44 


1838” 


9 


“Fuchsia radicans Nob., Organ Mts., Feb., 
and a small typeset label of “Herb. John Miers. 


Frutex 1-4 metralis aliquando in arboribus scandens 


isque, ca. 0. crassis, contiguis interdum 
facie nodosis. Tubi florales cylindrico-fusiformes (6-)7- 


—a. Flowering branch. —b. id — c. Detail of upper surface basal 


15 mm longi, sepalis (15-)19-34 mm longis, (4-)5-7 
mm latis, basi (5-)8-14 mm connatis, lobis patentibus 
saepe recurvatis vel raro reflexis. Bacca oblonga 16-26 
mm longa, 9-13 mm crassa, seminibus 2-2.5 mm longis, 
1-1.5 mm latis. Numerus gameticus chromosomatum n 


eee to climbing shrubs 1-4 m tall, or lianas 
to 15 m high in trees. Branchlets generally finely 
puberulent, sometimes subglabrous or densely short- 
pilose, the lateral ones often short and divaricate; 
older branches on large plants hanging, to 6 m 
long, with basal stems to 6 cm diam. Leaves op- 


Volume 76, Number 2 
1989 


Berry 575 
Fuchsia sect. Quelusia 


posite or in whorls of 3-4, coriaceous, (narrowly) 
ovate to elliptic, (35-)60-110 mm long, (15-)25- 
45 mm wide, apex acute to acuminate, base round- 
ed to acute and sometimes + uneven, smooth and 
lustrous dark green above, pale below, glabrous to 
lightly pubescent on both surfaces; margin sub- 
entire, secondary veins 5-8(-10) per side. Petioles 
1.5-3 mm thick, usually dull purple, canaliculate 
above, (7-)10-24(-35) mm long. Stipules prom- 
inent, crassate, 2-3 mm wide, 2-3 mm long, ca. 
0.5 mm thick, recurved, persistent, adjacent ones 
sometimes fused, drying tan-colored and giving a 
nodose appearance to the stems. Flowers solitary 
in upper leaf axils; pedicels 20-45 mm long, pen- 
dulous. Ovary cylindrical, 6-12 mm long, 2-4 mm 
wide. Floral tube cylindrical-fusiform, (6-)7-15 
mm long, 3-7 mm wide, glabrous to puberulent 
outside; nectary + smooth, 5-8 mm high. Sepals 
(15-)19-34 mm long, connate at the base for 
(5-)8-10(-14) mm and forming a long, usually 
4-ridged sepal tube, the free lobes (4-)5-7 mm 
wide at base and spreading to most often recurved 
or rarely reflexed at anthesis. Tube and petals 
bright red. Petals violet, obovate to spathulate, 11— 
20 mm long, 8-15 mm wide. Filaments red, 22- 
48 and 20-40 mm long; anthers red-purple, 3-4 
mm long, 1.5-2 mm wide. Style red, glabrous to 
pilose near base; stigma red, clavate, 2-4 mm long, 
1-2 mm wide, exserted 15-23 mm beyond the 
anthers. Berry oblong, 16-26 mm long, 9-13 mm 
thick, purple when ripe; seeds 2-2.5 mm long, 1- 
1.5 mm wide. Gametic chromosome number n — 


44. 
Distribution (Fig. 15). Typical of dense cloud 


forests on the crest and wet, coastal escarpments 
of the Serra do Mar and Serra Geral, in Rio Grande 
do Sul, Santa Catarina, Paraná, Sào Paulo and 
southern Rio de Janeiro states, from 500 to 1,450 
m. Also found at lower elevations on the top of 
isolated ““morros”” in the Atlantic coastal forest of 
Santa Catarina, or rarely occurring near sea level 
in coastal restingas in northern Rio Grande do Sul 
and southern Santa Catarina. Flowering mainl 
from November to March, occasionally in other 
months. 

Additional specimens examined. BRAZIL. RIO DE 
JANEIRO: Cachoeira de Macacü, road to Nova Friburgo, 
Sucre & Soderstrom 9044 (MO, RB); Angra dos Reis, 
Alto da Serra, 600 m, Sucre et al. 4908 (MO); RJ-16, 
Rio Claro, first tunnel, 550 m, de Lima 270 (RB); Organ 
Mountains, Gardner in Mar. 1841 (BM), Ramamoorthy 
& de Lima 1181 (MO); Hen of Rio de Janeiro, St. 
Hilaire B 2475" (P). ien PA lto da Serra, above 
Caraguatatubi — orthy E Vital 685, 686 (MO), 
688 (MO; n = 44); Paus Estadual de Caraguatatuba, 
Gentry & Zardini 49376 (MO); Alto da Serra de Pa- 


ranapiacaba, Emygdio 1520 (R); Serra de Cubatáo, Guil- 
lemin 470 in 1839 (P); Alto de Serra, Sào Paulo to 
Santos, Pabst A Pereira 5750 (HB, MBM); Vidal in 
Apr. 1937 (R #35043); Alto da Serra, Hoehne 7617 
A); Mattos 11714 (MBM); Munz 15415 (F, G, GH, 
, POM, US); Sandeman 2012 (K, OXF); Smith 1900 
(F, GH, S): Wettstein & Schiffner in May 1901 (W); 
Santos, we s.n. (G); Sallesopolis, d 314 (MO, 
RB); Est. Campo Grande, Hoehne 11894 (POM); Hor. 
raceia Exper. Station, Nascimento E ; Feb. 1943 (UC); 
road to Atibain, Monteiro et al. 4894 (F), near San 
Bernardo, Wettstein & Schiffner in July 1901 (W); Can- 
tareira, e 1477 (SP); Rio Grande, estrada do Mar, 
Hauff 120 (SP); Serra do Mar, between Mogi das Cruzes 
and Bertioga, Kirizawa et al. 552, 1015, 1038 (SP), 
km 74 of SP-98 from NE to Bertioga, Mun. Biritiba- 
Ucu, Pirani & Yano 7 ); Miracatu, Sitio Urupua, 
km 348.5 of = : 16, o 4 (SP); Reserva Capao 
Bonito, 70 km from Itapetininga, SP-139, Sakane 543 
(SP); Reserva em Botelho, between Sào Miguel Ar- 
canjos & Sete Barras, Prance et al. 687 1 (MBM), 6896 
(F); no locality, Burchell 3742 (BR, Ky; Kuhlmann s.n. 
(RB). PARANÁ: Santa Ana, Mun. Bocaiúva do Sul, Hatsch- 
bach & Guimaraes 23408 (M, MBM, MO); Jaguatirica, 
Mun. Bocaiúva do Sul, Ramamoorthy & Vital 124 (MO); 
Serro, Mun. Bocaiúva do Sul, Hatschbach 42976 (MBM); 
Serra Capivari, Mun. Campina Grande do Sul, Oliveira 
& Oliveira 867 (MBM); 7-12 km W of Sesmaria and 
Rio Capivari, Berry et al. 4446, 4447, 4449, 4450 (n 
— 44) (all MBM, MO); Sitio do Belizário, Mun. Campina 
Grande do Sul, Hatschbach 16426 (MBM); Papanduva, 
Mun. Campina Grande do Sul, Hatschbach 7091 (MBM); 
Curitiba, Hoehne 23109 (POM); Dusén in 12 Jan. 1914 
(F, GH, MO, NY, SP, US), in 20 Oct. 1928 (SP); Parque 
Nova Iguagu, Mun. Curitiba, Berry et al. 4455, 4456 
(MBM, MO); Kummrow 1294 (MBM); 10 km E of Cu- 
ritiba, Lindeman € Haas 2539 (MBM); km 34 road 
Curitiba- ray Fromm et al. 304 (R); Pabst et al. 
res (HB, MBM); Rio da Santa, Curitiba to Joinville, 
nrad & d du 2055, 2056 (MO); Rio Taquary, 
Mun. Quatro Barras, Berry et al. 4492 (MBM, MO; n 
= 44); Hatchback 19942 (MBM); Oliveira 577 (MBM); 
Serra de Aragatuba, Mun. Guaratuba, Hatschbach 7082 


orretes, Curiel in 14 Apr. 

1947 (MBM #39718); Hatschbach 751 (PACA); Mun. 

Morretes, Ramamoorthy & Hatschbach 828 (MBM), 

629 (MO; n = 44, nm = 88), 830 (MO; n = 44); Pilào 

de Pedras, Mun. Morretes, Kummrow 1917 (MBM); Pi- 

raquara, Hatschbach 188 (PACA); Serra do Emboque, 

Piraquara, A 24946 (MBM, UC); Rio 

o José dos Pinhais, Hatschbach 8019 
Sà 


, NA); 
. Sao José dos Pinhais, Hatsehbaa h 1765 57 
, UC); Guaricana, Mun. Sáo José dos Pinhais, Olive- 
ira 518 (MBM , MO); cae oe 34905 (MBM); Zinco, 
Mun. Sao José dos Pinhais, Oliveira 534 (MBM); Barro 
Preto, Mun. Sao José dos Pinhais, Oliveira 153 (MBM); 
Serra do Mar, Braga 550 (MO, RB, US); Alto da Serra, 
Mun. Sete Barras, Hatschbach 26892 (MBM, NA); Pirai, 
Mun. Tijucas do Sul, Hatschbach 40187 (BH, MBM); 


Mun. Tijucas do Sul, 
Mun. Tijucas do Sul, Haisohbach 40471 (MBM). SANTA 


576 


Annals of the 
Missouri Botanical Garden 


CATARINA: Rio Novo, Mun. Aguas Mornas, Klein & Bre- 
solin 10547 (HBR); Faxinal, Biguagu, Mun. António Car- 
los, Reitz C956 (HBR, MO, PACA, RB); Rambo 50392 
(PACA); Fazenda D E Bendito Novo, Zinco, 
Mun. Benedito Novo, Bresolin & Rocco 683 (HBR); 
Morro Spitzkopf, Mun. Blumenau, Reitz & Klein 9202 
(CM, G, HBR, MO); Smith & Reitz 6279 (HBR, R, RSA, 
US); Morro ido Barào, Mun. Botuverá, Reitz & Klein 
18015 (HBR, MO, RSA); Morro da Bateia, Mun. Brus- 
que, Reitz C1914 (HBR, MO, US); Morro do Iquererim, 
Mun. bus Alegre, Smith & Klein 7384 (HBR, MO, 
R, RSA, US); Morro do Ribeiráo, Mun. Florianópolis, 
Klein 7166 (FLOR, HBR, MO, RSA); Lajeado Alto, Gua- 
biruba, Souza et al. 609, 613, 620, 621, 625 (FLOR); 
Monte Crista, Sào Francisco do Sul, Mun. Garuva, Reitz 
& Klein 10027 (HBR, MO, RSA); Hórto Florestal, Mun. 
Ibirama, Klein 1975 (F, HBR, MO, PACA, RSA); Alto 
Cachoeira Grande, Aguas Mornas, Klein ] 
10751 (HBR); Morro do Baú, Mun. Itajai, Reitz 2104 
(HBR, MO, RSA, US); Morro da Fazenda, Mun. Ita 
Reitz & Klein 1740 (HBR, M 
Joinville, Schwacke 496 1 (R, RB); Joinville, ec Dona 
Francisca, Reitz & Klein 4220 (HBR , RSA); Morro 
Itajai, ism De (HBR, MO, 

a, Mun. Jacinto Machado, Reitz C318 
(GH, HBR, MO), 856 (PACA); rod do peine e 
acinto Machado, Reitz C1456 (GH 

MO); Encruzilhada a Serril, Mun. Lajes, Lourteig i 
(DS, HBR, NY, P, R, : Serra Geral-Encruzilhada, 
jes, Smith & Klein. 8020 ( SA 


Reitz & Klein 2849 (HBR, MO, UC, US); Bo 
da Serra, descent of Serra Geral to Lauro Müller, Berry 


Anitópolis, Mun. P , RSA 
; Morro do Cambirela, Mun. Palhoca, Klein & Bre 

solin 9297 Wr PA MO); Rambo 30332 (PACA); 

Serra a Boa o José, ancho Queimado, 


: € Klein E CBR, MO), Ec (HBR, MO, 


RSA ncho Pere ado, road to Boa Vista, Rama- 
m E Reitz 1159, 1160 (MBM, MO); Boa Vista, 
Cabins k Reitz 1163 (MBM, MO), 1167, 1168 


(MO); cial dia near Boa Vista, Ramamoorthy & Reitz 
1164, 1165 (MO); Rio dos Bugres, Schwacke in 5 July 
1885 (P); Serra do Matador, Mun. Rio do Sul, Reitz & 
Klein 7321 (G, HBR, MO, NY, UC, US); km 195-203 
of road Rio do Sul-Curitibanos, Mun. Pouso Redondo, 
Berry & Falkenberg 4501 (n — 44), 4502, 4504 (n — 
44) (all MBM, MO); Falkenberg 2318, 2321, 2322, 
2325 (FLOR); slopes of Serra Geral from Itaimbezinho 
to Praia Grande, Falkenberg 2370, 2376, 2377 (FLOR); 
Falkenberg et al. 3750, 3751 (FLOR), 3752 (n — 44; 
FLOR); above Turvo to Serra da Rocinha, Mun. Timbé 
do Sul, Matzenbacher 398 (ICN); 13-19 km above Timbé 
do Sul to Serra da Rocinha, Falkenberg et al. 3753 (n 
= 44), 3754, 3755 (FLOR); Serra da Rocinha, border 
RS and SC states, 22 km above Timbe do Sul, Falkenberg 
et al. 3756, 3757, 3760, 3761, 3766 (FLOR); Cerro 
Azul, Mun. Timbó, Berry & Falkenberg 4500 (MBM, 
MO); 0.5-4 km E of escarpment of Serra Geral between 
Uribici and Brago do Norte, Falkenberg et al. 3792— 
3796 (FLOR); 1-2 km W of escarpment of Serra Geral, 
towards Uribici, Falkenberg et al. 3788-3791 pra 
NE of Uribici, km 90 of road to Braco do Norte, Fal- 
kenberg et al. 3783, 3785 (FLOR); Sabiá, Mun. Vidal 


Ramos, Reitz & Klein 4274 (HBR, MO, NY, RSA, UC, 
US), 6657 (F, HB, HBR, MBM, MO, PACA, PEL, RSA); 
Sombrio, 10 m, Mun . Sombrio, Reitz C1131 (MO, RSA 

79111). RIO GRANDE DO SUL: Torres, Faxinal, 10 m, 
Waechter 827 (ICN #41273); Josafá, Mun. Cambara 
do Sul, Falkenberg 2339, 2341, 2343, 2345, 2347, 
2349, 2350, 2354, 2357, 2372, 2378 (FLOR); Itaim- 


li n r. 
( y O (PACA), 45524 (MO), 45529 
(PACA), 49363 LM 53983 (B, hos S), 58552 (B, 
HBR, MO, S); 20 km E of Tainhas RS-020 to Terra 
de Areia, Falkenberg et al. 3744(n= 44), 3745 (FLOR); 
dos da Serra, Rambo 


ula, 
36222 (ICN, PACA); Serra do Faxinal, near Sào Fran- 
cisco de Paula, s 32099 (PACA): Fortaleza, Mun. 
Cambara do Sul, m & Wasum 2122 (US); Wasum 
et al. 734 (US). 


Fuchsia regia subsp. serrae is the only taxon 
in sect. Quelusia besides F. brevilobis with the 
sepals connate for over one-third of their length. 
It is unique in its persistent, crassate stipules and 
lustrous, subcoriaceous leaves with thick petioles. 
Unlike the other subspecies of F. regia, the sepals 
are usually recurved at anthesis and four-angled 
in bud. 

This subspecies is most common along the crest 
and upper slopes of the ' the steep and 
very wet region on the eastern edge of the southern 
Brazilian planalto as it drops abruptly to the At- 
lantic coastal plains. The rugged terrain, with sheer 


“serras, 


drops o m or more, such as those found in 
the Aparados da Serra in northern Rio Grande do 
Sul, creates a broad diversity of microhabitats re- 
flected in the large degree of variability in different 
populations of this subspecies. For example, low, 
densely branched subshrubs characteristically grow 
in rock crevices on the steep drop-offs of the Apa- 
rados, while tall lianas with long, drooping branches 
are found in more protected, forested areas near 
the crest of the Serra. 

The degree of sepal connation varies consider- 
ably in different parts of the range of F. regia 
subsp. serrae. In northern Paraná, where it occurs 
together with the long-tubed F. brevilobis, the se- 
pals are often joined for half or more of their length. 
Along the Serra Geral in southern Santa Catarina 
and Rio Grande do Sul, however, the sepals are 
less joined than populations further north; in ex- 
treme cases, individuals like Falkenberg 3744 and 
3745 have the sepals joined for only 4-6 mm, but 
their thick, shiny leaves and nodose stipules are 
characteristic of subsp. serrae. 

ong the edge and just inland from the steep 
drop-offs of the Serra Geral in Rio Grande do Sul 


Volume 76, Number 2 
1989 


Berry 577 
Fuchsia sect. Quelusia 


and southern Santa Catarina, a sharp ecotone oc- 
curs between the drier, flatter planalto and the wet, 
steep coastal slopes. In this area, Fuchsia regia 
subsp. serrae occurs sympatrically with the east- 
ernmost populations of Fuchsia regia subsp. re- 
itzii, and the two subspecies intergrade extensively 
in areas such as Itaimbezinho, Serra da Rocinha, 
and the edge of the Serra Geral near Uribici and 
Bom Jardim da Serra in Santa Catarina. Further 
discussion and a list of intermediate, probable hy- 
brid individuals between these two subspecies is 
found in the Appendix. 

Towards the northern limits of the range of 
Fuchsia regia subsp. serrae, this subspecies also 
contacts populations of Fuchsia regia subsp. regia, 
especially in the Serra da Bocaina in northern Sào 
Paulo state. Populations of subsp. serrae predom- 
inate there on the lower, coastal slopes below 1,100 
m, while subsp. regia occurs at higher elevations 
and further inland. Extensive intergradation be- 
tween these two subspecies occurs on the upper 
coastal slopes of the Serra, where unusually pilose 
individuals are also found. Some of these individuals 
are octoploid, as in all the southern populations of 
subsp. serrae, while others are tetraploid, like some 
of the populations of subsp. regia from the Bocaina 
range and further north (Hoshino & Berry, 1989). 
Although F. regia subsp. regia reaches its southern 
limit at the Serra da Bocaina, populations of Fuch- 
sia with mixed ploidy levels and dense pubescence 
occur south of the Bocaina area as far as Ubatuba 
in the Serra do Mar of north-central Sao Paulo 
state (Hoshino & Berry, 1989). 

Fuchsia regia subsp. serrae also occurs sym- 
patrically with F. brevilobis in several localities in 
Paraná. Although F. regia is octoploid in this area, 

and F. brevilobis is tetraploid, the occurrence of 
be d intermediate individuals indicates 

at hybridization has taken place between these 
taxa (see Appendix). The unusually long sepal tubes 
of populations of F. regia subsp. serrae in this area 
are characteristic of F. brevilobis and may be the 
result of introgression with this species. 

Outside of its typical montane habitats, F. regia 
subsp. serrae has also been found in coastal res- 
tingas at sea level, near the Rio Grande do Sul- 
Santa Catarina border. Reitz (1961) first detected 
this apparent anomaly in the distribution of typi- 
cally montane taxa, and found it to occur in at 
least ten other species that normally occur along 
the crest of the Serra Geral. The presence of these 
species in the restingas must represent recent 
chance introductions from the nearby serras, as 
the coastal plains of this area are only of Quater- 


nary age (Klein, 1984). 


9c. Fuchsia regia subsp. reitzii P. Berry, subsp. 
nov. TYPE: Brazil. Santa Catarina: Municipio 
Sào Joaquim, Fazenda de Laranja, Sào Joa- 
quim, 1,400 m, 13 Jan. 1959, P. Raulino 
Reitz & Roberto Klein 8135 (holotype, UC 
#240288; isotypes, B, BR, G, GH, HBR, 
ICN, K, NY, U). Figure 18. 


Fuchsia pubescens Cambessédes in St. Hilaire, Fl. Brasil. 

Merid. 2(17): 275, tab. 134. 1830. TYPE: Brazil: 

"in fields near the Rio Yapó in the part of Sào Paulo 

Province called Campos pad currently in Pa- 
raná, Municipios Pirai do Sul and Jaguariaiva, May 


1820, Auguste de Saint Hilaire C 1545 inea 


as t pecimen escens, and Cam- 
bessédes's illustration shows. the verticillate leaves 
and flower and round fri f F. regi 


subsp. reitzii. The ed Sa from Minas Ger- 
ais cited by Cambessédes in his protologue is F. 
coccinea. 


Frutex erectus scandensve 1-4 metralis, ramulis pur- 
purascentibus nc nna vel dense puberulentibus vel pi- 
losis ferruginescentibus, ramis cito excorticatis. Folia 3- 

4(-5) ilie ci elliptica ad anguste ovato- -elliptica, apice 
acuta basi acuto-acuminata obtusave, nervis tertiariis con- 
spicuis, margine glan 
mm longis, stipulis lanceolatis 4 
longis. Flores verticillati, tubis floralibus cylindrico-fusi- 
ongis, sepa is anguste lanceolato- 
m longis, basi 2-6 mm connatis sub 
anthesi oa pone. obovatis 10-15(-20) mm lon- 
gis, 7-13 mm latis. Bacca globosa vel interdum ellipsoidea 
10-16 mm longa, 9-12 mm lata, seminibus 1.6-2 mm 
longis, 1-1.3 mm latis. Numerus gameticus chromoso- 
matum n — 44. 


Erect to scandent shrubs 1-3 m high, or climb- 
ers in trees or brush to 4 m tall. Young branches 
purplish, subglabrous to densely puberulent or pi- 
lose, the pubescence ferrugineous in some plants; 
older stems with reddish to pale tan, readily ex- 
foliating bark. Leaves in whorls of 3 or 4(—5), firmly 
membranous, (narrowly) elliptic to narrowly ovate- 
elliptic, 20-65(-90) mm long, 7-20(-40) mm wide, 
acute at the apex, acute to attenuate or obtuse at 
the base, dark dull green and subglabrous above, 
pale green and subglabrous to densely pubescent 
below with conspicuous tertiary venation, margin 
serrulate, usually with prominent glandular teeth 
ascending apically at a + 45° angle, secondary 
veins 5-8 per side. Petioles 3-7(-15) mm long. 
Stipules lanceolate, 0.8-1.4 mm long, 0.4-1 mm 
wide, thick at the base, purple, drying dark, di- 
vergent to recurved with age, with apiculate, semi- 
persistent tips. Flowers solitary or sometimes 2 per 
leaf axil, near branch tips; pedicels slender, 10- 


578 Annals of the 
Missouri Botanical Garden 
FIGURE 18. Fuchsia e subsp. reitzii. Flowering branch, detail of flowers, and detail upper surface of basal 


leaf. From Berry et al. 4 


35(-50) mm long, pendulous. Ovary oblong, (4—)5— 
9 mm long, 2-4 mm wide. Floral tube cylindrical- 
fusiform, 5-9(-11) mm long, 2-4(-6) mm wide, 
subglabrous to puberulent or loosely pilose outside, 
glabrous inside; nectary smooth, 2.5-5 mm high. 
Sepals 17-28 mm long, narrowly lance-oblong, 
connate at the base for 2-6 mm; free lobes 3-5(- 
6) mm wide at base, subacuminate at the apex, 
spreading at anthesis. Tube and sepals red. Petals 
purple, obovate, 10-15(-20) mm long, 7-13 mm 
wide. Filaments reddish, 25-45 mm and 20-38 
mm long; anthers red-purple, oblong-elliptic, 2— 
2.5 mm long, 1-1.5 mm wide. Style glabrous to 
pilose in lower half; stigma clavate, 2-2.5 mm 
long, 1-1.5 mm wide, exserted 5-15 mm beyond 
the anthers. Berry globose to ellipsoid, shiny dark 


purple when ripe, (10-)12-16 mm long, 9-12 mm 
wide; seeds tan, 1.6-2 mm long, 1-1.3 mm wide. 
Gametic chromosome number n — 44. 


Distribution (Fig. 15). | Occurring principally 
in low wooded areas and usually along streams or 
seepages, on the planalto west of the Serra Geral 
and Serra do Mar in south-central Paraná, Santa 
Catarina, and Rio Grande do Sul, at 880-1,730 
m. At the top of the escarpments of the coastal 
mountain ranges subsp. reitzii intergrades with F. 
regia subsp. serrae. Flowers mainly from Novem- 
ber to March. 


Additional specimens examined. BRAZIL. PARANÁ: 
General Carneiro, Hatschbach et al. 13670 (U); 10 km 
N of Palmas, Mun. Palmas, Hatschbach et al. 28175, 


Volume 76, Number 2 
1989 


Berry 579 
Fuchsia sect. Quelusia 


28237 (MBM); an dos Turcos, Mun. Guarapuava, 
Hatschbach 25602 (MBM 


HBR, MO, NY, RSA, SI, US); BR 153, Rio Roseira, 18 
m Horizonte, Mun. Agua Doce, Krapovickas & 
Cristóbal 33693 (CTES); 30 km S of Horizonte, Smith 
& Klein 13421 (HBR); 8 km N of Caçador, Mun. Ca- 
cador, Smith & Klein 8966 (HBR, MO, RSA, US); 12 
km N of Caçador, Smith & Klein 9145 (HBR, MO, R, 
RSA, US); 9-17 km W of Bom Jardim da Serra to Sao 
Joaquim, Mun. Bom Jardim da Serra, Berry et al. 4510 
(n — 44), 4511, 4512, 4513 (n = 44) (all MBM, MO); 
Falkenberg 2269, 2270, 2273, 2275, 2277, 2279, 
2280 (FLOR); 9 km E of Bom Jardim da Serra on road 
to Lauro Müller, Mun. Bom Jardim da Serra, Berry et 
al. 4514-4516 (MBM, MO); Falkenberg 2285, 2288 
(FLOR); Bom Jardim da Serra, Eskuche 1639-63 (SI); 5 
km E of Sáo Joaquim, Mun. Sáo Joaquim, Falkenberg 
et al. 3774-3776 (FLOR); road Sào Joaquim to Uribici, 
Falkenberg et al. 3777 (FLOR; n = 44); Uribici, slopes 
and near summit of Morro da Igreja, Falkenberg et al. 
3778-3782 (FLOR); Uribici, km 90 of road to Braco do 
Norte, Falkenberg et al. 3786 (FLOR); 0.5-1.5 km N 
of Várzea, road to Bom Jardim, Falkenberg et al. 377 1, 
3772 (FLOR); Marombos, Mun. Curitibanos, Klein 3287 
(HBR, MO, RSA); Curitibanos, Berry et al. 4506 (MBM, 
MO; n = 44); Falkenberg 2247 (FLOR); 5 km W of 
Curitibanos on BR-470, Berry et al. 4508 (MBM, MO); 
Falkenberg 2253 (FLOR); Campo dos Padres, Mun. Bom 
Retiro, Rambo 60135 (PACA); Reitz 2342 (G, HBR, 
NY, U, UC, US), 5213 (PACA); Smith & Klein 7652 
(HBR, MO); Moratá, Ponte Miriad, Reitz 4698 (RSA, 
S); 11 km E of Faxinal dos Guedes, Smith & Klein 11853 
(HBR, MO, R, RSA, US); descent of Serra Geral from 
Bom Jardim to Lauro Müller, Berry et al. 4518 (MBM, 
MO); Falkenberg 2305, 2307 (FLOR); 22 km above 
Timbé do Sul, Serra da Rocinha, Mun. Timbé do Sul, 
border with Rio Grande do Sul, Falkenberg et al. 3758, 
3759, 3763-3765 (FLOR); Bituruna, Fazenda Etienne, 
Mello Filho 752 (R); BR-2, km 280, Pabst et al. 6132 
da MBM, R); ads & Pabst 6305 (RB); near Lajes, 
n. Lajes, Lindeman & Haas 3703 (U), Texeira & 
Texeira 3 (SP); Rambo 49602 (MO, RSA, S); Campo de 
eào, Mun. Sta. Cecilia, Reitz & Klein 14185 (HBR, 
Mb RSA); Ponte Alta do Sul, Mun. Ponte Alta, Reitz 
& Klein 11294 (HBR, MO, US); Sào Francisco de Malta, 
Mun. Sáo Joaquim, Reitz 6662 (HB, HBR, MBM, MO, 
RSA); Fazenda de Laranja, Sao Joaquim, Smith & Klein 
7652 (HBR, R, RSA, US); 8 km NE of Sáo Joaquim to 
Cruzeiro, Smith & Klein 10277 (HBR, MO, R, RSA 
US) Poco Preto, near Rio Timbó, Mun. Porto Dita. 
Smith & Reitz 8670 (HBR, MO, R, RSA, US); 8 km 
SW of Fuck on BR-116, Davidse et al. 11076 (MO, 
SP). RIO GRANDE DO SUL: Itaimbezinho, Parque Nacional 
Aparados da Serra, Mun. Cambará do hu o 
361, 362, 2360 (FLOR); Jarenkow 
(FLOR); Rambo 49363 bp RSA), put OUS 
53981 (SI); Pirani & Yano 840 (SP); Tainhas, Mun 
Cambará do Sul, Vianna in 10 Feb. 1957 (ICN #1674); 
10-11 km N of Tainhas on RS-020, Falkenberg et al. 
3746 (n= 44), 3747, 3748 (FLOR); 1-4 km N of Post 


Haas ie (MBM); Serra da Ro- 
ci Bom Jesus to Turvo, Lindeman & 
Haas 9389 (ICN, T Rambo 53880 (B); al 4278 


(B, PACA, SI) Bom Jesus, Falkenberg 1234 (FLOR); 
Ausentes, Bom Jesus, Falkenberg 1182, 1183 (FLOR); 
Cambará do Sul, Falkenberg 263, 1119 (FLOR); Rambo 
36223 (ICN); Faxinal, Mun. Cambará do Sul, Bonatto 
Lima in 2 Apr. 1083 (FLOR); Stehmann 450 (FLOR); 
Sobral 2813 (FLOR); Esmeralda, Est. Ecol. Aracuri, 
Jarenkow 60 (FLOR); Bom Jesus, Aparados da Serra, 
Pabst 10201 (RB); Rambo 8787 (PACA); Rio dos Tou- 
ros, near Bom Jesus, Rambo 8552 (PACA); Capela de 
Sáo José, Aparados da Serra, Bom Jesus, Smith & Klein 
5914 (R, US); Cambará, Sào Francisco de Paula, Rambo 
36223 (PACA, S); Fazenda Bernardo Velho, Aparados 
da Serra, Bom Jesus, Rambo 34864 (PACA, S); Aparados 
da Rocinha, Bom Jesus, Rambo 45354 (PACA, S), 32487 
PACA); Passo do Guarda, Bom Jesus, Rambo 51857 
(PACA, S, US); Fazenda Englert, near Sào Francisco de 
Paula, Rambo 54635 (B, PACA); near Montenegro, 
Rambo 2204 (PACA); Agua Azul, near Caxias, Henz in 
3 Jan. 1947 (PACA #33277); Cerro do Umbu, near 
Osório, Rambo 1284 (PACA); Caracol, near Canela, Em- 
rich in 6 Mar. 1946 (PACA #33277), in 22 Mar. 1954 
(PACA #54247); Rambo 11998 (PACA); Vila Oliva, 
near Caxias, Rambo 31024 (PACA), 31270 (R), 53880 
(PACA), 56621 (PACA), 55021 (B, PACA); Passo do 
Socorro, near Vacaria, Rambo 51576 (PACA, S), 51665 
PACA); 25 km N of Vacaria, Pedersen 12739 (MO); 
m S of Vacaria, Lindeman & Haas 3734 (U) 
Fazenda da Ronda, near Vacaria, Rambo 34864 (PACA, 
SI); Taimbé, Sào Francisco de Paula, Sehnem 5125 (SI); 
Farroupilha, Camargo 980 (PACA); Santa Rita, near 
Farroupilha, Rambo 40172 (CAS, PACA, RSA), 45781 
bso PACA, S), 45782 (PACA); Flores de Cunha, Schultz 
7 (ICN); Soledade nan Passo Fundo, Rambo 50039 
ERN Jaquirana, S de Paula, Rambo 51968 
PACA); Gramado- eie do Caracol, Schweiger in Feb. 
1924 (ICN #44812); Laje de Pedra, Canela, Miotto 50 
(ICN); Arroio Tatiuí, near Vila Assis, km 193 of BR-386, 
Miotto et al. 184 (ICN); Sào Francisco de Paula, Ferreira 
et al. 628 (ICN); no locality, Waechter 1942 (ICN); 
Rambo 9943 (PACA). 


~ 


— = 
© 


— 


Fuchsia regia subsp. reitzii is distinguished by 
its whorled, membranous leaves with gland-serrate 
margins and a matte surface having more notice- 
able tertiary venation than the other subspecies of 
F. regia. Compared with F. regia subsp. serrae, 
the only other Fuchsia that grows near populations 
of subsp. reitzii, the sepals are much less joined 
and spreading (instead of recurved), the stipules 
are much less prominent, the leaf bases are more 
acute, and the berries are rounder. This subspecies 
is named in honor of P. Raulino Reitz of the Her- 
bário Barbosa Rodrigues, Itajai, Santa Catarina, 
who has made great contributions to the knowledge 
of the flora of Santa Catarina. 

Subspecies reitzii is restricted to the planalto 
west of the coastal mountain ranges of southern- 
most Brazil. Its overall distribution in this area 
parallels that of Araucaria angustifolia, although 
it does not tolerate the shade of the dense conifer 
forests. Both species extend as far east as the edge 
of the coastal escarpments but end abruptly there 


580 


Annals of the 
Missouri Botanical Garden 


and do not appear on the seaward slopes or in the 
lower, coastal forest (Hueck, 1953 

Along the eastern edge of the planalto, where 
sharp drop-offs usually occur down to the Atlantic 
lowlands, the range of F. regia subsp. reitzii over- 
laps with F. regia subsp. serrae, which extends up 
from the moist, coastal slopes of the serras. A great 
deal of intergradation and variation exists in the 
Fuchsia populations of these areas, especially at 
the crest itself and just inland of the steep drop- 
offs. In some instances, the extreme local variability 
is indicative of hybrid swarms, while in other sites, 
odd variants are found that can extend beyond the 
normal limits of variation of either subspecies (see 
Appendix). 

A number of unusually pubescent individuals, 
usually with conspicuously tan or ferrugineous hairs, 
occur in populations near the southern and western 
extremes of the range of subsp. reitzii in Rio Grande 
do Sul. Collections from the Farroupilha area, such 
as Rambo 45781, 45782, and 40172 and Ca- 
margo 980, are uniformly pubescent, while others, 
such as Rambo 31024 from Vila Oliva near Ca- 
xias, and Rambo 51576 from Passo do Socorro 
near Vacaria, occur together with more typical 
le individuals of the subspecies. 


LITERATURE CITED 


AITON, W. a Hortus Kewensis, Volume 2. London. 
AVERETT, J. E., W. J. Hann, P. E. Berry & P. 
RAVEN. a Flavonoids and flavonoid evolution 
in Fuchsia (Onagraceae). Amer. J. Bot. 73: 
1534. 


BaiLgv, L. H. 1900. P ln. sae of American Horti- 
culture. Macmillan, New 

Berry, P. E. 1982. The sys MN and evolution of 
Fuchsia a ee (Onagraceae). Ann. Missouri 
Bot. Gard. 1-198. 

1085. "The systematics of the apetalous fuch- 

sias ‘of South America, Fuchsia sect. ies 

(Onagraceae). Ann. Missouri Bot. Gard. 72: 

251. 


STEIN, S. CARLQUIST & J. W. NOWICKE. 
1988. - Fuchsia pachyrrhiza, a new tuberous — 
and section of Fuc hsia from northwestern Peru. Sys 
Bot. 13 9 
BORGMEIER, T. 1937. A história da Rod VE mimes: 
de Frei Velloso. Rodriguésia 3(9): 77-96 
BREEDLOVE, D. E. 1969. The GaU of Fuchsia 
sect. visuri die (Onagraceae). Univ. Calif. Publ. 
Bot. 53: 


P. E. RRY & P. H. Raven. 1982. The 
Mexican a Cond American species of Fuchsia 


(Onagraceae). Ann. Missouri Bot. Gard. 69: 209- 


CANDOLLE, A. P. DE. 1828. Pr — Systematis Na- 
turalis € Vegetabilis. Volume 3. Paris. 

CARAUTA, J. 1973. The UL M Vellozo's Flora 
ME. and its festa date of publication. 
Taxon 22: 281-284. 


CHAUDHURI, S. K. 1956. Cytogenetic studies in the 
ee Ph.D. Dissertation. Univ. Man- 
chester, Englan 
CROWDEN, R. K., J. Warcur & J. B. HARBORNE. 1977. 
Anthocyanins of Fuchsia (Onagraceae). Phytochem- 
istry 16: 400-402. 


Curtis, W. 1836. s discolor. Port Famine Fuch- 
sia. Bot. Mat., t. 3498. 
Don, G. 


1832. A General History of the Dichlamydeous 

Plants. 2: o 

FaEcRI, K. & L. VAN DER PUL. 1979. The Principles 
of Pollination es 3rd edition. Pergamon, Ox- 
ford. 


FEUILLÉE, L. E. 1725. Journal des Observations Phy- 
siques, Mathematiques et Botaniques, t. II. Histoire 
des Plantes Medicinales du Perou et Chily: 64, pl. 


GARDNER, G. c Fuchsia alpestris. Mountain fuch- 
sia. Bot. Mag., 999: 
HANSEN, A. 1969. " Checklist of the vascular plants of 


the pura of Madeira. Bol. Mus. Munic. Fun- 
chal 24: 5-7 

HARBORNE, J. B. den. Distribution of anthoryanins in 
higher plants. Pp. 359-388 in T. Swain (editor), 
Chemical Plant Systematics. Academic Press, Lon 


don. 
Harrison, J. 1841. Fuchsia cordifolia. m leaved 
uchsia. Fl. Cab. Fl. Mag. 9: 241, pl. 105. 

HauMAN-Merck, L. 1912. Observations d'éthologie flo- 
rale sur quelques PEpPres argentines et chiliennes. 
Recueil Inst. Bot. 9: 1-39. 

Henpy, M. D. € D. PENNY. 1982. Branch and bound 
algorithms to determine minimal evolutionary trees. 
Math. Biosci. 59: 277-290. 

HENNIG, W. ier eni Systematics. Univ. 
Illinois Press, Urbana, Illino 

On the true Fuchsia coccinea of 

61. 

1989. Observations on 

polyploidy i in Fuchsia sects. Quelusia and Kierschle- 

geria (Onagraceae). Ann. Missouri Bot. Gard. 76: 

585-592. 


Hueck, K. 


Distribuigáo e habitat natural do 
Pinheiro do Paraná (Arauc ay gee Bol. 
Fac. Filos. Univ. Sao Paulo 1 


. 1972. As Florestas da EE do Sul. Po- 
ligono & EDUnB, São Paulo. 
KLEIN, R. M. 1978. Mapa duni eaa do Estado de 
Santa Catarina. Flora Ilustrada Catarinense 5. 
Aspectos pida da vegetação do 
Sul de Brasil. Sellowia 36: 5-54. 
LINDLEY, J. Fuchsia gracilis. Slender fuchsia. 
Bot. ib 10: t. 847. 
27a. dr hsia conica. Conical-tubed fuch- 
sia. Bat Reg., 62. 
1827b. n hsia gracilis; Pip 6 Many- 
Bot. Reg 
i : Mes Pb. 
fuchsia. Bot. Reg., 256. 
1835. Fuchsia discolor. Port Famine fuchsia. 
Bot. Reg., t. 1805. 
. 1840. Standish’s fuchsia. Bot. Re 
1841. Fuchsia radicans. Rosine TA 
ot. Reg., t. 66. 
Looser, G. 1928. Botánica Miscelánea. Algunas es- 
pecies leñosas escasas en el interior de la provincia 


ilowered slender fuchsia. 


Hie dew 


Volume 76, Number 2 
1989 


Berry 
Fuchsia sect. Quelusia 


a pM Revista Univ. Católica Chile 5: 523- 


TEN W. 1981. Der Araukarienwald von Süd- 
brasilien. Natur. & Mus. 3(2): 46-53. 
909. The first Fuegian collection. 


, Toronto. 
1983. Flora of Tierra del Fuego. Missouri 
Pouca Garden, St. Louis, Missouri. 
Munz, P. A. 19 A revision of the genus Fuchsia 
oo Pio. Calif. Acad. Sci., Ser. 4, 25: 1- 
138. 


Navas B., L. E. 1976. Flora de la Cuenca de Santiago 
de Chile. A rdg II. Ed. Univ. Chile. 

NowICKE, J. W., J. J. SKvARLA, P. H. Raven & P. E. 

BERRY 1984. A palynological study of the genus 

Fuchsia (Onagraceae). Ann. Missouri Bot. Gard. 71: 


Nozzoutto, 2 1970. Flower color in Fuchsia culti- 
s. d. J. Bot. 47: 1215-1217. 

Mugs E 184 3. dics exoniensis. The Exeter fuch- 
sia. Paxton's Mag. 

PockNarL, D. T. D. c.  MILDENHALL. 

ligocene-early Miocene spores a 

Southland, New NE New 
Paleontol. Bull. 51: 1- 

PRAGLOWSKI, E i. Raves, J. J. SkvarLaA & J. W. 
NOWICKE. Angiospermae: Onagraceae Juss. 
Fuchsieae & ju er World Pollen & Spore Flora 


1984. Late 
and pollen from 
Zealand Geol. Surv. 


12: l- 
RAMBO, B. 1951. A migração da selva higrófila no Rio 
Grande do Sul. ies Bot. Herb. “Barbosa Ro- 
drigues" 3: 55- 
see. 1956. A Um dora dos Aparados 
riograndenses. Sellowia 7/8: 235-2 
P.H. 19 


RAVEN, 79a. A aa of reproductive biology 


in Onagraceae. New Zealand J. Bot 575-593. 
1979b. Plate papae hv rus Hemi- 
sphere biogeography. n K. Larsen & L. 


Holm-Nielsen (editors), E Blass Academic 


Press, London. 


Onagraceae as a model of plant e 

hilot In: L. D. un & S. K. Jain. d rs 
Evolutionary Biology. A Symposium Hono 
Ledyard Stebbins. ras & Hall, Lo Aa 

REITER, V., JR. 1944. Notes on the history of Fuchsia 
breeding. J. Calif. Hort. Soc. (Dec. 1944-Jan. 1945): 

-192. 

Rerrz, R. 1961. Vegetação da a marítima de Santa 
Catarina. Sellowia 13(13): 1 15. 

ROEMER, J. J. Re dut de Plantis Hispanicis, 
Lusitanicis, Brasiliensibus. Nur er 

Ruiz, H. € J. Pavón. 1802 Flora Peruviana et Chi- 
lensis, Volume 3. Madrid. 

Ruscut, A. 1982. Beija-flores do estado do Espirito 

nto. Rios, Sào Paulo. 

SALISBURY, R. A. 1791. Icones E Rariorum De- 
scriptionibus mu Lon 

SIEBERT, A. & A. Voss. 1896. pud A. Vilmorin, Blu- 

mengartnerei, 3rd ad 1: 332. 

SimPSON, B. B. 1973. Contrasting modes of evolution 
in two groups of Perezia (Mutisieae: Compositae) of 
era Pee America. Taxon 22: 525-536. 

Quaternary ido ud a K high 
a tees ie of South America. Pp. 1 89 in 


W. E. Duellman (editor), The South [duca Her- 


d in Its Origin, Evolution and Dispersal. Mono- 
useum Nat. Hist. Univ. Kansas, no. 7 

Sims, e E^ Fuchsia coccinea. Scarlet fuchsia. Bot. 

pid 


Mag 97. 
SkvahtA n J, F. H. RAVEN | & J. PRAGLOWSKI. 
Pp. 


479 in I. K. Keen & J. Muller (editors), "The 
Soe Mien Significance of the Exine. Academic 
ress, London. 


, W. F. Cuissok & M. SHARP. 1978. 

An ultrastructural study of viscin Aare in Ona- 
graceae pollen. Pollen & Spores 20: 5-143. 

STEBBINS, G. L. 1985. Polyploidy, hivbridizations and 
the invasion of new habitats. Ann. Missouri Bot. Gard. 
72: 824-832. 

SWOFFORD, D. L. 1985. PAUP, Phylogenetic Analysis 
Using Parsimony. Version 2.4. Illinois Natural His- 
tory Survey, Champaign, Illinois. 

SYTSMA, K. J. & J. F. SMITH. 1988. DNA and mor- 
phology : comparisons in the Onagraceae. Ann. Mis- 
souri Bot. Gard. 75: 1217-1237. 

TRUSWELL, E. M. Antarctica: a terrestrial P una 
history. /n: R. J. Tingey (editor), The Geology of 
Antarctica. Oxford Univ. Press. (in press). 

VUILLEUMIER, B. 1971. Pleistocene changes in the fau- 
na and flora of South America. Science 173: 771- 

0 


WATROUS, L. & Q. WHEELER. 1981. The out-group 
comparison method of character analysis. Syst. Zool. 
30: 1-11. 


WEBB, C. D. G. LLovp. 1986. The avoidance of 
interference between the presentation of pollen and 
stigmas in en II. Herkogamy. New Zea- 

: 163-178. 

; 197 9. The introduction of fuchsia. J. 

iiy Hort. Soc. 100: 440. 

79. Fuchsia, a garden history. The Plants- 

: 181-186. 

aña, Y 1976. Co-pigmentation and the color change 

with age in petals of Fuchsia hybrida. Bot. Mag. 
(Tokyo) 89: 45-57. 

K. HAvasHr. 1967. Analysis of flower colors 

in Fuchsia hybrida in reference to the concept of 

co-pigmentation. Proc. Imp. Japan. Acad. 43: 316- 

321. 


ZINMEISTER, W. J. 1987. Cretaceous paleogeography 
of Antarctica. Palaeogeogr., Palaeoclimatol., Palaeo- 
ecol. 59: 197-206. 

APPENDIX. Natural hybrids in 

Fuchsia sect. Quelusia 


The main criterion used in detecting hybrids was 
the morphological intermediacy of several char- 
acters. In all cases, the putative parents were known 
to occur nearby. 


l. F. magellanica x F. lycioides 


F. rere var. parviflora W. J. Hooker, Bot. 
isc. 3: 309. TYPE: Chile. Manus 1832, Bridges 
199 ra K—Benth.; isotype, CGE). 


ecimens examined. CHILE. V REGION (VALPA- 
RAÍSO): Valparaiso, Bertero 1006 (G, ie in valleys near 
Valparaiso, Bridges s.n. , K— Hook 


Representative cultivated specimens. "CHILE: Juan 


Annals of the 
Missouri Botanical Garden 


Fernández Islands, Más Atierra, Meyer 9551 (MO). U.S.A 
California: Berkeley, Bracelin 1422 (BM). ZIMBABWE. 
Salisbury, Biegel 5549 (K). 


Fuchsia lycioides belongs to a different section, 
Kierschlegeria, and differs widely 
gellanica in its small, alternate leaves, spinose leaf 
bases, smaller flowers with included stamens and 
reflexed sepals, and generally occurs in much drier 
habitats. Both species are tetraploid (Hoshino & 
Berry, 1989) and have been shown to be closely 
related by chloroplast DNA restriction site analysis 
(Sytsma & Smith, 1988). The ranges of the two 
species barely overlap in the hills around Valpa- 
raiso, where several natural hybrids were collected 
in the early 1800s. These plants are characterized 
by small flowers with larger, convolute petals than 
those of F. lycioides. The leaves are larger and 
more often opposite than in F. lycioides, with the 
characteristic dentation of F. magellanica. They 
also lack the spinose leaf bases typical of F. ly- 
cioides. The natural habitats where the early hy- 
brids were collected have probably been totally 
eliminated by urban growth. 

Hybrids of these two species appear quite often 
in cultivation and have sometimes been confused 


. ma- 


with the true F. lycioides, as in the cytological 
study of Chaudhuri (1956). 


2. F. regia subsp. regia X F. alpestris 
Specimens examined. BRAZIL. RIO DE JANEIRO: dada 


Dubois, Santa Maria Madalena, Santos Lima & Bra 
14240 (B, RB); Plowman & de Lima 12885 (F, M 


Both parental species occur locally on Pedra 
Dubois; the specimens cited above are intermediate 
in their short, fine pubescence, petiole length (6— 
10 mm), and flower size (floral tubes 10-12 mm 
long). 


3. F. regia subsp. serrae X F. brevilobis 
BRAZIL. SAO PAULO: Reserva 


iguel Arc- 
MBM). PARANÁ: 13 


Specimens examined. 
Carlos Botelho, between 


°, Lange train station and Prainhas, 310 
m, Cordero et a in 16 Jan. 1987 (MBM; n = 3 


Fuchsia brevilobis is sympatric with F. regia 
subsp. serrae over most of its range but usually 
grows at lower elevations. These taxa are unique 
in the section in having their sepals connate for 
more than one-third of their length, but F. brevi- 
lobis has more membranous leaves that are nar- 
rower, shorter-petioled, and more pubescent than 
F. regia subsp. serrae, while the latter has more 


prominent, persistent stipules. The flowers of F. 
regia subsp. serrae also differ from F. brevilobis 
in their four-ridged buds, glabrous, nitid surface, 
and flaring sepal tube. The intermediate individuals 
cited above are from three different localities where 
the parent species are locally present. In the Ca- 
pivari locality, the same hummingbird was observed 
visiting flowers of both species in succession. 

Further experimental work on artificial hybrids 
of these two taxa is desirable, since only tetraploid 
counts have been obtained from F. brevilobis, and 
all individuals examined of F. regia subsp. serrae 
have been octoploid (Hoshino & Berry, 1989). The 
count obtained from Cordeiro et al. in 16 Jan 
1987 was tetraploid, indicating that it may only 
be a variant of F. brevilobis, and that of Berry et 
al. 4448 was octoploid, as found in F. regia subsp. 
serrae. 


4. F. regia subsp. regia X F. campos-portoi 


Specimens examined. BRAZIL. RIO DE JANEIRO: Ita- 
tiaia, just above Hotel Brocken-Holsene, road to Abrigo 
Rebougas, 2,340 m, Berry et al. 4440 (MO, RB; n = 
red oe Agulhas Negras, Lanstyak 253 (RB 
#61360). 


Fuchsia campos-portoi is endemic to the high- 
altitude campos of Mount Itatiaia and is very dis- 
tinctive in its small, narrow, strongly serrate leaves 
and small, short-tubed flowers. Fuchsia regia is 
common in forests at lower elevations on the same 
mountain range and occurs together with F. cam- 
pos-portot in thickets near the treeline above 2,200 
m. Both specimens cited above are intermediate in 
leaf size, number of secondary veins, and floral 
features, such as the ratio of tube to sepal lengths. 
Meiosis in buds of Berry et al. 4440 was normal, 
with the same chromosome number as recorded in 
local parental populations. 


5. F. regia subsp. regia X F. coccinea 


cimens examined. BRAZIL. MINAS G ERAIS: Serra 


r (n 
— 22), 4547, 4548 (MO, RB); Falkenberg 3380 (FLOR); 
Mello Barreto 9168 (R); Serra da Caraga, Mello Barreto 
7157 MH). 


Fuchsia coccinea typically grows in thickets 
amid the rocky, open campo of the highest peaks 
in Minas Gerais. On the Serra do Itacolomi, fire 
has destroyed much of the woody vegetation on 
the highest slopes; F. coccinea is now very rare 
in that area, and intermediate forms with F. regia 
are common. In the series of Berry 4545-4550 
and Falkenberg 3380-3387, some of the individ- 


Volume 76, Number 2 
1989 


Berry 583 
Fuchsia sect. Quelusia 


uals were treated under F. coccinea, but several 
of them show signs of introgression with F. regia, 
such as leaves that are larger or more coriaceous 
and rounder-based than individuals of isolated pop- 
ulations of F. coccinea. The collections cited above 
are clearly intermediate between the two species 
in key characters, such as petiole length, degree 
of leaf pubescence, and leaf shape. Both species 
are tetraploid in this locality, and the one hybrid 
examined cytologically was tetraploid with normal 
bivalent formation. 

Both species are also known together from the 
Pico do Itambe, farther north in the Serra do 
Espinhago. Although the collections of Anderson 
et al. 35832 and Furlan et al. 3058 were treated 
under F. coccinea, they both have unusually thick 
leaves for that species, as well as rounded rather 
than cordate leaf bases; these traits may be t 
result of introgression with sympatric populations 


q) 


of F. regia. 


6. F. regia subsp. regia X F. glazioviana 


Specimens examined. BRAZIL. RIO DE JANEIRO: Mor- 
ro da Nova Caledônia, outskirts of Nova Friburgo, 1,810- 
1,950 m, Berry et a 4427 (MO, RB), 4429 
(RB), 4430 (MO, RB; n = 22). 

As in the previous two cases, F. glazioviana is 
a local, high-elevation endemic that is sympatric 
with F. regia near the former's lower altitudinal 
limits. Fuchsia glazioviana has small, densely 
packed leaves and smaller, narrower flowers than 
F. regia. Most of the hybrids were found along 
roadsides in cleared areas, and one large patch of 
variable individuals was seen in an exposed area 
near the upper limit of the cloud forest. Local 
populations of both species are tetraploid, as was 
the one intermediate individual counted. 


7. F. regia subsp. serrae X F. regia subsp. regia 


Specimens examined. BRAZIL. SÀO PAULO-RIO DE 
JANEIRO BORDER AREA: Reserva Florestal da Bocaina, Su- 
cre et al. 2932 (AAU, MO, RB, US); Serra da Bocaina, 
road to old Estação de Fruticultura iy IBDF park HQ), 
Duarte 7689 (MO, RB); Handro SP); 
thy & Vital 1176 (MO; n = baits bl Rio Bono: 
Serra da Bocaina, 1, Ha 1,100 m, Martinelli et al. 777 

nanal, Sertao do Rio Vermelho, E of Bocaina, 


(RB) Ba 
Brade 15269 (RB); bap Bocaina, Pals 4744 (MO). 


Populations of F. regia subsp. serrae and subsp. 
regia converge and intergrade extensively on the 
upper, seaward slopes of the Serra da Bocaina, 
close to the Rio de Janeiro-Sào Paulo border. 
Fuchsia regia subsp. regia reaches its southern- 
most limits in the Serra da Bocaina, occurring 
mostly at altitudes of 1,100-1,550 m on both sides 


of the Serra. Fuchsia regia subsp. serrae occurs 
exclusively on the lower, seaward slopes of the 
same range between 500 and 1,150 m, part of an 
almost continuous distribution down the coastal 
slopes of the Serra do Mar and Serra Geral as far 
as Rio Grande do Sul. 

Varied intermediate forms between the two sub- 
species occur between 1,150 and 1,350 m on the 
upper slopes of the Serra da Bocaina, along the 
road from Cunha to Paraty. Most of the fuchsias 
in this area, including those cited in the above list, 
are unusually pubescent for F. regia. The degree 
of sepal connation and the thickness and persis- 
tence of the stipules, which generally distinguish 
these taxa, vary widely between individuals in this 
area, whereas populations higher up on the Serra 
and further inland are largely glabrous and fall well 
within the normal range of variation of F. regia 
subsp. regia. Hoshino = Berry (1989) reported 


duals of Fuchsia from 


tetraploid and octoploi 
this area, further ee the intergradation of 
the basically octoploid subsp. serrae and the mostly 
tetraploid subsp. regia. Hexaploid hybrids should 
be sought in mixed populations. 

Although F. regia subsp. pae is not found 
a popra ations of 


ove Ubatuba ap- 
proximately 60 km south of e Bocaina massif. 
Tetraploid and octoploid individuals occur side by 
side at the Alto da Serra near the crest of the 
mountain (Hoshino & Berry, 1989), again with the 
unusual pubescence for F. regia and considerable 
variability in such traits as the degree of stipule 
development. In most other features, however, these 
individuals resemble F. regia subsp. serrae more 
closely than subsp. regia. 


8. F. regia subsp. serrae X F. regia subsp. reitzii 


BRAZIL. qu CATARINA: top 
1 denm Lauro 

m, Berry 
et al. 4517, 4519 (MBM, MO); Falkenberg 1302 (FLOR); 
Rio Caveiras a Lajes, Lutz in 29 Dec. 1949(R #116687); 
Serra da Rocinha, border of Santa Catarina and Rio 
Grande do Sul, 1,250-1,280 m, 22 km above Timbe do 
Sul, Falkenberg et al. 3762 (FLOR, MO; the whole series 
of Falkenberg et al. 3756-3766 is a variable series of 
mostly intermediate variants); Rambo 4604 (PACA); NW 


ecimens examined. 


SUL: Itaimbezinho, Aparados da Serra, Falkenberg 261, 
264, 275, 354, 357 (FLOR); Pirani & Yano 839 (SP); 
Rambo 49363, 50150 (PACA); Caracol, near Canela, 
Emrich in Feb. 1951 (PACA 50187); Lindeman & Haas 

n. (ICN #21790); 18 km N of Canela, Salto da Fer- 
radura, Schultz s.n. (ICN #21032); Bom Jesus, Fal- 


Annals of the 
Missouri Botanical Garden 


pA ña 150 (FLOR); Taimbe, Sáo Francisco de Paula, 
Rambo 49363 (SI); Fazenda Bernardo Velho, Aparados 
da E Bom Jesus, Rambo 34864 (SI). 


Both of these two subspecies are morphologically 
and ecologically distinct over most of their ranges, 
with F. regia subsp. reitzii confined to thickets in 
the southern planalto and F. regia subsp. serrae 
inhabiting the wet, seaward slopes of the coastal 
mountain ranges. Fuchsia regia subsp. serrae has 
nitid, subcoriaceous leaves with thick stipules and 
flowers with strongly connate, recurved sepals, while 
subsp. reitzii has thinner, more whorled leaves with 
serrate margins, thin stipules, and flowers with 
spreading, slightly connate sepals. 

À narrow ecotone occurs all along the crest of 
the Serra do Mar and the Serra Geral in Santa 
Catarina and Rio Grande do Sul as the high rolling 
plains end and the steep coastal ranges drop abrupt- 
ly to the Atlantic lowlands. Along this belt, which 
occurs sometimes just at the crest of the ridges 
and other times extends a few hundred meters or 
several kilometers inland, the two subspecies of F. 
regia converge and intergrade in a bewildering 
variety of variants, some of them exceeding the 
normal limits of variation of either parental sub- 
species. 

The narrowest ecotone observed was at the Serra 
da Rocinha, on the Santa Catarina- Rio Grande do 
Sul border. In this area, the planalto is covered 
right to the edge by grassy campos, followed by 
steep drop-offs covered by a special “Aparados” 


flora adapted to the persistent rains and mists from 
the rising coastal air masses (Rambo, 1956). Along 
the rim of the serra cliffs, between 1,250 and 1,280 
m, hybrid swarms of F. regia subsp. serrae and 
reitzii occur together with occasional plants that 
can be distinguished as parental, such as those 
included in the collection series of Falkenberg et 
al. 3756-3766. 

Other areas where similar series of intermediates 
have been observed and collected include the Apa- 
rados da Serra at Itaimbezinho, in Rio Grande do 
Sul, and the area around the crest of the Serra 
Geral between Bom Jardim da Serra and Lauro 
Müller in Santa Catarina. Near the town of Urubici, 
Santa Catarina, plants of F. regia subsp. serrae 
occur farther inland from the crest of the Serra 
Geral, around the base of the Morro da Igreja and 
Campo dos Padres, the highest points in Santa 
Catarina. Where the two subspecies occurred sym- 
patrically, as in the collection series of Falkenberg 
et al. 3787-3791 (FLOR, MO; just W of the crest 
of the ridge between Urubici and Brago do Norte, 
at 1,030 m), some unusual variants of subsp. ser- 
rae were found with flowers less than half the 
normal size (43790). 

oth subspecies are octoploid and are charac- 
terized by variable numbers of multivalents during 
meiotic metaphase (Hoshino & Berry, 1989); the 
one intermediate individual examined had the same 
chromosome number and type of meiotic config- 
uration. 


OBSERVATIONS ON 
POLYPLOIDY IN FUCHSIA 


Takuji Hoshino? and Paul E. Berry? 


(ONAGRACEAE)! 


ABSTRACT 


Chromosome numbers are reported for all eleven taxa currently recognized in Fuchsia sect. Quelusia and for F. 
lycioides, the sole member of sect. AS Results are based on the study of individuals from 103 native 
populations and 13 cultivated plants. Most species and sections of Fuchsia are diploid (n = x = 11), but sects. 
Quelusia and Kierschlegeria are entirely de ba including the only naturally occurring octoploids in the genus. 
Polyploidy in these sections is always associated with 3(4)-aperturate pollen grains, instead of the normal 2-aperturate 
condition in the genus. All ona counts obtained from F. lycioides and from seven of the nine species in sect. 
Quelusia were tetraploid, but intraspecific variation in ploidy level was found in two Brazilian species of sect. Quelusia. 
The only wild individual examined of F. alpestris was tetraploid, whereas a cultivated specimen of this species from 

urope was octoploid. In the widespread, polytypic F. regia, all individuals examined of subspecies reitzii and serrae 
were octoploid, but both tetraploid and octoploid populations were found in subsp. regia. Both ploidy levels were also 
d in the extreme northern range of subsp. serrae, in individuals that were morphologically intermediate with 


. This situation suggests that octoploidy has evolved recently and possibly repeatedly in s 


sect. Quelusia, 


whereas tetraploidy probably originated in a common ancestor to sects. Quelusia and Kierschlegeria. 


Fuchsia is one of the largest genera of the 
Onagraceae, with over 100 species occurring most- 
ly in tropical montane cloud forests. It is placed 
in its own tribe, Fuchsieae, on the basis of its fleshy 
fruits and mostly 2-aperturate pollen. Chromo- 
somes in Fuchsia are large and are heterogeneous 
in size for the family, with interphase nuclei having 

ycnotic areas that are variable in number, size, 
and density (Kurabayashi et al., 1962); these are 
unspecialized features shared with Lopezia and 
Circaea. All species examined in the genus have 
the basic chromosome number of the family, x — 
11 (Raven, 1979), and none have the translocation 
systems or aneuploidy that are characteristic of 
other onagraceous genera such as Oenothera and 
Clarkia. 

In recent efforts to improve our understanding 
of the relationships of the different species and 


sections of Fuchsia, the distribution of chromosome 
numbers has been determined for as many species 
as possible in the genus, together with extensive 
surveys of the palynology and foliar flavonoid 
chemistry of most taxa (Nowicke et al., 1984; 
Averett et al., 1986). To date, diploid chromosome 
counts (n — 11) have been obtained for 69 species 
and tetraploid counts (n = 22) for 10 species 
(Breedlove, 1969; Breedlove et al., 1982; Berry, 
1982, 1985; Berry et al., 1988; Hoshino & Berry, 
1988). Of the ten sections currently recognized in 
the genus (Berry et al., 1988), six are entirely 
diploid. The two largest sections, Fuchsia ( 

species) and Hemsleyella (14 species), are pre- 
dominantly diploid, but sect. Fuchsia has six known 
tetraploid or partly tetraploid species (Berry, 1982), 
and sect. Hemsleyella has two tetraploid species 
(Berry, 1985). This paper presents the results of 


' Supported by grants to P. Berry by 


the American Philosophical Society, the Consejo Nacional de Investigaciones 


Cientificas y Tecnológicas (CONICIT, Venezuela) = Conselho Nacional de Desenvolvimento Científico e Tecnológico 


(CNPq, Brazil), as well as a U.S. Nat 
to 

extensive transect in Chile to collect seeds of m 
and P. Raven for helpful comments on a E beni this 


? Departamento de Biologia de Organ 


eiman-Dietiker, n 2 Ri 


We are especially grateful 


making ie initial pe counts in this study and to C. Marticorena, who made an 
any Fuchsia populations. Live seeds or fixed buds were also kindly 


uelme. We also thank D. Falkenberg, P. Goldblatt, 


y of LAN p 1 Ridai-Cho, Okayama 700, Jap 
mos, dp m rsidad Simón Bolivar, Apartado 89000, Ph AP 1081, Venezuela. 
S.A 


Current address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U. 
ANN. MISSOURI Bor. GARD. 76: 585-592. 1989. 


586 


Annals of the 
Missouri Botanical Garden 


ABLE | reamed of chromosome numbers in Fuch- 
sia sect. Quelusia 


Gametic 
Chrom 
some 
Taxon Number 
F. alpestris Gardner 22, 44 
F. bracelinae Munz 22 
F. brevilobis P. Berry 22 
F. campos-portoi Pilger & Schulze 22 
F. coccinea Dryander 22 
F. glazioviana Taubert 22 
F. hatschbachii P. Berry 22 
F. magellanica Lamarck 22 
F. regia (Vellozo) Munz subsp. regia 22, 44 
F. regia subsp. reitzii P. Berry 
F. regia subsp. serrae P. Berry 44 
€ 
y 
t 
a y 
4 
4 > 
a 31 
* d 
^ t 
wW’ c 
€b 
"pm 
e 
* 
A oe 
e 
^ 
Sa 
=< o f 
y. v x 
"d 
‘ > 5 * 
e. o y? 
MA a 
, a EF» 
FIGURES 1-4. 


a comprehensive survey of the cytology of all 11 
taxa of sect. Quelusia, concurrently with a sys- 
tematic revision of the section (Berry, 1989). It 
also presents the first published counts for Fuchsia 
lycioides, the sole member of sect. Kierschlegeria. 
The only previous reports on chromosomes in 
sect. Quelusia were from F. magellanica. The 
first report by Warth (1923) was n = 22 for F. 
coccinea, but photographs in a subsequent publi- 
cation (Warth, 1925) showed that this was a mis- 
identification of F. magellanica. Johanssen (19292, 
b) reported diploid counts of n = 11 for two hor- 
ticultural selections of F. magellanica, but no 
vouchers or illustrations were provided to verify 
the identity of his plants, which were very likely 
misidentified (P. Raven, pers. comm.). A report of 
n — 44 in F. magellanica by Haque (1952) also 
lacked vouchers. Chaudhuri (1956) made counts 
of n — 22 in F. magellanica and noted that it had 


u 


A > r a 
L4 


A « 
e Ñ " , 
PLA 
— , 
EJ » 
6 > 
0 * 
e 
es e 2 
* 


Photomicrographs of meiotic chromosomes of Fuchsia sect. Quelusia. —1. Metaphase I in F. 


bracelinae (Berry et al. A Rum = 44 (2211). — 2. Diakinesis in F. campos-portoi (Berry et al. 4435). 2n = 44 


(2211). — 3. Metaphase I in F. 


cinea (Berry et al. 4458). ca = 44 (2211). —4. Metaphase I in F. regia subsp. 


regia (Berry et al. 4544). ee = a (22II). Scale bars = 10 u 


Volume 76, Number 2 


Hoshino & Berry 587 


1989 Polyploidy in Fuchsia 
Pd 
» een T 
e M ES I p : 
[4 . r 
dt^ j d \ Vis s w ' ; 
A- 
1 x Wy ^ Ew: x 
T $ 
& A 4 i 
5 ó 
2 L^ 4 
r ; E HIT oí 
‘ 8 s. 
r yn » 3 
9 cc ; Fog x 
$ $} ^ e 
- =~ 4^ v i RA he 
/ $ i 
2% 
s 4 x ` 
; + AE * 
7 ¿A 8 
FIGURES 5-8. Photomicrographs of meiotic metaphase chromosomes of Fuchsia sect. Quelusia. —5. Fuchsia 


regia PPS Eip (Berry & o a 2n = 88 (15IV € 14ID.— 
. Fuchsia regia 


UL regia subsp. a n hybrid with subsp. serrae, from Berry & Falkenberg 4517). 2n 


et al. 4 = 88 (9IV € 


101). — 
= 88 n & 14ID. Scale bars = 


distinctly shorter chromosomes than several diploid 
species he examined in the genus. Kurabayashi et 
al. (1962) examined two different collections of F. 
magellanica, which they also found to be tetraploid 
with short chromosomes. Although Chaudhuri 
(1956) reported a tetraploid count for F. lycioides, 
no voucher specimens were provided, and his de- 
scription of this entity clearly indicates that it was 
a hybrid, probably of F. magellanica x F. ly- 


cioides parentage. 


MATERIALS AND METHODS 


Meiotic chromosomes were examined in pollen 
mother cells of young buds fixed in Carnoy's so- 
lution (3:1 absolute ethanol to glacial acetic acid) 
for 24 hours, then transferred to 70% ethanol for 
storage. Anthers were squashed in 1% aceto-or- 
cein. 

Somatic chromosomes were studied in root tip 
apices, which were pretreated in 0.002 M aqueous 
8-hydroxyquinoline for 6 hours at 5?C, then fixed 


6. Fuchsia regia subsp. serrae (Berry 
. serrae (Berry et al. 4493). 2n — 88 (17IV & 


subsp 


in Carnoy's solution and macerated in 10% HCl 
for 6 minutes at 60°C. The root tips were stained 
in 1% aceto-orcein for 10 or 20 minutes and then 
squashed. 

To establish pollen fertility, pollen grains were 
stained with Alexander's solution (Alexander, 1969). 


RESULTS 


Chromosome counts from 40 different popula- 
tions of native and cultivated individuals of F. ma- 
gellanica were consistently tetraploid (n — 22; 
Table 1, Appendix). The populations sampled cov- 
ered most of the range of this widespread species 
of southern Argentina and Chile, leaving little doubt 
that it is cytologically uniform throughout its range. 

Counts of all other taxa in sect. Quelusia rep- 
resent their first validated chromosome reports (Ta- 
ble 1). Seven Brazilian species, including the newly 
described F. brevilobis and F. hatschbachii (Ber- 
ry, 1989), are all tetraploid, based on at least two 
counts for each species. The Chilean F. lycioides 


588 


Annals of the 
Missouri Botanical Garden 


om 


= 


Chromosome counts 
in Fuchsia regia 


* Tetraploids 
O Octoploids 


ae so 
de n 


RE 9. Geographical distributions of the different cytotypes of Fuchsia regia in southeastern Brazil. Pairs of 
capital letters refer to state abbreviations, RJ — Rio de Janeiro, etc. Contour lines indicated at 800 m. 


(sect. Kierschlegeria) is also tetraploid. Normal 
bivalent formation was found in all tetraploid col- 
lections examined (Figs. 1-4). In the Brazilian F. 
alpestris, the single naturally occurring individual 
counted was tetraploid, but a plant of this species 
cultivated in Europe was octoploid. 

Fuchsia regia, the most widespread species of 
the genus in Brazil, has tetraploid and octoploid 
populations (see Appendix). Populations from the 
southern half of its range were entirely octoploid, 
including all populations counted of subspecies 
reitzii and serrae and one naturally occurring hy- 


brid between them. In most of the octoploids, nu- 
merous quadrivalents and bivalents were formed 
(Figs. 5-8). 

Populations of F. regia subsp. regia with both 
tetraploid and octoploid individuals were found in 
northern Sào Paulo and southern Rio de Janeiro 
states (Appendix, Figure 9). The southernmost tet- 
raploid individuals of F. regia came from north- 
central Sào Paulo State. These collections are 
morphologically intermediate, probable hybrids be- 
tween subspecies regia and serrae. They also 
occurred sympatrically with octoploid individuals 


Volume 76, Number 2 
1989 


Hoshino & Ber 


ry 
Polyploidy in Fuchsia 


FicunES 10, 11. 
serrae (2n = 88) (Ramamoorthy & Hatschbach 830 e 
(2n — 66), artificial hybrid between Ramamoorthy 6 


ru regia, 2n — 4 


Photomicrographs of pollen grains viu Fuchsia sect. n ies Fuchsia regia subsp. 


x F. regia subsp. serrae 


. Fuchsia regia subsp. r 
4) and | a thun Hatschbach 


830 (subsp. serrae, 2n — 88), cultivated at Missouri j| e Cadea, M2588. Scale bars = 150 u 


of F. regia subsp. serrae, but no hexaploid indi- 
viduals were found in these areas. In Minas Gerais 
and other parts of Rio de Janeiro, all populations 
of subsp. regia sampled were tetraploid. The dis- 
tribution of the different cytotypes of F. regia is 
shown in Figure 9 

Although meiosis was not observed in an artificial 
hexaploid hybrid between a tetraploid plant of F. 
regia subsp. regia and an octoploid one of subsp. 
serrae, more than 50% of the pollen grains of the 
hybrid were aborted. Both parents, however, showed 
fully era and apparently viable pollen grains 

l 


(Figs. 


DISCUSSION 

Sections Kierschlegeria and Quelusia are the 
only entirely polyploid sections of Fuchsia. In both 
groups, polyploidy is associated with 3-aperturate 
pollen, considered to be derived from the more 
common 2-aperturate condition in the genus (Now- 
icke et al., 1984). Despite differences between the 
two sections in viscin thread type (Nowicke et al., 
1984), seed number and anatomy, sexual systems, 
and other morphological features (Berry, 1982), a 


preliminary chloroplast DNA restriction site anal- 
ysis by Sytsma & Smith (1988) has shown the two 
sections to be more closely related to one another 
than to any other section of the genus. The results 
of a separate cladistic analysis of the species of 
these two sections by Berry (1989) were consistent 
with the hypothesis that tetraploidy evolved in a 
common ancestor. Since the other eight sections 
of the genus are predominantly diploid, e 
the South Pacific sect. Skinnera and al zen- 
tral American sections, it is clear that no exiis 
member of sect. Quelusia or sect. Kierschlegeria 
could have directly given rise to those groups. 
Octoploidy in natural populations of Fuchsia is 
known only in sect. Quelusia, where it has been 
detected primarily in F. regia, a species that also 
has tetraploid populations. Quadrivalents are com- 
mon in the octoploid individuals examined, indi- 
cating a close chromosomal homology between the 
genomes that came together. Both the presence of 
different ploidy levels and the varying numbers of 
multivalents in F. regia indicate a recent origin of 
octoploidy in the genus. Whether the octoploidy 


detected in F. alpestris originated naturally or 


590 


Annals of the 
Missouri Botanical Garden 


under cultivation is not known, but the extent of 
the two different chromosome numbers needs to 
be examined further in nature. Cases of intraspe- 
cific polyploidy are widespread in the Onagraceae 
and have been recorded in over 40 species and 11 
genera (Lewis, 1979). 

The high level of pollen abortion in the artificial 
hexaploid between a tetraploid and an octoploid 
subspecies of Fuchsia regia is indicative of irreg- 
ularities in meiotic pairing and disjunction, which 
may explain the absence of naturally occurring 
hexaploids among the individuals sampled in areas 
where tetraploids and octoploids both were found. 
Areas where tetraploid and octoploid populations 
of F. regia are known to occur sympatrically, such 
as the Serra da Bocaina and Campos do Jordào in 
northern Sao Paulo state, and the Itatiaia massif 
and the Serra dos Orgaos in Rio de Janeiro, should 
be sampled more intensively for naturally occurring 
hexaploids. 

Within the Onagraceae, the pattern of poly- 
ploidy in Fuchsia sect. Quelusia is most similar to 
that of Ludwigia, where polyploidy has been a 
frequent and important part of the evolutionary 
strategy in most of the genus, with numerous cases 
of intraspecific polyploidy (Raven & Tai, 1979). 
Unlike the situation in tribe Onagreae, where poly- 
ploidy is associated with autogamy and aneuploidy 
in annual species, aneuploidy is unknown in Fuch- 
sia and Ludwigia, and polyploid taxa are perennial 
and predominantly outcrossing. This clearly shows 
that the effects of polyploidy can vary widely in 
related genera, depending on factors such as the 
longevity of the species and the habitat in which 
they occur. 


LITERATURE CITED 

ALEXANDER, M. P. Differential e of aborted 
and nonaborted pollen. Stain Technol. 44: 117-122. 

AVERETT, J. E., W. H. HAHN, E & P.H. 
RAVEN. 1986. Flavonoids dti flavonoid ore 
in iius (Onagraceae). Amer. J. Bot. 5- 
15 


BERRY, P E. 1982. The systematics and evolution of 
dg sect. Fuchsia (Onagraceae). Ann. Missouri 
Bot. 9 198. 

The systematics of the apetalous fuch- 

sias ets south America, Fuchsia sect. oo 

(Onagraceae). Ann. Missouri Bot. Gard. 72: 

251. 


1989. A systematic revision of Fuchsia sect. 
mess (Onagraceae). Ann. Missouri Bot. Gard. 76 
584. 


——— . STEIN, S. CARLQUIST & J. W. NOWICKE. 
1988. Fuchsia pachyrrhiza, a new tuberous species 
and gue of Fuc hsia from northwestern Peru. Syst. 
Bot. 92. 

LE >». E. 1969. The systematics of Fuchsia 
sect. d (Onagraceae). Univ. Calif. Publ. 
Bot. oe 

RRY & P. H. Raven. 1982. The 

Mexican and ue American species of Fuchsia 

t for sect. Encliandra. Ann. Mis- 

ri Bot. Gard. 69: 209-234 

nus DHURI, S. K. 1956. Cytoger retic studies in E 
genus Fuchsia. Ph.D. Thesis. University of Mar 
chester, U.K. 

HAQUE, A. 1952. Chromosome counts of species and 
varieties of garden plants. Annual Rep. John Innes 
Hort. Inst. 41: 47-50 

Hosuivo, T. & P. E. Berry. 1988. Chromosomal ob- 
reir on Fuchsia species and artificial hybrids. 

. Missouri t Gard. 75: 1153-1154 

TN edd D. A 929a. New chromosome numbers 
in the TUM Amer. J. Bot. 16: 595-597. 

929b. Proposed phylogeny of the Onagra- 
ceae based erg on number of chromosomes. 
Proc. Natl. Acad. U.S ey 14: 882-885. 

KURABAYASHI, M., H. Lewis & P. H. Raven. 1962. 


comparative study of mitosis in the Onagraceae. Amer. 
6. 


. Bot. 49 2 
Lewis, W. H. 1979. Polyploidy in pao piae dicot- 


yledons. Pp. 241-268 in W. 
ploidy. Biological Relevance. 


(editor), Poly- 


e Press, New 


OrK. 

Nowicke, J. W., J. J. SkvarLa, P. H. Raven & P. E. 
BERRY. 1984. A palynological survey of the genus 
d (Onagraceae). Ann. Missouri Bot. Gard. 71: 


RAVEN N, P. H. 1979. A survey of purs end 
in Onagraceae. New Zealand J. B : 575-59: 
—— Ar W. TAL 1979. a al chromo- 
somes in Ludwigia (Onagraceae). Ann. Missouri Bot. 

Gard. 62-879. 

SytsMa, K. J. & J. F. Smith. 1988 [1989]. DNA and 
morphology: pg in the Onagraceae. Ann. 
Missouri Bot. Gard. 75: 1217-1237. 

WARTH, G. TN "Über Fuchsien mit verschieden ge- 
emus Pollen und ps ie romosomen- 
zahl eutsch. Bot. . 41: 281-285 

925. To. histologische und stam- 

DRESS tliche Fragen aus der Gattung Fuchsia. 

Z. Indukt. Abstammungs- Vererbungsl. 38(3): 200- 
251. 


APPENDIX I. mosome ia in Fuchsia 
sects. Quelusia and Kierschlegeri 


Fuchsia alpestris Gardner. M with 7 
RAZIL. RIO DE JANEIRO: Teresópolis, Berry et lal Ar 
Fuc hsia alpestris Gardner. Populations with n = 44. 
ULTIVATED: The Netherlands, from plants long in cul- 
varian $ in Europe, Berry & Brako 001-80. 


‘All counts were meiotic, except somatic counts indicated by *2n.^ 


both plants of F. lycioides; es seria 


aven counte 
d a. nce. 


. Ramamoorthy. 
except tho ose with a cite 


nly confirmed sai 
included. Unless otherwise indicated, voucher specimens are 


* Counts of Ramamoorthy do were made 
counts were made by the authors, 
icd orts wit chers or identifiable vie siis were 
the Missouri Botanical Garden (MO). 


Volume 76, Number 2 
1989 


Hoshino & Ber 591 


ry 
Polyploidy in Fuchsia 


Fuchsia bracelinae Munz (n = 22). 

AZIL. ESPÍRITO SANTO: Pico da Bandeira, Berry et 
al. 4: pe , 153 2. MINAS GERAIS: Pico da Bandeira, Plus 
et al. 


Fuchsia brevilobis P. Berry (n = 22). 
E of Sao Paulo- Paraná 
R-116, Davidse et al. 10902 (n — 


of [nen Berry 
Porto de Cima, Berry & Juarez 4495; above Prainhas, 
Cordeiro et al. 215 (MBM); below Estagao Eng. Lange, 
road to Prainhas, Cordeiro et al. 218 (MBM); Mun. 
Antonina, Cacatu, Hatschbach et al. 50788 (MBM). 


Fuchsia campos-portoi Pilger & Schulze (n — 22). 
BRAZIL: RIO DE JA NEIRO: ink Berry et al. 4435, 
Ramamoorthy & de 


Fuchsia xU SPORE X F. regia subsp. regia (natural 
hybrid, n — 
BRAZIL: RIO DE JANEIRO: Itatiaia, Berry et al. 4440. 


Fuchsia coccinea Dryander (n = 22). 

Bra ZIL: MINAS GERAIS: Serra da Piedade, Berry et al. 
4553, 45 
Fuchsia pw Taubert (n = 22). 

BRAZIL: RIO DE JANEIRO: Morro da Nova Caledónia, 
Nova Friburgo, Berry et al. 4423, 4430 (probable in- 
trogressive hybrid with F. regia subsp. regia). 


Fuchsia hatschbachii P. Berry (n = 22). 

BRAZIL: PARANÁ: Mun. Campo Largo, Serra Sào Luis 
do Puruná, Berry et al. 4458, 4461; Mun. Bocaiüva do 
Sul, Bacaetava, Kummrow et al. 2998 (MBM 

CULTIVATED: The Netherlands, from seed of Berry 
4464, from Paraná, Brazil, Berry & Brako 008-86. 


Fuchsia lycioides Andrews (n — mm 

CULTIVATED: University o 
Berkeley, UCBG H 53.1303-S2, from seed of ian ig 
in 6 Oct. 1953, from Panta "Mollas Chile; University of 
California at Los Angeles, seedling from seed c ollected by 
D. M. Moore in Coquimbo, Ch le (2n = 44; without 
voucher). 


] G d 


Fuchsia 1 magellanica Lamarck (n = 22). 
: NE sy EN: Villa gays Nahuel Huapi, 
Diem 3614 " 44), 3615 (2n — 

CHILE. Minn m Región): Concepts Barrio Uni- 
versitario, Riquelme s.n. (2n — as Canoas, Con- 
cepción to Chaimavida, Riquelme s.n. (2n — 44). Los 
: (X Región): S of Valdivia, Marticorena & Quezada 
1671 (2n = 44), 1672 (2n = 44); Puente Rahue, Prov. 
Osorno, Marticorena & Quezada 1676 (2n = 44); Fru 
tillar Bajo, Prov. Llanquihue, Marticorena & Quezada 
1677 (2n = 44); Frutillar Bajo to Frutillar Alto, Marti- 
corena & Quezada 1078 ph = 44); 2 km W of Llan- 

ezada 1679 (2n = 44); 2 
Morteoreno & Quezada 1680 


Z 


> 
= 
2 
S. 
— 


& Quezada 1682 (2n — 44) Prov. 
Ancud, Marticorena & Quezada 1663 (2n — 44); Par 

anqui bo Curamo, Chiloé, Marticorena & Quezada 1684 
(2n = 44); 4 km from Castro to Quellón, Chiloé, Mar- 
ticorena & Quezada 1686 (2n = 44), 1687 (2n = 44); 
Lago Natri, Chiloé, Marticorena & Quezada 1689 (2n 
— 44); Rio Mollueco, Castro to Quellón, Marticorena & 
Quezada 1692 (2n — 44); n past Coinco Alto, Castro 
to Quellón, Marticorena & Dues ada 1693 (2n — 44); 


Quellón, Marticorena & Quezada 1694 (2n — 44); Dal- 


cahue, Castro to 


iid (2h = 44); 2 
Mar ena & Quezada 1703 (2n = 44); Lago TM 


ago Mun p Mid Vx icai & Quezada 1706 (2n = 
44); Puerto Varas to Ensenada, Lago Llanquihue, Mar- 
ticorena & a 1707 (2n = 44); Saltos de Petrohué, 


= 44); fub bois 


Marticorena & is 1712 (2n = 44), 5 km S of 
aut autin, Marticorena & Quezada 1713 
(2n = 44); Corral, Valdivia, Godoy s.n. (2n = 44). MA- 
GALLANES rg Región): Punta Hatley, Seno Otway, Pis 


ano s.n. (2n — 44) Fuerte Bulnes, Magellan South, 
Pisano s.n. (2n = 44). 
CULTIVATED: Ireland, Nelson 576 (2n = 44), 1162 


2n = 44. previously living at MO); California, at Univ. 

aca Botanic Garden, Berkeley (n = 22, 2n = 44; 

aay et al., y oes (n — 22; photograph 
”). 


in Warth, 1925, as “F. cocc 


Fuchsia regia (Vellozo) Munz subsp. regia. Populations 
with n = 22 


BRAZIL: RIO DE JANEIRO: Morro da Nova Caledónia, 
Friburgo, Berry et al. 4425, 4433, 4424 (probable 
hybrid with F glaz ioviana); Itatiaia, road to Planalto, 
Berry et al. 072, 074, 076, 082. MINAS GERAIS: Tron- 
queiras to Vale Verde, Alto Caparaó, Berry et al. 4543; 
Serra do Itacolomi, Ouro Preto, Berry et al. 4544, 4546 
(probable i qui aa Pacha with F. coccinea). SAO PAULO: 
do Jordao, Ramamoorthy 683 (2n 
= 44); Campos do Jordão. Berry et al. 089, 090, 091; 
Serra da Bocaina, Berry et al. 083, 085. 
The Netherlands, from seed of Ber 
4425, ‘tow Rio de Janeiro, Brazil, Berry & Brako 019. 
80. 


Fuchsia Ed (Vellozo) Munz subsp. regia. Populations 
with n = 

BRAZIL. RIO DE JANEIRO: Teresópolis, Berry et al. 4419; 
Itatiaia, Planalto, pil et al. 4439; Serra dos Órgãos, 
Falkenberg & al. 3743 (FLOR). são PAULO: Campos do 
Jordào, Ramam M & Vital 673, 676, 680; Serra 
da Bocaina, Ramamoorthy & de Lima 1178. 


Fuchsia regia subsp. reitzii P. Berry (n = 44). 
BRAZIL. SANTA CATARINA: Curitibanos, pes & Fal- 
kenberg 4506 (multivalents common); Bom Jardim to 
z rr d 4510 M 
rra to Lauro Müller 
Berry & Falkenberg 4517 (arbe hybrid with F. nid: 
subsp. se errae; multivalents common); Mun. Uribici, SW 
of Uribici, Falkenberg et al. 3777 ( ). RIO G py 
bo SUL: E of Tainhas on RS- 486, Falkenberg et al. . 
(FLOR); Posto Fiscal, W of Serra de Rocinha, loaded 
et al. 3767 ( ). 
CULTIVATED: The Netherlands, from seed of Berry & 
Falke a EP lm from Santa Catarina, Brazil, Ber & 
Brako € 


Fuchsia regia al serrae P. Berr 4). 
BRAZIL: SAO PAULO: road between Mogi das Cruzes and 


592 


Annals of the 
Missouri Botanical Garden 


Bertioga, Pirani & Yano 640 (SP). PARANÁ: re Bo- 
caiúva do Sul, 9 km W of Sesmaria, Berry & Juarez 
4448 (possible dr with F. brevilobisk 12 km W of 
Sesmaria, Berry & Juarez 4450; Estrada da Graciosa, 
Mun. Morretes, Berry et al. 4492 milan common), 
3; Mun. Morretes, Ramamoorthy & d 
829 ln = 44, 2n = 88), 830. SANTA CATA : Rio 

Sul to Curitibanos, Berry & Falkenberg 1501 (multi. 


Sul, below Serra ds Rocinha, Falke nberg et al. 3753 
(FLOR). RIO GRANDE DO SUL: of Tainhas on RS- 
486, Falkenberg et al. E (F LOR : 

CULTIVATED: The Netherlands, from seed of Berry 
4450, from Paraná, Brazil, Berry & Brako 023-86; the 
Netherlands, from seed of Berry 4504, from Santa Ca- 
tarina, Brazil, Berry & Brako 024-86 


Fuchsia regia: plants intermediate between subsp. re- 
gia and subsp. serrae (n — 22, 44). 


plants intermediate between subsp. regia and subsp. ser- 
ae (n 44). 

BRAZIL. RIO DE Mines above Parati to Cunha, Berry 
et al. 097 (n = 44), 100 (n = 22). SAO PAULO: Alto da 


previously living specimen at MO); Serra da Bocaina, 
Ramamoorthy & de Lima 1176 (n = 44) 


Fuchsia regia subsp. regia X F. regia subsp. serrae (2n 
= 66). 


Artificial hybrid between Med d 083 (subsp. 


regia, n — 22) and Ramamoorthy & Hatschbach 830 
(subsp. serrae, n — 44), redu at Missouri Botanical 
Garden, M2588. 


NOTES ON THE 

CORDIA PANAMENSIS 
COMPLEX (BORAGINACEAE) 
AND A NEW SPECIES 
FROM COLOMBIA 


Cordia is a large pantropical genus of about 
300 species, divided by most recent authors into 
six sections (Nowicke & Miller, in press). Section 
Myxa, with over 150 species, is the largest in the 
Neotropics and includes all but one of the paleo- 
tropical species (Miller, 1985). It is difficult tax- 
onomically because most of the groups that can be 
recognized within the section have obscure bound- 
aries. 

One such group is the Cordia panamensis Riley 
group (Miller, 1985), which is widespread in Cen- 
tral America, the West Indies, and northern South 
America. This group is particularly problematic in 
terms of its definition. within sect. Myxa, as well 
as species delimitations within it. It is loosely de- 
fined and may very well not be monophyletic, yet 
the species all share a similar appearance and are 
often confused in herbaria. The C. panamensis 
group is characterized by ovate to nhe sca- 
brous to scabrid, usually anisophyllous leaves that 
generally dry brown. All of the species except C. 
anisoph ylla J. S. Miller are dioecious. Perhaps C. 
bicolor A. DC. and C. sellowiana Cham. should 
be included, but C. bicolor does not dry brown, 
and both species differ in being homostylous. 

Sculpturing on the surface of the endocarps 


KEY TO THE SPECIES OF THE CORDIA PANAMENSIS GROUP 


appears to be useful in distinguishing species of 
Cordia sect. 
yet been used e this. Dried fruits of various species 
of the C. panamensis group were soaked in 5% 
NaOH for two days to soften the exocarp and 
mesocarp, then washed in running water to remove 


yxa (Miller, in prep.) but has not 


outer tissues and expose the cleaned, outer surface 
of the endocarp. Cordia gentryi has an endocarp 
with raised, quadrangular areas (Fig. 1C); in C. 
anisophylla the raised areas are much longer (Fig. 
1 D). Other members of the group have rugulose 
(C. panamensis, Fig. 1E) to spinulose (C. Aebe- 
clada I. M. Johnston, Fig. 1F) endocarps. 

Recent studies of members of the C. panamensis 
group have helped clarify the relationships of species 
in Central America (Miller, 1985, 1988) and Ven- 
ezuela (Gaviria, 1987). The group is defined here 
as consisting of ten species, but problems still exist 
with differentiating C. panamensis and C. macro- 
phylla. | have seen little material of C. macro- 
phylla and have maintained it as separate following 
Johnston (1949). As part of a review of the 
panamensis group, several novel collections from 
Pacific coastal Colombia appear to represent an 
undescribed species. 


la. Leaf apices acute to obtuse. 
2a. Stems, petioles, and inflorescence branches with echinate = o C. cymosa (J. D. Smith) Standley 
2b. Stems, petioles, and inflorescence branches with simple hai 
3a. Petioles longer than 15 mm; twigs puberulent b os South America ............. C. tetrandra Aubl. 


3b. Petioles shorter beds 15 mm; twigs velutinous to hirsute; West Indies „nn 


E 


Leaf apices acuminate to ca 
4a. Twigs strigillose, the hass 
5 


C. sulcata DC. 


e. 
appressed, short, white or transluce 
a. Flowers distylous; corollas longer than 10 mm; mature fruits ellipsoid, longer than 8 m 


pc ae J. S. Miller 


5b. Flowers dioecious; corollas shorter than 5 mm; mature fruits ovoid, shorter than 8 mm 


4b. Twigs with erect or curly hairs or densely short-velutinous, or if strigillose, the hairs brown. 
ou C. 


7b. iocis of t 
ems hide 


On Twigs e lutinous; Jamai 
9b. Twigs hirsute; Mexico to South ae Trinidad, and Tobago ... 


C. sericicalyx DC. 
gentryi J. S. Miller 


nt. 
mentose; Gui = A C. toqueve Aubl. 


n 
inflorescence velutinous to ci fruits glabrou 
o hispid. 


C. macrophylla L. 


. C. panamensis Riley 


ANN. MISSOURI Bor. GARD. 76: 593-595. 1989. 


594 Annals of the 
Missouri Botanical Garden 


FIGURE 1. A-C. Cordia pene eee et al. 17874 (MO), Colombia). — A. Fruiting branch. — B. Fruits with 


persistent calyces.—C. Dorsal view of endocarp. — D. Cordia anisophylla (Mori & Kallunki 6373 (MO), Panama), 
dorsal view of endocarp. — E. Co rdia panamensis (Miller & Miller 961 (MO), Panama), dorsal view of endocarp. 
F. Cordia hebeclada (Gentry et al. 19686 (MO), Peru), dorsal view of endocarp. 


8b. Stems iue ent to densely short velutinous. 
Fruits white; twigs hirsute to velutinous; upper leaf surface scabrous; "s Indie 
C. sulc ia DC. 
9b. Fruits yellow; twigs short and evenly ie brown sou upper leaf surfac 
glabrous to scabrid; Colombia, Ecuador, Peru ....................... . hebeclada 1. M. p 


Volume 76, Number 2 
1989 


Miller 595 


Cordia panamensis 


Cordia gentryi J. S. Miller, sp. nov. TYPE: Co- 
lombia. El Valle: Bajo Calima, Dindo area, 
100 m, 11 July 1984, 4. Gentry, M. Mon- 
salve & D. Wolfe 47874 (holotype, MO 
3521285). 


ad 8 m alta, ramunculis Wer strigillosis ad 
puberalentis, Folia persistentia, petiolis 1.7-2.6 mm lon 
gis; laminae anisophyllae, folia dus ovatis, 30-35.3 
cm longis, 10.7-13 cm latis, apice € caudas, basi obtusis, 
minoribus orbicularis, s ida, pagi 
grosse pubens. Flores ignoti. Es drupaceus, auran- 
tiacus, endocarpo osseo, inaequilateraliter ovoideo, 16 mm 
longo, 10 mm lato. 


Tree 8 m tall, the twigs densely puberulent with 
erect, brown hairs. Leaves persistent, dimorphic; 
petioles 1.7-2.6 
puberulent, with erect, brown hairs; the larger blades 


mm long, canaliculate adaxially, 
narrowly ovate, 30-35.3 cm long, 10.7-13 cm 
wide, the apex caudate, 3-4 cm long, the base 
obtuse; smaller blades orbicular, 11-15.5 cm long, 
9.5-14 cm wide, the apex abruptly short acumi- 
nate, the base rounded to acute, both types with 
the margin entire, the upper surface scabrous, the 
hairs translucent, appressed, the midrib raised, pu- 
berulent, the lower surface coarsely pubescent, with 
6-10 pairs of secondary veins. Flowers unknown. 
Infructescence broadly cymose, ca. 13 cm broad. 
Fruits drupaceous, orange-yellow, the fruiting ca- 
lyx slightly accrescent, unevenly 5-lobed, 9-11 
mm broad, appearing torn along the margin, uni- 
formly strigillose, somewhat sericeous inside, the 
mesocarp mucilaginous, the endocarp bony, inequi- 
laterally ovoid, 16 mm long, 10 mm broad, the 
surface reticulate with raised quadrangular areas. 


Distribution. Cordia gentryi is known from 
the Colombian provinces of Chocó and El Valle at 


elevations below 100 m in wet tropical forests. 


Additional specimens examined. COLOMBIA, CHOCÓ: 
area of Baudó, on the right side of Río Baudó, about 13 
km upstream from the estuary of Quebrada Carpio and 
the shore opposite Quebrada Cola Barquita, 11 Feb. xe i 

Fuchs & Zanella 21294 (MO). EL VALLE: Bajo 
ca. 10 km N of Buenaventura, agn his Colombia 
concession, transition between tropical wet and pluvial 
forest, ca. 50 m, 10 Dec. 1981, Gentry 25508 (MO). 

Cordia gentryi is most easily distinguished by 
its caudate leaf apex and large, orange-yellow fruits. 
It most resembles C. panamensis and C. aniso- 
phylla J. S. Miller. Cordia panamensis differs in 
having hirsute twigs and white fruits shorter than 
7.5 mm. Cordia anisophylla differs in having thin- 
ner ellipsoid fruits 6-7.5 mm broad and less-pro- 
longed leaf apices. 


LITERATURE CITED 

GAVIRIA, J. 1987. Die Gattung Cordia in Venezuela. 
Mitt. Bot. Staatssamml. München 23: 1-279. 

JOHNSTON, L 1949. Studies in the Boraginaceae 

VIII. Boraginaceae of the southern West Indies. 
J. Arnold ie : 111-138. 

MiLLER, J. S. 985. pei of the genus Cordia 
(Boraginac her in Mexico and Central America. Ph.D. 
Dissertation. St. Louis Univ., St. Louis, Missouri. 

1988. A revised treatment of b rupe 
for Posi. Ann. Missouri Bot. Gard. 75: 456-521 

NoWICKE, J. W . S. MILLER. Pollen Bec of 
the Cordio idis ae (Boraginaceae): Auxemma, Cordia, 
and Patagonula. Pl. Evol. Syst. (in press). 


—James S. Miller, Missouri Botanical Garden, 
P.O. Box 299, St. Louis, Missouri 63166, U.S.A. 


NOTES 


A NEW ASPIDOGYNE 
(ORCHIDACEAE) FROM 
VENEZUELAN GUAYANA 


The genus Aspidogyne Garay is one of the 
recent segregates from Erythrodes Bl. (Garay, 
1977) and comprises 27 species, widely distributed 
in the American tropics, primarily Brazil. They are 
small to medium-sized terrestrial, humicolous, to 
(rarely) subepiphytic herbs growing in mostly shady 
places of rain or cloud forests. 

Aspidogyne belongs to the pantropical subtribe 
Physurinae, of seven genera. Among them, As- 
pidog yne can be recognized by the following com- 
bination of features: horizontal stigmata; elongate, 
erect column; and entire emarginate rostellum. 

A revision of the Aspidogyne species of the 
Venezuelan Guayana for the orchid treatment in 
Steyermark's Flora of the Venezuelan Guayana 
has revealed a different identity for the species that 
appeared in Foldats 70) as Erythrodes picta 
(Lyndley) Ames, a synonym of 4spidogyne ar- 
gentea (Vell.) Garay. Foldats had already noted 
that the Venezuelan material cited under £. picta 
differed from the type, and suggested that it might 
represent a new variety. New evidence shows that 
the differences are sufficient for recognition of a 
new species, described below. 


Aspidogyne steyermarkii Carnevali & Foldats, 
sp. nov. TYPE: Venezuela. Bolivar: cabeceras 
del Rio Chicanán, Sierra de Lema, 80 km SW 
de El Dorado, 500 m, 6%5'N, 62°W, 22 Aug. 
1967, Steyermark 89371 (holotype, VEN; 
isotype, MO). 

Species 
dorsali acuto, petalis non lobulatis, calcar 
belli lobo centrali transvere rhombico net 


s haec 4. argentea (Vell.) Garay saat at sepalo 
cylindrico, la- 


Terrestrial herbs, 9-19 cm high. Stems basally 
rhizomatose, creeping at the base, aphyllous, api- 
cally ascendent to erect, 2—6-foliate. Leaves deep 
olive-brown with red streaks on both sides of midrib 
and with the same colored spots near margin above, 
pale lavender with pale streaks and dots below, 


ovate to ovate-elliptic, acute, basally rounded to 


ANN. MISSOURI Bor. 


2156, Caracas 1010-A, Venezuela. 


somewhat truncate, 7-28 mm long, 8-12 mm 
wide; petioles about 12 mm long, basally dilated 
and sheathing. Inflorescences of erect, laxly pauci- 
to pluriflorous racemes, 4.5-11 cm long, the pe- 
duncle densely pubescent. Flowers resupinate, small, 
with white tepals and labellum. Tepals uninnervate, 
subfleshy. Dorsal di. nne acute, 4 mm long, 
te-elliptic, subacute, 


.2 mm wide; lat epals o1 
basally attenuate, parc oblique, 5 mm long, 
2 mm wide; petals narrowly obovate-elliptic, acute, 
3.8 mm long, 1.1 mm wide. Labellum elliptic in 
overall outline, 5.1 mm long, 2.7 mm wide, con- 
tracted above its apical third, obovate below the 
contraction, dilated in a transversely rhombic ter- 
minal lobe above, 0.8 mm long, 1.2 mm wide, 
subacute. Spur cylindric, acute, slightly arched, 

.2 mm long, 0.7 mm thick. Column clavate, 
mm long; rostellum aristate, acute. 


This new species belongs in sect. Argenteae 
Garay. It is similar to Aspidogyne argentea (V ell.) 
Garay but differs by having acute (vs. rounded or 
obtuse) dorsal sepals, cylindric (vs. fusiform) spur, 
petals apically not lobulate, and the labellum with 
a transversely rhombic, subacute (vs. transversely 
obreniform, truncate, apiculate) apical lobe. 


We are grateful to Lic. Bruno Manara for crit- 
icism of the Latin diagnosis and for the drawing. 


LITERATURE CITED 


Forbpars, E. l Erythrodes. kA ru In: T. 
asser (editor), Flora de Venezuela 15(1): 263-267. 
GARAY, A. 77. Systematics of the ve haere 


(Orchidaceae) in the New World. Bradea 2(28): 1 


—Germán Carnevali and Ernesto Foldats, both 
authors: Jardín Botánico de Caracas, Herbario 
Nacional de Venezuela, INPARQUES, Apartado 
: Carnevali’s 
current address: Missouri Botanical Garden, P.O. 
Box 299, St. Louis, Missouri 63166-0299, U.S.A. 


GARD. 76: 596-597. 1989. 


Volume 76, Number 2 Notes 597 


" 5 cm 4 


FICURE l. Aspidogyne steyermarkii. —a. Flowering habit. — b. Lateral view of labellum and column. — c. Perianth 
segments and labellum flattened. — d. Pollinia. —e. Labellum 


THREE NEW SPECIES 
OF BOMAREA 

(ALSTROEMERIACEAE) 
FROM MESOAMERICA 


of the treatment of Bomarea 
Mirbel is the ‘Fore Mesoamericana, I found three 
previously undescribed species in the Missouri Bo- 
tanical Garden herbarium (MO). The types of all 
three species are from Panama, from where no 
new species have been described since the treat- 


ment of the genus in the Flora of Panama (Killip, 
1945). 

Characters of these new species common to all 
Alstroemeriaceae (Dahlgren et al., 1985) are: se- 
pals 3; petals 3; stamens 6 with anthers pseudo- 
basifixed (the tip of the filament inserted in a deep 


FIGURE 1. 


scale as E (From holotype, with one leaf from Sytsma et 


ANN. MISSOURI Bor. GARD. 


Bomarea caudatisepala. —a. Plant showing two ce ond rays. — b. Petal, inside surface (same 
1.) 


t al. 4 


76: 598-601. 1989. 


Volume 76, Number 2 
1989 


Notes 


pit) and introrsely dehiscent; style 1 with 3 stig- 
matic branches. Like all other Mesoamerican species 
of Bomarea, the new species have fully inferior 
ovaries and loculicidally dehiscent, capsular fruits. 
Stem diameter is measured between the first and 
second leaves below the umbel; length of compound 
umbel rays is measured from the involucre to base 
of the uppermost ovary. 


Bomarea caudatisepala Gereau, sp. nov. TYPE: 

Panama. Chiriqui: 8°46'N, 82°25'W, upper 

NE slopes of Cerro Pate de Macho and along 

soggy ridge of Continental Divide, 1,900- 

2,000 m, 19 June 1987, Croat 66496 (ho- 
lotype, MO; ae PMA). Figure 1. 

Herba volubilis caule glabro ad parce puberulo sub 


umbella. FOLIUM: lamina ovato-lanceolata, 6.9-12.5 cm 
longa x 2.7-4.5 cm lata, superne glabra, inferne secus 


nervos glabra ad parce crispato-pubescente. UMBELLA 
bracteis involucralibus 5- 1-3.8 cm longis x 0 
5 tis, glabris; radiis 6-14, 17.1-26.2 cm longis, 


saltem versus apicem parce fulvo-puberulis, simplicibus 
ebracteolatisque. FLOS: v oblongo-lanceolatis, 3.6- 
4.0 cm longis x m latis, externe aurantiacis, 
tpequoque sepalo cornu viridi 7-10 mm longo sub apice 
cm 


longis X 1.6-1.7 cm latis, sepala per 2 
tibus, externe aurantiacis, interne flavo-aurantiacis ad vi- 
ridibus, purpureo-maculatis; ovario anguste obconico, sub 
anthesi 7-10 mm longo x 3-5 mm lato, parce fulvo- 
puberulo 

Vine; stem 2.0-2.5 mm diam., glabrous 
throughout to sparsely puberulent just below the 
umbel. Petiole 10-13 mm long; leaf blade ovate- 
lanceolate, 6.9-12.5 x 2.7-4.5 cm (L/W ratio 
— 2.6-3.0), glabrous above, glabrous to sparsely 
crisp-pubescent on the nerves beneath. Involucral 
bracts 5-8, 3.1-3.8 x 0.4-0.5 cm, glabrous; rays 
of the umbel 6-14, 17.1-26.2 cm long, sparsely 
brown-puberulent at least toward the tip, un- 
branched and ebracteolate. Sepals oblong-lanceo- 
late, 3.6-4.0 X ca. 0.9 cm, externally orange with 
a green horn 7-10 mm long inserted 2-4 mm 
below the tip; petals obovate, 3.8-4.3 x 1.6-1.7 
cm, exceeding the sepals by 2-5 mm, orange with- 
out, yellowish orange to green with purplish spots 
within; CEN 2.9-3.0 mm long; anthers 4.0- 
6.0 x .4 mm; ovary narrowly obconic, 7— 
10 x a 5 at les sparsely brown puberulent; 
style 10-14 mm long; stigmatic branches 1.4-1.9 
mm long. 


1] t 


Known only from three Cerro Pate 
de Macho, in cloud forest and elfin forest at 1,700— 
2,100 m elevation. 


Additional collections examined. PANAMA. CHIRIQUÍ: 
Cerro Pate Macho NE of Boquete, S slope along path to 


B 

o 

FIGURE 2. Bomarea suberecta. —a. Apex of plant 
with inflorescence. —b. Involucral nk 


(From Mc- 
Pherson 9333, with one bud from holotype.) 


Finca Serrano, near the crest at ca. 2,000 m, 26 May 
1981, Andersson & Sytsma 1307 (GB, MO); SE slopes 
and summit Cerro Pate Macho, trail from Rio a 
4 km NE of Boquete, 1,700-2,100 m, 26 May 
Sytsma, Knapp & Andersson 4871 (MO). 

s new species of uncertain affinity is easily 
distinguished from all other Mesoamerican Bom- 
area by the presence of a horn just below the tip 
of each sepal. Andean species with horned sepals 
all have much smaller flowers and further differ 
from B. caudatisepala in having very short umbel 
rays (B. caudata Killip, B. cornigera Herbert) or 
compound inflorescences (B. cornuta Herbert). 


Bomarea suberecta Gereau, sp. nov. TYPE: Pan- 
ama. Chiriqui: slope of Cerro Respinga above 
town of Cerro Punta, 2,500 m, 9 Aug. 1972, 
W. G. & J. J. D'Arcy 6587 (holotype, MO). 
Figure 2. 


Herba volubilis vel plerumque suberecta ad erecta, ubi 
erecta usque ad 2 m alta foliis Pa E ad squamas 
reductis; rhizomate radices fibrosas in tumores tuberosos 
terminantes emittente; caule Ere Al puberulo pilis mul- 
ticellularibus sub umbella. FOLIUM: lamina late ovata ad 

ovato- lanceolata, (6.0-)8.2-15.3 cm longa x 3.3-7.2 


trifidis lobo centrali lateralibus multo longiore, glabris; 
radiis 5-12, 2.2-4.3 cm longis, dense fulvo-puberulis, 
Fg eed cbracteolai vel (in typo tantum) supra me- 
dium bracteolam ca. 5 mm longam gerentibus. FLOS: se- 
palis wi is lanceolatis, 1.7-2.0 cm longis x 0.5-0.7 
cm latis, base rubris ad aurantiacis sursum flavescentibus 


600 Annals of the 
Missouri Botanical Garden 
z 
Oo 
1 | ux 
BS 
M. hy 
| |. 
AN AR 
PN jh 
A 
FIGURE 3. Bomarea quem Apex of plant showing one inflorescence ray. (From holotype.) 


petalis obovatis, 1.8-2.2 cm longis x 0.7-1.0 cm latis, 

sepala per 1-3 mm excedentibus, flavis wa aurantiacis; 

ovario late obovoideo, sub anthesi 4-7 mm x 3- 
mm lato, dense fulvo-puberulo. 


Vine or more often a suberect to erect herb to 
2 m from a creeping rhizome with fibrous roots 
terminating in tuberous swellings, the lower leaves 
reduced to scales when plant erect; stem 1.9-4.9 
mm diam., glabrous to short-puberulent with mul- 
ticellular hairs just below the umbel. Petiole 7-16 
mm long; leaf blade broadly ovate to ovate-lan- 
ceolate, (6.0-)8.2-15.3 x 3.3-7.2 cm (L/W ratio 
= ].7-2.7(-3.0), glabrous above and beneath. 
Involucral bracts 4-5, 0.9-2.0 x 0.4-0.6 cm, 
deeply trifid with central lobe much longer than 
lateral lobes, glabrous; rays of the umbel 5-12, 
2.2-4.3 cm long, densely brown puberulent, un- 
branched, ebracteolate or bearing (only in type) 
above the middle a bracteole ca. 5 mm long. Sepals 


oblong-lanceolate, 1.7-2.0 x 0.5-0.7 cm, red to 
orange at the base, shading to yellow above and 
green at the tip; petals obovate, 1.8-2.2 x 0 
1.0 cm, exceeding the sepals by 1-3 mm, yellow 
to orange; filaments 10-14 mm long; anthers 2.3- 
3.5 Xx 1.0-1.6 mm; ovary broadly obovoid, 4-7 
X 3-5 mm at anthesis, densely brown puberulent; 
style 7-15 mm long; stigmatic branches 1.2-3.1 
mm long. 

Known from the vicinity of Cerro Punta and 
Cerro Pando (on Costa Rican frontier) in Panama, 
and from the Atlantic slope of the Cordillera de 
Talamanca in adjacent Costa Rica, in montane rain 
forest, Quercus forest, and elfin forest at 1,900- 
2,750 m elevation. 

Additional Por dd d PANAMA. CHIRIQUÍ: 
Cerro Punta, 2,100 3 July 


Aug. 
1984, Croat 26454 (MO), 26472 (MO, PMA); above 


Volume 76, Number 2 
1989 


Notes 601 


Cerro Punta, 2,000 m, 9 May 1971, D'Arcy 5365 (MO); 
8°55'N, 82*44'W, E slopes of Cerro Pando, 2,000-2,300 
m, 15 Oct. 1981, Knapp 1650 (MO); 8°50'N, 82°35'W, 
road from s Punta village towards Boquete near Bajo 
Grande, ca. 2,100 m, 5 June 1986, McPherson 9333 
(MO); 8*52' N, 82°33'W, Cerro Punta, along ridge to 
watershed to Bocas del Toro, 2,200 m, 26 Jan. 1985, 


van der W 


er 
LIMON: 9%00'-12'N, 82*57'-59'W, unnamed clica 
between Rio Terbi and Rio Sini, 2,300-2,500 m, 11 Sep. 
1984, Davidse, Herrera Ch. & Grayum 28957 (MO), 
2,400-2,750 m, 13 Sep. 1984, 29046 (CR, MO); 9?14'- 
15'N, 82°59 W, os massif, ridge between Rio Tar- 
aria and NE-most páramo, 1,900-2,300 m, 16 Sep. 
1984, Davidse & Herrera Ch. 29206 (MO). 


This new species is distinguished by its deeply 
trifid involucral bracts; these organs are entire in 
all other known members of the genus. In Me- 
soamerica, the new species could only be confused 
with B. acutifolia (Link & Otto) Herbert. In ad- 
dition to its entire involucral bracts, B. acutifolia 
has larger flowers (petals 2.6-3.9 cm long) than 
B. suberecta and a consistently twining or trailing 
habit. The suberect to weakly vining habit of B. 
suberecta is otherwise only known in a few Andean 
species, most notably B. ovata (Cav.) Mirbel. 


Bomarea bracteolata Gereau, sp. nov. TYPE: 
Panama. Chiriquí: 8%45'N, 82?15'W, Fortuna 
Dam, above Gualaca, on forested slopes be- 
hind forestry experimental station, ca. 1,200 
m, 9 March 1985, McPherson 6706 (holo- 

e, MO; isotype, PMA, not seen; other iso- 
types distributed as B. allenii). Figure 3. 


Herba ETUR caule glabro. FOLIUM: lamina ovato- 
lanceolata, a -17.5 cm longa 4.9-6.6 cm lata, 
f s hyalino cena pilis 
wel lar nar complanatis, inter nervos puberula. 
UMBELLA: bracteis a ee a 10, 4.2-8.8 cm longis 
x 1.8-3.7 cm latis, superne puberulis; radiis 5-14, 14. 
42.3 cm longis, glabris, bivel trifurcatis, AM ramo 
bracteola foliacea suffulto, ima 3.6-7.0 cm longa. FLOS: 
sepalis oblongo-lanceolatis, 4.1-4.8 cm bos x 1.2-1.6 
cm latis, rubris vel glauco-salmoneis; Tg anguste ob- 
3.8-)4.6-5.1 cmlongis x (0. .2-1.3 cm latis, 
sepalis 3 mm brevioribus ad 3 mm Meca externe 
rubellis, interne pallide viridibus ad flavo-albescentibus, 
maculis nigris vel atro- Land obsitis; rie obconico, 
sub anthesi ca. 7 mm -4 mm lato, glabro. 


Vine; stem 2.3-4.9 mm diam., glabrous. Petiole 
16-41 mm long; leaf blade ovate-lanceolate, 13.0- 
17.5 x 4.9-6.6 cm (L/W ratio = 2.5-2.7), gla- 

above, hyaline-hirsutulous with flattened 
multicellular hairs on the nerves and puberulent 
etween the nerves beneath. Involucral bracts 5- 


10, 4.2-8.8 x 1.8-3.7 cm, puberulent above; 
rays of the umbel 5-14, 14.9-42.3 cm long, gla- 
2- or 3-branched, each branch subtended 
by a leaflike bracteole, the lowest bracteole 3.6- 
7.0 cm long. Sepals oblong-lanceolate, 4.1—4.8 x 
1.2-1.6 cm, red or glacous pink-salmon; petals 
narrowly obovate, (3.8-)4.6-5.1 x (0.8-)1.2-1.3 
cm, from 3 mm shorter to 3 mm longer than the 
sepals, reddish without, pale green or yellow-white 
with black or dark purple spots within; filaments 
ca. 36 mm long; anthers ca. 5.6 x 2.1 mm; ovary 
obconic, ca. 7 X 3-4 mm at anthesis, glabrous; 
style ca. 36 mm long; stigmatic branches ca. 2.7 
mm long. 


brous, 


Known from the provinces of Chiriqui and Ve- 
raguas in Panama, in premontane wet forest at 
750-1,350 m elevation. 


Additional collections examined. PANAMA. CHIRIQUÍ: 
type Mg : Mar. 1985, McPherson 6704 (MO, PMA); 
8?45'N, 5'W, Fortuna Dam, in valley S of lake, 
1,200-1, pn m, 25 Dec. 1986, McPherson & Aranda 
10126 (MO, PMA). VERAGUAS: e of Santa Fé, slopes 
of Cerro Tute, 750 m, 25 Mar. 1947, Allen 4367 (MO); 
Cerro Tute, trail ur agricultural school near Santa Fé, 
1,200 m, 17 Sep. 1979, Antonio 1853 (MO); 8°32'N, 
81%07'W, trail to Tots Tute, above Escuela Agricola 
Alto de Piedra just W of Santa Fé, 800-1,350 m, 5 June 
1982, Knapp & Dressler 5456 (MO) 

Bomarea bracteolata is probably most closely 
related to B. allenii Killip, the only other large- 
flowered Mesoamerican Bomarea with branched 
umbel rays. These species differ in sepal length 
(4.1—4.8 cm in B. bracteolata, 5.3-6.5 cm in B. 
allenii), lower leaf surface (pubescent in B. brac- 
teolata, glabrous in B. allenii), and length/ width 
ratio of leaf blades (2.5-2.7 in B. bracteolata, 

4.0-5.4 in B. allenii). The inflorescence and flow- 
ers of B. bracteolata are very similar to those of 
B. kranzlinii Baker of Colombia (cf. Stein & 
McDade 3205, MO), but the latter species has 
smaller bracts and bracteoles and narrower leaves 
that are glabrous beneat 


LITERATURE CITED 


DAHLGREN, R. M. T., H. T. CLIFFORD & P. F. Yeo. 19 
d mune of Monocotyledons. Springer- Verlag, 


Kiki. E P. 1945. Bomarea Mirb. In: R. E. Woodson 
& UE Flora of Panama. Ann. Missouri 
-17. 


Bot. Card, 32: 


—Roy E. Gereau, Missouri Botanical Garden, 
P.O. Box 299, St. Louis, Missouri 63166-0299, 
U.S.A 


HOSTA YINGERI 
(LILIACEAE/FUNKIACEAE): 
A NEW SPECIES 

FROM KOREA 


The genus Hosta Tratt. includes ca. 22-25 
species of rhizomatous herbaceous perennials from 
Japan, China, and Korea. The genus is well known 
to gardeners in the United States, Europe, and 
New Zealand, since the species and cultivars are 
widely grown in shady gardens for their attractive 
foliage and flowers (Aden, 1988). They are also 
grown as ornamentals and eaten cooked in China, 
Japan, and Korea. Traditionally Hosta has been 
placed in the Liliaceae (e.g., Cronquist, 1981) or 
in the Funkiaceae (Dahlgren et al., 1985) along 
with Hesperocallis A. Gray and Leucocrinum Nutt. 
ex A. Gray, both from western North America. 

Bailey (1930) and Stearn (1931) noted that 
Hosta is a taxonomically difficult genus with con- 
fused nomenclature. Although a number of papers 
have been published on Hosta during the past 50 
years, there has been little improvement in our 
understanding of the genus. The matter has been 
largely complicated by broad (Fujita, 1976) or 
narrow (Maekawa, 1940) species concepts and 
confusion caused by cultivars. The plants have 
relatively few good diagnostic features that can be 
observed in pressed and dried herbarium specimens 
(Hylander, 1954). In an attempt to improve our 
knowledge of Hosta, my associates Carleen Jones, 
Myong Chung, and Haynes Currie and I have been 
attempting to assemble a collection of living hostas. 
Several years ago Barry R. Yinger, then of the 
U.S. National Arboretum, supplied seeds of several 
accessions of Hosta collected from Taehuksan Is- 
land, a remote island off the southwestern coast of 

orea. Examination of my cultivated plants and 
field-collected vouchers at NA reveals a new species, 
named in honor of the collector, Barry Yinger. 


Hosta yingeri S. B. Jones, sp. nov. TYPE: Korea. 
holla Namdo, Shin An Gun, Huksan Myon, 
Taehuksan Island, east side of Yeri village, 
34.40N, 125.6E. Among rocks on northwest- 
facing talus slope, shade, on cut-over hillside, 
ca. 60 m, common, 23 Sep. 1985, B. R. 
Yinger, T. R. Dudley, J. C. Raulston, A. P. 
Wharton & Y. J. Chang 3616 (holotype, NA). 
Figure 1. 


PARIS folii elliptica- -lanceolata vel anguste-ovata, semi- 
erecta; apex acuminatus, petiolus valde sulcatus, basis 


tum, oler in id i stamina tria + tria, libera, 
exserta, declinata, in base perianthii affixa; tria filamenta 
elongata. 


Plants glabrous, herbaceous perennials from 
short, clumpy rhizomes. Leaves ascending oblique- 
ly, spirally arranged at base of stem; petioles 3- 
7(-15) cm long, 2-5 mm wide at the middle of 
the petiole, greenish or sometimes with purple spots, 
winged; blades 4-15(-21) cm long, 2.2-7(-11) 
cm wide, elliptic-lanceolate or narrowly ovate, rigid 
and heavy-textured, acuminate at the apex, grad- 
ually narrowed at the base into the petiole, having 
a V-type fold in the base of the blade extending 
down into the petiole, the veins of upper leaf surface 
inconspicous when fresh, the veins of lower leaf 
surface in 3-4(-6) pairs of prominent, smooth 
lateral veins on either side of the midrib. Scapes 
2-4 times longer than the leaves, erect, smooth, 
usually with 1-2 linear-lanceolate bracts below the 
inflorescence, 1-2 cm long, 2-3 mm wide; raceme 
20-25-flowered, 
around the central axis of the raceme; pedicels 
1.2-2 cm long, longer than the subtending bracts; 
bracts flat, greenish, papillose at the apex, 8-12 
mm long, 2-3 mm wide. Perianth (fresh) whitish 
purple, tubular-funnelform, ca. cm long; 
tube slender, ca. 1.8-2 cm long, ca. 5 mm wide, 
white inside; throat white with purple between each 
lobe, veins not intensely colored; limb ca. 1.8-2 


the flowers equally arranged 


cm long, lobes 6, ca. 5 mm wide, inner surface 
light purple; tops of the flower buds purplish. Sta- 
mens distinct, 3 + 3, conspicuously exserted; fil- 
aments white, attached to the base of the perianth 
tube, strongly declined, then curving upwards, one 
set ca. 3.1 cm long, the other ca. 4.4 cm long, 
the filaments attached to the connective of the 
anthers in a groovelike pit; the anthers ca. 3 mm 
long, white beneath, black above. Style filiform, 
ca. 4.5 cm long, exserted beyond the stamens; 
stigma small, capitate, moist. Capsules cylindric, 


ANN. Missouni Bor. GARD. 76: 602-604. 1989. 


Volume 76, Number 2 Notes 603 
1989 


FIGURE l. Hosta yingeri. — A. Habit. — B. Downward view of the raceme axis showing the flowers spread evenly 
around the central axis of the inflorescence. — C. Flower viewed from front. — D. Flower viewed from the side. (Drawing 
prepared by Carol L. Gubbins Hahn from fresh material of garden-grown seedlings of holotype collection.) 


604 


Annals of the 
Missouri Botanical Garden 


2.5-3 cm long, 4.5-6 mm wide; seeds black, flat- 
tened, winged, ca. 8.4 mm long, ca. 3.4 mm wide; 
cotyledon small, undifferentiated. 


Flowering in August and September in Korea 
and in my garden located in northeast Georgia in 
August; fruiting and maturing seed in September 


and October. 


Paratypes. | KOREA. TAEHUKSAN ISLAND: garden-grown 
material of same collection as holotype, Yinger et al. 
(GA); Yinger et al. 3585 (NA) and garden-grown material 

); Yinger et al. 3244, 3610 (NA) and garden-grown 
material (GA); Sohuksan Island, Yinger et al. 3164 (NA). 


Hosta yingeri is distinct from other species of 
Hosta in its relatively thick, adaxially dark green 
leaves. This feature may be an adaptation to its 
coastal habitat, growing at 2-60 m above sea level. 
It is also distinguished by its delicate raceme of 
flowers spread evenly around the central axis of 
the inflorescence; typically, Hosta has subsecund 
racemes. An additional diagnostic feature of H. 
yingeri is the exceptional length of the second set 
of stamens, with both sets exserted well beyond the 
perianth. Our living material has been compared 
with representative accessions of all species of Hos- 
ta in our garden and with loans of herbarium ma- 
terial. Hosta yingeri is an attractive and interest- 
ing new species with horticultural potential. It 


appears to be associated with Maekawa's (1940) 
subgenus Bryocles, section Tardanthae. Hosta 
yingeri does not resemble our garden-grown ma- 
terial of H. tsushimensis obtained from the Na- 
tional Arboretum. 


LITERATURE CITED 


ADEN, P. id The Hosta Book. Timber Press, Port- 
land, 

BalLey, L. H £1930. Hosta: the Plaintain Lilies. Gentes 

erb. 2: 117-142. 

e IST, A. An Integrated System of Classi- 
n < Flowering E ts. Columbia Univ. Press, 

New York. [Pp. 112 

DAHLGREN, T . T., H.T. ae P.F. Yor. 1985. 
The Families of xs E cat Springer-Ver- 
lag, Berlin. [Pp. 187-188.] 

Fuyita, N. 1976. The genus Hosta in Japan. Acta 
Phyt totax. Geobot. 27: 66-96. 

Horn 1954. The genus Hosta in Swedish 

gardens. Acta Horti Berg. 16: 331-420. 

MAEKAWA, F. The genus Hosta. 2 Fac. Agric. 

v. Tokyo, Sect. 3, Bot. 5: 317-4 

STEARN, W. T. 1931. The hostas or a a revision 
of the on lilies. Gard. Chron. 2: 27, 47-49, 
88-89, 

— Samuel B. Jones, Jr., Department of Botany, 

The University of Georgia, Athens, 

30602, U.S. A. 


Georgia 


LINDSAEA MESARUM, A 
NEW FERN SPECIES FROM 
THE RORAIMA SANDSTONE 


When the present author published his revision 
of the New World species of Lindsaea (Kramer, 
1957), the collection Tate 430 (NY) was cited as 
a probably distinct but as yet insufficiently known, 
undescribed taxon. This taxon is now known by 
over 20 collections and is therefore described as 
new. 


Lindsaea mesarum Kramer, sp. nov. TYPE: Ven- 
Bolivar: Distr. Piar, summit of Muri- 
sipan-tepul, Aparamá Range, 5?53'N, 
62?03'W, sandstone mesa, cracks of rock, 
2,200 m B. K. Holst 3548 (holotype, MO; 
isotype, Z). Figure 


ezuela. 


Rhizoma breviter repens, squamis parvis, fuscis vesti- 


ginis, vel continui ad pinnulas minores; indusium firmum, 
marginem subaequans. Sporae triletae. 


Rhizome rather short-creeping, 1.5-2.5 mm 
diam.; scales fuscous, small, narrowly triangular 
with broad base, acuminate, ca. 1 x 0.2 mm, ca. 
8-seriate at base. Leaves a few-several mm apart; 
petiole usually lustrous, atropurpureous to black, 
smooth but the basal part bearing some small wart- 
like bases of shed scales: adaxial (sometimes also 
abaxial) side flattened (near the apex) to faintly 
sulcate, abaxial side otherwise rounded; length ca. 
12-22 cm, equaling the lamina to much exceeding 
it. Lamina narrowly oblong, ca. 10-20 cm long, 
paucijugate-bipinnate, or a few once-pinnate or 
subbipinnate sterile leaves present beside the bipin- 
nate fertile ones; primary rachis dark reddish to 
blackish brown, abaxially faintly biangular to dis- 
tinctly sulcate. Pinnae from 1 odd one to more 
often 1—3 pairs and a conform terminal one, mod- 
erately to very strongly ascending, hence the width 
of the lamina quite variable; lateral pinnae with a 


petiolule of ca. 8 mm, 7-10(-16) cm long, up to 


15 mm wide, somewhat narrowed near the base, 
gradually and strongly tapering to the apex. Sec- 
ondary rachises medium to dark brown, dull, adax- 
ially sulcate, the groove bordered by rounded ridges, 
abaxially subacutely biangular. Pinnules close to 
contiguous or even overlapping, coriaceous, often 
blackish when dry, dull or adaxially sublustrous, 
dimidiate, spreading or somewhat ascending, up to 
ca. 25 well-developed ones to a side, above these 
some much-reduced pinnules, a few finally con- 
fluent into a short, pinnatifid pinna-apex. Larger 
pinnules ca. 6-9 mm long, 3.5-5 mm wide, about 
1.5 times as long as wide, semiovate to subtrian- 
gular, base cuneate, posterior margin straight or 
convex, anterior margin approximately straight, 
entire or more often distantly and shallowly incised 
to ca. 0.5 mm, the lobes flanking the incisions 
mostly touching to overlapping, thus the incisions 
inconspicuous. Inner margin of pinnules straight, 
diverging from the rachidule, an outer margin not 
developed; all margins, especially the posterior, 
sclerotic, apex subacute to obtuse but not rounded. 
Sterile pinnules very shallowly and rather irregu- 
larly crenate. Veins immersed but somewhat evi- 
dent, adaxially often slightly impressed in dry spec- 
imens, free, simple or the inner forked above the 
base, ca. 5-8 vein-ends per pinnule present. Sori 
occupying the anterior margin, interrupted, or in 
small pinnules continuous, (1-)2-5 per pinnule, 
each on 1-4(-6) vein-ends, ca. 1.5-4 mm long; 
indusium firm, entire, often somewhat undulate (not 
sinuate!), about equaling the margin, ca. mm 
wide. Spores trilete, subtetrahedral, tawny, smooth, 
with prominent laesura, ca. 35 um (proximal) x 
26 um (lateral view). 


Paratypes. | VENEZUELA. BOLÍVAR: Distr. Piar, Apa- 
rama Range, Camarcaibarai-tepuí, 2,400 m, shaded ledge, 
B. K. Holst 3632 (MO, Z); Murisipán-tepui, 2,350 m, 

eroded sandstone mesa, Holst, Steyermark & Liesner 
2947 (MO, Z); summit of Auyan-tepui, 2.140 m, sandy 
ground under large rock, Steyermark, Carreño E., 
McDiarmid & Brewer-Carías 116035 (UC); Chimantá 
Massif, Apacará-tepui, ca. 2,200 m, Steyermark, Huber 
& Carreño E. 128445 (MO, UC). amazonas: Dept. Ata 
bapo, Cerro de Marahuaco above Salto Los Monos, 2,255 
m, base of rock, Liesner 17973 (MO), 18003 (MO, Z); 
same locality, 2,520-2,650 m, Steyermark & Holst 


ANN. Missouni Bor. GARD. 76: 605-607. 1989. 


606 


Annals of the 
Missouri Botanical Garden 


LS 


<> 


FIGURE 1. Lindsa 
—C. A single pinnule, x 20. AL from Holst, Steyermark & Liesner 2947 (Z). 


aea mesar 


130808 (MO, Z); same a 2,580-2,600 m, e 
ermark & Delascio 129253 (UC); same locality; sum 


, Ho cia pda & Mori 126072, 125979 
(NY); same locality, 2,580 m, same collectors 725947 
(NY); Cerro Verde Shiho, summit, 2,450-2,480 m, 
same le 126351 (NY); same locality, 2,480 m, 
same collectors 726289 (NY); same locality, 2,450 m, 
same collectors 726351 (NY); Cerro Duida, Ridge 25, 
Tate 430 (NY); Cerro de la Neblina, 3 km ENE of Pico 
Zoloaga, 1,900-2,000 m, Stein & Gentry 1606 (MO), 
same locality, S slopes of Canon Grande, 1,770-1,850 


. Two laminas, x Y. 


— B. Part of secondary rachis with two pinnules, x13. 


m, Croat 59509 (MO); Dept. Rio Negro, Cerro de la 
eblina, n on Venezuela- Brazil border, 2,000 m, W. 
W Thomas & Plowman 3073 (MO, NY, UC). 


The following, almost sterile collection may also 
belong to this species: Cerro de la Neblina, Liesner 


16023 (MO, UC). 


All collections are from cracks and crevices of 
sandstone rocks, from sandy soil in open thickets 
or other vegetation, or from open forest with Bon- 
netia roraimae (where recorded), between ca. 


Volume 76, Number 2 
1989 


Notes 607 


1,800 and 2,800 m. The species is often described 
as being locally common. 

The interrupted sori were at first taken for an 
abnormality due to incomplete fertility of the leaves, 
as is often seen in the genus. The constancy of the 
character, however, leaves no doubt about its being 
a distinctive feature of the species. 

Lindsaea mesarum resembles L. stricta (Sw.) 
Dryander var. jamesoniiformis Kramer in habit, 
but this is certainly due to convergence. The two 
taxa occur in the same habitat and are sometimes 
collected next to each other. The closest relative 
of L. mesarum is L. klotzschiana Moritz ex Et- 
tingsh., which has thinner, much more pronounced- 
ly incised pinnules and larger rhizome scales. 
Another probable relative is L. parkeri (Hooker) 
Kuhn subsp. steyermarkiana Kramer, a little- 
known taxon of somewhat uncertain status; it has 
thinner pinnules that are much less narrowed to- 
wards the apex. If the inconspicuous incisions of 


the anterior margin of the pinnules of L. mesarum 
are overlooked, it will key out to £. stricta (Sw.) 
Dryander in the present author's key (Kramer, 
1957); from this it can be distinguished by its 
smaller rhizome scales, and the forms of L. stricta 
with dark leaf axes have strongly rounded pinnules. 


The author is indebted to the Curators of the 
Herbaria MO, NY, and UC for lending and/or 
donating material, and to Mr. A. Zuppiger, Zürich, 
for preparing the illustration. 


LITERATURE CITED 
o K. U. 1957. A revision of the genus Lindsaea 
n the New World with notes on allied genera. Acta 
Bot. Neerl. 6: 97-290. 
—K. U. Kramer, Institut für Systematische Bo- 
tanik der Universitat Zurich, Zollikerstrasse 107, 
CH-8008 Zürich, Switzerland. 


TWO NEW SPECIES 

OF PASSIFLOR A 
SECTION DECALOBA 
(PASSIFLORACEAE) FROM 
COSTA RICA 


Living collections of two species of Passiflora 
L. section Decaloba DC. were made in Costa Rica 
and subsequently studied in cultivation. Both were 
found to represent undescribed species. The study 
of the living plants as well as herbarium specimens 
has allowed the following detailed descriptions. Col- 
or names follow the standardized English names 


system of the ISCC-NBS (1965). 


Passiflora nubicola MacDougal, sp. nov. TYPE: 
cultivated at Duke University, 1980, from 
stem of MacDougal 220 collected 27 Aug. 
1977 in: Costa Rica. Puntarenas: Cordillera 
de Tilarán, road through Monteverde Cloud 
Forest Reserve, Pacific slope just below con- 
tinental divide, ca. 1, m, MacDougal 
220GR (holotype, DUKE; isotypes, BM, CR, 
MEXU, MO). Figure 1. 


Passiflora scandens; petioli eglandulosi; folia bilobata 
vel trilobata, lobis obtusis vel truncatis, angulo inter lobos 
laterales 70-90(-100P, lobis lateralibus quam centrali 
subaequalibus vel longioribus, marginibus integris; brac- 
teae ad 1.4 mm longae; flores parvi albidi subpendentes; 
coronae filamenta uniseriata vel biseriata, filamentis ex- 
terioribus luteolis 2-3 mm longis; ia plicatum; 
ovarium glabrum; semina 8-10 sulca 


Gracile climbing perennial herb 1.5-3(-4) m 
long, lightly and minutely puberulent throughout 
at shoot tip (except laminas) with antrorsely ap- 
pressed trichomes 0.04-0.10 mm long, becoming 
very sparsely puberulent or glabrescent on mature 
structures. Stem 1.5-3 mm diam., perennial with 
little secondary growth (several-year-old stems 3 
mm diam., dark green, not woody), obtusely sub- 
5-angulate (drying acutely sulcate-striate) and 5- 
6-carinate, the carinae becoming verrucose and 
somewhat scabrous below; posture of p b 
cernuous; phyllotaxy 2/5. Stipules 1-2(-2 
0.15-0.25 mm, linear-triangular, falcate at e 
necrescent. Petioles 0.8-2(-2.8) cm, eglandular. 
Laminas 2.2-4(-4.6) x 4-6(-9.2) cm at fertile 
nodes, depressed-obovate to strongly depressed- 
obovate or semicircular in general outline, adaxially 


ANN. 


glabrous, often with slight variegation consisting of 
a few discontinuous patches of white between (not 
along) the primary and secondary veins, abaxially 
very sparsely puberulent with trichomes restricted 
to primary and secondary veins, sometimes flushed 
purplish (especially new growth), margins entire, 
sparsely puberulent to nearly glabrous, shallowly 
(2-)3-lobed less than 0.15 the distance to the ob- 
tuse base, the lateral lobes obtuse to rounded or 
truncate, the central lobe (obsolete to) obtuse or 
truncate and sometimes emarginate, the angle be- 
tween the primary lateral veins 70-90(-100P, the 
ratio of lateral to central vein lengths 1.0-1.4, the 
ratio of laminar width to length 1.5-2.1; laminar 
nectaries 8—14(-18) per leaf (to 26 on immature 
plants), borne abaxially between the primary veins, 
often appearing light yellowish adaxially, the most 
proximal pair only occasionally strictly axillary but 
not enlarged; leaves of juvenile plants similar, var- 
iegated, vein ratio close to 1.0; seedlings unknown. 
Tendrils straight during development at shoot apex. 
Prophyll of vegetative ramifying bud 1, lanceolate, 
acuminate. Peduncles 1.0-2.0 cm, geminate, un- 
iflorous; bracts 3, 0.7-1.4 x 0.05-0.25 mm, usu- 
ally borne near apex of peduncle, linear-triangular, 
often involute near base and appearing narrowly 
lanceolate, early necrescent, stramineous. Flowers 
-2 cm diam., subpendent, whitish with green 
and yellow corona, slightly sweetly odorous; floral 
stipe (3.0-)4-6.0 mm (to 5-9 mm in fruit); hy- 
panthium 5-5.5 mm diam.; sepals 7.5-11 x 2.6- 
4.4 mm, oblong to oblong-lanceolate, rounded, 
ecorniculate, pale yellow-green abaxially, whitish 
adaxially; petals 5.0-7.5 x 2.4-3.4 mm, oblong 


to ovate-oblong, rounded, white; filamentous co- 


ca. 1.7 


rona in 1-2 series, the outer filaments ca. 23-25, 
2-3 mm, simply curved and spreading, thickest 
near the base, abaxially flattened, slightly laterally 
compressed, the apex obtuse or rounded, light yel- 
lowish green basally, bright yellow toward apex; 
inner series (if present) 1.8-2.2 mm, capillary, 
clavate or dilated at the apex, inclined toward an- 
drogynophore; operculum 0.9-1.5 mm, membra- 


Missouni Bor. GARD. 76: 608-614. 1989. 


Volume 76, Number 2 Notes 
1989 


609 


FIGURE 1. Passiflora nubicola. — A. Habit A O 220GR).— 
leaf (Brenes 6158). — D. Seed with raphe (Dryer 460). 


— B. Flower (MacDougal 220GR).— C. Large 


610 


Annals of the 
Missouri Botanical Garden 


nous, plicate, pale yellow-green; nectary-trough 
without a raised annulus; limen (disk) ca. 2.5-2.7 
mm diam., not colored or spotted; staminal fila- 
ments connate for 4.7—5.9 mm along the very pale 
green androgynophore, the free portions ca. 3-4 
mm, pale yellowish green; anthers 2.2-2.5 mm, 
not marked with purple; ovary 2.0 x 1.8-2.0 
mm, widely obovoid to widely ellipsoid, glabrous, 
light yellow green; styles ca. 4-5 mm, light yellow 
green; stigmas 1.5-2.4 mm diam., capitate. Fruit 

5 X 1.8-2.1 cm, widely ellipsoid to very 
widely obovoid, obtusely trigonous in cross section, 
exocarp purplish or bluish black, glaucous, meso- 
carp spongy, white; arils no longer than seeds, 
clear, gelatinous; seeds 3.6-4.2(-4.5) mm long, 
2.5-2.9 mm wide, 1.5-1.6 mm thick, obovate (to 
widely obovate), very slightly obcampylotropous (or 
nearly symmetric), transversely sulcate with (7-)8- 
10 sulcae, the ridges verrucose or knobbed, the 
chalazal beak erect to slightly inclined away from 
raphe. Chromosome number n — 


Additional specimens examined. Costa Rica. 
ALAJUELA: Finca Johanson, Los Angeles de San Ramón, 
[ca. 10%09'N, 84°29'W], 11-12 Mar. 1928 (fl), Brenes 
6019 (F); camino de Finca Johanson, Los Angeles de San 
928 (fl), Brenes 6158 (F); road to Penas 
km from continental 
divide, Atlantic slope, 1,400 m, 13 Apr. 1981 (fl, fr), 
Knapp & Mallet 857 (TEX) road to Peñas Blancas ca. 
3-4 km from divide, Atlantic slope, 1,300 m, 13 Apr. 
1981 (fl), Knapp & Mallet 858 (TEX). CARTAGO: Ta- 
panti, 1,300-1,700 m, 26 Oct. 1983 (fl), Chacon et al. 
1530 (CR); ca. 1 km N and downstream of the junction 
of the Rio Dos Amigos and the Río Grande de Orosi on 
the road to Tapanti, 1,500 m, 10 Aug. 1980, MacDougal 


1244 (DUKE). PUNTARENAS: vertiente Pacifico, Reserva, 
Monteverde, 1,520-1,580 m, 17 July 1976 (fr), Dryer 
460 (CR, F); 14 June 1977 (fr), Dryer 1449 (F); 27 


Aug. 1977, MacDougal 220 (DUKE). 


This small and inconspicuous passionflower is 
known from four areas of lower montane rainforest 
in Costa Rica. Passiflora nubicola grows at edges 
of primary forest, in treefalls, or along trails, where 
its thin wiry dark green stems trail over fallen logs 
and among grasses, bushes, and small trees. The 
leaves are similar in shape to those of a juvenile 
P. biflora Lam., and their flecks of white varie- 
gation are similarly distributed. The flower also 
resembles that of a miniature P. biflora, but in the 
Orosi valley possesses only one row of coronal 
filaments. The species is closely related to P. bi- 

ora. 

At the Monteverde Reserve, the main herbivore 
of this species is Heliconius clysonimus (J. Mallet, 
pers. comm 

The clone on the same plant as the holotype 
of this species was grown at Duke University 1977- 


1983, and the chromosome count was determined 
from the clones of this plant. Floral anthesis was 
around 0830 hrs. in the greenhouse, with the flow- 
ers closing in the early afternoon. No fruits were 
set by autogamy during cultivation, and more than 
40 attempts at self -pollination also failed to produce 
fruit. Passiflora nubicola was very fertile in both 
directions with P. biflora (MacDougal 260GR & 
275GR), yielding well-seeded fruit and vigorous F, 
progeny. Those progeny, generally morphologi- 
cally intermediate in vegetative and reproductive 
character states, had variously malformed flowers 
and were sterile, producing no viable pollen. Fruits 
(with hybrid embryos) produced on P. nubicola 
matured in 45-48 days and contained 35-64 seeds 
per fruit (N — 13 fruits). 

The epithet means *'cloud-dweller," 
to its habitat of misty forest. 


in reference 


Passiflora gilbertiana MacDougal, sp. nov. TYPE: 
osta Rica. San José: along roadcut 18.2 km 

S of Villa Mills on Pan-American Highway, 
7,700 ft, 10 Aug. 1977, L. E. Gilbert 8 
(holotype, LL; isotypes: F, MO, MSC). Figures 


Passiflora scandens; petioli eglandulosi; folia supra vit- 
tata, infra purpur el atrovinos bata vel tri- 
lobata, lobis lateralibus acuminatis vel acutis, interdum 
deu lobo centrali acu 

nterdum obsoleto, angulo inter lobos laterales 13-21", 
lobis lateralibus quam centrali subaequalibus vel brevio- 
ribus, marginibus integris; bracteae 0.8-4.5(-5.7) mm 

A i iusculi, odorem 
menta uniseriata vel 


ten ex alentes; coronae fila 
ata ,aureis, 5-7 mm 


pue operculum plicatum; ovarium pubescens; semina 
6-8 sulcata. 

Perennial climber 2-4 m long, minutely + pu 
berulent throughout (except laminas) with (0.06-) 
0.10-0.15(-0.20)-mm suberect or slightly an- 
trorsely appressed trichomes, the shoot tip densely 
minutely puberulent, becoming moderately (to 
sparsely) puberulent below. Stem 1.5-5 mm diam., 
with normal secondary growth below, subterete 
(drying angular), striate; posture of shoot tip cer- 
nuous; phyllotaxy 2/5. Stipules (1.7-)2.5- 
4.5(-6.0) x (0.2-)0.3-0.5(-0.7) mm, falcate, 
narrowly lanceolate to linear-triangular, very dark 
purplish, distally stramineous, yellow, or brown and 
necrescent. Petioles 0.8-3.5 cm (to 5 cm in ju- 
veniles), eglandular, adaxially puberulent, abaxially 
glabrous or sparsely puberulent. Laminas (4-)6- 
13(-17.5) x (1.7-)2.5-6.5 cm at fertile nodes, 
elliptic to narrowly obovate or oblong-obovate in 
general outline, adaxially glabrous or nearly so with 
a few trichomes proximally on primary veins, olive 


Volume 76, Number 2 Notes 611 
1989 


wwe 


uoce 


A 


FIGURE 2. Passiflora gilbertiana. —A. Habit (MacDougal 655).—B. Flower (MacDougal 655).—C. Seed 
(Gilbert 8). —D. Leaf of seedling (MacDougal 1432). —E. Large leaf from below reproductive nodes (MacDougal 
1431). 


612 


Annals of the 
Missouri Botanical Garden 


green or flushed purplish, striped with white or pale 
greenish yellow the length of the 2 lateral veins 
and sometimes along the basal part of the central 
vein, abaxially sparsely puberulent to glabrescent 
with trichomes mainly restricted to the primary 
and secondary veins, very deep purplish red to 
very dark reddish purple, becoming greenish on 
older leaves, the margins entire, puberulent to 
sparsely puberulent with 0.06-0.10-mm tri- 
chomes, very shallowly 3-lobed or shallowly 2-lobed 
for 0.04-0.15(-0.25) the distance to the cuneate 
to obtuse (or narrowly rounded) base, the lobes 
acuminate to acute, sometimes rounded at the very 
apex, or the central lobe obtuse to obsolete, the 
angle between the primary lateral veins 13-21^*, 
the ratio of lateral to central vein lengths 0.94— 
1.2(-1.3), the ratio of laminar width to length 
0.35-0.50; laminar nectaries (5-)9-14(-17) per 
leaf (fewer on seedlings), borne abaxially between 
the primary veins, llate with. purple border, 
appearing light yellowish adaxially, the most prox- 
imal pair not strictly axillary or enlarged; leaves 
of juvenile plants similar, to m, with 
slightly larger lobe angle and relative width; leaves 
of seedlings with lobe angle 40—60°, central lobe 
on first leaves longest, then subequal in length, 
nearly as wide as long. Tendrils straight during 
development at shoot apex. Prophyll of vegetative 
ramifying bud 1, broadly lanceolate and often tri- 
dentate, acuminate, very dark purple. Peduncles 
2.4-3.8(-4.4) cm, geminate, uniflorous; bracts 3, 
(0.8-)1.7-4(-5.7) x (0.1-)0.2-0.4 
triangular, very dark purplish proximally, stra- 
mineous, yellowish, or brownish and necrescent 
apically. Flowers ca. 2 cm diam., suberect, pale 
greenish with a golden corona, with a strong putrid 
odor of scatole; floral stipe 4.0-6(-8.0) mm; hy- 
panthium 6 mm diam.; sepals 11-15 x 3.0- 
4.5(-6.0) mm, narrowly triangular-oblong or lan- 
ceolate-oblong, rounded, ecorniculate, lightly but 
evenly puberulent to apex, very light yellow green 
on both sides; petals 2.0-3.5 x 1.0-1.5 mm 
narrowly oblong-ovate to ovate, rounded, very light 


mm, linear- 


yellow-green; filamentous corona in 1-2 series, the 
outer filaments ca. 46-50, 5-7 mm, strongly 
spreading and only slightly and simply curved, 
cylindrical and gradually slightly ampliate toward 
the rounded apex, pale yellow, yellow-green, or 
whitish basally, the remainder vivid yellow; the 
inner filaments (if present) 0.6-1.2 mm, expressed 
mainly in the sepal sectors, slightly clavate, erect; 
operculum 1.8-2.4 mm, membranous, plicate, 
cream to yellow; nectary-trough without raised an- 
nulus; limen (disk) ca. 2 mm diam., pentagonal, 
bright yellow; androgynophore pale at base, dark 
pink to grayish purplish red distally; staminal fil- 


aments connate (6.2-)7.5-8(-9.0) mm along an- 
drogynophore, the free portions 5-6 mm, dark 
pink at base, light yellow-green distally; anthers 
2.0-2.2 mm, not marked with purple; ovary 1.8- 
2.5 x 1.4-2.1 mm, widely ellipsoid to subglobose, 
densely puberulent, dark green; styles ca. 6 mm, 
light yellow green; stigmas ca. 1 mm diam., clavate- 
capitate. Fruit 1.2-1.4 cm diam., subglobose to 
widely ellipsoid, lightly puberulent, purplish or bluish 
black; arils translucent white, length unknown; seeds 
(2.9-)3.1-3.3 mm long, 1.9-2.1 mm wide, 1.1- 
1.2 mm thick, obovate, nearly symmetric to very 
slightly campylotropous, transversely sulcate with 
(6-)7-8 sulcae, the ridges coarsely verrucose or 
knobbed, the chalazal beak very slightly inclined 


toward raphe. 


Additional specimens examined. COSTA RICA. SAN 
José: clone of Gilbert's type collection cultivated at Duke 
University 1980-1984, MacDougal 655 (BM, CR, 

JKE, HUA, MEXU, TEX); from type locality, Au p? 
1980, MacDougal 1431 (CR, DUKE, MO), 
DUKE); side roads along the eric y. 
10-15 km N of San Isidro de General, near kms 18. 
116, at the crossing of the jen Paymer, and N along the 
road from there, 1,500-1,600 m, 8 Aug. 1981, Taylor 
1120 (DUKE); along the nie Hwy. above San 
Isidro de General, ca. 15 km N at river crossing, 1,500 


m, 5 Mar. 1985, Taylor & Skotak 4771 (DUKE, MO). 


~ 


Passiflora gilbertiana is remarkable for its var- 
iegated reddish purple leaves and stinking golden 
flowers. It is a member of a species group of Central 
American passionflowers in section Decaloba char- 
acterized by bilobed variegated leaves, greenish to 
yellow perianth, filiform unbanded outer coronal 
filaments, and notable reduction in the size of the 
petals relative to the sepals. This group is part of 
section Decaloba but was not recognized by Killip 
(1938), who scattered its component species among 
of section Decaloba. 


several of his invalid *'series" 
Within the group, the new species is most closely 
related to P. jorullensis H.B.K. of Mexico, P. 
apetala Killip of Costa Rica and Panama (Fig. 3), 
and an undescribed relative of P. apetala from 
the mountainous border of Mexico and Guatemala. 
Its distinctive characteristics include its long nar- 
row leaf with a relatively small interlobe angle and 
more strongly expressed red to purple coloration, 
retention of conspicuous laminar variegation at ma- 
turity, and yellow flower with small petals and 
second (inner) series of coronal filaments. The inner 
series of coronal filaments varies as to presence or 
absence within the species. In Taylor & Skotak 
4771 the series is well developed, numbering ca. 
40; in the type it is absent. However, the cultivation 
of somatic clones (MacDougal 655) from the type 
plant revealed that the expression of the second 


Volume 76, Number 2 
89 


Notes 613 


FIGURE 3. 


368). — B. Passiflora gilbertiana m 655). —C. Passiflora apetala (MacDougal 467).— 
0). 


—B. 
apetala, flower only (MacDougal 45 


filamentous series was variable within this individ- 
ual, with (0-)5-12(-20) filaments per flower. The 
expression changed gradually along the stem, at 
times stronger, at times weaker, perhaps dependent 
on environmental conditions. In certain cases the 
expression seemed to be related to the position of 
the flower on the vine, with smaller flowers near 
the end of flowering axillary branches completely 
lacking the second series. 

On the label of the type is the collector's note 
"odor of pigpen." The flowers of this new species 
share this strong odor (described as between “ster- 
coraire et putride" by Planchon, 1849) with P. 
jorullensis (pers. obs.). Gilbert also noted on the 
type label **visited by hummingbirds,” but the same 
day he also observed a single wasp flower 
(pers. comm.). It is thus probably significant that 
P. jorullensis, with a nearly identical odor, is pri- 


at a 


marily wasp-pollinated at least in some part of 
Mexico (MacDougal, unpubl.). 

The habitat at the type locality is treefalls and 
partially shaded thickets along a dirt road on steep 
slopes in the lower reaches of Alnus forest. Plants 
there climbed over bushes and trailed into small 
Alnus and Piper trees. An important herbivore at 
the type locality is a yellow flea beetle that is an 


Comparison of leaves and flowers of three closely related species. — A. Passiflora Dino (MacDougal 


— D. Passiflora 


undescribed species of Monomacra (from Mac- 
Dougal 1431, voucher identified by C. Duckett, 
deposited in the Cornell University Insect Collec- 
tion). Several of the cited specimens show char- 
acteristic damage by flea beetles. 

A clone from the type collection was cultivated 
at Duke University and from there was distributed 
to several commercial greenhouses in California 
and made available for sale to the public by 1980 
In cultivation the plant bloomed from the morning 
to afternoon, with anthesis around 0930 hrs. and 
flower closing in mid to late afternoon. The stigmas, 
although lowered to anther level into the afternoon, 
were not receptive after ca. 1230 hrs. No fruits 
were set by autogamy during cultivation, but during 
limited hand-pollinations, 6 of 11 self-pollinations 
before 1230 hrs. produced mature fruit containing 
19-31 seeds per fruit. 

Mr. Patrick Worley, a commercial plant hy- 
bridizer, crossed a clone of the type with P. biflora, 
P. ornithoura Masters (MacDougal 623), and P. 
jorullensis (MacDougal 368); many F, progeny 
were grown to maturity and all flowered. Pollen 
viability was not examined. In each case the un- 
pleasant odor was conserved. One especially or- 
namental individual of the cross with P. jorullensis 


614 


Annals of the 
Missouri Botanical Garden 


was named *'Sunburst" and has been available to 
the public from several nurseries since 1983. 

I take great pleasure in naming this species for 
its discoverer, Lawrence E. Gilbert of The Uni- 
versity of Texas at Austin, whose seminal contri- 
butions to the understanding of the ecology of the 
Passifloraceae and their herbivores are well known. 
Dr. Gilbert first suggested the study of passion- 
flowers to me and has aroused and guided an in- 
terest in these plants in many other students. It is 
particularly gratifying to name this plant because 

had the privilege of being with Dr. Gilbert the 
day he discovered it and gathered the type collec- 
tion. He immediately suspected it to be undescribed 
and his exuberance kindled in me what has become 
a career. 


Many of the observations given here were made 
at Duke University, and I gratefully acknowledge 
Donald E. Stone for his inspiration and guidance 
during my graduate studies. Thanks are due the 
Organization for Tropical Studies for their summer 


field course that introduced me to Lawrence Gilbert 
and to these Costa Rican passionflowers. The staff 
of the Duke University greenhouses provided many 
years of favorable conditions for passionflower vines, 
and the Jessie Smith Noyes Foundation generously 
provided postdoctoral support for research on this 
family at the Missouri Botanical Garden. The draw- 
ings were prepared by John Myers. 


LITERATURE CITED 
ISCC-NBS (Inter-Society Color Council — National Bu- 


The American species of Passi- 


loraceae, Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 


BR, J. E. 1849. Fl. Serres Jard. Eur. 5: 528. 


— John M. MacDougal, Missouri Botanical Gar- 
den, P.O. Box 299, St. Louis, Missouri 63166, 
U.S.A 


PASSIFLORA MAYARUM, 
A NEW SPECIES RELATED 
TO P. PROLATA IN 
SUBGENUS PASSIFLORA 
(PASSIFLORACEAE) 


Collections from Central America made since 
the time of Killip's (1938) monumental monograph 
of the American Passifloraceae have allowed a 
clearer delineation of a number of taxa, and in 
some cases have confirmed his suspicions that cer- 
tain specimens might represent undescribed species. 
Such is the case with Passiflora prolata Masters 
and its apparent sister species newly described here. 


Passiflora mayarum MacDougal, sp. nov. TYPE: 
Mexico. Quintana Roo: roadside secondary 
scrub 3 km N of San José de la Montaña on 
road N of Tomás Garrido (ca. 75 km W of 
Chetumal), 100 m, 9 May 1982, Chater et 
al. 89 (holotype, MO; isotype, BM). Figures 
1A, C, 2. 


Passiflora (subgenus Passiflora) scandens, _ 
trichomatibus 0.05-0.25 mm longis; stipulae 10-22 x 
.0 mm; petioli 2(-4)-glandulosi; folia trilobata, lobis 
acuminatis vel caudatis, marginibus serratis vel serrulatis; 
bracteae 4.8-6.6 x 2.0-3.4 cm, ova 
npe coronae filamen i 
apicem in lacinias divi 
puberulum. Passiflora prolata affinis est. 


Vine, size unknown, puberulent throughout ex- 
cept glabrous or nearly so on tendrils, adaxial sur- 
face of lamina, flowers, and sometimes adaxial sur- 
face of bracts. Trichomes 0.05-0.20(-0.25) mm, 
erect and straight or curved. Stems subterete to 
obtusely subangular, striate, apparently not hollow. 
Stipules (10-)13-18(-22) x 0.4-1.0 mm, linear- 
narrowly oblanceolate (or linear-triangular to lin- 
ear-narrowly lanceolate), long-attenuate, tardily 
deciduous, glandular-denticulate; petioles 2(-4)- 
glandular 4—% the distance to the appex (or if 

-glandular, rarely 1 pair near the apex), the nec- 
taries sessile or up to 1 x (0.7-)1-2.2 mm, (cra- 
teriform to) patelliform; laminas 10-13 x 9-14 
cm, glandular-serrate(-serrulate), not variegated, 
glabrous adaxially except the primary veins pu- 
berulent at least half their length, 3-lobed for 0.4— 
0.7 the distance to the base, the lobes lanceolate 
to narrowly ovate-oblong, acuminate to short-cau- 


date; laminar nectaries marginal, 2-5 slightly en- 
larged teeth borne in each lobe sinus, enlarged 
teeth absent at base of lamina. Prophylls of veg- 
etative ramifying bud 2, 1-2 mm long, lanceolate, 
equal or one smaller. Peduncle 1 per node, 1.5- 
4(—5.0) cm, uniflorous; bracts 4.8-6.6 x 2.0-3.4 
cm, valvate in bud, free to the base, ovate-elliptic, 
conspicuously 2—4(—6)-glandular at the margin ba- 
sally, otherwise entire or the basal Y2 obscurely 
glandular-denticulate, acuminate to long-acumi- 
nate, yellow green, deciduous in fruit, adaxial sur- 
face puberulent or glabrous with puberulent main 
veins and a narrow band of tomentum along mar- 
gin, the venation basally acrodromous with prom- 
inent intramarginal veins. Perianth white, longest 
coronal filaments purplish to violet, white at the 
apex, banded white basally; floral stipe ca. 12-14 
mm (to 20 mm in fruit); hypanthium ca. 5-6 mm 
long, ca. 13-14 mm diam., puberulent; sepals 24— 
34 x 7-12 mm, lanceolate-triangular, puberulent, 
sometimes only sparsely so, subapically with an 
arista (8-)10-18 mm; petals 24-34 x 6-9 mm, 
oblong-lanceolate; coronal filaments in 6-8 series, 
the 2 outermost similar, often apically tortuous, 
the outer 26-32 mm, the second series broader 
at their base and ca. 3-6 mm shorter, the next 
(3-)4-5 series tuberculate or 1-5(-7) mm, the 
innermost series 6-10 mm, banded purple and 
white; operculum basally connate for 2-4.5 mm, 
membranous, the margin erose or short-fimbriate 
and often slightly plicate, with a subapical and 
slightly deflexed series of fimbriae or filaments l- 

(-5) mm long; nectar ring (annulus) present in 
the middle of the nectary; limen (disk) ca. 6-8 mm 
diam. around the widely broadened base of the 
androgynophore, ca. 3 mm above the floor of the 
nectary; staminal filaments connate 10-13 mm 
along androgynophore, the free portions 8-9 mm, 
glabrous; anthers 10-12 mm; ovary 5-6.5 x 2- 
3.2 mm, narrowly ovoid-ellipsoid, glabrous (rarely 
sparsely puberulent); styles 10-11 mm including 
stigmas, glabrous; stigmas 4 mm wide, deeply bi- 


lobed. Fruit 7.5-9 x 4.5-5 cm, ellipsoid, greenish 


ANN. MISSOURI Bor. GARD. 76: 615-618. 1989. 


616 Annals of the 
Missouri Botanical Garden 
A 
E 
E 
e 
FIGURE 1.— A. Leaf of Passiflora mayarum with detail of marginal teeth (Téllez & Cabrera 2429). — B. Leaf of 


P. diri with detail of marginal teeth (MacDougal 596). — 


yellow to yellow, estipitate; arils longer than the 
seeds, color and flavor unknown; seeds 7.2-8.0 
mm long X 4.0-4.5 mm wide X 1.9 mm thick, 
obovate, strongly compressed, nearly bilaterally 
symmetric with the chalazal end squared and shal- 
lowly 3-dentate, brown, reticulate-foveate with 35— 
46 foveae. 


Habitat and distribution. This species grows 
in the Caribbean lowlands of northern Guatemala, 


92 w 


FIGURE 2. Map of Guatemala, Belize, and neighboring 
countries showing distributions of Passiflora mayarum 
rolata. 


and P. p 


C. Seed of P. mayarum (Téllez & Cabrera 2429). 


western Belize, and the southern part of the Yu- 
catán Peninsula (Fig. 2). The collections come from 
elevations of 20-250 m, except for two collections 
from one station at 550 m in Belize. The habitat 
is usually open or partially open vegetation. Al- 
though the information is not present on the spec- 
imen's label, Bartlett (1935: 20) stated that his 
collection 12840 was collected in the wooded swamp 
vegetation called escobal, which is a vegetation 
type dominated by the escoba palm, Cryosophila. 
This edaphic climax occurs in the Petén and Yu- 
catán on wet soil surrounding water holes (““agua- 
das") and as the transitional zone between the 
inundated areas of the w 
and the well-drained **high bush” uplands (Bartlett, 
1935; Lundell, 1937). However, most of the col- 
lections are from disturbed and more well-drained 
habitats, namely, roadsides and secondary areas in 
low forest, selva mediana subperennifolia, or sel- 


va alta subperennifolia (Rzedowski, 1978). 
Phenology. 


The few collections known sug- 
gest that reproduction occurs primarily during the 
wet season from April through November. One 
sheet, however, was collected in flower at the end 
of December. Although the type has nearly mature 
fruit, fully mature fruit and seeds are known only 


from Téllez & Cabrera 2429. 
“Granadilla” (Tún O. 372). 


Vernacular name. 


Volume 76, Number 2 
1989 


Notes 617 


Additional specimens examined. MEXICO. CAM- 
PECHE: 30 km al S de Xpujil, 22 Apr. 1982, Cabrera & 
Cabrera 2482 (MEXU, MO). QUINTANA ROO: 8 km al S 
de Tres Garantias, sobre el camino a Tomás Garrido, 22 


pad pe Cabrera & Cabrera 4174 ae MO); 24 


T 


vas, 
1973, Dwyer 10823 (MO); vic. Millionario, 1,800 ft., 
30 May 1973, A. Gentry 7733 (MO). GUATEMALA. PETÉN: 


of Tikal National Park, 19 June 1973, Dwyer 11269 
(MEXU, MO); S of Tikal National Park, 19 June 1973, 
A. Gentry 8338 (MO—2 sheets, US); camino de El 
Remate en el Km 59, Parque Nacional de Tikal, 24 Oct. 
1969, Tún O. 372 (F, NY, US); Santa Elena, en el camino 
para San Andrés, Km 4, lado saliente, 4 Oct. 1969, Tün 
O. 1374 (F) 


Passiflora mayarum is most similar to and is 
undoubtedly very closely related to P. prolata from 
the premontane to montane broad-leaved forests 
of the Pacific watershed of Guatemala and Chiapas, 
Mexico. Except for the vestiture, the two have 
similar leaves and nearly identical glandular bracts. 
At the time of his 1938 monograph, Killip knew 
Passiflora prolata only from the type. He knew 
the related species newly described here from a 
single collection by Bartlett (12840). He had iden- 
tified that sheet as P. prolata (Killip, 1936), but 
later expressed some misgivings. In his monograph 
(1938: 389) he stated, “I am referring the Bartlett 
collection to P. prolata with some hesitation, as 
the stem and under surface of the leaves are pu- 
berulent and the petiolar glands are not quite as 
in the type. It comes, moreover, from the low- 
lands." The more recent collections of both species 
confirm that the two are separate morphologically 
and ecologically. Neither fruit nor seeds are known 
from P. prolata, so it is impossible to compare the 
two species on these points. They may be distin- 


guished by the following key. 


la. Stems and leaves glabrous except for slight pu- 
e h 


long, acuminate. Petioles glandular i in the distal 
half, the glands 0.3-0.8 mm diam. and usually 
longer than wide. [da lobes conspicuously 

ng-acuminate; laminar rulate, often 


t the junction of the petiole. Bracts glabrous 
abaxially. Hy m an als glabrou 
Habitat idu 700-2,000 m ooo... P. prolata 


_ 
c 
un 
er 
O 
d 
un 
pf 
a] 
a 


P - 
berulent. Stipules 10-22 mm long, long-atten- 
uate. Petioles glandular at a variable position, 
the glands 0.7-2.2 mm diam. and wider than 


long. Laminar lobes acuminate to short-caudate; 
laminar margin uniformly serrate (to serrulate), 
not nectariferous at the junction of the petiole. 
Bracts puberulent abaxially. Hypanthium pu- 
berulent, sepals puberulent or only sparsely so. 
Habitat elevation 20-550 m „u P. mayarum 

Not having seen the type of P. prolata (Leh- 
mann 1630—G-BOIS), I have relied on the pro- 
tologue and five more recent collections, including 
one of my own, that match the protologue. Three 
of these collections were identified as P. prolata 
by Killip (who saw the type), and one was collected 
within 30 km of the type locality at a similar 
elevation. The flowers of P. prolata and P. may- 
arum appear to be rather similar, apparently borne 
above the horizontal plane, with two prominent 
series of purplish outer coronal filaments, several 
reduced intermediate series, and a well-developed 
inclined inner series. The operculum in both is 
similar, although that of P. mayarum has shorter 
subapical filaments. I have seen the flower of P. 
prolata only from Matuda 16479 (F), in which 
the intermediate coronal series are reduced to var- 
iably expressed series of tubercles. It is not possible 
to determine clearly the relative length of the two 
outer series except to say that they are roughly 
similar in this poorly pressed specimen. Given the 
lack of comparative reproductive material and the 
coronal variation that occurs within related species, 
I have for now distinguished the two species mainly 
by vegetative characteristics. 

he leaves of P. mayarum tend to be less deeply 
lobed, but there is considerable overlap with P 
prolata. The quality and quantity of the indument 
differ: the few trichomes that occur on the leaves 
and nodes of P. prolata are ca. 0.2-0.4 mm in- 
stead of 0.05-0.25 mm long. Despite the variation 
in their vestiture, the bracts of both species always 
have a narrow band of dense tomentum along the 
margin adaxially. The trichomes are crispate, and 
those of adjacent bracts are interlocked during the 
development of the bud, keeping the bracts valvate. 
This marginal band of crispate tomentum occurs 
in number of species of Passiflora and is correlated 
with foliaceous valvate bracts. The ovary of 
mayarum is glabrous in all collections exiled 
except Tün O. 1374, in which it is sparsely pu- 
berulent. 

Some of the Mexican collections of the new 
species were distributed as P. platyloba Killip and 
are cited as such in Sousa & Cabrera (1983). 

Passiflora mayarum and P. prolata belong to 
subgenus Passiflora, where they share character- 
istics with species that Killip (1938) included in 


his informal *'series" Serratifoliae and Incarnatae. 


618 


Annals of the 
Missouri Botanical Garden 


Killip placed P. prolata next to P. incarnata L. 
as one of six species in /ncarnatae, which was 
MSE n = Passiflora (as sub- 
nus “Granadilla”) as having unwinged stems, 
free te narrow stipules, lobed leaves, eglan- 
dular sepals, filamentous corona in more than two 
series, petiolar glands present, and denticulate or 
fimbriate operculum. That delineation is fraught 
with difficulties since most of those character states 
are undoubtedly plesiomorphic for the subgenus. 
Except for the lobed leaves, it also fits the species 
put in the Serratifoliae group. An understanding 
of the relationships among these species and the 
granadillas as a whole awaits a revision of subgenus 
Passiflora. Passiflora mayarum is probably ge- 
netically close enough to the commercially impor- 
tant P. edulis Sims to warrant investigation into 
using it as a rootstock or as a source of genetic 
rovement for the commercial passionfruit. It is 
especially notable that P. mayarum has been col- 
lected growing in warm wet soil, the kind of con- 
ditions that induce fungus-related failure of the 
commercial crop 
The epithet mayarum refers to the native dis- 
tribution of this species, centered in the heart of 


the Mayan Old Empire 


I am grateful to the Jessie Smith Noyes Foun- 
dation for postdoctoral support for research on 
passionflowers, to the curators of the herbaria listed 
above, and to John Myers for the drawing. 
LITERATURE CITED 
BARTLETT, H. H. 19 A method of procedure be 

field work in tropical American phytogeography ba 


Botany of the Maya Area: Miscellaneous uei I 

I. da Carnegie Inst. Wash. 461(1): 

KILLIP, E. P. WR e of the Mayan region. 
In: Bam of the Maya Area: Miscellaneous Papers. 
I-XIII. Publ. Ca Inst. Wash. 461(13): 299- 


1938. The American species of Passiflora- 
ceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 1 
613 
Lunpett, C. L. 1937. The mm of Petén. Publ. 
negie Inst. Wash. 478: 1-244. 
Ruxnows J. Vegetación de México. Editorial 
, S.A., D.F., 
. & E. F. al C. 1983. Listados 
s de México. II. Flora de Quintana Roo. 


Dn de Biologia, Univ. Autónoma de México, 
D.F., México. 


—John M. MacDougal, Missouri Botanical Gar- 
den, P.O. Box 299, St. Louis, Missouri 63166- 
0299, U.S.A 


TWO NEW SPECIES OF 
TOURNEFOR TIA 
(BORAGINACEAE) 
FROM COLOMBIA 


Recent collecting efforts in western Colombia 
have yielded two very distinct and undescribed 
species of Tournefortia L. Both are members of 
sect. Tournefortia, a pantropical group with its 
center of diversity in Andean South America. 
Species of Tournefortia from this area remain con- 
fused, since the most recent taxonomic treatment 
available is de Candolle's (1845). Macbride (1960) 
covered the species from Peru, but his treatment 
is regional and incomplete. Killip prepared an un- 
published treatment of the Andean species. The 
following two species are new. 


Tournefortia brantii James 5. Miller, sp. nov. 
TYPE: Colombia. Antioquia: Mpio. Valdivia; 
road to Briceno, 1.1 km W of the Medellin- 
Caucasia highway, pasture, and secondary for- 
est adjacent to road, 7%03'N, 75?28' W, 1,800 
m, 4. E. Brant & O. Escobar 1293 (holotype, 
MO 3531269). Figure 1. 


Arbor parva ad 3 m alta, dr aed — dee 
opposita, ovata ad elliptica, a bi m longa, 5-9.5 
lata, apice acuminato, mg 


ay dec — aaa a 
pubescentia, infra villos , bis 


M oe cymos di- 


lobis 5, ovatis, 1 mm longis, tubo 4-5 mm longo, strigoso. 
Fructus non visi 


Small tree 3 m tall, the twigs villous, the hairs 
erect, 1-2 mm long, brown. Leaves opposite; pet- 
ioles 5-12 mm long, villous, the hairs erect, 1-2 
mm long, brown; leaf blades ovate to nearly elliptic, 
9-17 cm long, 5-9.5 cm wide, the apex acumin- 
ate, the base cuneate to decurrent, the margin 
entire but appearing ciliate from hairs of the upper 
surface, the secondary veins 8-9, ascending, aris- 
ing from the midrib at 30°-50°, the tertiary and 
quaternary veins clearly visible on the lower sur- 
face, ramifying reticulately, the upper surface pu- 
bescent, the hairs erect, ca. 1 mm long, translucent 
white to clear, approaching villous along the midrib, 
the lower surface villous, densely so along the mid- 
rib and secondary veins. Inflorescence a twice di- 
chotomously branched cyme, the peduncle 4-11 


ANN. 


mm long, villous, the hairs erect, 1-2 mm long, 
brown, the fertile branches 3-9 cm long, the flow- 
ers borne densely at early anthesis, 1-2 mm apart, 
the fertile branches later becoming more lax, the 
flowers then 3-5 mm apart. Flowers bisexual, with 
the stigma borne beneath the anthers; sepals 5, 
free to the base, linear-lanceolate, 3-5 mm long, 
unequal in length, densely brown strigose; corolla 
white, tubular with 5 spreading lobes, 5 mm long, 
the lobes ovate, 1 mm long, the tube 4-5 mm 
long, brown strigose, densely so near the apex, 
sparsely so near the base; stamens 5, the anthers 
borne just beneath the mouth of the corolla tube, 
nearly sessile, ellipsoid, 0.8 mm long; ovary ovoid, 
] mm tall, the style 3 mm long, the stigma conical. 
Fruits unknown. 


Tournefortia brantii is a member of sect. Tour- 
nefortia and belongs to a group of about 10 poorly 
defined species that have opposite leaves. These 
species occur from Costa Rica through Colombia, 
Ecuador, Peru, and into Bolivia and are centered 
in the northern Andes. While problems exist with 
species delimitation among the opposite-leaved 
species of Tournefortia, T. brantii is distinct among 
the South American species by having dark brown 
indument and sepals to 5 mm long. It is most similar 
to T. johnstonii Standley, a species of Panama 
(Miller, 1988), which differs in having sepals longer 
than 6 mm and a corolla tube 8-10 mm long. The 
other species of Tournefortia with opposite leaves 
in Colombia, 7T. pauciflora Nowicke and T. sca- 
brida Kunth, differ in having shorter calyces and 
less prominant indument on the twigs (Nowicke, 


1974). 


Tournefortia spicata James 5. Miller, sp. nov. 
TYPE: Colombia. Narino: Mpio. Tumaco, 80 
km from Tumaco along road to Pasto, in wet 
area of primary forest, near Orbignya palms 
and streams, “Guayacana,” 19 July 1984, 
M. J. Balick, L. E. Forero P., S. R. King & 
N. Hernández 1667 (holotype, MO 3531270). 
Figure 


Missouni Bor. GARD. 76: 619-622. 1989. 


620 Annals of the 
Missouri Botanical Garden 


N? 3531269 


2 COLOMBIA 
= AG INA‘ 
N 7 
i o 
D = 3 l. « t l 1 = 
i griw . Pasture ind sé )ndatr 
- t t ro 
~ r 1 ^ 
ir 1293 
URI TANICAL ARDEN HERBARIUM (MO) 


FIGURE 1. Tournefortia brantii (holotype). 


Volume 76, Number 2 Notes 621 
1989 


N2 3531270 


PLANTS OF COLOMBIA 


No. 16€ Pa has 
Her wers y w t É (dm 
rea primary f st r Qrbignya 
6 treams 
bi | Nar ifc un. T 80 k 
3 along ad t Paste 'Guay ac an 
Forer R. K 
54. 
w Pi Age f r 
veloj t e Mertz-Gils F r 
tt r the ar n B a 


FIGURE 2. Tournefortia spicata (holotype). 


622 


Annals of the 
Missouri Botanical Garden 


Frutex ad 3 m altus, ramunculis veluntinis ad Meri 
Folia de ovata, 14.5-21.5 cm longa, 8-10 cm lata, 
acuta, supra et infra 


Sepala [eet lanceolata, m longa; corolla lutea, 
ps nigra, 8 mm m longa, Jobis 5, ovatis, 2 mm longis, 
4-5 mm longo, hirsuto. Fructus non visi. 


Soft-stemmed shrub to 3 m tall, the twigs ve- 
lutinous to villous with erect golden-brown hairs 
ca. 1 mm long. Leaves alternate; petioles 11-28 
mm long, velutinous, the hairs erect, ca. 1 mm 
long, golden brown; leaf blade ovate, 14.5-21.5 
cm long, 8-10 cm wide, the apex acuminate, the 
base obtuse to widely acute and decurrent along 
the petiole for 0.5-1 cm, the margin entire, sec- 
ondary veins 13-15, arising from the midrib at 

09-80? near the base of the leaf, 35°-40° near 
the apex, the tertiary and quaternary veins clearly 
evident, ramifying in a reticulate pattern, the upper 
surface hirsute, the hairs erect to spreading, arising 
from the aeroles only giving a slightly bullate ap- 
pearance to the leaf at maturity, the midrib villous, 
the lower surface hirsute, villous along the midrib 
and y veins. Inflorescences densely spicate 
borne in pairs from a dichotomously branched pe- 
duncle 9-14 cm long, this velutinous to villous, 
the hairs erect, ca. 1 mm long, golden-brown, the 
spikes 5-8 cm long, 1.5-2.5 cm broad, the flowers 
borne 1 mm or less apart, a few scattered, 3-9 


mm apart at the base. Flowers bisexual with the 
style borne beneath the anthers; sepals 5, free to 
the base, linear-lanceolate, 4-5.5 mm long, uneven 
in length, hirsute; corolla yellow, later turning black, 
tubular with 5 spreading lobes, 8 mm long, the 
lobes ovate and acuminate at the apex, ca. 2 mm 
long, the tube 4-5 mm long, hirsute outside; sta- 
mens 5, inserted just beneath the mouth of the 


corolla tube, the anthers nearly sessile, lanceoloid, 
ca. 1 mm long; ovary ovoid, ca. 0.6 mm tall, the 
style ca. 5 mm long, the stigma conical with a 
thickened annular ring at the base. Fruits unknown. 


Tournefortia spicata stands apart from the oth- 
er species of sect. Tournefortia, and its relation- 
ships are unclear. It is unusual in its soft-stemmed, 
short growth habit, dense inflorescence, and ve- 
lutinous to villose peduncle. The flowers are dis- 
tinctive in having linear-lanceolate sepals 4-5.5 
mm long and a corolla tube 8 mm long, It is known 
only from the area in which the type was collected, 
and fruits remain unknown. 


Additional specimens examined. COLOMBIA. NARIÑO: 
Mpio. Tumaco, La Guayacana, 20-100 m, Soejarto 958 
(US). 


I thank the Department of Botany at the Smith- 
sonian Institution, particularly Joan W. Nowicke, 


for providing E. P. Killip's unpublished manuscript. 


LITERATURE CITED 


CANDOLLE, A. P. DE. 1845. Prodromous Systematis Na- 
turalis Regni Vegetabilis Volume 9. 
KiLLip, E. P. The Andean species of Tournefortia. Smith- 
sonian Institution Archives (unpublished). 
Mac pa e as 1960. Boraginaceae. /n: Flora of Peru. 
. Nat. Hist., Bot. Ser. 13(5)2: 539-609. 
ji LER, Ls " 1988. A revised treatment of Boragina- 
for Panama. Ann. Missouri Bot. Gard. 75: 456- 
NOWICKE, J. W. 1974. Three new species of Tourne- 
idis ius ce from the Andes and comments 
RUP of E. P. Killip. Bull. Torrey Bot. 
Club 101: "229-2 


—James S. Miller, Missouri Botanical Garden, 
P.O. Box 299, St. Louis, Missouri 63166, U.S.A. 


A NEW SPECIES OF 
URGINEA (LILIACEAE) 
IN NIGERIA 


The genus Urginea Stein. is represented i in | West 


Kunth, U. ensifolia (Thonn.) Hepper, U. 
pauciflora Bak., and the variable U. altissima (L.) 
Bak., which was recently discovered to resolve into 
a complex of three species: U. altissima (L.) Bak., 
sensu stricto, U. gigantea (Jacq.) Oyewole, and U. 
viridula Bak. (Oyewole, 1975). Samples of wild 
populations continue to reveal the inadequacy of 
the existing taxonomy of the genus. For instance, 
complexes such as in U. indica (Oyewole, 1987) 
or entirely new variants are being discovered. 
Hence, a revision of the genus is due. 

The present report describes a recently discov- 
ered wild population of a new species within the 
main campus site of the University of Ilorin, Kwara 
State, in west central Nigeria. 


EcoLoGY 


The flora of a rock outcrop located about 1.5 
km to the east of the academic area of the main 
campus site of the University of Ilorin, about 20 
km outside the city of Ilorin, was first sampled in 
May 1982. The site of the rock was to be part of 
a proposed botanical garden. 

About 100 tiny bulbs of an evidently new rep- 
resentative of Urginea were collected between May 
1982 and February 1983, from the shallow soils 
of the outcrop. The soil is brown and lateritic, with 
occasional dark humus in depressions. The vege- 
tation is low grassland, with few small trees of 
Entada africana, Erythrina pulcherrima, and 
other legumes interspersed with stands of Butyro- 
spermum paradoxum and Lophira lanceolata, 
and low stands of Cochlospermum sp. Herbaceous 
species of Commelinaceae and Asteraceae are com- 
mon among low grasses. Seasonal marshes and 
ponds support the growth of various Cyperaceae, 
Anthericum nigericum, and many low, herbaceous 
Acanthaceae and Scrophulariaceae. 

ulbs of U. nana occur in an unshaded area 
with low grass and forb cover. Other bulbous geo- 
phytes occurring in the same habitat and inter- 
spersed with U. nana are Dipcadi tacazzeanum 


(Hochst. ex A. Chev.) Bak. (D. filamentosa Mor- 
ton) and Pancratium hirtum A. Chev 


TAXONOMY 


Urginea nana ~ sp. nov. TYPE: Nigeria. 
Ilorin: near the academic area, University of 
Ilorin, Soof2111 dope IUH; isotype, 

HI). Figures 1- 


Bulbus ovoideo-globosus, circa 3 cm altus, 4 cm dia 
collis bulborum 0.5-4 cm, tunicae albae, tunicis eei 
rioribus maroribus; folia tenuia, Bliformia Fendi aut 
lineari-lanceolata, erecta, 8-20 cm longa, 1-3 mm lata; 
pedunculus erectus, 1.5-4 cm longus; racemus com- 
pressus umbellatus, floribus 1-6, pedicellis crassis sed non 
rigidis, postremo in fructu reflexis; tepala subpunice- 
subflavida, aut subviridula flavida, 6-10 mm longa, 1-2 
mm lata; maginibus flavescentibus, antherae 1-2 mm 
longae; capsula globosa vel ovoidea fulva, apice mucro 

8-12 mm longa, nas ne sr iuis fulvis, 
lo ongis, usque 4 


compresso e Puit foliis fliformibus a aut lineari- 
lanceolatis differ 


Bulb small, ovoid-globose, up to 3 cm high and 
4 cm diam., usually with bulb neck, 0.5-4 cm 
long and consisting of rolled bases of old leaves; 
scales white, outermost tunics membranous; leaves 
thin, filiform and spreading, or linear-lanceolate 
and erect to acutely arching, 8-20 cm long, 1-3 
mm wide, not borne with the flowers; peduncle 
stifly erect, 1.5-4 cm high, naked; inflorescence 
compressed, 1—6-flowered, all arising from about 
the same level terminally; pedicel thick but not 
stiff, reflexed during fruit development; tepal 6, 
pinkish and tinged yellow at the margin, or greenish 
yellow with hyaline margin, in 2 whorls of 3 each, 
alternating and free to the base, 6-10 mm long, 
mm broad, outer segments reflexed at anthe- 
sis; flament hyaline, anthers bicelled, dorsally med- 
ifixed and versatile, up to 2 mm long, completely 
fertile; ovary pyramidal, 3-4 mm high at anthesis; 
style short and stiffly erect, about as long as the 
ovary. Fruit a 3-chambered capsule, brownish at 
maturity, ovoid-spherical, with mucronate apex, 
only slightly trigonal from the exterior, 8-12 m 


ANN. Missouni Bor. GARD. 76: 623-625. 1989. 


624 


Annals of the 
Missouri Botanical Garden 


d e 


REl. Growth habits of U. nana. —a, b, c. Fruit- 
ing plants of Morphs A, B, and C, respectively. —d. Ma- 
ture flower before anthesis. — e. Dehisced fruit. — f. Seed. 


long as well as in circum.; seeds dark brown to 
black, 7-8 mm long, 4-6 mm across (including 
the membranous wing), uniseriate in each loculus. 


Paratypes. NIGERIA. ILORIN: about 1.5 km to 
the northeast of the main buildings of the Univer- 
sity of Ilorin, 00/2121, 500/2229, 500/2230, 
500/2231 (IUH, FHI). 


These bulbs resemble U. pauciflora in being 
few-flowered, and the depauperate morphs of U. 


indica in having linear-lanceolate, stiffly erect leaves 
and few-flowered inflorescences. They differ, how- 
ever, in the form and width of the vegetative leaf, 
and in the size of the scape and the positioning of 
the flowers on the inflorescence. Examination of 
the herbarium specimens at the Herbarium of the 
Forest Research Institute of Nigeria (FHI), Ibadan, 
and at the University of Ibadan Herbarium (UIH) 
revealed that such specimens have not been col- 
lected previously in West Tropical Africa. Earlier 
examination of materials at Kew also showed that 
specimens of this taxon have not been recorded. 


Certain variations were observed in the floral 
and vegetative morphology, on the basis of which 
the population was segregated into three morphs 
designated A, B, and C. This informal categori- 
zation was necessary for the purposes of karyotypic 
examination and eventual cytogenetic studies. 


Morph A. Bulbs ovoid-spherical, 3-4 cm 


o” 


$ 


> ^ 


FIGURE 2. Vegetative plants of U. nana. —a, b, c 
from Morph: A, B, and C, respectively. 


diam., 2-3 cm high, usually with relatively short 
bulb neck. Leaves thin, narrow, coiled (filiform), 
10-15 cm long, 1-2 mm broad, dark green, cres- 
cent-shaped in transection. Peduncle up to 2.5 cm 
high outside of the bulb neck, thick and stiffly erect, 
green, bearing 2-3 flowers terminally. Tepals 8- 
10 mm long, 2-3 mm broad, yellowish green with 
hyaline margin. 

rph B. Bulbs ovoid-spherical, 2-3 cm high 
and diam., with a conspicuous bulb neck as high 
as 4.5 cm. Leaves thin, narrow, coiled, light green, 
crescent-shaped in transection, 8-12 cm long, about 
l mm broad. Peduncle stiffly erect, 3-4 cm high 
outside of the bulb neck, pinkish to purple, usually 
bearing 3-6 flowers terminally. Tepals pinkish tint- 
ed green on the margin, 7-8 mm long and ca. 2 
mm broad. 

Morph C. Bulbs small, oval-globose, 0.75- 
1.5 cm diam., 1.5-2.5 cm high, with relatively 
short bulb neck. Leaves light green, acutely erect, 
linear-lanceolate with open transcection and shortly 
apiculate apex, 15-20 cm long, 2-3 mm broad. 
Peduncle thin, erect but not firm, up to 2.5 cm 
high outside of the bulb neck, bearing one terminal 
flower usually. Tepal greenish with hyaline margin, 
6-7 mm long, about 1 mm broa 

The floral dimensions of the three morphs over- 
lap so that they are not easily distinguished, but 
morph C differs distinctly in vegetative morphol- 
ogy. Plants of U. nana must have escaped rec- 
ognition for so long because of their habit of growth: 
the fruiting shoot is hardly seen above the heap of 
burnt litter; the inflorescence begins to emerge 
from the soil less than 36 hours after the first 
burning. The flower and fruit colors merge with 
that of the dark brown soil surface once the charred 


Volume 76, Number 2 
1989 


Notes 625 


FiGURE 3. 


remains are blown away by the wind, and the 
vegetative leaves pass easily for the leaves of Bul- 
bostylis fimbristyloides and related sedges. The 
leaves are not evident when the plant flowers but 
rather are produced when the surrounding vege- 
tation is luxuriant. 

Following is a taxonomic key for separating the 
three morphs and their close relatives within the 
genus: 
la. Pedicel 3 cm long or longer; inflorescence var- 

ious in height and number of flowers, but a lax 
raceme; capsule globose U. indica 


lb. Pedicel shorter La 2 cm lon 
2a. Scape 6-10 cm tall; a — a ra- 
ceme of 6-10 flowers; fleshy tunics pur- 
plish violet J. pauciflora 
2b. Scape less than 5 cm tall; inflorescence o 
1-6 flowers borne terminally; fleshy tunics 
white a U. nana, morphs A, B & C 
3a. Leaves pia flat in transection, 
cm long or dat inflorescence sin- 
gle-flowered, terminal, or with two 
flowers, one of which is subterminal 
U. nana, morph 
3b. Leaves obtusely arching or spreading, 
coiled, crescent-shaped in transection, 
up to 12 cm wit inflorescence 2-6- 
flowered, umbellat 
4a. e Pink. 3-6 per inflores- 
NCE AN U. nana, morph B 
4b. Peer greenish, 2-3 per inflo- 
rescence s. U. nana, morph A 
DISCUSSION 


rginea is probably of recent origin in West 
Tropical Africa (Oyewole, 1975), arising probably 
like its very close relative Albuca L. from southern- 
southeastern Africa and migrating across central 
Africa into the west (Oyewole, 1971). It is therefore 


Fruiting plants of U. nana. —a, b, c from Morphs A, B, and C, respectively. 


conceivable that, faced with myriads of ecological 
niches in this region, the number of representative 
species and infraspecific taxa is likely to be much 
higher than Hepper (1968) recognized. The distinct 
morphology of U. nana, with its peculiar habitat, 
sets it apart from previously known species of the 
genus. However, the existence of different morphs 
within a seemingly uniform ecological niche sug- 
gests that the population of U. nana may not 
entirely represent one genetic system. It is hoped 
that investigations now in progress on aspects of 
the biosystematics of this species will resolve this 
problem before long. 

The discovery of U. nana in Nigeria underscores 
the fact that the latest treatment of Liliaceae in 
West Tropical Africa (Hepper, 1968) is due for 
revision. It also indicates that Urginea may be 
represented in this subregion by many more taxa 
than are now known (Oyewole, 1975). 


LITERATURE CITED 


"us F. N. 1968. des P UM African mono- 
otyledons. Kew Bull. 1-38. 

OYEWOLE, S. de studies in the 
genu Albuca Lin. with special reference to those 
ee occurring in Nigeria. Ph.D. Thesis. Univer- 

sity of Ibadan, Nigeria. 

axonomic treatment of the Urginea 

altissima (L.) B comple A 2 West Africa. Bol. 

Soc. Brot. 49(2e un 163- 

l Anco PR in the g 
fri 


nus 
ca. IV. Population. dif- 


Pe a Kunth, Ann. Missouri Bot. Gard. 74: 


—S. O. Oyewole, Department of Biological Sci- 
ences, University of Ilorin, Nigeria. 


| 
4 


c— 


Julian Alfred Steyermark 


sue of the Annals of the Missouri Botanical Garden is dedicated with respect and admiration to Julian 
Alfred Bud who began and ended his astoundingly productive life in St. Louis, Missouri. The following 
set of recollections s will help paint a Vien of one of the world's great botanists and plant explorers. A memorial 
service for Dr. Steyermark was held at the Missouri Botanical Garden, where he passed the final years of his 
career. The first « contribution, by Dr. Pe er H. Raven, Director of the Missouri Botanical Garden, is an overview 
of Dr. Steyermark’s life an eas and was presented at the memorial service. Several remembrances 
generously contributed by Dr. Steyermark’s friends and colleagues follow. 


The frontispiece: Julian Steyermark, 1987, Warren County, Missouri. 


JULIAN ALFRED STEYERMARK 


Born, 27 January 1909, St. Louis, MO. 

Graduated from Soldan High School, St. Louis, MO, 1925. 
Married Cora Shoop, 1 September 1937, no children. 

B.A., Washington University, St. Louis, MO, 1929. 

M.S., Washington University, St. Louis, MO, 1930. 

M.A., Harvard University, Cambridge, MA, 1931. 

Ph.D., Moe ame Henry Shaw School of Botany, 
St. Louis, MO, 

Research Assistant to Robert E. Woodson, Jr., 

Missouri Botanical Garden, 1934-1935. 

Biology Instructor, U opui City Senior High School, 
University City, MO, 1935-1937. 

Taxonomist-Ecologist, U.S. Forest Service, summers 1936-1937. 
Assistant Curator, Field Museum of Natural History, 
Chicago, IL, 1937-1947. 

Associate Curator, Field Museum of Natural History, 
Chicago, IL, 1947-1950. 

Curator, Field Museum of Natural History, 

Chicago, IL, 1950-1958 

Professor of Botany, Southern Illinois University, 
Carbondale, IL, April-June 1958. 

Professor of Botany, University of Missouri, Columbia, MO, 
June-August 1958. 

Botanist, Instituto Botánico, Ministerio de Agricultura y Cría, 
Caracas, Venezuela, 1959-1984. 


Curator, Instituto Botánico, Caracas, Venezuela, 1975-1981. 
Curator, Missouri Botanical Garden, St. Louis, MO, 1984-1988. 
Died, 15 October 1988, St. Louis, MO. 


OD" 


132,223 is the last number in Julian Steyermark's collecting book. 
Julian Steyermark described more than 1,000 new species 
of plants during his career. 


New Books 
from the Missouri Botanical Garden 


Thesaurus Dracularum, 1 & 2 
Carlyle A. Luer with Rodrigo Escobar R. German translations by Fritz Hamer. Illustrations by Stig Dalstróm. 


Thesaurus Dracularum is a monograph of the genus Dracula. Each of the 80-odd known species is illustrated 
by a watercolor painting accompanied by descriptions, discussions, and distributions in both English and 
German, with distribution maps and black and white line drawings. In elephant-size format. Standing orders 
available. The first two of six fascicles. $40.00 each. 


The Moraeas of Southern Africa 
Peter Goldblatt 


Morea is a genus of 119 currently recognized species. Th pht hose found south of the Zambezi 
and Cunene Rivers, some 103 species. In addition to the DAR il in a rigorous taxonomic 
monograph, conservation, cultivation, taxonomic history, and floral and pollination biology are treated. ye 
of the taxa treated are illustrated with full-size, full-color illustrations by Fay Anderson. xi + 224 pp., 64 
color plates. National Botanic Diras Kirstenbosch, and the Missouri Botanical Garden. ISBN 0- 6550-09974- 
7. 1986. $35.00. 


. Modern Systematic Studies in African Botany 
- Peter Goldblatt & P. P. Lowry, II, Editors. 


This monograph contains abat 60 papers dealing with systematics, ethnobotany, and conservation, presen nted 
at the Eleventh Plenary Meeting of the Association for the Taxonomic Study of the Flora of Tropical Africa 
.. held at the Missouri Botanical Garden, St. Louis, on 19-14 June 1985. 110 PP- 1988. nia 


To place an order, use this form or a photocopy of it. US; shipments: add $1 .50 for one book and $.50 | 
for each additional book; non-U.S. shipments: check or money order in U.S. funds, payable to U.S. bank, 
add $2.50 for one book, and $.50 for each additional book. Orders should be prepaid; a $1.00 fee will b. 
added to orders requiring invoices. No shipments are made until LÁ is received. Mail form with you —— 
Check or money order, payable to Missouri Botanical Deia to: | à bz 
"dete Department Eleven — 
Missouri Botanical Garden 
P.O. Box 299 
‘Bt. Louis, MO 63166-0299, 1 US. Es 


j xd send the pu circled above to: 


: "Bi Pome, enclosed. 


Ere CI Sed inicios ($1.00 e wie d 
xe HH added to total). 25. ^ 


Pol Cale “Couey D Ln oH NM 


Prices, ARE Secr ToC dies Wirioor Nonce | 


CONTENTS 


New Species and Combinations in Tropical American Polystichum (Dryopterid- 
. aceae) David S. Barrington 
New or Noteworthy Orchids for the Vepssüelán Flora. VII. Additions in Maxillaria from 
the Venezuelan Guayana Germán Carnevali & Ivon Ramirez | 


Contribución al Genero Mimosa (Miinosaaose) Renee H. Fortunato .. 
A Revision of Mesoamerican Psychotria Subgenus Psychotria (Rubiaceae), Part II: Species 

l Clement W. Hamilton 
New Species and Other Taxonomic Matters in the New World es (Melastomat- 
. aceae) omas Morley 


: Study of the Flora of Rucamanque, Cautin Province, Chile Carlos Runipit purger 
Hauenstein, José San Martín & Domingo Contreras . 


AER Novelties i in the Orchid Flora of Southern Venezuela Gustavo A. Romero & Germán 


Carnevali 


; Novelties i in the EFR from. Vehituelan Guayana - EN van der Waf- VP ME 
‘Taxonomic Studies i in the Micónieae (Melastomataceae). HI. Cladistic Analysis of Axillary. 


- flowered Taxa — ‘alter S. Judd 
^ Survey of Reproduce. Biology in Nene Melastomataceae End 
- Memecylaceae — - Susanne - Renner .. 


E Nectaries in ‘Mehstoniatacess and Their ‘Systematic and Evolutionary FN 
5 


Implications. Bruce A. Stein & Hiroshi Tobe 


B A ciendo Revision of Fuchsia Section Quelusia E -Paat E. Berry. em = 
in Fuchsia Sections es and Kiershlegería (Onag: na 


| * Observations | on : Polyploidy 
: raceae) - poe Hoshino & Paul E Berry. 


NOTES - 


: : Notes on the Cordia panamensis e Gs [o and « a New Si 
SEPA bin : : 


365 


374 
381 


386 


430 


444 


Annals 
of the 

Missouri - 
Dotanical 


vos cos 


olume 76 


Volume 76, Number 3 
Fall 1989 


Annals of the 
Missouri Botanical Garden 


The Annals, published quarterly, contains papers, primarily in systematic botany, con: 
tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside. the 
Garden will also be accepted. Authors should write the Editor for information concerning 
arrangements for publishing in the ANNALS. Instructions to Authors are printed in the back 


of the last issue of each volume. 


Editorial Committee 


George K. Rogers 
Editor, Missouri Botanical Garden 


Amy Scheuler 
Editorial Assistant, 
Missouri Botanical Garden 


- Glenda Nau 


Magdalen Lampe 
. Publications Staff 


Henk van der Werff 


Marshall R. Crosby 


Missouri Botanical Garden 


Gerrit Davidse 
Missouri Botanical Garden 


John D. Dwyer 
Missouri Botanical Garden & 
Saint Louis University 


Peter Goldblatt 


Missouri Botanical Garden 


- Dale E. hiena 


Missouri Botanical Garden 


Missouri Botanical Garden 


For subscription information contact Department i 
- Eleven, P.O. Box 299, St. Louis, MO 63166. Sub- 


scription price is $75 per volume U.S., $80 Canada 


and Mexico, $100 all pi countries. + Four i issues 


; per Meg 


B Missouri Botanical Garden 1989 Am : 


- Eleven, P.O. Box 299, St. Louis, MO 63166. 


-The ANNALS OF THE ps DEN GARDEN 


(ISSN 0026-6493) is published quarterly by. the 
Missouri Botanical Garden, 2345 Tower Grove Av 


. enue, St. Louis, uo: RAI o Second class postagt 


] mailing offices 


paid at St. Louis, M 


POSTMASTER: Send address changes to ANNALS o 


THE Missouri BOTANICAL GARDEN, Departmei 


Volume 76 
Number 3 
1989 


Annals 
of the 
Missouri 
Botanical 


Garden 


NA 


People were Julian Steyermark's first love, but 
flowers were his second. He was a passionate man, 
and some of his passions were shaped by early trips 
out of a St. Louis that was crowded and smoky 
into the gentle Missouri countryside, where he and 
his friends first sought unusual birds. Later he 
reveled in the autumnal drifts of asters, in enjoying 
the purple-brown trilliums and delicate orchids of 
early spring, and in collecting samples from the 
legions of sedges that he could see only dimly 
through the torrential cascades of sweat that poured 
from his brows as he crouched over gravelly riv- 
erbars during the torrid Missouri summers. Even 
the long-exserted anthers of the ribworts that ap- 
peared in early summer must have fascinated this 
unusual young man, who grew in knowledge about 
the diversity of plants that nurtures all who take 
the time to stop and appreciate it. 

Everywhere he went Julian was recognized for 
his achievements: the governments of Guatemala, 
Ecuador, and Venezuela; his alma maters at Wash- 
ington University and here at the Garden; the 
Missouri Department of Conservation; and the Sier- 
ra Club all soon came to know him as an inspiring 
and unique individual, and gave him abundantly 
of their prizes and awards. Everyone knows that 
Julian collected more plant specimens than anyone 
else has ever done, an incredible 132,000 speci- 
mens, and that he was a prolific writer and her- 
barium worker. In fact, if we hadn't known him 
personally, we might think —reading about his ac- 
complishments—that he was some sort of drudge, 
dreadful, demanding, and uninteresting. Nothing 
could have been farther from the truth, however; 
those of us who had the privilege of enjoying his 
company, whether in the field, listening to his fa- 
vorite music, or simply exchanging views on some 
current topic, know far better. 

Julian certainly did enjoy hard work, and ap- 


preciated all of those wonderful manifestations of 
recognition that were accorded him for his accom- 
plishments—to be included in the Guinness Book 
of World Records as the world's champion plant 
collector was for him an absolute delight. In his 
life, however, he was sustained to a far greater 
degree by the love of his many friends. Boyhood 
friends who ranged through the Missouri woods 
together searching for rare warblers retained the 
joys of their mutual admiration into the years of 
maturity. Those who had the privilege of climbing 
the steep, rocky slopes and breaking through the 
interlocking branches of the forests of Venezuela 
with him never forgot the experience, and ex- 
pressed their deep concern during his final illness. 
Once you knew Julian, you simply did not forget 
im. 

Returning here to his boyhood home at an age 
when most have slipped into the relaxation and 
hazy days of retirement and gentle decline, Julian 
undertook the preparation of an account of the 
plants of what is perhaps the least-known part of 
the world botanically, the interior of Venezuela, 
where the tepuis and gorges that he loved so deeply 
make up the spectacular fabric of the fabled Lost 
World. As he did everywhere he lived and worked, 
Julian renewed sturdy old friendships, and forged 
new ones, a process that continued all his life. Bruce 
and Luther, Gerrit and Jeany, Tamra and Bruno, 
and so many others—you each gave him much, 
especially during his final days of great physical 
discomfort, but you got far more than you gave. 
Julian's open, happy smile; his sweet, joyous, and 
dedicated personality; his attentiveness to the task 
at hand; and his unremitting, dignified friendship 
will form memories that will be with each of us 
until our own final days. 

For someone who spent his childhood amongst 
the beauties of bellflowers, gerardias, and desmo- 


ANN. Missouni Bor. GARD. 76: 627-651. 1989. 


628 


Annals of the 
Missouri Botanical Garden 


diums, gumweeds might have seemed a strange 
choice as the subject of a doctoral dissertation. 
These sunflower relatives exude a white, sticky sap 
that fills their unopened flower heads before they 
expand to reveal the golden-yellow inflorescence 
hidden within each. Gumweeds are relatively un- 
common in Missouri, but range widely over the 
American West and Mexico. Julian's graduate ad- 
visor, Jesse Moore Greenman, who trained an en- 
tire generation of American taxonomists here in 
St. Louis, was determined to complete the great 
task of John Torrey, George Engelmann, and Asa 
Gray —to make known the variety and complexity 
of the plants of our continent, and he assigned 
doctoral problems accordingly. For this purpose, 
)r. Greenman enlisted Julian, who was afire with 
his love of plants, into his army; and the young 
man set out along the dirt tracks and across the 
fields of the West, publishing his work ultimately 
in the Garden’s Annals. 

the 28-year-old Julian Steyermark, just 
married, there followed 21 years in Chicago, where 
he ultimately became Curator of Botany at the 
Field Museum. That institution, and his contact 
with the famous botanist Paul Standley, provided 
his introduction to the plants of Guatemala and 
brought home to him just how little we knew about 
the plants of the American tropics. Missouri's flow- 
ers remained his favorites, however, and he cul- 
tivated them lovingly, with his wife Cora, in their 
own wildflower garden near Chicago, a sanctuary 
now maintained by The Nature Conservancy. For 
30 years Cora and Julian crisscrossed Missouri 
searching for plants, driving thousands of miles 
back and forth from their home, camping in riv- 
erbottoms, by dusty roadsides, or in forests, so that 
they would come to know more fully the detailed 
distributions and characteristics of the plants o 
this state. Their cultivation of many of these plants 
in their home garden was not merely a scholarly 
exercise; it was also an act of love, a celebration 
of their natural beauty and their individual char- 


acter, as anyone who has turned the pages of 


Julian’s exemplary 1963 Flora of Missouri will 

now. What memories those explorations provided 
Julian and Cora, and what times they shared to- 
gether! To know about some of them, read Cora's 


Behind the Scenes, a testimony to the beauties of 


our state and the joy that is in store for those who 
take the time to appreciate, understand, and enjoy 
them. Cora wrote: 


The morning was cold and clear. Pine trees and 
shrubs, gray and bent with ice and snow, lent variety 
to the icicled oaks. At the top of the hill we came 
upon the sunrise. The whole view for miles across 


the hills was a pinos blaze of crystalline purity. 

Against the sun, the tops of trees turned to polished 

gold, dadag i to silvered lower limbs rising above a 
b 


soft setting of snow-covered shrubs. 
Julian stopped the car. "I'm stuck with a view," És 
he m "Im afraid Pll break En if I move.’ 


t whole view looks as if it might break if you 
breathed on it. This is something you see once in a 
lifetime. I'm glad we stalled last night," I said and 
really meant it. 


From Chicago, Cora and Julian moved on to 
Caracas, where they traded goldseal, gentians, and 
Chimantaea, Sterig- 
The exchange 


puccoon for Brocchinia, 
mapetalum, and Heliamphora. 
proved a rewarding one, and Julian went on to add 
more to our knowledge of the plants of Venezuela 
than anyone had before. A quarter of a century 
was enough—filled with friends from Venezuela, 
new friends from the States, wonderful new plants, 
many of them extravagantly beautiful, and hundreds 
named in Julian’s honor, hundreds of them. After 
Cora’s death, Julian came back here to his home- 
town and began to write in earnest about the in- 
credible plants of the Lost World. His last work 
will be completed here, and will stand as a final 
testament to his industry and to his scholarship. 
As for any human being, however, what will prove 
to be even more important to all of us will be our 
memories, for Julian Steyermark was an inspira- 
tional person, one who projected his own values 
without trumpeting them, and thereby enriched all 
our lives, as he did the literature of botany. 
In concluding her book, Cora wrote: 


t was a long exploration = more than 30 years 
from which we have related merely a few experiences 
representing each “os of he la season. So 

we see —a book is not only written and rewritten, it 
is lived through a life of sweat, privations, and joys 
behind the scenes, before the final word is ever put 
on paper 
at was her affectionate term for Julian 
in us book] and I, hereby will the state to the next 
botar 


Is that not a reflection worthy of us here today? 
We all toil in a common field, inspired by early 
memories, contributing to knowledge, doing what 
we can for one another and regretting what we 
cannot do. In the plants that support and nurture 
us, enrich our souls with their beauty, and inspire 
our actions, we have found a common bond. Here 
today we have found another: the remarkable and 
full life of a truly luminous human being, our friend 
Julian Steyermark, whose fellowship we enjoyed so 
deeply, and whose life we will remember with grat- 
i eter H. Raven, Director, Missouri Bo- 
tanical Garden; from Julian Steyermark's me- 
morial service. 


Volume 76, Number 3 
1989 


Steyermark Recollections 629 


During the 1940s in my beginnings as a botanist, 
I started to understand the importance of Dr. Jul 
Steyermark's works on the taxonomy of the Vane 
ezuelan flora. The National Herbarium already pos- 
sessed numerous collections made by the remark- 
able scientist. By the 1950s, our friendship had 
grown stronger and lasted for more than 30 years. 
His long and exhaustive research of the Venezuelan 
geography created a major collection that has con- 
tributed in a decisive way to enhance Venezuela's 
floristic inventory. 

His humility and generosity were the most out- 
standing features of his personality. A few years 
ago, before departing for Venezuela's Guayana, | 
asked Julian to inform me on the area I was going 
to visit, a place he knew and had explored. Among 
the helpful and valuable facts he supplied was the 
name of a man who knew the region well and could 
act as a “baqueano” [guide]. Arriving in the area, 
I found the person Julian had mentioned. I told 
him my mission in the area and that Dr. Steyermark 
had recommended him to me. To my suprise he 
“Mr. Steyermark? I have never heard 
> However, I insisted and slowly re- 


answered, 
such a name.’ 
peated, “Are you sure you don’t know Dr. Julian 
Steyermark?" His facial expression suddenly 
changed and he replied smiling, you mean 
Julian, of course I know him and I am ready to 
help you." 

This story helps to illustrate Julian's sense of 
humility, generosity, and harmony. His legacy as 
a scientist goes hand in hand with the personal 
attributes that made him a great human being and 
a beloved friend to those who knew him.— Leandro 
Aristeguieta, Universidad Central de Venezuela, 
Jardín Botánico de Caracas. 


When I arrived at the Instituto Botánico in 
Caracas in the summer of 1975 looking for a 
project to help me learn about tropical plants, I 
was brought into the office of one very busy, grum 
Dr. Julian Steyermark. Faced with a botanical neo- 
phyte, he suggested that I begin collecting plants 
from the hills surrounding the Instituto—a patc 
of secondary, deciduous forest in the heart of the 
city. Years later this project led to a paper we 
published together on the vegetation of the decid- 
uous forests of Caracas, but only after many jour- 
neys in each other's company to the few wooded 
areas near the city not yet overtaken by apartment 
buildings or “ranchos.” Following up our finding 
of five species of the coca genus, Erythroxylum, 
growing side by side in the hills above the Botanical 
Garden, a student of mine at Simón Bolivar Uni- 
versity in Caracas completed a wonderful thesis on 


FIGURE 1. 


in southern Ecuador 


Fuchsia steyermarkii, from cloud forest 


the comparative reproductive biology of the group. 
Another result of our study was the recognition of 
two species new to science, Eugenia mcvaughii 
and Erythroxylum undulatum, both based on plants 
found less than 100 meters from the front door 
of the Instituto. It was the shortest trip I know of 
for specimens to be placed in the type collection 
of the National Herbarium. 

Now years later I sit in Julian's former office— 
sometimes in his chair, picking up where he had 
left his unfinished project, the Flora of the Ven- 
ezuelan Guayana, more "iren known as 
the “Flora of the Lost World.” Julian was a *'ba- 
queano” (guide, in sas to me, for he opened 
up many paths that I was to follow. 

My botanical tribute to Julian was the naming 
Just after his death, I 
was to return to the original site where the plant 
was found, in lush cloud forest on the steep slopes 
south of the Nudo de Sabanilla, 


of Fuchsia steyermarkii. 


in southernmost 


630 


Annals of the 
Missouri Botanical Garden 


Ecuador. | got there in a rented taxi, a few hours 
by car from the city of Loja; he got there by foot 
and mule, on a fortnight trek through endless rains, 
in October of 1943. His Fuchsia is a most unusual 
species, with needlelike leaves, and if it survives 
in cultivation it will be highly esteemed. 

Many simple folk all over Venezuela knew and 
remembered Julian. I met some of them, usually 
in remote places, like “Horacio,” a Makiritare 
Indian who spoke no Spanish but had climbed many 
a tree for him; and José Mendoza, an Andean 
homesteader from San Vicente de la Revancha, 
whose senile father had guided Julian years ago 
far into the páramos towering over the valley. Both 
reacted similarly when I mentioned Julian's name: 
a big, amused grin accompanied by signs of respect 
for a man so dedicated to his botanical mission. — 
Paul E. Berry, Missouri Botanical Garden. 

I had been at the New York Botanical Garden 
as a doctoral student for all of three weeks in 1980 
when I first met Julian Steyermark. He was visiting 
the New York Botanical Garden to consult our 
Guayana Highland collections and to collaborate 
with Bassett Maguire on several taxonomic treat- 
ments. Of course, I already knew of him by rep- 
utation. | was a novice to tropical botany, so in an 
attempt to say something intelligent to him I dug 
back into my M.S. thesis on /soetes and located 
the citation of Steyermark 85179, which I had 
identified as /. melanopoda Gay & Dur. | reported 
this to Julian, and to my amazement he immediately 
told me where in Missouri the specimen had come 
from and the month during which he collected it 
23 years previously! This was not to be the last 
time over the following eight years years that Ju- 
lian's prodigious memory would impress me. 

| also became aware of Julian's capacity for 
work as I observed him during his subsequent per- 
iodic visits to New York. Occasionally, he would 
share my office because of our mutual interests in 
the Guayana Highland flora. He stayed in the Gar- 
den's Snuff Mill apartment during his visits. By 
7:30 A.M. he would be at the herbarium. With a 
single-mindedness he worked on specimens until 
about 6:00 P.M. with only a half-hour break at the 
lunch table. Julian was not one for idle chat, and 
he clearly did not like to be disturbed while working. 
At six, and also with a single-mindedness, he would 
round up any remaining staff and students and go 
have a huge dinner, followed always by a stop at 
the local ice cream establishment. Occasionally, he 
would take in a movie, but more often than not he 
would head back to the herbarium for several more 
hours of work before retiring for the night. 


And thus, day after day, he would make his way 
through huge piles of specimens, writing descrip- 
tions of new taxa on the spot in his nearly illegible 
scrawl. Julian confessed that he “over-described”” 
occasionally, and that perhaps he split excessively. 
lowever, he justified it by saying that he viewed 
his role as one of describing biotic diversity as he 
saw it. He would leave it to following generations 
of botanists to determine just how “good” his de- 
cisions were. How history will judge his taxonomic 
decisions does remain to be seen, but there can be 
no doubt that Julian had an excellent "eye" for 
I also know that I trust his 
specimen annotations more than those by just about 


seeing differences. 


anybody else. During the first half of this century, 
when a researcher needed quick, reliable deter- 
minations for neotropical specimens, the solution 
was "send them to Standley." Julian was heir to 
this role. His magnum opus, Flora of the Vene- 
zuelan Guayana, will aptly attest to his unparal- 
leled knowledge of the neotropical flora. In an age 
of few heroes, Julian was one for many of us.— 
Brian M. Boom, The New York Botanical Gar- 
Len. 

Hace cuatro anos, el 7 de junio de 1984, tuve 
el honor de darle con alegria y en nombre de los 
botánicos de Venezuela, un adiós, un hasta luego, 
a Julian Steyermark. Esta despedida era transito- 
ria, ya que por su propia decisión, volvia a su tierra 
natal, Saint Louis, Missouri. A pesar de su ida, 
Julian no podia resistir la tentación de permanecer 
mucho tiempo allà, solo en el norte, razón por la 
cual, con frecuencia hasta principios de este ano, 
regresaba a Venezuela, pues como el decia: amor 
a primera vista," fue su reacción al ver por vez 
primera a este pais en 1943. Y, cual enamorado, 
exclamó: “Mi amor a Venezuela, se debe no solo 
a sus flores, a su fauna, a sus paisajes, sino tambien 
a mis amigos quienes poseen una alta sensibilidad, 
capacidad, y deseo de trabajar en la botánica y en 
la conservación de sus recursos naturales. Por eso 
mi amor por Venezuela fue instantáneo.” 

Julian Steyermark, nació en Saint Louis, Mis- 
souri, el 27 de enero de 1909. 
prana edad sintió el reto de conocer la naturaleza 
que lo rodeaba. Allá por el año de 1925, cuando 
era Boy Scout en su ciudad natal, se interesó en 
la zoologia, fijando primeramente su atención en 


esde muy tem- 


las aves. En ese mismo ano ingresa al club de 
Biologia de su colegio y su espiritu naturalista se 
concentró en el fascinante mundo de las plantas. 
Desde entonces, sólo vivió en función de ellas. 
Combinando sus estudios con sus labores de Guia 
Naturalista en los parques de Saint Louis y sus 


Volume 76, Number 3 
1989 


Steyermark Recollections 


FIGURE 2. 


alrededores, fue formando una buena colección de 
plantas la cual fue adquirida por el Herbario de 
Missouri en 1934 por 84 dólares. Este dinero fue 
la base para comprar su primer vehiculo, para 
poder continuar con mayor facilidad sus trabajos 
botánicos. 

Julian llegó a Venezuela en diciembre de 1943, 
cuando en misión del gobierno de su patria de 
vino a buscar la corteza del árbol de la 
Mundial para combatir el paludismo. Sus primeras 
incursiones botáncas en el pais las efectuó en lo 
que es hoy el Parque Nacional El Avila, de alli 
pasó al Oriente de nuestro territorio, regresando a 
Estados Unidos en 1945 con una colección de 
8,000 muestras botánicas. 

La belleza escénica de los diferentes ambientes 
venezolanos, el contacto con las plantas, coló tan 
hondo en la sencilla humanidad de Julian, que 
sonaba que Venezuela debería ser su pais. Este 
sueño, cual regalo de navidad, se convirtió en 
realidad cuando en diciembre de 1959 recibió la 
invitación de Tobias Lasser, por recomendación de 
William H. Phelps, para que formara parte del 
personal del Herbario Nacional de Venezuela, donde 
permaneció activamente como estudioso de la flora 
Venezolana. Por su larga trayectoria y aportes al 
conocimiento floristico del pais recibió entre otras 
las siguientes distinciones: Orden Andrés Bello 
(1974); Orden al Mérito en el Trabajo, Primera 
Clase (1974); Orden Henri Pittier (1979); Premio 
William H. Phelps, otorgado por la Sociedad Ve- 


Julian Steyermark (lower right) at ca. age 20 as counselor at summer camp in Maine. 


nezolana de Ciencias Naturales; Miembro Corre- 
spondiente Nacional de la Sociedad de Ciencias 
Naturales de la Salle; Miembro Titular de la Fun- 
dación Terramar; Investigador Emeritus del Jardín 
Botánico de Caracas, INPARQUES (1986). 

Julian Steyermark, después de sesenta y nueve 
años de búsqueda y trabajo activo en su mayor 
obra botánica, Flora of the Venezuelan Guayana, 
murió en Saint Louis, Missouri U.S.A, el 15 de 
octubre de 1988. Por eso hoy siento la necesidad 
de darle en nombre del personal del Jardín Botánico 
de Caracas un adiós lastimoso, con dolor y nos- 
talgia. 

En este momento recuerdo, que un día en fe- 
brero de 1984 Julian me dijo, sentado en la cima 
del Aratitiyope (Templo Sagrado de Las Guaca- 
mayas), Territorio Federal Amazonas, en un mo- 
mento de descanso y contemplando la majestuosi- 
dad reflexiva del paisaje, “Francisco mi mayor 
deseo es emplear todas mis fuerzas y mi dinero, 
para seguir explorando este maravilloso pais. Tam- 
eni quisiers que al morir, me enterraran en un 

” Julian, si bien es cierto, que tu cuerpo no 
re en la arenisca de los tepuyes, tu imagen 
caracterizada por el panuelo rojo, con el cual cubria 
tu cabeza, quedará guardada en estas mesetas y 
en la memoria de todos los que salimos al campo 
contigo. Tü estarás presente en cada planta del 
tepuy, pues quién de nosotros no se reirá de tus 
ocurrencias cuando nos topemos por ejemplo con 
una Steyerbromelia de Marahuaca; o cuando sa- 
boreemos una base foliar tierna de una Rapateaceae, 


632 


Annals of the 
Missouri Botanical Garden 


FIGUR Co cement from Washington University, St. 
left to de Marion. C Child (Moss), cme y eni Julian Steyermark, 
t E. y Fuller. Photo contributed by Mildred Mathia 


Mildred Mathias, Rober Woodson, Jr., 


Ile! 


bien sea del Roraima, Auyantepuy, Chimanta, ¢ 
Huachamacari.— Francisco Delascio Chitty, Di- 
rector, Jardin Botánico de Caracas, 
Memoria del l. 


Nov. 


Herbario 
Nacional de Venezuela; de la l 
Congreso Venezolano de Botánica, 13-18 
1988 


I first met Julian Steyermark 60 years ago. He 
was painting an orchid in the old Floral Display 
House of the Missouri Botanical Garden during the 
annual orchid show. At that time he was an un- 
dergraduate student at Washington University. 

Later Julian became chairman of the botany 
group of the Webster Groves Nature Study Society. 
He wrote articles for Nature Notes, which is the 
monthly publication of the society. He also spoke 
at the meetings and led botany group field trips to 
local areas. 

Since Julian didn't have the use of an automobile 
while he was a student at Washington University, 
he depended on his friends to take him to areas 
that he couldn't reach by streetcars or trains. He 
and I went on a field trip to northwestern Missouri 
to the loess mounds of Atchison and Holt counties 
during the Labor Day weekend in 1932. This area 


“=== 


Louis, Missouri, June 1929. Graduates in botany, 
Alexander Buchholz, e Allen, Unidentified, 


contains plants that are found only in this part of 
Missouri. We located most of these species, and 
Julian took specimens of them for the Missouri 
Botanical Garden herbarium. 

On several occasions Julian and I took street- 
cars to South Broadway and found many intro- 
duced plants along the railroad tracks near the 
Mississippi River. Some of these plants were very 
seldom found elsewhere in our area. On the way 
home from these streetcar trips, oeil identified 
plants in vacant lots along the city stre 

On one of our field trips to the Oaks Julian 
saw a sedge that he wanted to reach on the other 
side of a stream. We walked through the shoulder- 
deep water to reach the sedge. He climbed the 
steepest slopes and descended into the deepest val- 
leys in his search for plants. Julian never seemed 
to tire, and if he was tired, he continued on anyway. 

ne of Julian's favorite places was Pickle Springs, 
where many rare plants were found. This area is 
now Pickle Springs Natural Area. Another favorite 
Mis- 


rare 


area was near the Meramec River at Pacific, 
This area, 


plants, is now known as Pacific Palisades. 


souri. which also contains many 


Although Julian was so dedicated to collecting 


Volume 76, Number 3 
89 


Steyermark Recollections 633 


and identifying plants, he did take some time off 
to play tennis at Tower Grove Park. Also he at- 
tended some of the Webster Groves Nature Study 
Society's parties. 

On the day that Julian received his Ph.D. degree 
from the Henry Shaw School of Botany of Wash- 
ington University in 1933, I happened to meet 
him with his parents at the Missouri Botanical 
Garden. He couldn't stay away from the garden 
that meant so much to him, even on his graduation 
day.— Arthur Christ, St. Louis, Missouri; from 
Julian Steyermark's memorial service. 


I am very sad with the passing of Julian, a 
taxonomist extraordinaire and a dear personal 
friend. I feel very privileged to have known him 
closely during the last two decades of his life. Julian 
was truly one of the greats in his field and his 
contributions to taxonomy, exploration, collecting, 
and conservation will long be remembered and rec- 
ognized— and use 

Of all the people I have known personally, he, 
perhaps as well as anyone, exemplified a person of 
whom it can be said without any exaggeration that 
his vocation was also his avocation. I will, above 
all else, remember him as a person totally devoted 
to his work and one for whom his work held a 
never-ending fascination. 

I would like to recall several personal experi- 
ences and incidents that illustrate some aspects of 
his personality and career. 

first got to know Julian personally in 1971, 
when as a graduate student | went to Venezuela 
to study and collect plants for my thesis. I was 
already well aware of his many accomplishments 
and his international reputation. As with almost 
any other novice going to a foreign country to 
collect, I was having difficulties in making the nec- 
essary arrangements. With some trepidation, I 
asked for his help, and I was very grateful that he 
took time to arrange contacts that made my work 
much easier. 

I later learned that he guarded his time very 
jealously, surely a necessity for someone who re- 
ceived little secretarial help at the time, who pub- 
lished thousands of pages, and who was unrivaled 
as a plant collector. However, if you were serious 
about plant collecting, he would often try to help. 
He was very well aware that the next 20 or 30 
years might be the last chance for any of us to 
study and collect the undisturbed vegetation in 
many areas of Venezuela that had never been bo- 
tanically surveyed, and he always felt only a small 
portion of the country had been adequately sur- 


veye 


After I joined the Missouri Botanical Garden in 
1972, a number of other staff members, including 
Tom Croat and Al Gentry, collected in Venezuela, 
and we seemed to pass muster as plant collectors 
since Julian proposed in 1975 that the Garden and 
the Instituto Botanico organize a cooperative pro- 
gram of joint expeditions to areas in which the 
vegetation was being severely damage 

fter we secured funds for this work, Julian and 
I made a number of field trips together, and it was 
then that I really got to know him well. 

Julian was a very goal-oriented person who 
doggedly pursued his objectives until they were 
accomplished, whether it concerned writing the 
Flora of Missouri during his free time or whether 
it concerned collecting plants from a tepui never 
before explored. 

en he went to the Field Museum in 1937, 
he had already collected over 25,000 plants. There 
he worked under and was greatly influenced by 
another of the great plant collectors, Paul Standley, 
who was to collect well over 100,000 numbers by 
the end of his own career. Julian early on, as he 
told me early during our association, determined 
in his own mind that he would eventually surpass 
Standley’s numbers, and he 

ven on a daily basis, a had Standley 
before him— would set a goal of a certain number 
of collections to be made that day, and he would 
challenge his companions to do the same. He would 
invariably have a very precise recall of what he 
collected each day, and he loved to challenge you 
to guess the totals gained by each person, spicing 
the proceedings by betting a milkshake on who 
could guess the closest. I soon learned to keep 
closer track of what I was collecting! 

Although Julian was a modest person who lived 
a quiet, simple life, he, like all of us, appreciated 
recognition. After several years of working togeth- 
er, I knew that he was proud of the large number 
of plants he had collected, and I began to think of 
ways in which this collecting record might become 
more widely recognized. 

I finally hit upon the idea of nominating him as 
a champion plant collector to the Guinness Book 
of World Records. Because I was a bit uneasy that 
he might consider this a trivialization of his accom- 
plishments, I checked out the idea with him be- 
forehand, and he approved. Unfortunately, it took 
the editors of Guinness several years to decide 
what plant collecting was all about and whether it 
really had a place in their book. In the meantime, 
Julian would ask me every time I came to Vene- 
zuela what had happened to this proposal, and | 
had to tell him to be patient, that it was still in the 


Annals of the 
Missouri Botanical Garden 


works. The record was finally included in the 1986 
edition, and Julian seemed to enjoy the additional 
attention it brought him from the popular media. 
I earlier mentioned that Pll always remember 
Julian as a person totally devoted to his work. This 
perception comes from watching him put in long 
hours of fiercely concentrated work each day, sev- 
en days per week, and 52 weeks per year, but it 
was also brought home to me in a much more 
amusing way on our first trip together in 1978. 
Our objective on that trip was to collect in a 
small range of mountains known as the Cerros de 
Bachiller in the Coastal Cordillera of northern Ven- 
ezuela, where new agricultural developments seemed 
certain to destroy much of the virgin vegetation 
in the near future. Close to a small mountain stream 
at the foot of the mountains, we had constructed 
a nice base camp, from which we collected every 
day. As was customary, we slept in hammocks 
strung beneath a plastic sheet to keep out the rain. 
Julian would always wake just before dawn, and as 
we were getting ready for the day's collecting, he 
would tell me his dreams of the night before. Every 
morning during the two weeks of that trip, the 
dream he recalled would be enjoyable and botan- 
Their s e "Lost 
World," and he would discover the most ailing 


ical. setting would often be in t 
new plants, which often combined bizarre char- 
acters from several families, so that he would be 
very puzzled and strongly challenged on how to 
interpret these strange plants. After a couple of 
days of this, I began to think to myself, here is an 
even more devoted man than I had realized, not 
only does he devote most of his working hours to 
botany, but even his sleeping hours are devoted to 
it. 

So these are some of the ways Pll always re- 
member Julian, a valued friend, a man of great 
accomplishments, a man truly devoted to the good 
things of this world, and one who leaves an enor- 
mous legacy to us all.— Gerrit Davidse, Missouri 
Botanical Garden; from Julian Steyermark's me- 
morial service. 


Omnia fluerit. Everything flows. Fifty-five years 
ago Julian Steyermark, then 24, and taking his 
Ph.D. here, wrote to me in the spring of 1933— 
I was at UCLA 


of Grindelia collections that I had made, gum- 


at the time—asking for the loan 


weeds of the Compositae (as we called them then). 
On June 21st he wrote again, that the loan was 
being returned and added that he was tending a 
34-acre plot of grindelias, and rushing his getting 
a Ph.D. He wrote, “I shall be starting off soon on 
a 2 or 3 weeks botanical expedition into unexplored 


parts botanically of Missouri, getting into many 
remote sections." Some readers may wonder when 
Julian really began botanizing: on October 13th, 
1930, he collected Grindelia lanceolata along the 
railroad tracks bordering the Washington Univer- 
sity campus. That collection was numbered 1506. 
By September 11, 1932 he had numbered to 7/64, 
or 5,658 field numbers in less than two years. 
Now and then I encountered Julian at the Gray 
Herbarium or the New York Botanical Garden. 
Then, with deep pleasure, I came here in 1986 to 
work in the library, as Julian before me had come 
to forage in the hortus siccus, the dry garden. In 
researching the vismias and the shrubby gentians, 
I came to know Julian's detailed collections and 
his keen interest in plant geography and those 
“unexplored parts botanically” 
their specimens you shall know them!— so true of 
letter of 1933: 
**One derives the chief or almost the chief pleasure 


of Venezuela. By 
botanists. Let me return to Julian's 


from studying plants in their natural habitat, ex- 
ploring for them in remote places which add to the 
zest and thrill of discovery. At least I have always 
felt that way and most field naturalists hold similar 
opinions." The tepuis used to be called the "Lost 
World" but that was before Julian. Omnia fleu- 
rit.— Joseph Ewan, Missouri Botanical Garden; 


from Julian Steyermark's memorial service. 


My introduction to Julian Steyermark was 
through the Flora of Guatemala while working as 
a Peace Corps volunteer in Honduras on a floristic 
survey of one of the central valleys. Honduras does 
not have any written flora, so the next best thing 
was the Flora of Guatemala of which many vol- 
umes were written by Standley and Steyermark. 
Filled with useful information, excellent keys, and 
descriptions, it taught me a great deal about trop- 
ical botany. When I was finally preparing to leave 
Honduras with some of my more interesting col- 
lections, | was directed here to the Missouri Bo- 
tanical Garden as the best place to have them 
identifie 

When 1 finally made it here some months later 
and luckily was able to get a job, I was told that 
Julian Steyermark had just come up from Vene- 
zuela and was on the staff! One of the first things 
that I wanted to do was meet him, and you can 
imagine my surprise and pleasure upon learning 
that I was going to be working for him, and how 
much even more surprised I was when three months 
later | was in Venezuela on a major expedition to 
the Lost World! 

It was a real thrill and a tremendous learning 
experience to work in the field with Julian. He had 


Volume 76, Number 3 
989 


Steyermark Recollections 635 


RE 4. Field trip, 1929-1930. Left to E Charles Leo Hitchcock, ra Allen, Julian Steyermark, 
nde Darlington, Mildred Mathias, Dorothy Francis, George Goodman. Photo contributed by Mildred Mathias. 


an "eagle eye" for spotting plants and if you didn't 
see something that he did, he would let you know 
it. When I first went out on my own to collect 
some specimens I found a beautiful plant that I 
thought was very unusual; it had large bright red 
bracts surrounding a dense cluster of flowers. When 
I returned to camp, I pulled it out of my collecting 
bag with pride to show him. But when he saw it, 
he took it from me in disgust and threw it to the 
ground, saying sternly “this is Psychotria poep- 
pigiana subsp. barcellana, one of the most com- 
mon and weedy plants of South America, don't 
ever collect it again." I was a little bit embarrassed, 
so the next day I made even more of an effort and 
risked my neck to get up on some diffici 
bluffs. When I got back to camp, I sheepishly pulled 


another member of the same family out of the bag 


lt sandstone 


to show him. This time, however, he looked at it 
with a great deal more interest, a puzzled look 
came over his face, and he admitted that he didn't 
know what it was, and that it likely 
something new. From then on I learned that when- 
ever he gave that look at a plant, it meant some- 
thing interesting had turned up. In this particular 
case it was a genus of the coffee family new for 


was very 


science. 

Fieldwork was very special for Julian—rather 
than “vacations,” he went on field trips. It was 
impossible not to be caught up by his enthusiasm 
for collecting plants. He always likened collecting 


to a “treasure hunt." Even with his two artificial 
hips, he would plow through brush, crawl over and 
under logs, fall down, laugh about it, and push on. 
He truly liked collecting all kinds of plants, but I 


think that the Bromeliaceae were his favorite fam- 


Julian was involved in the first helicopter ex- 
pedition in Venezuela, which took place in 1967 
to the isolated Cerro Jaua. He quickly realized the 
advantages of that mode of transportation and found 
that it would increase even his amazing productiv- 
ity. He would use it frequently in the following 
years to get to some of the most inaccessible of 
the Lost World summits. To think that in all the 
years prior, when he made so many of his famous 
expeditions to some of the most remote areas in 
the country, that everything was by foot, including 
long and hazardous climbs to the tops of several 
of the more prominent tepuis, such as Chimantá 
and Auyan-tepui. Once I understood the complex- 
ities involved in these huge expeditions back then, 
my admiration for him increased even more. 

e was a great friend to the Indians of the 
region. They enjoyed his sense of humor and spirit 
of adventure. He is somewhat famous in certain 
Indian tribes. Two years ago, when we were plan- 
ning a trip to a series of unexplored mountains in 
the heart of the Lost World, we were surprised to 
be met by the son and grandsons of an Indian who 


had worked with Julian back in the 1940s. They 


636 


Annals of the 
Missouri Botanical Garden 


had heard that **Hulian Esteyermark" was to be 
back in the area, and were eager to volunteer their 
help. 

On a personal side, it took a while to get to 
know him, and he did have some unusual ways. 


I thought that he was kidding when he told me 
about his liking for rock & roll music until he 
showed me his more than 800 records ranging 
from Elvis Presley to the Rolling Stones. He also 
loved Venezuelan and Andean folk music. 


— He loved to bet milkshakes on whether his friends 
could identify a plant or not. I paid out my fair 
share to him and to Ron Liesner (an expert at plant 
family identifications). 


-He would eat sparingly for breakfast or lunch, 
but there were few who could match him at the 
dinner table. Buffets at nice restaurants were his 
favorite, and he would go back time after time 
before he was ready for dessert. 


—Other than fancy buffets, he thrived on fruits, 
vegetables, and ice cream and was a great exper- 
imenter with native edibles. Flower fragrances were 
we would have contests in the 
field to see who could find the most fragrant flower, 


irresistible to him; 
and he smelled every one with great pleasure. 


— Swimming was a favorite pastime of his. The 
only time that he would take off from the herbarium 
or field (or let any of his assistants do so) was to 
go to the beach. He loved floating in the waves 
and would do so for hours at a time. 


Julian was a most ardent and effective conser- 
vationist. He worked hard in Venezuela through 
the National Park Service to protect the forests in 
the North that were, and still are, rapidly being 
destroyed, and was instrumental in identifying many 
protection. In late 
88, we were told that the President of Venezuela 


areas in the South worthy o 
was signing into legislation a new National Park in 
the delta of the Orinoco River. It was planned with 
Julian's help. 

Finally, I would like to share some parts of a 
letter that was sent to Julian a few weeks ago that 
illustrates his impact upon others around him, not 
just as a botanist, but as a person who could inspire 
others: 


was a student of your Biology Class at University 
City High School during the 1937-38 period at the 
time E K to your leaving for the Field Museum 
in Ch . My memories of deed are very strong and 
ae e as a teacher and as a ou provide 
a great ia model for a i anali interested in sci- 
ence. And | remember how impressed I was with the 


fact that my teacher was leaving to become Curator 
of a portion of the famous Field Museum. You and 
several of the other faculty members at U. City H 
provided great inspiration to accomplish at that point 
in my life. Thanks. 


Fifty years later, another student of his feels the 
same way. 

I am deeply saddened to lose a great friend and 
a teacher. I will never forget him and what he has 
done for botany, for conservation, and for me.— 
Bruce K. Holst, Missouri Botanical Garden; from 
Julian Steyermark's memorial service. 


The expression on his face was rather eloquent 
when Julian saw the pile of newspapers that I had 
put in front of him on his desk. He knew that | 
was an ecologist starting to do research in the cloud 
forests of Rancho Grande, in National Park Henri 
Pittier some 100 km west of Caracas. My "bo- 
tanical collections" consisted mainly of large, 
labeled leaves pressed between five or even ten 
sheets of newspaper in order to assure their 
drying. “What am I supposed to do with this?" 
Julian asked me impatiently. “Well, I think that 
you can identify them," I answered a little bit 
scare fter a few silent minutes, during which 
Julian lane ed through the "specimens," he re- 
turned the pile to me and said, shaking his head, 
“These specimens are unidentifiable, they are all 
sterile; there is nothing I can do with them.” And 
with these words he continued typing labels on his 
old OLIVETTI typewriter, and I suspected that | had 
done something wron 

After this episode, vids h occurred sometime in 
early 1974 in Julian's office at the Instituto Bo- 
tánico in Caracas, | realized that plant collecting 
in a tropical cloud forest was rather different from 
what I had done before on a very limited scale in 
the Mediterranean macchia of southern ltaly or 
the island of Sardinia. Under Dr. Steyermark's 
guidance I learned how to make proper plant col- 
lections and how to identify them to family, genus, 
or even species. I soon realized, too, how fortunate 
I was to have such an outstanding teacher: every 
passing day | was increasingly impressed by the 
astonishing taxonomic knowledge of this man, who 
seemed to be omniscient and blessed with a re- 
markable memory of the most complicated plant 

mes! 

After having been invited to my first chili con 
carne at Julian's home (his favorite dish expertly 
prepared by his wife Cora), 
strengthened, partly because in the meantime I 
had improved the quality of my botanical speci- 
mens. À couple of vears later, we engaged in a 


our relationship 


joint venture, resulting in the publication of the 


Volume 76, Number 3 
1989 


Steyermark Recollections 


Flora del Avila. Julian wrote most of the keys of 
that book at home while recovering from his two 
hip operations and while I was in Austria concluding 
my Ph.D. thesis. At my return to Venezuela in 
September 1976, we then had a rather hard time 
finishing the rest of the manuscript (some 1,600 
pages), since my Tyrolean stubbornness was not less 
persistent than Julian's Missouri stubbornness! This 

ook, however, constituted a tremendous experi- 
ence for me, and since then we had become col- 
leagues, working together very closely in the In- 
stituto Botánico. From then on, Julian was always 
anxious to be the first to see my collections from 
the Venezuelan Amazonas and later from the many 
tepuis of the Guayana Highlands. 

In 1982 we decided to start another new project 
titled Flora and Vegetation of the Venezuelan 
Guayana. We realized that such an enterprise 
would take years of most intensive work, but we 
both decided to dedicate all our efforts and re- 
sources to that project. Although destiny later di- 
vided our paths, Julian and his collaborators have 
advanced relentlessly towards completion of that 
flora, whereas I have continued to contribute by 
collecting interesting plants from the many places 
visited during my ecological and phytogeographical 
fieldwork in that region. 

It was a great satisfaction for me to invite Julian 
to an expedition into the Chimanta massif, one of 
the richest Guayana mountains, where he and John 
Wurdack had done such pioneering botanical ex- 
ploration in 1953 and 1958. When we established 
camp near a huge colony of Chimantaea mirabilis, 
a most striking member of the Compositae discov- 
ered by him in 1953, Julian could not hide the 
emotion caused by this reencounter after 25 years! 
And then, there he was, the " Julian Stey- 
ermark, sitting at his camping table labs and 
pressing plants and plants and plants, 
amazingly numerous plants every night (at that 
time he was at his number 128,000). And every 
two minutes he would call me and exhibit another 
exciting find, or explain another variation or another 
novelty. One year later we would be again on the 
same mountain and discover a new genus (4co- 
panea in the Theaceae), and then get flooded at 
our camp at 2 o'clock in the morning with icy 
water half a meter deep from the nearby cano! 

Julian Alfred Steyermark, the great botanical 
explorer of the Guayana and many other places of 
the Venezuelan country, has now left us, but his 
memories, as numerous as his botanical collections, 
will always accompany us from tepui to tepui, to- 
gether with the gratitude of having had the privilege 
to be one of his students!— Otto Huber, Caracas, 
Venezuela. 


I first heard of Julian Steyermark when I was 
a student at the University of Wisconsin. Dr. Hug 
Iltis said he wanted to prepare a flora of Wisconsin 
that was as good as Steyermark's Flora of Mis- 
souri. Julian enjoyed hearing that. 

I first met Julian when I went to Venezuela in 
1977 as part of the joint Missouri Botanical Garden 
and Herbario Nacional de Venezuela project spon- 
sored by NSF and CONOCIT. When I arrived, 
Julian was skeptical and concerned and told me 
about the difficult conditions: rain, heat, 
snakes, and other perils. Julian, Francisco Delascio, 
and I went to the Delta Amacuro (delta of the 
Orinoco River) for a month. The collecting and the 
trip were exciting and one was able to ignore the 
difficult conditions. I quickly learned Julian was 
very good at spotting plants to collect. He first 
started using clippers when he saw me using them 
and we sometimes joked that he learned to collect 
plants from me (he had 114,000 collections when 
we met) 

Numerous field trips with Julian were a privilege. 
I learned many plants from him and how to find 
specimens to collect. We frequently bet a milk- 
shake if we thought we could catch the other person 
not knowing a plant to family. Even a sterile plant 
was game if that person collected it. Usually I could 
catch him only if it was a genus that was unusual 
for the family and the genus was new for Venezuela. 
We also bet milkshakes for guessing how many 
plants we had collected before we pressed and 
numbered them. I, also he, sometimes would collect 
numerous extra mosses so the competitor's guess 
would be too low. Once, after Julian had guessed 
right three days in a row, he stated his guess aiming 
at a fourth win, then I guessed one lower, and 
Ángel González guessed one higher. Julian had to 
do something else. Ángel and I pressed the plants 
and wrote the field notes. It came out to Julian's 
number! Ángel and I decided to trick him. One 
species of herb had both flowering and fruiting 
individuals. We split them into different collection 
numbers. But Julian had the last laugh. Not only 
were they different species, they were different 
genera! We also guessed at arrival times for a 
milkshake. Cora, Julian's wife, said they had bet 
ice-cream cones when they did fieldwork in Mis- 


insects, 


ouri. 

Julian loved to try different restaurants and go 
to the movies, and many of his friends enjoyed 
going with him. For a long time, Julian was my 
closest friend. Julian had a good sense of humor 
and it was fun to tease him. But it did annoy him 
when I said I was from Missouri and to show me. 
Gerrit Davidse did the same. For a long time he 
idn't consider us Missourians. 


un 


638 


Annals of the 
Missouri Botanical Garden 


e often corrected my Spanish. I sometimes 
joked that Julian taught me Spanish. He vehe- 
mently denied it, but everyone else liked it. 

When he came to Missouri, he had to get his 
driver's license, and I had to teach him to drive 
my car. Later, I joked I had taught him how to 
drive. He would mention the first two things the 
driving examiner asked were the lights and horn, 
and I hadn't shown him those. 

Julian claimed that he had shaved every morning 
since the thirties. We could be short of water and 
he would still shave. He, Bruce Holst, and I were 
almost stranded on top of a tepui without our things. 
Bruce and I kidded him that we would see him 
unshaved. But then we decided he would shave 
with his machete. The helicopter came so we didn't 
find out. 

Julian and I were in some very risky, potentially 
dangerous places in collecting. But the most wor- 
ried and most concerned I ever saw him was when 
we arrived back in Caracas and he couldn't find 
his hairpiece. It frustrated him when women saw 
him without his hairpiece the first time in the field 
and told him they thought he looked better without 


When we were stranded on Cerro Duida, wet, 
with no shelter or supplies, in a potentially fatal 
situation, Julian joked about dying on a tepui. We 
assured him that was fine as long as it was some 
other tepui. 

In the field Julian often remembered collecting 
the same rare species 30 years previously. An 
amazing number of species were either described 

him or named after him. I wonder what level 
Venezuelan botany would be at if it were not for 
Julian. He worked fast, long hours, and accurately. 

One could go many places in Venezuela and 
local people would remember Julian when he was 
there 20 to 40 years previously. 

Identifying specimens, I worked on many dif- 
ferent plant groups. Julian was a good friend be- 
cause I could talk about many different groups and 
he had worked on them also. 

For quite a while when I ran into a problem 
with a plant group or if I had a plant I couldn't 
identify to family, my first reaction was still to go 
ask Julian, only to remember Julian was no longer 
here.— Ronald Liesner, Missouri Botanical Gar- 


den. 


I first met Julian Steyermark in October, 1963, 
at a difficult time in my life, and I am proud to 
say that since that time, he has helped me more 
than anyone else. 


At that time I was an unemployed and com- 
pletely frustrated young man 24 years old; and 
just “to kill time," I decided to draw flowers. So 
I began to go around with some paper and a box 
of color pencils, and devoted myself to drawing 
flowers in Caracas near where I lived in a student 
dormitory. After a month or so, I had checked all 
the gardens, houses, and whatever natural vege- 
tation there was around and had exhausted inter- 
esting subjects for drawing and so expanded into 
surrounding neighborhoods, eventually reaching the 
Botanical Garden of Caracas, where there was a 
large variety of exotic and beautiful flowers. 

remember that one day, while concentrating 
on trying to reproduce the colors of a 
paradise" flower (Strelitzia reginae), a man came 
up to look at my work. I had seen him occasionally 
on previous days, but we never had spoken to each 
other. He was August Braun, the head gardener. 
After exchanging some comments, he told me that 
there was no artist at the Botanical Garden, since 
the former artist (the late Mr. Ebersteins) had 
retired four months earlier; and he suggested that 
I show my drawings to Dr. Lasser, the director. 

The following day I took some of my best draw- 
ings and showed them to Dr. Lasser. He liked them 
but said that drawings needed for scientific publi- 
cations had to be more “objective” and made with 
black india ink. As I had no experience with that 
kind of drawing, he introduced me to Dr. Julian 
Steyermark and asked him to give me some ori- 
entation. So | met the man, and got the job! 

After I had gained some confidence, in 1963 
Julian asked me to draw a new species that he was 
describing, Palicourea chimo, and 1 complained 
that I had only herbarium specimens, not live plants. 
Julian, who also began his botanical career as an 
artist, understood and helped me look for a solution 
and invited me along on a trip to Naiguata, on the 
northern side of the coastal mountain range of 
Venezuela, where he had not been before. The 
Palicourea was encountered and drawn fresh. 

On the two-day expedition we found a new 
species, which Julian named Heisteria olivae (later 
placed into the synonymy of H. latifolia) to honor 
our guide, the young architect Francisco Oliva 
Esteva [see recollections in this volume J. 

Julian was so impressed by the rich vegetation 
there that two weeks later we went back to explore 
further. As Julian had begun his botanical activity 
in Venezuela in search of quinine plants and was 
an adviser to a large drug company in the U.S., 
he was always interested in potentially useful plants 
as suppliers of drugs. On our second expedition to 


Volume 76, Number 3 
1989 


Steyermark Recollections 639 


the coastal range, we found a big Lacmellea tree, 
called **vaco negro" (black cow) by our guide, who 
stated that its sweet, milky sap was used as a 
remedy for asthma. Since we found only flowers 
and fruits on the ground, Julian felt frustrated, but 
I returned a few weeks later, climbed the tree using 
and collected several complete 
specimens. I illustrated these, and Julian named 
Lacmellea costanensis as a new species. 

So delighted was Julian with the scenery of those 
slopes facing the sea, completely carpeted by virgin 


a liana 


cloud forests, that he soon made yet another ex- 
pedition there. On this third trip, we went with Mr. 
G. . Dunsterville, the great orchid artist, and 
his wife. Nora. Cora Steyermark also came with 
us, but, as she was suffering from some bad health 
and could not match our pace, Julian went ahead 
with a bunch of yellow and red woolen threads, 
dropping them from time to time along the trail 
for her to follow. So, late in the afternoon, Cora 
reached us bright with joy and carrying all the 
threads she picked up along the trail. 

n this same trip, Victoriano Carreno Espinoza 
joined us. Julian had met him a month earlier on 
a previous expedition to Cerro Humo in the eastern 
peninsula of Paria. He was so satisfied with Victor’s 
knowledge of plants, skill, and enthusiasm, that 
Julian brought him to Caracas and helped him get 
a job in horticulture at the Central University. 
Victor continued to work with Julian and quickly 
became his faithful and efficient assistant on many 
expeditions throughout Venezuela during a fifteen- 
year period. 

When I met Julian, he was living in an apartment 
complex called **Residencias Magnolia" located in 
Los Palos Grandes, Caracas. The apartment had 
a terrace where he kept plants that he brought 
back from his botanical expeditions. He would lead 
visitors on a ritual tour around the garden to give 
an explanation of the plant names and the regions 
they came from, with considerations about their 
color, shape, or scent. But, Julian's expeditions 
were so frequent and fruitful that the apartment's 
terrace became too crowded, and about 1971 he 
moved to the suburb of Santa Eduvigis. There he 
had more room to display his plants and on the 
back side of the house he created a miniature jungle 
under a mango tree. In some cases Julian collected 
sterile plants, which he brought home until they 
flowered and then made herbarium specimens from 


m. 
Besides the small garden space, noise from traf- 
fic, and the fact that their house was broken into 


bothered Julian and Cora, so in mid-1975 they 


found a new house in the neighborhood of Santa 
Fe Sur. They were particularly pleased with it, 
because from it they enjoyed a complete view of 
Avila National Park across the Caracas Valley, 
and it was close to a place officially listed as a 
“green area," where supposedly no other buildings 
could be built. But several months later, in spite 
of all the official statements, construction began 
on a huge apartment complex whose wall reached 
right up to the Steyermark's garden. He felt very 
frustrated, and in his field book he remarked sadly, 
“Nov. 20, 1975. Destruction of green area back 
of our home." 

Yet the new house still had room for gardening, 
and the Steyermarks displayed the beautiful col- 
lection of bromeliads on the porch and put the 
aroids, orchids, begonias, peperomias, and ferns 
under a plastic net. In the center, there was a 
planter of red roses, surrounded by several Hip- 
peastrum lilies, and another plant of rosemary 
surrounded by the thorny Euphorbia splendens. 
In back, towards the new apartment complex's 
wall, there was a Chrysalidocarpus lutescens palm, 
a Tibouchina semidecandra with big purple flow- 
ers, a Mussaenda erythrophylla, a healthy Clusia 
minor with waxy rose flowers, a terrestrial Cyr- 
topodium orchid, and other miscellaneous herbs. 
On the right side, climbing on the fence was Aris- 
tolochia ringens, Heliconia caribaea, Nicolaia 
elatior, and Lafoensia with white flowers pollinated 
by bats. In a pot, there was also a “lily of the 
valley" with very fragrant white flowers n Julian 
especially enjoyed. On the side of the garden close 
to the porch grew a row of [ris confusa that never 
bloomed. Once a month, Victor went to the house 
to cut the grass and clean the garden. 

Cora, a member of the Audubon Society, en- 
joyed observing birds, and every day would put 
bananas and oranges in a feeder to attract them 
to the garden, and sugar water for the humming- 
birds. She kept a diary of the garden and recorded 
the exact date of blooming for every plant, days 
of rain and sun, and other data for the garden's 
management. She was also one of the founders of 
a garden club, which in 1973 created a scented 
garden for the blind along the avenue Rómulo 
Gallegos in Caracas. 

On many occasions Julian would honor me with 
an invitation to dine in his house, where I thor- 
oughly enjoyed Cora's excellent cooking, especially 
the pies and brownies. Sometimes when we came 
back late at night from some expedition, he invited 
me to sleep in their house in the guest room. The 
following day, of course, I would get a delicious 


640 Annals of the 
Missouri Botanical Garden 
breakfast. | remember that when we sat at the At that time Julian was preparing a monograph 


breakfast table, Julian and Cora would recount the 
dreams they had the night before. 

On a trip to Puerto Cabello in 1973 with Julian 
and Victor, I found out that Julian had a love for 
snakes. We walked along an old canal breached 
in areas and useless due to many rocks and earth 
that had fallen into it; 
little waterfall, it was full of clear and fresh water. 
Julian, who loved to drink from sparkling streams, 
put his right hand on the edge of th 
bent over to take a drink (with the left he was 
holding the plant press). When he finished drinking, 
and before he stood up, he spotted a poisonous 
snake out of the corner of his eye, a fer-de-lance 


but in a corner, under a 


e water and 


(Bothrops atrox), just between his hand and his 
cheek. Victor hurried there with the machete to 
kill it, but Julian didn't let him; instead, he pushed 
and chased the snake away into the high vegetation. 

In 1983, Julian returned to that area again with 
Paul Berry and me to gather specimens of Sela- 
ginella gigantea, but the old trail was completely 
lost. Paul and I managed to reach the Selaginella 
site, but just as we arrived, a heavy thunderstorm 
came up. Paul hurried to make a collection of the 
rare Selaginella, and then we headed straight down 
the mountain, since we lost our trail very quickly. 

‘hen at last we reached Julian half an hour later, 
the rain still falling, we found him sitting on his 
press, trying to protect himself with a completely 
wet piece of paper but he regained his spirits as 
soon as he saw Paul's collection. All that area where 
the Selaginella is found now belongs to the San 
Estaban National Park, created in 1986 thanks to 
Julian's enthusiastic report 

In October 1970, Julian joined the Venezuelan 
Brazilian Frontier Commission in a survey of the 
Cerro de la Neblina, the highest mountain in the 
Guayana, and he collected as intensely as ever. 
While participating in that trip, the expeditionaries 
were sent supplies from San Carlos de Rio Negro 
(Venezuela) and Cocui (Brazil) by helicopter, so 
Julian decided to send his first lot of collections 
ahead to San Carlos, preserved in formol in plastic 
bags; but, due to confusion, the plants were un- 
loaded in Cocui. When Julian returned to Caracas, 
he realized that a lot of his plants were missing, so 
in March of the following year he sent me down 
to San Carlos and Cocui to see if I could find the 
lost collections and bring them back. When I met 
the Brazilian officer in charge at Cocui, he told me 
that he had been there only since January, and 
that the soldiers, cleaning the warehouse, found a 
lot of very bad-smelling plants and threw them into 
the Rio Negro. 


of the Rubiaceae for the Flora of Venezuela to 
which I contributed as an artist, and in 1973, when 
I received my degree at the Central University, I 
resigned my job at the Botanical Garden and turned 
to teaching. Julian complained about my decision 
and told me that he had a lifelong dream to write 
a flora of the Guayana, for which he would like to 
employ me as the artist; but at that time I didn't 
see how it would be possible. Anyway, I kept in 
contact with him, and from time to time, on the 
weekends or during vacation, he would invite me 
to join him on some botanical expeditions, even 
though at that time he was plagued by arthritis 
of the hips. As Julian could not walk like he used 
to, he needed extra help on his collecting trips. 
In 1973 Julian was asked b 
agency to make botanical surveys of several coastal 


y a governmental 


areas recently designated as national parks, and 
he invited Victor and me along. We would rent a 
boat for the day, and Victor and I would go out 
with a bag and a machete and collect samples from 
every plant we saw. Then, we would go back to 
the boat, where Julian cleaned the specimens and 
put them in his press. In the evening, we returned 
to the hotel with the plants, and Julian, keeping 
with his usual routine, worked for hours to number 
the specimens and to make notes on habit, habitat, 
and other peculiarities that he observed about the 
living plants. On the following day when the em- 
ployees went to clean the apartment, they would 
complain loudly about the mess. 

In February 1974, Julian joined the great ex- 
pedition directed by Charles Brewer-Carias to ex- 
plore the biggest sinkhole on top of Cerro Sarisa- 
rinama in southern Venezuela. Victor went with 
him since he was having difficulty walking at that 
time. Even with the problems getting around, he 
managed to collect more than 1,000 plants in 20 
days, many of them rare, including 50 new to 
science. From the bottom of the sinkhole, Charles 
sent up a pretty little melastome with white flowers 
and a shaded rose-colored ring around the calyx. 
It turned out to be a new species that John Wur- 
dack of the Smithsonian Institution named Phain- 
antha steyermarkii. For his part, Julian felt so 
satisfied and grateful with Victor's work, that he 
named three new species for him: Psychotria car- 
renoi, Phyllanthus carrenoi, and Sloanea car- 
renoi. 

During the several months before the expedition 
to Sarisarinama, Julian had. been corresponding 
with people who had had hip operations to correct 
arthritis problems. They told him that they had felt 
much better after the surgery, so at last he decided 


Volume 76, Number 3 
1989 


Steyermark Recollections 


to undergo the operation. Since he had to be op- 
erated on one side first, and then six months later 
on the other side, he had to stay at home for a 
whole year. But how was anybody going to convince 
Julian to rest? That word was unknown to him, 
and in fact I suspect he was afraid to rest. For 
that reason, it did not surprise me that he soon 
found a way to keep busy at home. 

His good friend Otto Huber had been preparing 
a list of all of the plants in the national herbarium 
ever collected on the south slopes of Avila National 
Park, the mountain that rises above the valley of 
Caracas to the north. Since Otto was an ecologist, 
he proposed to Julian that they coauthor a book 
to be called the Flora of 4vila. Julian was fasci- 
nated with the idea, but he insisted that the value 
of such a flora would increase if it also covered the 
less-explored northern slopes. 

As he was recovering in a hospital room that 
had a view of Avila high in front, he was inspired 
to write the following phrase in the introduction to 
the flora in reference to me fantastic view of the 
mountain from the city: e see this beautiful 
green panorama as we look through the windows 
of our houses, offices, colleges, hospitals or fac- 
tories.” 

Surely when Julian planned the Flora of Avila 
he had in mind the style of his huge and successful 
work, Flora of Missouri. Since that was lavishly 
illustrated, he wanted me to draw as many species 
as possible in every plate, just like in his previous 
work. He didn’t want to prepare a strictly scientific 
work, but one that was usable by many different 
people. One of the ways that he did this was to 
insist that the publisher include a metric scale on 
the back cover, with the hope that the flora would 
be carried along up the mountain and used on the 
spot to identify plants. A hopeful assumption, in- 
deed, but one that demonstrated Julian’s interest 
in making plants known and loved. 

So, we began work on the Flora of Avila! I had 
to make drawings from live material, and then bring 
them back to Julian to be identified. As the work 
progressed, he began to complain that I had too 
many drawings of orchids and other showy plants 
but few or nothing of several families; and as time 
passed, we reached a serious impasse. Luckily for 
us, there was Otto Huber, who could throw water 
on the flames, and the work went on. It was hard 
for Julian to type, due to the semireclined position 
in which he had to remain, so he would turn his 
manuscripts over to Cora, who was the only person 
able to read and interpret his thick scratches. Thus 
she also was a member of the flora team. Julian 
was later awarded the prestigious "Order of Henry 


Pittier" by the Ministry of the Environment for 
the flora. 

One peculiarity that I observed in Julian's char- 
acter was that during his fieldwork, he never dared 
to identify a plant to species, only to genus. For 
example, once I asked him, “Julian, how do you 
like that nice Palicourea fendleri?" Immediately 
he criticized my confidence, which he termed as 
an expression of ignorance and presumption, and 
pointed out that P. fendleri was known only from 
the coastal range of Venezuela, and now we were 
in the Andes .... 
Garden in Caracas, would he make a decision on 
the identity of a plant by checking it against her- 
barium specimens or sending it to one of his col- 


Only later, at the Botanical 


leagues. Once, I told him that I felt that he was 
too insecure about identifying plants. He answered 
sharply that he was a scientist and I was only a 
rank amateur, that ignorance is petulant, and that 
I should be more careful in my opinions. 

In those years, Julian also devoted his weekends 
to a survey of the plants of the wild areas around 
his neighborhood of Santa Fe, which at that time 
was being encroached upon by housing develop- 
ments. That work was later included in a paper by 


Paul Berry and Julian entitled “Florula of the 


deciduous forests of Caracas.” 

As a scientist, Julian was absolutely objective, 
and he made no statement without proof to back 
it up; but sometimes, I thought him to be too 
objective. For example, when we were working on 
the Flora of Avila, I was fascinated with a beautiful 
orchid, Oncidium papilio, growing in Julian's gar- 
den, which in the thirties had been collected 
to Caracas" according to the label. I argued to him 
that ^ 
but Julian cut me short and said that he, as a native 
show-me-man,” and therefore, 


"close 


close to Caracas" might mean "in Avila," 
of Missouri, was a “* 
he wouldn't include any species in his treatment if 
not supported by a more precise collecting locality. 
But, if I were to make a collection of that orchid 
from the mountain itself, he would include it in the 
flora .... I did look for it indeed, but I couldn't 
find a specimen; that is, until 1985 in the northern 
side of the national park, seven years after the 
flora was published. 

In September 1979, Julian went back to Mis- 
souri for a couple of weeks for the dedication of 
the Steyermark Woods. Cora stayed in Venezuela 
alone in their house, and an American lady who 
lived close by kept in contact with her and took 
her to the market whenever she needed to go. Cora 
felt very uneasy and alone in Venezuela, and Julian 
was making plans to bring or send her back to 


Missouri. In fact, since some time earlier, he had 


642 


Annals of the 
Missouri Botanical Garden 


been in contact with Dr. Peter Raven, Director of 
the Missouri Botanical Garden, and had told him 
of his desire to write a Flora of the Venezuelan 
Guayana. Dr. Raven was interested, and they be- 
gan to consider ways to produce it. Although Julian 
had become a Venezuelan citizen, he realized that 
the only way to obtain the help that he needed was 
to return to the large herbarium in Missouri, and 
so he made final plans to leave Venezuela with Cora 
in the spring of 1 

All of his careful plans changed abruptly in 
October of 1983. One night, when he arrived home 
from the botanical garden, he found Cora tied to 
a Chair, very scared. What had happened? That 
afternoon, somebody rang the bell at the entrance 
and when she opened the door, there were two 
young men at the gate. One of them pointed a gun 
at Cora and ordered her to open up. She did as 
they said, and when they entered, they tied her to 
a Chair with a blanket and searched the house. 
They found very few things of interest and soon 
left, leaving Cora tied to the chair, where she stayed 
until Julian found her hours later. He called the 
police immediately, and when the officers arrived, 
and as a matter of routine, they asked Cora to 
show them her passport. She went upstairs, still 
very upset, and on her way back down lost her 
footing and fell, breaking a bone in her right leg. 

er leg was set in a cast, and she was released 
from the hospital, but a couple of weeks later she 
contracted pneumonia, and a week later, on No- 
vember 4, 1983, she passed away at age 92. Her 
last wish to return and see her country again was 
unfulfilled. 

I visited Julian frequently in the following weeks. 
He spoke very little about what had happened, 
perhaps trying to forget as quickly as possible. He 
kept busy working and asked me seriously about 
contributing the drawings for the flora of Guayana 
that he had planned. In order to motivate me, he 
placed at my disposition the thousands of color 
slides he had taken during many years of explo- 
ration. I started by choosing close-ups of flowers, 
which would be the most useful for my work; but 
as I went on, I was impressed by the many beautiful 
slides of landscapes, and an idea came to my mind: 
to write a book on the Guayana using many of 
Julian's slides. So, without telling him anything, I 
wrote an essay on the history of the search for E 
Dorado, entitled “En la Tierra del Oro," and added 
to it 100 of the slides. When I had the manuscript 
completed, I showed it to Julian, who was at first 
confused, because he didn't understand that I had 
tricked him a little bit. Anyway, after he read the 


manuscript and made some corrections, he agreed 
that I should carry out the plan and went along 
with me to an editor, who promised to publish it 
as soon as possible, though it was not completed 
until December of 1988, too late for Julian to see 


- 
"p 


In December 1983, I began the drawings for 
the Flora of the Venezuelan Guayana using the 
provisional survey of the herbarium in Caracas that 
Julian made to find out which families were known 
from the area. In April of 1984, he moved back 
to his hometown of St. Louis, but only a few 
months later he came back to Caracas for two or 
three months to begin checking the herbarium sys- 
tematically to record all of the plants collected in 
the area covered by the intended flora. And in 
February of 1985, Julian was in Caracas again. 
This time, he had with him a young man, his 
efficient assistant Bruce Holst, who helped in the 
general inventory of the herbarium; he was admired 
for his ability and correct Spanish. 

Until then, except for the short trip in 1971 in 
search of Julian's lost plants in San Carlos and 
Cocui, I had never been on any expedition south 
of the Orinoco River. Julian wanted to give me an 
opportunity to see as much of the Guayana as 
possible now, to help me in preparing the drawings 
for the Flora by taking pictures, or making draw- 
ings from live plants, as I preferred. So, in Easter 
week of 1985, Julian, Bruce, and I headed for the 
Gran Sabana, in Bolivar State, with the intention 
of going as far as possible in the ten days that we 


On that trip, we rented a car called a "Zephyr," 
a clumsy, two-wheel drive that, when weighed down 
with all of our gear, barely cleared the ground. 
Because of the busy vacation week, all of the four- 
wheel drive vehicles were already rented. On the 
rough roads of the south, everybody looked at us 
in astonishment since it was the only two-wheel 
drive in the whole Gran Sabana, but Bruce man- 
aged to sort out the numerous ruts in the road and 
cross the streams to make it all the way to Kamá 
g the way, we collected mostly along 
the road since Julian didn't want Bruce or me to 
go too far into the woods and use up too much 
time, but we managed to find numerous plants to 
sketch and photograph that would be valuable for 
my illustrations. 

On our way back down to the lowlands, we had 
a funny incident with a group of university students 
camped by the side of the road who were in charge 
of checking the cars returning to the north and 
helping the tourists with information. As we slowed 


Volume 76, Number 3 
1989 


Steyermark Recollections 643 


down to pass them, the guard on watch blew his 
whistle for us to stop, and when he saw some plants 
that we had in the back of the car, he called for 
the leader to come out of his tent. He asked for 
our identification and told us that it was forbidden 
to remove plants from the Canaima National Park. 
Julian answered that he was Dr. Julian Steyer- 
mark, working for the Ministry of the Environment 
and... 

“Are you Dr. Steyermark, the author of the 
Flora of Avila?" he asked in delight. “I have 
wanted to meet you for many years to congratulate 
you on such a beautiful book!” 

Immediately he called all of his helpers to come 
out and insisted that all of us line up for a picture 
to be taken with our arms around each other. When 
the photographer apologized for a delay in taking 
the picture, Bruce mentioned that there was no 
hurry since he was standing next to the most beau- 
tiful girl of the group. 

In August of the same year, Julian and Bruce 
were in Caracas once more. On this visit, Julian 
received his retirement payment and immediately 
purchased a Toyota Land-Cruiser for use on ex- 
peditions by both the Caracas Garden and Missouri 
Botanical Garden. We took it on its maiden voyage 
to Puerto Ayacucho in the Federal Territory of 
Amazonas (1-13 September 1985). We had some 
trouble at the crossing of the Orinoco since a fierce 
storm was coming up that made the pilot of the 
ferry think twice about crossing the wide river. But 
there was a long line of cars waiting, and at last, 
some people convinced him to make the trip, hop- 
ing to beat the storm. As soon as we were well out 
into the river, it started to rain, and big waves 
splashed over the floor of the ferry, obliging us to 
take refuge under the covered area. Many pas- 
sengers began regretting having started the trip, 
and we were beginning to think that we might not 
make it to the other side. Apparently Julian was 
not at all concerned, even though his new car was 
perched on the front of the ferry, since he sat 
quietly under the overhang seemingly enjoying the 
roller coaster. But after one hour or so of being 
tossed around and getting soaked by the waves, 
we reached the town of Caicara on the other side. 
Bruce overheard one of the ferry attendants say 
that our trip was the roughest that he ever made. 

On that expedition, I was fascinated by the many 
strange flowers that we found, especially with a 
half-dozen species of Mandevilla, a vining genus 
of the Apocynaceae family with showy yellow flow- 
ers (a couple of new species would be described 
later from those collections), and different species 


of purple morning glories and mallows. Of course, 
when I saw a pretty flower, I asked Bruce to stop 
to take a picture; but Julian started getting bothered 
with so many stops, and he ordered Bruce to follow 
his instructions, not mine, and to go ahead without 
stopping any more to make better time. Julian 
looked very bothered when he said that, and we 
went on a short time without speaking, all of us 
feeling tense. I knew Julian as a mild-mannered 
man, who only on a few occasions would lose his 
patience, and usually for just a short time. So, I 
began to tell him in a humble voice: “Julian, sup- 
pose we see a donkey on a bicycle. Shouldn't that 
be a reason to stop and take a picture? ...” (No 
answer). Again, a bit later: “Julian, and if we saw 
a mammoth flying over that boulder over there, 
wouldn't it be a good reason to stop and take a 
picture? . . .” No answer! I understood that maybe 
went too far with my stupidities, and | decided 

o shut up and resign myself to his wishes .... 
Suddenly Julian cried: 

“Bruce, stop! . . . Bruno, did you see those big 
yellow flowers in that bush over there? ... Have 
you already taken a picture of them?" 

n the way back to Caicara a few days later, 
Bruce was driving slowly on the muddy road to 
avoid the many potholes, but Julian asked him to 
speed up, thinking that when we arrived in Caicara 
at the hotel, he had to record in his field book all 
the plants he collected that day. Bruce tried to 
obey, but when he came across any rut, he would 
slow down again. At last Julian lost his patience 
and ordered him to turn over the wheel; and the 
Land-Cruiser started away at full speed, jumping, 
splashing, and shaking us up. We arrived at Cai- 
cara early that evening, and we were the first 
customers in the restaurant, as was usual with 
Julian. 

The following day, early in the morning, we 
crossed the Orinoco River again, this time with a 
nice sun and good weather, and by evening we 
were in Caracas. 

As a man who loved nature, not only to enjoy 
it but also to make it known and be loved, he 
supported with his own money several expensive 
expeditions to Guayana. He also paid for some 
botanists from Venezuela to visit herbaria in the 
United States and for some U.S. botanists to go to 
Venezuela to work on treatments for his flora. As 
he was demanding with himself, he used the same 
standard for others. I was in the office one night 
when Luis Marcano Berti, a Venezuelan botanist 
whose trip to the New York Botanical Garden was 
funded by Julian, called. “Are you also working at 


644 


Annals of the 
Missouri Botanical Garden 


night and on the weekends?" Julian asked. “Yes!” 
"Oh, very good,” answered Julian contentedly. 

In my case, although he wanted me to work 
hard on the drawings, he wished too that I would 
be able to see and enjoy from time to time the 
natural beauty of Missouri and especially of the 
Ozarks. To help me out, he asked Bruce to take 
me camping some weekend, which we did several 
times. So, with their help, I was able to get to know 
some of that wonderful country of Missouri, and 
also gaze at the stars through Bruce's telescope. 
Bruce, for his part, would tell me that in late 
October, when Julian was better, we would all go 
out again together to see the fall colors, which 
delighted Julian so much. 

The weekend after Julian passed away, Bruce 
took me out to the Ozarks. The sun was bright 
that afternoon, and the many colors of the falling 
leaves were spectacular. We regretted that Julian 
was not with us to enjoy one of his favorite shows 
of nature. 

That night, Bruce and I sat a long time looking 
at the fire. The words came out with difficulty, and 
several times I had to bite my lips not to weep. 
Both had lost a great friend, and I missed a man 
who for me was like a father. At that moment we 
felt closer than ever before, feeling the responsi- 

ility we now had to bear of completing Julian's 
great dream, but glad to have been with Julian in 
his last hours. 

A few days later, while writing down these mem- 
ories, a letter came from Victoriano Carreno, dated 
Caracas, October 4, 1988. It said: "Dear friend, 
I heard that you were sick, and I am praying to 


God for your quick recovery. All of us remembe 


you, and we hope that you can visit us again... .”” 

Surely, Julian would have enjoyed receiving this 
letter with the sincere wishes of Victor, the faithful 
field assistant who during many years helped him 
get a bit closer to his dearest dream.— Bruno 
Manara, Caracas, Venezuela. 


The return of Dr. Julian A. Steyermark to Mis- 
souri during the last week in September [ca. 1978] 
provided me with a chance to talk again to my 
esteemed friend and to reminisce about my first 
experiences with him over two decades ago. 

I had grown up on Steyermark's Spring Flora 
of Missouri, which I found very useful in adjacent 
southern Illinois. While I 
the Missouri Botanical Garden 1 —56, working 
on my Ph.D. at Washington University, I was 
constantly reminded of Dr. Steyermark because of 
his vast collections in the MBG herbarium, because 


was doing research at 
9 


of his prodigious writings, and because of the stories 


retold about him by Edgar Anderson, Bob Wood- 
son, and others. 

Following my Ph.D., I began my professional 
career in the Department of Botany at Southern 
Illinois University in January 1957. It was my 
great fortune that during the three-month spring 
quarter in 1958, Julian Steyermark was offered a 
Visiting Professorship in the Department of Botany 
at SIU. The impact that those three months had 
on my professional career cannot be overstated. 

Our department at that time was in cramped 
quarters. | already was sharing a 16 by 8-foot 
office with Professor John Voigt, the departmental 
ecologist. Since Dr. Voigt's interests and mine were 
more closely aligned to Dr. Steyermark's interests, 
the three of us were crowded into this small office. 

Steyermark and I spent every day and partway 
into nearly every evening working side by side. He 
was putting the finishing touches on his Flora of 
Missouri and I, along with my wife, Beverly, was 
revising Jones's Flora of lllinois. Often Steyer- 
mark would quiz me on some specimen he was 
working on or sketching. Many species previously 
unknown by me were quickly and eagerly learned. 

While much of the three months was devoted 
to serious botany, Dr. Steyermark would take a 
little time off now and then to come to our home 
for dinner. On one occasion he took my wife and 
me to the local movie theater to see War and 
Peace. I recall the incident vividly because Julian 
insisted on picking us up in his vehicle, an Inter- 
national Scout built high to enable him to negotiate 
Missouri's back roads. The Scout was so high that 
I had to climb awkwardly. My poor wife stood 
helplessly at the door until Dr. Steyermark boosted 
her up into her seat! 

One of my most pleasant recollections was an 
overnight visit to Steyermark's estate near Bar- 
rington, Illinois, north of Chicago. Julian had talked 
so often about the wildflower preserve he had built 
that my wife and | were thrilled to be invited to 
see it. We arrived early one afternoon. Julian guid- 
ed us over about half of the property, pointing out 
with great enthusiasm the many species transplant- 
ed from the Ozarks. That evening we talked of 
plants, browsed through his extensive library, and 
listened to classical music. 

We awoke the next morning to the delightful 
smell of bacon frying. Cora, Julian’s wife, hac 
prepared a feast, including the most delicious baked 
eggs my wife and I had ever eaten. The rest of 
the morning was spent walking over the remainder 
of the grounds before we all began the long drive 
back to Carbondale. 


From mid-March to late April 1958, Steyer- 


Volume 76, Number 3 
1989 


Steyermark Recollections 


mark enthralled me with stories of his experiences. 
I was particularly excited about his rediscovery in 
1957 of the very rare and little-known Geocarpon 
minimum. This species had been found originally 
in Jasper County [Missouri] in 1913. This re- 
mained the only known station until Steyermark 
had rediscovered it in St. Clair County [Missouri] 
in 1957. My nagging at Dr. Steyermark to show 
me Geocarpon paid off. He agreed to take Dr. 
Voigt and me with him on a three-day excursion 
to see if we could discover additional stations for 
this dwarf rarity. * 

We left Carbondale on May 2, heading in the 
general direction of Springfield, Missouri. Leaving 
Cape Girardeau, we stopped briefly at the Bollinger 
Mill and then headed into the Ozarks. Up and down 
lettered and double-lettered county highways, past 
shortleaf pine woods, over clear, rock-bottome 
streams we went, Steyermark pointing out species 
right and left. We made frequent stops, usually 
“Wait! 


There's a county record." I tried to absorb all the 


after an enthusiastic comment such as 


species as fast as I could, but Steyermark never 
relented. We screeched to a halt in Bollinger Coun- 
ty, pulling off the road next to a steep, wooded 
cherty slope. Julian asked if any of us had ever 
seen the little sedge Scirpus verecundus. We re- 
plied in the negative, and off we went down the 
slope. Scirpus verecundus was not known from 
Bollinger County, but Julian was optimistic, saying 
that this slope looked just right for the little plant. 
In what could not have been more than five min- 
utes, the woods echoed Steyermark's voice: “Whee! 
Over here. I’ve found it!” Scrambling over the 
treacherous chert, Dr. Voigt and I made it to the 
kneeling botanist who crouched over a four-inch- 
tall tuff of green. I had seen my first Scirpus 
verecundus, and I was never to forget the habitat. 
Some 15 years later, while exploring in Alexander 
County, Illinois, I camp upon a similar steep, cherty 
slope. Recalling my earlier experience, I began to 
comb the woods and was rewarded by making the 
first discovery of Scirpus verecundus in Illinois. 

By late afternoon we had reached Dallas County 
and were heading along Route 43 when Steyermark 
observed that the little chickweed on the road shoul- 
der looked peculiar. On closer look, the plant in- 
deed was different from anything we had ever seen. 
We learned later that the plant was Cerastium 
pumilum, a new record for Missouri. 

We persisted beyond supper and into the dusk- 
iness of twilight. By dark, we were still on the road. 
I began to understand how Steyermark had been 
able to canvass every square foot of Missouri in 
30 years—he never stopped to rest. I saw my first 


1958) 


STEYERMARKMOBILE (ca. 
IGURE 5. Steyermarkmobile. Contributed by Robert 
Mohlenbrock. 


Ozark tarantula, scurrying aross the road in the 
glow of our headlights. Finally we stopped for camp 
at about 9:30, reeling from the vast amount of 
information imparted by Dr. Steyermark. 

e arose early on May 3 in anticipation of 
seeing Geocarpon. We had entered Polk County 
[Missouri] near Graydon Springs when Dr. Stey- 
ermark announced that we had arrived at a likely 
looking spot for Geocarpon. It was a glade on a 
west-facing sandstone escarpment next to Coates 
Branch, a tributary of the Little Sac River. We 
quickly found /soetes butleri, Talinum parviflo- 
rum, and Arenaria patula, species known to be 
associated with Geocarpon at its other stations. 
Then, in a small, moist depression on the glade, 
we spotted it—dwarf, two inches tall, semi-suc- 
culent. At this stage, the plants were wine-purple. 
Once we knew what we were looking for, we began 
to spot another, and another and yet another. Our 
discovery marked the third known location for Geo- 
carpon in the world! On the same glade, I also 
saw for the first time in my life Selenia aurea, 
Saxifraga texana, and Collinsia violacea. 

Our spirits were buoyed by these discoveries. 
We hastened into Green County and stopped at a 
likely looking sandstone glade near Pearl. We no- 
ticed Selenia aurea and Saxifraga texana from 
a distance and, more closely, /soetes butleri. It 
certainly would be just a matter of time. Sure 
enough, the wine-purple plants of Geocarpon min- 
imum became evident after we had wandered into 
an adjacent glade. I don't recall how many spec- 
imens there were but I would think there were 
dozens of them 

s we were hopping over the glade in our 
crouched positions, we happened upon a small plant 
previously unknown from Missouri. It was Scler- 
anthus annuus, the awlwort of the Caryophylla- 
ceae. 


646 


Annals of the 
Missouri Botanical Garden 


FIGURE 6. 


Julian Steyermark. Date unknown. 


With our success in finding Geocarpon, we 
headed into Dade and Cedar counties, stopping at 
several glades. Much to consternation, we 
couldn't find Geocarpon at any of them. By late 
afternoon, 


our 


we started back to the east where we 
had planned to camp at beautiful Alley Springs. 
Darkness caught us again while still on the rolling 
hills of the Ozarks. The night scarcely slowed 
Dr. Steyermark as he continued to name roadside 
plants as our headlights struck them. 
gon elliottii, 


" Andropo- 
' he shouted once, as we whizzed by 
a clump of two-foot tall grasses. Most people have 
trouble identifying this species in the daylight with 
the specimen in front of them, but Steyermark's 
identification was right on the money. 

We got to Alley Springs at last. The long, eventful 
day was conducive to good sleeping, and I scarcely 
noted that I was spending the first night of my life 
atop a hard, slatted picnic table! 

Next morning we were soon on the road, finding 
more goodies. About two miles west of Piedmont 
along Highway 34, we found Cerastium brach y- 
petalum, the third collection of it in Missouri. Near 
Cape Girardeau, we discovered our second station 
for Cerastium pumilum. Since the afternoon was 
young, Dr. Steyermark expressed a desire to go 


into Perry County to a deep ravine he had had his 
eyes on for years. The ravine was new collecting 
ground for Julian, and he took his plant press with 
him. He is the only botanist I have ever been in 
the field with in the United States who carries his 
press with him. Most of us use plastic bags or tin 
cans (vascula), and our semiwilted specimens are 
proof of it. Steyermark had custom-made his own 
press with a hinged back that opens like a giant 
book. Our visit to the ravine netted nearly a dozen 
records for Perry County. 

We got back to Carbondale early in the evening, 
finishing a three-day red-letter trip I will never 

orget.— Robert Mohlenbrock, Dept. of Botany, 
Southern lllinois University; Ain with per- 
mission from Missouriensis 1(3): 


My first meeting with Julian Steyermark was at 
the Caracas Botanical Garden in 1960, after re- 
turning from landscape architecture studies at the 

hode Island School of Design. At that time, there 
were very few books in Spanish about local orna- 
mental plants. As a landscape architect, | felt it 
would be interesting to write a book about tropical 
trees that would be useful for landscape projects. 
Gradually, my visits to the Botanical Garden be- 
came more frequent. Julian was always ready to 
answer my many questions about trees and other 
plants, as well as to identify living material that | 
brought in. 

In 1963, we made a three-day expedition to the 
mountains above the coastal town of Naiguatá in 
a very rich rainforest at 1,000 m elevation and 
only about 60 km by road from Caracas. We found 
many new species there including huge trees, and 
we collected the bromeliad Guzmania sanguinea 
for the first time in Venezuela. At later dates we 
went on other trips, and even met at the top of 
different tepuis with a group from the Terramar 
Foundation. 

n 1962, when I began to write and illustrate 
a color edition of ornamental trees in Spanish, 
Julian kindly supervised my work. We also main- 
tained close contact over the next eight years as 
I worked on my second book called Garden Plants 
of the Tropics (printed in English and Spanish). 
Julian wrote the introduction for me. 

Despite his intensive work in families such as 
the Rubiaceae, Julian was always intrigued with 
the beauty and wonderful colors of the Bromeli- 
aceae. The enchanting forms and the extravagance 
of the flowers fascinated him, something we cer- 
tainly shared. I began to fancy the idea of making 
a book on Bromeliaceae along with him. The first 
step was to compile photographs, including ones 


Volume 76, Number 3 
1989 


Steyermark Recollections 647 


Julian occasionally took of the Bromeliaceae on 
the tepuis. Because of the difficulties of getting to 
the Guayana, his pictures were particularly valu- 
able. Soon he agreed to write the text for the book 
on Bromeliaceae of Venezuela with me. It took 
about five or six years to organize the text and 
pictures. From 1980 to 1983, Julian was already 
very involved with the Flora of the Venezuelan 
Guayana project, so I decided to do most of the 
text in order to facilitate his work. I would generally 
give him eight to ten pages of text to check and 
translate; he would review these and return them 
to me after a few days. He was very adept at 
translating text from Spanish to English and vice 
versa. Once, Julian became upset with me because, 
according to him, I was including too many species, 
and he had no more time to keep reading, cor- 
recting, and translating. Although there were about 
230 species prepared, I went on against his wishes 
to include a few more that I had collected at the 
last minute. I insisted that he go over these too. 
He always wanted to please people, so he did the 
final effort to finish the last translation and cor- 
rections for our work. I can still remember hearing 
him along the corridors of the Botanical Institute 
saying, “I am very busy with the Flora of the 
Guayana and have no time to do anything else. 
Please can you stop adding new data to the book? 
Our manuscript was handed to the printer just dio 
before Julian left Caracas to move to St. Louis, but 
then the book took about two years to be printed. 
Before leaving his post at the Caracas Botanical 
Garden, Julian was awarded diplomas and medals 
together with a reception and a farewell party, by 
the National Parks Institute in recognition for his 
tremendous work accomplished in Venezuela. He 
was most happy and pleased by these honors. After 
Julian's departure from Venezuela, we kept in touch 
by mail about the last details and corrections of 
the proofs of our book. He returned many times 
to Caracas with his assistant Bruce Holst to work 
and collect for his monumental Flora of the Ven- 
ezuelan Guayana. Although he was expecting to 
finish this project, destiny had other plans.— Fran- 


cisco Oliva-Esteva, Caracas, Venezuela. 


Libro de la Sabiduria de Salomon: 


Mas las almas de los justos están en la mano de 


acia, y como un aniquilamiento su partida 

de entre nosotros: mas ellos, a la verdad, reposan en 
paz. 

Julián Steyermark no está hoy entre nosotros, pero 

no podemos decir que está muerto, no podemos 


decir que ha desaparecido; no existe la muerte para 
un hombre que hizo tantos caminos. Su obra como 
su bondad son imperecederas. 

Su alma se separa de su cuerpo ya inmóvil el 
sabado 15 de octubre, allá lejos, en tierras norteñas, 
en la ciudad de Saint Louis. Allá mismo donde nació. 
Pero la mayor parte de su vida, los mejores años, 
los pasa con nosotros. 

o ha habido hombre que haya estudiado más 
nuestra flora. La pasión máxima de Julián Stey- 
ermark fue la Guayana, con sus imponentes te- 
puyes, pero también recorrió con sus ojos bien 
abiertos a toda Venezuela, viendo y colectando 
muestras botánicas. 

Deja una valiosa obra. Era infatigable. Fueron 
múltiples sus contribuciones: las Rubiáceas, las Pi- 
peráceas, exploraciones botánicas de las regiones 
orientales, nuestra vegetación, la Sierra [mataca y 
altiplanicie Nuria, la flora de la Sierra de San Luis 
en Falcón, el Cerro Autana, Cima del Macizo Jaua, 
Cerro Patao en Paria, el Roraima, el Auyantepuy, 
la flora del Avila. 

Julián Steyermark fue un excelente botánico. 
Un incansable admirador de nuestra naturaleza. 
Coleccionó miles de muestras para herbario. Clasi- 
ficó una cantidad igual. Atendió centenares de con- 
sultas. Pero lo más importante es que fue un hombre 
bueno. Honesto, noble, muy sencillo, extremada- 
mente sencillo, cordial, amable, siempre dispuesto 
a responder cualquier pregunta. El gran comple- 
mento de su erudición era la sencillez. 

No podemos decir que Julian Steyermark ha 
muerto. Tal como está escrito en la Biblia: las almas 
de los justos están en la mano de Dios, y no llegará 
a ellos el tormento de la muerte eterna. Ellos a la 
verdad, reposan en Mauricio Ramia, Uni- 
versidad Central de Venezuela; palabras pro- 
nunciadas en el acto de apertura del IX Congreso 
Venezolano de Botánica, el día 13 de noviembre 
de 1988 en Caracas. 


Julian took my wife and me for our first look 
at the Grand Savanna in the Guayana Highland, 
and with us the Dunstervilles looking for orchids 
and to whom the place was an old story. As we 
entered the Grand Savanna, Julian and I stopped 
on an area of dry open sand and discovered a tiny 
sundew with one or two sessile flowers in the center 
of the rosette. Julian named it Drosera felix be- 
cause of the happy event of our visit. 

Later on our trip we were camping one night 
by a small stream, and while Julian was preparing 
supper, I went out exploring and found an aquatic 
aroid in a deep pool that the stream had hollowed 
out of the solid rock. Julian came when he was 


648 


Annals of th 
Missouri ES Garden 


free and with his longer arm was able to collect it. 
It proved to be a new genus, and George Bunting 
named it Jasarum, compounding it with Julian's 
initials and Arum because Julian had discovered 
so many new genera that such tactics were nec- 
essary. Later I contributed with the bromeliad Ste y- 
erbromelia.— Lyman B. Smith, Smithsonian In- 
stitution. 


I participated in a number of field trips to Mis- 
Jr. Steyermark was accumulating 
vouchers for his flora. The average trip down [from 
Chicago] was about six hours, and he would con- 
tinuously quiz me on botanical questions. 

Typical questions would be: “What type of pla- 
centation occurs in the Caryophyllaceae?" or “What 
monotypic genus in the Cruciferae is endemic to 
the Kerguelen Archipelago?" or “Name all of the 
phanerogamic genera native to Antarctica." In 
many cases he was actually thinking about the 
second question while | was working on the answer 
to the first one. So you get some idea how many 
questions were involved on a given trip. 

would say that he, perhaps more than any 
other person, stimulated my interest in taxonomic 
botany.— Floyd Swink, Morton Arboretum. 


Julian knew Missouri as did few others, and he 
had a remarkable memory of people, sites, and 
plants. The Department of Conservation bought 
the land where Julian found pondberry [ Lindera 
melissifolia] from a man called O. K. Hamlett. 

e were discussing this site one day and Julian 
not only remembered O.K.'s name but he described 
in detail where several of the rare species were 
located on the property. I had been on the property 
a week earlier and Julian talked as if he had — not 
25 years before. 

When the Natural History Section first started 
to locate and buy significant sites (often with rare 
or endangered species) we used a list Julian had 
prepared 20 years before. The same list went into 
the Natural Heritage Database prepared by The 
Nature Conservancy and the Department of Con- 
servation. 

Thanks to Julian, a number of these areas are 
now State Natural Areas and protected for those 
who will come after us. Sand Ponds (the pondberry 
site), Pickle Springs, and Barn Hollow are exam- 
ples. 

Every time we walk in these places Julian's shade 
will be there too.— John Wylie, Retired Natural 
History Officer, Jeflerson City, Missouri. 


Members and friends of the Garden were sad- 
dened by the death of Dr. Julian Steyermark on 
Saturday, October 15, 1988. Steyermark was one 
of the Garden's best-known research botanists, 
famous locally and around the world for his bo- 
tanical collections and publications, and for his 
strong support for conservation of the environment. 

Born in St. Louis on January 27, 1909, Julian 
Alfred Steyermark enrolled in 1925 in the Henry 
Shaw School of B 
between the Garden and Washington University, 
where he received A.B. (1929), M.S. (1930), and 
Ph.D. (1933) degrees. He also earned an 
degree (1931) from Harvard University. During 
his early years at the Garden, Julian indulged his 
other great love, painting watercolors of native flora 
and plants in the Garden's orchid collection. At 
that time he began collecting herbarium specimens 
in Missouri and other states, and by graduation in 
1933 he had already surpassed the quantity most 
botanists collect in a lifetime. 

After college, Steyermark assisted Robert 
Woodson, a research botanist at the Garden. This 
led to his first tropical expedition, a collecting trip 
to Panama in 1934. He then taught high school 
in the St. Louis area for two years before moving 
to the Field Museum of Natural History in Chicago 
as an Assistant Curator. For the next 21 years 
Steyermark worked on the Flora of Guatemala 
and other tropical projects during the week, but 
continued his Missouri studies in his spare time. A 
revealing and delightful look at those weekend ex- 
peditions from Chicago to the wilds of Missouri was 
provided by his wife, Cora, in her book, Behind 
the Scenes, published by the Garden in 1984 (and 
still available in the Garden Gate Shop). 
efforts culminated in the massive Flora of Missouri 

3). This widely acclaimed manual is often re- 
ferred to as the state's “botanical bible" and has 
sold out six printings. 

During World War II, Steyermark was sent to 
South America as part of the government's pro- 


otany, a cooperative program 


These 


gram to locate new botanical sources for quinine. 
At that time he first visited the Venezuelan 
yana region, which he lovingly referred to as *“The 
Lost World.” (See the Missouri Botanical Garden's 
Bulletin, November /December 1986 and July, 
August 1987.) His interest in this picturesque area 
was so great that in 1958 he left the Field Museum 
to take a position at Instituto Botánico in Caracas, 
, Julian returned to the Mis- 


Gua- 


Venezuela. In 
souri Botanical Garden, where he strove to com- 
plete his last major project, a flora of the more 


than 9,000 plant species in the Venezuelan Guay- 


Volume 76, Number 3 
1989 


Steyermark Recollections 


649 


FIGURE 7. Cora and Julian Steyermark in front of their home at Santa Eduvigis, Caracas, Venezuela, 1965- 


~ 


1970 


ana. This task remains for his friends and col- 
leagues to complete. 

Bruce Holst, project coordinator for the Flora 
of the Venezuelan Guayana, said, “It was a real 
thrill and a tremendous learning experience to work 
in the field with Julian. He really had an “eagle 
eye' for spotting plants, and if you didn't see some- 


ET = 


FIGURE 8. 


thing that he did, he would let you know it . 
ieldwork was very special for Julian. He never 
took any ‘vacation,’ he went on field trips instead. 
He was just so excited about collecting plants, it 
was impossible not to be caught up by his enthu- 
siasm. He always likened collecting to a ‘treasure 
hunt’! Even with his two artificial hips, he would 


Julian Steyermark on El Avila, outside of UB. Venezuela. Date unknown. 


650 


Annals of the 
Missouri Botanical Garden 


plow through brush, crawl over and under logs, 
fall down, laugh about it and push on....” 
Steyermark received numerous honors during 
his eventful life, including awards and citations 
from the governments of Guatamala and Vene- 
zuela, and the 
1979 


Sierra Club awarded him a special citation for his 


ardens Henry Shaw Medal 
That same year, the Ozark Chapter of the 


conservation efforts, and the Missouri Department 
of Conservation named the 73-acre Julian A. Stey- 


ermark Woods, near Hannibal, in his honor. 

To commemorate Steyermark's achievements, 
the Garden has established a fund to promote bo- 
tanical research and expeditions in Latin America, 
particularly in the Venezuelan Guayana region, and 
to support research on the natural history of Mis- 
souri. In addition, the Missouri Department of Con- 
servation and the Garden are revising Julian's best- 
known book, which will be retitled Steyermark's 
Flora of Missouri, a lasting memorial to a giant 
among botanists. 


FIGURE 9. 


(holding papers). 


A memorial service was held for Julian Stey- 
ermark at the Garden on October 25, 1988. In 
the eulogy, Dr. Raven said, . His last work will 
be completed here, and will stand as a final tes- 
tament to his industry and to his scholarship. As 
for any human being, however, what will prove to 
be even more important to all of us will be our 
memories, for Julian Steyermark was an inspira- 
tional person, one who projected his own values 
without trumpeting them, and thereby enriched all 
our lives, as he did the literature of botany.” 
George Yatskievych heads the Garden's joint 
project with the Missouri Department of Con- 
servation to revise Steyermark's Flora of Mis- 
souri.— Luther Raechal assisted Dr. Steyermark 
on the Flora of the Venezuelan Guayana and is 
continuing his work on the staff of this ongoing 
project; reprinted with permission from the Bul- 
letin of the Missouri Botanical Garden 77 (1): 5 
1989. 


: a 4 
M. 


e 
V Im 


Geoffrey C. K. Dunsterville (fourth from left); Nora Dunsterville (sitting in front); Julian Steyermark 


Volume 76, Number 3 Steyermark Recollections 651 


FicunE 10. ora and Julian Steyermark on their wedding day, 1 September 1937, in Hannibal, Missouri. Photo 
contributed by aa Lippold. 


FIGURE 11. “A dream come true, but Mr. Peterson is responsible for picking it. The rain had broken the stem, 
and it was no crime, therefore, for Mr. Peterson to take the flower to paint," Steyermark wrote on the back of this 
photograph, dated 5 May 1929. 


FIGURE 12. Julian Steyermark. 


PLANT TAXA DESCRIBED BY 
JULIAN A. STEYERMARK' 


Mary Susan Taylor? 


ABSTRACT 


In his 62 years of actively collecting plants (1925-1987), Julian Steyermark described 2,392 taxa, including 1 


family, 38 genera, an species. He named plants 


ountries and from 16 states in the U.S.A. Types 


om 26 c 


s fr 
of taxa that he named are listed alphabetically by family, with author, literature citation, collector, collection number, 


date of collection, locality, elevation, and depo 
names, SAS names, and some 
include 

are oe ed. 


sitory. 


comme 
current nomenclature (which are not referenced in the bibliography). Appendices 
nymy, collectors index, and family index. The bibliography includes 395 references. Statistical summaries 


nts are supplied, where pertinent, on never-published 


Julian Steyermark collected his first plant in 
1925, his last in 1987, both in Missouri. He col- 
lected so many plants, over 130,000 collections, 
that he received recognition from the unusual source 
of the Guinness Book of World Records. Julian 
was most known for his Flora of Missouri, and his 
work in Guatemala and Venezuela, yet he described 
plants from 26 countries and from 16 states in the 

His statistical record is astonishing. Julian Stey- 
ermark described a single family, 38 genera, and 
1,864 species. Including other ranks, he named at 
least 2,392 taxa. Taxonomic Literature (second 
edition) cites an as yet unobtainable reference 
edited by H. Villanueva, 1977 (Bibl. Alfredo Jahn, 
Bol. Inf. 9: 9-12) which purports to contain the 
‘most important of over 350 publications’ by Julian 
Steyermark. It is estimated by the present author 
that this compilation lacks about 80 references, 
although the exact number is unknown. 

In the list of types families are arranged alpha- 
betically, and species alphabetically within families, 
then by descending rank within species. Data in- 
clude the name, author(s), place of publication, 
collector, collection number, date of collection, lo- 
cality, elevation, and depository. Current nomen- 
clature (which are not referenced in the bibliog- 


chiche, Quebrada 
Jac olini umbrosa (Benth.) S. F. Blake 


raphy) and accession number of the Missouri 
Botanical Garden sheet are indicated when found 
serendipitously. New combinations are not includ- 
ed. A few never-published names are included and 
correct nomenclature given for sheets known to 
have been distributed bearing the unpublished name. 
Author abbreviations follow Draft Index of Author 
Abbreviations Compiled at The Herbarium, Roy- 
al Botanic Gardens, Kew and Taxonomic Liter- 
ature (second edition). Publication abbreviations 
are standardized according to Botanicum-Perio- 
dicum-Huntianum. 


STEYERMARK TYPES 
ACANTHACEAE 


Aphelandra e Lindl. f. lutea Standl. & Stey- 
erm., Field. Mus. . Hist., Bot. Ser. 23(5): ah 
1947. P. C. Ds 89560; 16 Mar. 1 941; ( 
temala, Dept. Escuintla, Río Gavilán, NE of Escuintla. 
720 m; F (holo) = 4. gigantiflora Lindl. var. gigan- 
tiflora, 
Bravaisia integerrima (Spreng.) Standl. var. pilosa Stey- 
. 1984. Steyer dect et al. 
198 1; Vene vuelo. Yarac , Dept. 
qur 50m; VEN (i job: 
f. erythrantha 
Standl. 2 n ist. Field. uS us. Nat. Hist., Bot. Ser 
23(5) 2 . P. C. Standley 60082; 15 De. 
1938; opina Dept. Chimaltenango, above Las 


uum 


' Julian never kept a list of as he described or maintained a HCM bibliography. He was always helpful with 
d 


suggestions on where to look for more of his 
seek me out to ask if I had lo 


about his life and his wor 


I spent re on the 


list 
? Missouri Botanical Carden: P.O. Box 299, St. Louis, 


work, and soon I didn't e to prompt 
died at certain journals. I shall i bo his generous gift of time in talking 
was a is er ue dan and gracious with advic e when 


o Bruce Holst, Paul Ber nd Lu 


u, Myra Guzman- Teare, and ne Pool gave me support and encouragement during the two years 


im with questions, he would 


e Holst, 


Missouri 63166, U.S.A. 


ANN. Missouni Bor. GARD. 76: 652-780. 1989. 


Volume 76, Number 3 Taylor 653 
989 Plants Described by 
Julian A. Steyermark 
TABLE l. Statistics. Numbers of Steyermark taxa TABLE 2. Family and genera described by Julian Stey- 
named within each rank, and within each country and — ermark. 
state. 
FAMILY Tepuianthaceae 
TARA ENER 
G A 
Families: l Genera: 3 del de 
Tribes: 1 Species: 1.864 C ecd steraceae 
00 : Acopanea Theaceae 
Sections: 10 Subspecies 54 
M : Prin d Ochnaceae 
Subsections: 4 Varieties 241 . : 
i Ahozolia = Sechium Cucurbitaceae 
Series 5 Forms: 175 : 
: a Aphanocarpus Rubiaceae 
Subgenera 3 New names 33 m 
bri » Aratitiyopea Xyridaceae 
Bartholomaea Flacourtiaceae 
Total taxa: 2,392 known to date Bonifazia = Disocactus Cactaceae 
Publications: 379 known to date Brewcaria Bromeliaceae 
Epo : 229 known to date Cardonaea Asteraceae 
COUNTRIES FROM WHICH TAXA WERE NAMED se phalodandron Rubiaceae 
, Chimantaea Asteraceae 
Argentina l Belize 6 rococcus = Coccochondra Rubiaceae 
"enis : im » p Cory adds Rubiaceae 
ARMS na: Croizatia Euphorbiaceae 
Costa Rica: 1 Cuba: 2 Cuatrecasasiodendron Rubiaceae 
Dominica: 1 aes i oum l lgrenia ata iid 
Ecuador: 23 El Salva 2 Dioicodendron Rubiaceae 
Il Guiana: 19 Cantera i" Holsitänthts Ruhine 
;uvana onduras 
) EN v 
H - 
pa E Panama z Maguireocharis Rubiaceae 
aragul teen Maguireothamnus Rubiaceae 
pS Rico: 1 South Africa: 1 | Rubiaceae 
E S POPMA - Trinidad: 4 Mortoniodendron Tiliaceae 
U.S.A.: 124 Venezuela: 1,420 Neblinal AA 
STATES IN THE U.S.A. FROM WHICH TAXA WERE NAMED Orthion Violaceae 
amia: i. Arkansas: 2 Pachecoa Fabaceae 
Arkona: V Colorado: 3 Pagameopsis Rubiaceae 
Califonia: 28 Illinois: 3 Perissocarpa Ochnaceae 
daho: € Missoüri: 62 Pittierothamnus = Amphidasya Rubiaceae 
anti + Ohio: 1 Rojasianthe Asteraceae 
New Mexica 9 "unnésséi 1 blina = Rutaneblinaria Rutaceae 
Oregon: 2 Utah 1 Senefelderopsis Euphorbiaceae 
oa 13 Washington: 1 Sipaneopsis Rubiaceae 


Calderas; 1,800-2,100 m; F (holo) = Justicia aurea 
Schlecht. f. erythrantha (Standl. & Steyerm.) D. Gib- 
son, 1973. 

Poikilacanthus dg p Standl. & WA Field Mus 
Nat. Hist., Bot 23(5) 246. 1947. H. Linen 
523; 15 Aug. FA Guatemala, Dept. Alta Verapaz, 
between Chamá and Cobán; 900 m; F (holo). 

Ruellia cr (Walt.) Steudel f. alba Steyerm., 
Rhodor : 585. A. Steyermark 6058; 23 
June n U. S.A., Missouri, Caldwell Co., along Long 

reek, 1.5 mi. S af Kingston; F (holo), MO (1174668) 
= - R. humilis Nutt. var. Md (A. Gray) Fern. f 
alba (Steyerm.) Steyerm. 

Ruellia pedunculata Torr. f. que ri Steyerm., Rhodora 
51: 118. 1949. B. Bauer s.n.; 20 June 1948; U.S.A., 
Missouri, St. Francois wr 

Ruellia strepens L. f. a y E 

2. J. A. dons: 70292; 26 Aug. 1950; U.S.A., 
Missouri, Schuyler Co., tributary to Chariton River, NE 


Tecunumania Cucurbitaceae 
Tepuianthus 


Tepuianthaceae 
Thymocarpus 'ae 


larantaceae 
Vesicarex Cyperaceae 


Yutajea Rubiaceae 


of Livonia, 6 mi. SW of Glenwood; F (holo), MO 
(1703638). 

taa hondurensis Standl. & Steyerm., Ceiba 

191, 1952. P. C. Standley s 18 d 1948; 
Honduras, Dept. Morazán, región é sla, cerca 
de Suyapa; F (holo) = CON MM e ginis (Jacq.) 
D. Gibson, 1973. 
ACTINIDIACEAE 

Saurauia ei cog uei Standl. & Steyerm., Field 
Mus. Nat. Hist., Bot. 23(5) 215. 1947. J. 4. 
Steyermark 49810; ^w p 1942; Guatemala, Dept. 


Huehuetenango, Cruz de Limón, Sierra de los Cuchu- 


matanes; 2,600-3,000 m; F (holo). 


Annals of the 
Missouri Botanical Garden 


Saurauia meridensis Steyerm., Fieldiana Bot. 28(2): 368. 
1952. J. A. Steyermark 56344; 6 May 1944; Ven- 
suele: Mérida, Río Canagua, near Canagua; 1,065- 
1,820 m; F (holo). 

Saurauia “perseifolia”” Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 216. 1947. J. A. Steyer- 
mark 41748; 24 Dec. 1941; Guatemala, Dept. Izabal, 
along Río Tamejá, Cerro San Gil; 50 m; F (holo). Note: 
S. persifolia is the correct spelling. 

"aurae veneficorum Standl. & d 

Hist 


Bot. Ser. 23(5): 217. 1947. J. . 


F me Mus. 


qui imula, Montana Norte to El Taa. Cerro Brujo; 1,700- 
2,000 m; F (holo). 


AGAVACEAE 
Agave d a Standl. & Steyerm., Field Mus 
Hist., Es r. 23(1): 4. 1943. P. C. Standley 82030; 
2 Jan B le Dept. Huehuetenango, along 


a road E of B ee 1,950 m; F 
(holo). 


ALBOLBODACEAE — Xyridaceae 
AMARANTHACEAE 


C ‘ladothrix lanuginosa Nutt. var. ae Steyerm., Ann. 
Missouri Bot. Gard. 19: 389. . J. A. Moore & J. 
E s m 3795; 26 le iu 

3 mi. E of Study Butte; 762 m; GH 
i "MO veneers Tidestromia lanuginosa var. 
caruosa (Steyerm.) Cory. 


AMARYLLIDACEAE 


is ratium m Standl. & Pug Field Mus 
t. Hist., Bot. S . 4. ossium: 
39288; 13 Apr. "1940; ped Dept. zabal, be- 
tween Escobas and Montaria Escobas, across bay from 
Puerto Barrios; sea level; F (holo). 


ANACARDIACEAE 


d a ipiis ille "orum pue iin & Wurd., 
Bol. Soc. Ven . 26: 159. 1965. J. A. Stey- 
BRE 90883: 4 dune 1962; Venezuela, Miranda, 
Oritopo, above Los Guayabitos did Baruta; 1,500 m; 
VEN (holo). 


ANNONACEAE 


4nnona primigenia Standl. E us Field Mus. Nat. 
Hi . C. L. Lundell 2213; 
Mar. 1933; oben. im Para. Gavilán, Falla- 
bón-Yaxhá road; F e 


Desmopsis 


l. & Steyerm., Pg Mus. 

Nat. Hist ., Bot. Ee. 233) 156. 1944. J. A. Steyer 

es 34304 ; 17 Jan. 1940; Guatemala, Dept. Que- 
zaltenango, Montaña a Volcán Santa María; 
1,500 m; F (holo). 

decis s a Standl. & Steyerm., 
Nat , Bot. Ser. 23(4): 157. 1944 
mark 41543 s Dec. 

o Frío, Cerro San Cil; 

Sapranthus onu — " a S Field Mus. 
Nat. Hist., a Ser. 2 . P. C. Standley 
59219; 8 1938; B. ‘Depi. Guatemala, 
Estancia Grande; 600 m; F (holo). 


Field Mus. 


APOCYNACEAE 


AR decussatum Woodson subsp. jauaensis 
Ste ag nos Maguire, Mem. New York Bot. Gard. 23: 
. oJ. ^i Steyermark 98086; Mar. 
1967; Cod dn Bolivar, Cerro Jaua; 1,922-2,100 
m; VEN (holo). 

s dar costanensis Steyerm., Bol. Soc. Moss Ci. 
Nat. : 240. 1964. J. A. Steyermark 92131; 15 
19 Toi 1963; Venezuela, Distr. Federal, m Nai- 
gu 00-1,300 m; NY, US, VEN (holo). 

Mandevilla subcarnosa (Benth.) Yaoi var. angus- 
tata Steyerm. a Bot. Venez. 2: 318. 1967. J. A. 
Stey ermark 93737; ; 11 May 1964; Venezuela, Bolivar, 
Auyan-tepui; 1,660 m; VEN (holo). 

Peltastes manarae Steyerm., Acta Bot. Venez. 3 
1968. B. J. Manara s.n.; 11 July 1966; Venezuela, 
Distr. Federal, a a Guayoyo, Cerro Naiguata; 
1,000 m; VEN (holo 


t. 
22-97 


4 Ai 


AQUIFOLIACEAE 


llex abscondita Steyerm., pee Missouri Bot. Gard. 75: 
320. 19 i 


0. 8 Croat 59478; 30 Nov. 1984; Vene- 
zuela, E Amazonas, a ro i^a la Neblina, Camp 7; 
1,900 m; MO (holo— ei NY, VEN. 


Hex iaa Steyerm., Ann. Missouri Bot. Gard 
75: 321. 1988. ne Tier m 12 Mar. 1985; 
Venezuela; T.F. s, De Cerro 

Marahuaca; 1,500- T 650 m; MO " holo — 2446418) 

VEN. 


Ilex M ei Steyerm., Ann. Missouri Bot. Gard. 75: 
3: 988. J. 4. Steyermark 111402; 4 Jan. 1975; 
Venezuela, Bolívar, a b Ignacio de Yuruani 

and S ancisco de Yur ; 1,200 m; VEN (holo). 
lex ea Standl. & B on. Field Mus. Nat. Hist., 


22 Feb. 7 ] 
Tonana, between Canjula and La Unión Juárez, near 
SE portion of Volcán Tacaná; 2,000-3,000 m; F (holo). 
Ilex aracamuniana Steyerm., Ann. Missouri Bot. Gard.: 
R. 


in press . Liesner & G. Carnevali 22473; 
26 Oct. 1987; Venezuela, . Amazonas, Dept. Rio 
de , Cerro Aracamuni, summit, Proa Camp; 1,400 


O (holo), 

lle ae Steyerm., Ann. Missouri Bot. Gard. 75: 

322. 1988. N. T. Silva & U. Brazao 60873; 24 Jan. 
1966; Brazil. Amazonas. Sierra Pirapucú, Comissão de 
Limites BR-1; 1,250 m; G, INPA, MO (holo— 
3283928), NY, VEN. 

Ilex brevipedicellata Steyerm., Ann. Missouri Bot. Gard 
15: 322. 1988. J. 4. Steyermark & B. Holst 130531; 
21-22, 24 Feb. 1985; Venezuela, T.F. Amazonas, 
Dept. Atabapo, Cerro Marahuaca, Sima Camp; MO 
(holo — 3446417), VEN. 

Ilex brev vipetiolata sa es & Wurd., Mem. nee York 

1957. J. A. Steyer mark 
Wurdack 347; 2 27 ea 948; Venezuela, Boliv r, Ua al- 
pán-tepui; | ,400 m; NY (holo) = /. py eta lla 
Steyerm., 1952 

Ilex Pekin Steyerm., Ann. Missouri Bot. Gard. 
323. 1988. F. Cardona 1480; 16-24 Feb. 1946; 
Venezuela, T.F. Amazonas, orillas del Rio Castanho, 
afluente del Padauiri que del Río Negro y éste ültimo 
al Amazonas; 100-140 m; VEN (holo). 

llex ciliolata Steyerm., Ann. 


88. R. L. 


issouri Bot. Gard. 75: 


Liesner 19479; 4 Nov. 1985; Ven- 


Volume 76, Number 3 
1989 


Taylor 655 
Plants Described by 
Julian A. Steyermark 


ezuela, Bolivar, 0.4 km N of El Pauji on trail to Uaiparú; 
800-900 m; MO (holo — 3446424), VEN. 
llex culmenicola Steyerm., Fieldiana Bot. 28(2): 322. 
1952. G. H. H. Tate 424; 26 Nov.-16 Dec. 1928; 
Venosieli: T.F. Amazonas, Mt. Duida; 1,830-2,000 
m; F, NY (holo). 
Ilex davidsei Steyerm., Ann. Missouri Bot. Gard 75: 324. 
1988. G. Davidse 22719; 23 July 1984; Venezuela, 
Negro, lower part of the Rio 


Ilex glabella Steyerm., . Missouri Bot. Gard. 75: 
: . R. L. Liesner 16688; 15-16 Mar. 1984; 
Venezuela, T.F. Amazonas, Dept. Rio Negro, Cerro de 
la Neblina, Camp 4; 780 m; MO (holo— 3524663), 
VEN. 


Ilex glaucophylla Steyerm., Bol. Soc. Venez. 15: 176. 
1954. L. Aristeguieta 2295; June 1954; Venezuela, 

Miranda. Los Guayabitos; 1,200 m; F (holo), VEN. 

llex gleasoniana Steyerm., Fieldiana Bot. 28(2): 322. 

952. G. 1. Tate 526; 26 Nov.-16 Dec. 1928; 

Venezuela, T F. Amazonas, Ridge 25, Mt. Duida; 2,000 
m; F, NY (holo). 

llex gransabanensis Steyerm., Ann. Missouri Bot. Gard. 
75: 325. 1988. T. Croat 54158a; 26 July 1982; 
oe Bolivar, rd. from Ikabarü to Santa Elena, 

N of Ikabaru; 750 m; MO (holo— 3524662), 

Ilex guaiquinimae Steyerm., Ann. Missouri Bot. Gard. 
15: 325. 1988. J. 4. Steyermark et al. 117276; 24 
May 1978; ml Bolivar, Cerro Guaiquinima; 
900 m; MO, VEN (holo). 

Ilex holstii ji Ann. Missouri Bot. Gard. 75: 326. 
1988. J. A. Steyermark & B. Holst 130481; 21-22, 
24 Feb. 1985; Venezuela, T.F. Amazonas, Dept. Ata- 
bapo, Cerro Marahuaca, Sima Camp; 1,140 m; MO 
(holo — 3446423), VEN. 

Ilex a Steyerm., Ann. Missouri Bot. Gard. 75: 
2 A. Steyermark et al. 130112; 24-28 
Feb. 1984; Venezuela, T.F. Amazonas, Dept. Rio Ne- 
gro, Cerro Aratitiyope, 70 km SSW of Ocamo; 990- 
1,100 m; MO (holo— 3226838), VEN. 

Hex jauaensis Steyerm., Ann. Missouri Bot. Gard. 75: 
327. 1988. J. A. Steyermark et al. 109294; 22-28 
Feb. 1974; Venezuela, Bolívar, Meseta de Jaua, Cerro 
Jaua, SW summit; 1,750-1,800 m; VEN (holo), MO 
(2582368) 

Ilex karuaiana Steyerm., Fieldiana Bot. 28(2): 323. 1952. 

4. Steyermark 60660; 27 Nov. 1944; Venezuela, 


Ilex liesneri Steyerm., Ann 
987. R. L. leer 18469; 8-9 Mar. 
zuela, T.F. Amazonas, Dept. 
huaca, 1-2 km i of Sima Camp; 1,100 m; MO (holo — 
3462076), VE 

Ilex ie er Aa erm., 
327. R. L. Liesner 19878; 
m Bolívar, 3 w q^ of El Pauji, S slope and 
summit of El Abismo; 800-1,050 m; MO (holo— 
3446416), VEN. 

llex magnifructa Edwin var. minor Steyerm., Ann. Mis- 
souri Bot. Gard. 75: 328. 1988. F. Cardona 2943; 
Feb. 1962; Venezuela, T.F. Amazonas, Cerro Ualipano; 
1,760 m; MO, US (holo 

Ilex marahuacae Steyerm., Ann. Missouri Bot. Gard. 75: 
328. 1988. J. 4. Steyermark & B. K. Holst 130425; 
21-22, 24 Feb. 1985; Venezuela, T.F. Amazonas, 


sa Missouri Bot. Gard. 
78; 11 Nov. us 


Cerro Marahuaca, Sima Camp; 1,140 m; MO (holo— 
3524661), VEN. 
llex nervosa Triana var. glabrata Steyerm., Fieldiana 
Bot. 28(2): 324. 1952. J. A. Steyermark 56653; Ven- 
ezuela, Mérida, above mp 2 285- 2, 745 m; F (holo). 
Ilex paruensis Steyerm., Ann : 
330. 1988. O. Huber 4437; 7 
.F. Am pana y Atabapo, a del Parú (Ar 
ko); 1,100 m; VEN (holo). 
Ilex ed Steyerm. , Ann. Missouri Bot. Gar 
330. Ts. 
sta of El Pan 
900 n m; MO (holo— 3524660), VEN. 
Ilex n Steyerm., Ann. Missouri Bot. Gard. 75: 331. 
1988. G. Davidse & 0. Huber 22679; 25-27 Nov. 


28(4): 959. 1957. J. A. Steyermark 58123; 31 Aug. 
194.4; Venezuela, T.F. Amazonas, Cerro Duida; 1,075 
m; F (holo). 
Ilex m Klotzsch f. glaucescens Steyerm., Fieldiana 
E 28(4): 959. 1957. J. A. Steyermark 58349; 4 
1944; Venezuela, T.F. Amazonas, Cerro Duida; 
l, 20- 2,075 m; F (holo). 
e retusa Klotzsch f. major Steyerm., Pesa E. 
28(4): 960. 1957. J. A. Steyermark 58127; 3 
1944; oo T.F. Amazonas, Cerro Duida; T 675 


m; F (ho 
Ilex spathulata Steyerm., Ann. Missouri Bot. Gard. 75: 
331. 1988. R. L. Liesner 17599; 17-18 Feb. 1985; 


azonas, Cerro icm 1,095- 
Y - ge retusa Klot 
s Subrotundifolia pedir! ña Fieldiana Bot. Ya 328. 
2. J. A. Steyermark 5 6 Oct. 1944; Ven 
cal, Bolivar, E (be T ae m; E 
ns ma Steyerm., Ann. Missouri Bot. Gard. 75: 332. 
988 ^ K. Holst 3617; 26 Mar. 1987; Venezuela, 
Bolívar, Dist. Piar, Camarcaibarai- -tepui, summit; 2,400 
; MO (holo— 3446422), V 
Ilex atea Steyerm., Fieldiana Bot. 28(2): 330. 1952. 
J. 


1,520 m; 


A. Steyermark 58022; 26 Aug. 1944; Venezuela, 
T.F. Amazonas, Cerro Duida; 1, 095- 1,520 m; F (holo). 
Ilex tepuiana Steyerm. ex Edwin, Mem. New York Bot. 
Gard. 12(3): 138. 1965. J. 4. Steyermark 60290; 
15-17 Nov. 1944; Venezuela, Bolivar, mesa between 


Ptari-tepui and Sororopan-tepui; 1,600 m; F (holo), 
NY 


llex venezuelensis Steyerm., Fieldiana Bot. 28(2): 330. 
95 . A. Steyermark 58340; 4 Sep. 1944; Ven- 
ezuela, T.F. Amazonas, Cerro Duida; 1,820-2,075 m; 


F (holo). 

Ilex vesparum Steyerm., iom Bot. 28(2): 332. 1952. 
J. A. Steyermark 61659; 21 Mar. 1945; Venezuela, 
Anzoátegui, above Santa e 2,000-2,300 m; F (holo). 

Ilex wurdackiana Steyerm., Ann. Missouri Bot. Gard 
75: 332. 1988. J. A. Steyermark & S. Nilsson 449; 
2] Apr. 1960; Venezuela, Bolivar, Cerro Venamo, 
slopes; 1,100-1,140 m; VEN (holo). 


656 


Annals of the 
Missouri Botanical Garden 


ARACEAE 


4nthurium Ed Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 209. 1947. J. A. Steyermark 
42212; 9 oni 1949; Guatemala, Dept. Zacapa, Río 
Sitio Nuevo, Sierra de las Minas; 1,200-1,500 m; F 
(holo) = A. subcordatum Schott, 1958 

Anthurium duidae Steyerm., Fieldiana Bot. 28(1): 92. 
1951. J. A. Steyermark 58099; 31 Aug. 1944; Ver 
ezuela, T.F. Amazonas, Cerro Duis 1,675 m; F (holo) 

— A. ptarianum Steyerm., 1979. 
Peru krauseanum Steyerm., A Bol; 28(1): 
5 A. Steyermark 60847; 4-5 Dec. 1944; 
Venezuela, Bolivar: SE base of Co -tepul; 1,460- 
1,615 m; F (holo). 

Anthurium mansteroidi s Steyerm., Fieldiana Bot. 28(1): 
93. 1951. J Steyermark 56727; 23 May 1944; 
Venezuela, Bolivar, N of Mesa Bolivar, along Rio Onia; 
pun "m m; F (holo) = 4. clavigerum Poepp. & Endl., 


n ptarianum Steyerm., Fieldiana Bot. 28(1): 
94. 1951. Steyermark 59509; 29 Oct. 1944; 
Venezuela, Davit- Ptari- -tepui; 1,810 m; F (holo). 

Anthurium radic yas er & Steyerm., Field Mus. 
Nat. Hist., pa Ser. 23(5): 210. 1947. J. A. Steyer- 
mark 44776; 6 Mar. 1942; Guatemala, Dept. Alta 
Verapaz, ab Río levolay, N and NW of Finca Cu- 
bilquitz to Quebrada Diablo; 300-350 m; F (holo) — 
A. pittieri Engler, 1958. 

Anthurium retiferum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 211. 1947 Steyermark 
48021; 14 July 1942; Guatemala, Dept. Huehuete- 
nango, iuro de los Cuc aate Cerro Huitz; 1,500 
2,600 m; F (holo). 

pe ne a titanium Standl. & EE Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 211. ak lcd 
37963; 16 Mar. 1940; ama. Dept. S . San Marc 
Volcán Tajumulco along Rio Cabüs; 1,300- 1.500 1 m; 
F (holo). 

Monstera di re pee & Power Field Mus. 
Nat. Hist., Bot. 23(5) 212. 1947. Steyer- 
mark 45423; AS Mar. 1942; Cuatemala, Dept. Petén, 
m Finca Yalpemech and Chinajá; 50-100 m; F 
(ho 

Philodendron duidae To pes Bot. 28(1): 9 
1951. J. eyermark 5 . 1944; Ven. 
e Mie T.F. der mE im E RS 25-1,200 m; 
F (holo) = P. englerianum Steyerm. subsp. duidae 
(Steyerm.) Buntin 18, 1988. 

Philodendron engle rianum e erm., E Bot. Tr 

* Lave ME A: ırk 6086 


> 


Venezuela, Bolivar, Cart ao-tepui; i: (holo A 

Philodendron krauseanum Steyerm., pape Bot. 28(1): 

.1 . Steyermark 60. 25 Nov. 1944; 
Venezuela, Bolivar, Quebrada O- n ma; 915-1,065 
m; F (holo). 

o rco dig a Sie yerni ., Fieldiana Bot. 28(1): 
99. A ao 59682; 1 Nov. 1944; 
eu Bolivar. Prari-te ; 1,600 m; F (holo) = 
P. callosum subsp. ois inner Bunting, 
1988. 

Philodendron rojasianum Standl. & Steyerm., Field Mus 
Nat. Hist., Bot. Ser. 23(1): 3. 1940. J. 4. Sieverinark 
33244; Dec. 1939; Guatemala. Dept. Quezaltenango, 
Finca Pirineos, Volcán Santa Maria; D 1,500 m; 


F (holo) = P. guatemalense Engler, 1 
cero williamsii Steyerm., Fieldiana Bot. 28(1): 
9 . L. Williams 13922; 23 Jan. 1942. Ven- 


ezuela, T.F. Amazonas, Yavita; 128 m; F (holo) = 
Se hismatoglotiis spruceana (Schott) Bunting var. wil- 
liamsii (Steyerm.) Bunting 8. 

Sc pénis in Crete Bunting & Steyerm., Brit- 
tonia 21(2): 187. 1969 . Steyermark 97811; 17 
29 Mar. 1967; Veneriala; Boliv ar, del Rio Kanarakuni, 
N and NE of Mision de Campamento Sanidad del Rio 
Kanarakuni; 400 m; NY, VEN (holo). 

m arpa duidae Steyerm., Fieldiana Bot. 28(1): 101. 

1951. J. A. Steyermark 57994; 25-26 Aug. 1944; 
NBN T.F. Amazonas, Caño Negro, Cerro Duida; 
F (holo). 

Stenospermation ammiticum Bunting, Bol. Soc. Venez. 
Ci. Nat. 25: 42. 1963. J. 4. Steyermark 89543; 27 
Aug. 1961; Venezuela, Bolívar, cabeceras de Río Chi- 
canan; 900 m; BH, ien (holo). Note: This is the 
accepted spelling of the genus. 

Stenospermatium pittiert Steyerm., Fieldiana Bot. 28(1): 

51 yermark 57455; 18 July 1944; 
Venezuela, Táchira, Alto de Lirio, batwan Bramon and 
Las Delicias; F (holo). Note: Stenospermation is the 
correct spelling for the genus. 

Urospatha savannarum RAE Fieldiana Bot. 28(1): 
102. 1951. J. A. Steyermark 57853; 23 Aug. 1944; 
Venezuela, T.F. Amazonas, Eus of Cerro Duida, Sa 
bana Grande; 200 m; F (holo) = U. angustiloba Engl., 

1988. 


ARALIACEAE 
Dendropanax quw Steyerm., Fieldiana Bot. 28(2): 
441. 1952. J. 4. Steyermark 55434; 11 Feb. 1944; 


Venezuela, Lara, Me Ee Buenos Aires and Páramo 


as Rosas; 2,285-3,290 m; F (holo) 
Dendropanax neblinae ur e, Steyerm. & Frodin, 
Mem. New York Bot. Gard. 38: 82. 1984. B. Maguire 


. 42062; 
Amazonas, just S o 
m; MO (3401817), NY 
Dendropanax veillonii SHAA Pittieria 
J.- ; 1971; Venezuela, Mérida, San Eusebio, 
2,350 m; VEN (holo). 
a eae ti auyantepuiensis Steyerm., Acta Bot. 
Vene 271. 1967. J. A. Steyermark 93974; 15 
May 1964; Venezuela, Bo 


14- ie Nov. 1957; Venezuela, T.F. 
amp i ead de la Neblina; 700 


E 


lívar, Auyan-tepui; 2,050 
2,300 m; VEN (holo) = Schefflera MUT O 
(Steyerm.) Maguire, Steyerm. & Frodin, 

Didy mopanax chimantensis Steyerm. & Nonis Mem. 
New y 


Gard. 17(1): 456. 1967. J. A. Ste 


ermark 75788 953; V ARE Bilan 
Apacara-tepui, C himantá Massif; 2 2,000-2 2,150 
(hola = Schefflera chi Maguire) 


Maguire, Steyerm. & Frodin, 1984. 

o eco , Acta Bot. Venez. 
2: 273. 1967. J. A. Stevermark 9. 3926; 15 May 1964; 
edid Bolivar, Auyan- -tepui; 2,050-2,300 m; VEN 
(holo) = Schefflera Pee nsis (Steyerm. & Maguire) 
Maguire, Steyerm. & Fro 

aa capitatus (Jacq.) Decne. & Planch. var. mi- 
nor Steyerm., ot. 28(2): 442. 1952. J. A. 
Steyermark 56242; 3 May 1944; srl: Mérida, 
between La Cumbre, San Jose and Mucutuy; 1,820 
2,590 m; F (holo). 

Oreopanax venezuelense Steyerm., A Bot. Ys 2) 
4 A. Steyermark 62017; 11 Apr. 1945; 
Venezuela, Monagas, Cerro En. l, 100- 1 300 
m; F (holo). 

Schefflera ayangannensis Maguire, Steyerm. & Frodin, 


~ 
~ 


Volume 76, Number 3 
1989 


Taylor 657 
Plants Described by 


Julian A. Steyermark 


Mem. New York Bot. Gard. 38: 63. 1984. S. S. Tillett 
et al. 45178; 1 Aug. 1960; Guyana, Mt. Ayanganna; 
900-1,100 m; NY (holo), VEN. 
Schefflera coriacea subsp. neblinae Maguire, Steyerm. 
New York Bot. Gard. 38: 79. 1984. 
j . 1966; 
V suena Pirapucu, = da Neblina, Rio Negro, Rio 
Cauaburi, Rio Maturaca; 1,300 m; NY (holo). 
Schefflera aia Steyerm., Fieldiana Bot. 28(2): 
443. 1952. J. A. Steyermark 57336; 15 July 1944; 


; 25-26 Aug. 1944; 

Venezuela, T.F. Amazonas, d Duida; 305-1,095 
m; F (holo). 

Schefflera globulifera Maguire, wr & Frodin, Mem. 

Bot. Gard. 38: 69. 1984. B. Maguire et 
al. 37354; 23 Jan. 1954; Venezuela, T.F. Amazonas, 
Cerro de la Neblina, Cumbre Camp; 1,700-2,000 m; 
NY (holo). 

Schefflera gracillima ges & Maguire, Bol. Soc 
Venez. Ci. Nat. 32: 371. 1976. J. A. Steyermark et 
al. 109373; 22- 28 yis 1974; Venezuela, Bolivar, 
Meseta del Jaua, Cerro Jaua; 1,750-1,800 m; NY, 
VEN (holo). 

Schefflera guanayensis Maguire, Steyerm. & Frodin, 
Mem. New York Bot. Gard. 38: 66-67. 1984. B. 
Maguire et al. 31740; 2 Feb. 1951; Venezuela, T.F. 


Wurdack 31405; 12 Feb. 1951; Venezuela, T.F. Ama- 
zonas, Cerro Parú, Serranía Parú; NY (holo). 
Maguire, Steyerm. & Frodin 


L. Politi 27756; 17 Dec. 1948; Venezuela, T.F. Ama- 
zonas, Cerro Sipapo (Paraque), Phelps Camp to N Sa- 
vanna; 1,400 m; MO (3401819), NY (holo). 
Schefflera huachamacarii Maguire, Steyerm. & Frod 
Mem. New York Bot. Gard 38: 76. 1984. B. Maguire 
et al. 30253; 15 Dec. 1950; Venezuela, T.F. Am 
zonas, Cerro Huachamacari, vicinity of Summit jan 
1,620 m; NY (holo). 
Se M digi monosperma Maguire, ed rm. & Frodin 
. New York Bot. Gard. 38: 51. 1984. Z. iur 
n ena uela, Roraima; K ( om 


Schefflera — odin, Mem. 
: 


= 
"no 
- 
oz 
= 
w 
ec 


rodin 

984. B. Meam 

.F. pas Cerro 
Sipapo, Upper E Basin; 1,900 m; NY (holo). 

Schefflera pedicelligera Maguire, Steyerm. & Frodin, 
Mem. New York Bot. Gard. 38: 63. 1984. B. d al 
et al. 29546; 19 Nov. 1950; Venezuela, T.F. 
zonas, Culebra Creek drainage, Rio isi. Ce. 
rro Duida; 1,500 m; NY (holo). 

Schefflera pimichinensis Maguire, Steyerm. & Frodin 
Mem. New York Bot. Gard. 38: 55. 1984. B. Maguire 
& J. J. Wurdack 35635; 14 Apr. 1953; Venezuela, 
T.F. Amazonas, Sabana El Venado, left bank of Caño 
Pimichin, above Puerto Pimichin; 140 m; NY (holo), 
VEN. 


Schefflera pn Maguire, Steyerm. & Frodin, 
w York Bot. Gard. 38: 52. 1984. N. T. Silva 
& U. jen 60941; 29 Jan. 1966; Brazil, Amazonas, 


463; 23 Apr. 1960; Venezuela, Bolivar, leidas 
of Rio Cuyuni; 1,300 m; VEN (holo). 

Schefflera ea Steyerm., Bol. Soc. Venez. 
Ci. Nat. 25: 46. 3. J. A. Steyermark & S. Nilsson 
429; 21 Apr. m Venezuela, Bolivar, Cerro Vena- 
mo; 1,100-1,140 m; VEN (holo) = S. duidae Steyerm. 
subsp. quinquestylorum (Steyerm.) Maguire, Steyerm. 
& Frodin 84. 

Schefflera reiiculatà (Gleason) Maguire, Steyerm. & Fro 

Frodin, Mem. 


98078; 22-27 Mar. 1967; aaa, Estado Bolívar, 
Meseta de Jaua; 1,922-2,100 m; NY (holo), VEN. 
Schefflera reticulata (Gleason) Maguire, Steyerm. 

din subsp. yutajensis Maguire, Steyerm. & Frodin, 

em. New York Bot. Gard. 38: 59. 1984. B. Maguire 

& C. K. Maguire 35154; 11 Feb. 1953; Venezuela, 

T.F. Amazonas, Serrania Yutaje, Rio Manapiare; NY 
(holo). 

Schefflera rugosifolia Maguire, n & Frodin, Mem. 

Bot. Gard. 38: 71. 1984. McConnell & 

Quelch 663; Venezuela, Bolivar, Mt. Roraima; K (holo). 

Sc hefflera M eS "emn & Holst, Ann. Missouri Bot. 

1988. A. Gentry & B. Stein 46694; 

e Verla T.F. Amazonas, Cerro de 
la Neblina: 1,900-2,100 m; MO (holo — 348100), VEN. 

ie i sipapoensis Maguire, Steyerm. & Frodin, Mem. 

w York Bot. Gard. 38: 59. 1984. B. Maguire & 
L. “Politi 28471; 20 Jan. 1949; Venezuela, T.F. Ama- 
pem pa uv (Paraque), Upper E Basin; 1,500- 
1,650 m; NY (holo 

Schefflera . as Steyerm pd Bot. 28(2): 444. 
1952. J. A. Scd. 57321; 15 July 1944; Ven- 
ezuela, Táchira, base of Paane de Tamá, 4-10 km 
above Betania; 2,500-2,895 m; F (holo 

Schefflera tamatamaensis Maguire, Steyerm. & dowd 
Mem. New York Bot. Gard. 38: 55. 1984. J. J. W 
dack & L. S. Adderley 43148; 23 s 1959; Ven- 
i T.F. Amazonas, Cano Tama Tama, just above 
Tama — Tama, right bank Rio Orinoco; 150 m; K, MG, 
NY (holo), US, VEN. 

Schefflera tremuloidea Maguire, Steyerm. & Frodin, Mem. 
New York Bot. Gard. 38: 61. 1984. B. eso et 
al. 42379; 13 Dec. 1957; Weneandla. T.F. Am 
Summit, Cerro Neblina, E escarpment of upper Cañon 
Grande Basin; 2,000 m; NY (holo), MO (2911090, 
3401829), 

Schefflera tremuloidea Maguire, Steyerm. & Frodin var. 

marahuacensis 

; Yo 


1981; Venezuela, T.F. Ama- 


me Cerro Marahuaca: 2,500 m; MO, NY (holo), 


Sc hafñera balla (N.E. Br. ) Viguier subsp. duidana 
Maguire, Steyerm. ot 
r 4, 


mit, slopes of ridge; 1,650-1,800 m; NY (holo). 
gS ae yutajensis Steyerm. & Holst, Ann. Missouri 
Gard. 75: D 1988. B. K. Holst & R. En 
id 3426; ll Mar. 1987; Venezuela, T.F. 
zonas, Dept. Pen valley of Rio Coro-Coro, a of 


658 


Annals of the 
Missouri Botanical Garden 


Serrania de Yutaje; 1,100 m; MO (holo — 3485098), 
MO. 


ARECACEAE 


isi d yaracuyense Henderson & Steyerm., Brit- 

a 38: 309. 1986. J. A. Steyermark et al. 100311; 

9-10 Nov. 1967; Venezuela, Yaracuy, Cerro de Cha- 
pa; 1,200-1,400 m; BH dcn VE 

, Fieldiana Bot. 28(1): 71. 


of Bergantin, tributary to Rio 

Neveri, along Rio Leon by Quebrada Danta; 500 m; 

F (holo). 

Bactris duidae Steyerm., Fieldiana Bot. 28(1): 73.1951. 
J. A. Ste eyer rmark 57908; 23 Aug. 1944; Venezuela, 
T.F. Amazonas, near Base River (Caño Negro) at SE 
base of E. Duida: 215 m; F (holo). 

— kamarupa Steyerm., Fieldiana Bot. 28(1): 7 

51. J. A. ea 605 42; 25 Nov. 1944; Xa 
d Bolivar, Quebrada O-paru-ma, tributary of Río 
Pacairao, below Santa Teresita de Kavanayen; 915- 


1,065 m; F (holo) 

Bactris ptariana Steyerm Fieldiana Bot. 28(1): 77. 
1951 A. Steyermark 60046; 10-11 Nov. 1944; 
ae Bolivar, Ptari-tepui; '1,585- 1,600 m; F 


(holo). 

Bactris sororopanae Steyerm., dede Bot. 28(1): 78. 
1951. J. A. Steyermark 60781; 29 Nov. 1944; Ven- 
ezuela, Bolívar, along Rio onn base of Sororopan- 
tepui, W of La Laja; 1,220 m; F (holo). 

Bac tris fab nite Steyerm., Fieldiana Bot. 28(1): 80. 

5 A. Steyermark 57933; 23 Aug. 1944; Ven- 
malk, T. F. Amazonas, SE base of Cerro Duida, along 
Base River (Caño Negro); 225 m; F (holo). 

gi ein a Standl. & Steyerm., Field Mus. 

at. , Bot. Ser. 23(50): 196. 1947. J. A. 2 


ib 51666; 31 Aug. 1942; Guatemala, s Hue 
huetenango, Cerro Negro; 1,600-2,080 m; E (halo). 
Field 


Chamaedorea aguilariana Standl. & Steyerm., 
Mus. Nat. Hist., Bot. Ser. 23(5): 197. 1947. P. la 
Standley 86890; 9 Feb. 1941; Guatemala, Dept. 
zaltenango, between Finca Pirineos and Patzulin; 1,200. 
1,400 m; F (holo). 

Chamaedorea brachypoda Standl. & Steyerm., Field 
Mus. Nat. Hist., Bot. Ser. 23(5): 198. 1947. J. A. 
ace at 39185; 9 Apr. 1940; Guatemala, Dept. 
Izabal, between Bananera and La Presa, base of Sierra 
del Mico; 150 m; F (holo). 

Chamaedorea carchensis Standl. & Steyerm., Field Mus. 

Pie Ser. 23(5): 199. 1947. P. C. Standley 

. 1941; Guatemala, Dept. Verapaz, along 

ie Coban and San Pedro Carcha; 
1,350 m; F (holo). 

i cdi ee digitata Standl. & Steyerm., Field Mus 

. Hist 


, Bot. Ser. 23(5): 200. 1947. J. A. Steyer. 
ud 516 83; 31 Aug. 1 1942; Guatemala, Dept. 
peon Cerro Bg l, -2,000 m; F (holo). 

Chamaedorea fus andl. & Steyerm., Field Mus. Nat. 


Hist., Bot, < hes 201. 1947. J. . 

25 oS 0; Guatemala, Dept. liba. Sierra 
del Mico; 300 ; F (holo). 

Chamaedorea nu ELA Standl. & Steyerm., Field Mus. 
Nat. Hist Tue Ser. 23(5): 202. 1947. J. A. Steyer- 
mark 43583a; 5 Feb. 1942; Guatemala, Dept. El 
Progreso, sid de las Minas; 2,900 m; F (holo). 

Chamaedorea pachecoana Standl. & Soyir Field 
Mus. Nat 


. Hist., Bot. Ser. 23(5): 203. 1947. P. C. 


Standley 63104; 20 Jan. 1939; Guatemala, Dept. 
Guatemala, cultivated, said to come from Dept. Toto- 
nicapan, but probably from some other part of the 
Occidente; F (holo). 

Chamaedorea quezalteca Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 204. 1947. P. C. Standley 
87159;9 Feb. 1941; Guatemala, Dept. Quezaltenango, 
between Finca Pirineos and Patzulin; 1,200-1,400 m; 
F (holo). 

Chamaedorea rojasiana Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 205. 1947. J. A. Steyer- 
mark 33479; 1-2 Jan. 1940; Guatemala, Dept. Que 
zaltenango, Quebrada San Gerónimo, Finca Pirineos, 
Volcán Santa Maria; 1,300-2,000 m; F (holo). 

Chamaedorea skutchii Standl. & Steyerm., iu iem 

ae Ser. 23(5): 206. 1947. A. F. 

1934; Guatemala, Dept. Re ida, 
Volcán Zunil; 2,400 m; F (holo) 

Chamaedorea stenocarpa Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 206. 1947. J. A. Steyer- 
mark 41893; 25 Dec. 1941; Guatemala, Dept. Izabal, 
Cerro San Gil; 650-900 m; F (holo). 

Chamaedorea stricta Standl. & Steyerm., Field Mus 
Nat. Hist., Bot. Ser. 23(5): 207. 1947. J. A. Steyer- 
mark 37381; 9 Mar. 1940; Guatemala, Dept. San 
Marcos, Volcán Tajumulco, up Cerro de Mono; 1,400- 
1,700 m; F (holo). 

Chamaedorea vu is Standl. & Steyerm., Field Mus. 
Nat. Hist., Dot, Ser. 23(5): 208. 1947. J. A. x odi 
mark 37735; 14 Mar. 1940; Guatemala, Dept. Sar 
Marcos, Volcán Tajumulco, 9 mi. NW of El Pine 
1,300-2,000 m; F (holo). 

rds Steyerm., Pedana Bot. 28(1): 82. 1951, 


Dahlgren Jaen Steyerm., Fieldiana Bot. 28(1): 82. 

4. Steyermark 60044; 10-11 Nov. 1944; 

eee ‘Bolívar, SE facing slopes of Ptari-tepui; 

1,585-1,600 m; F (holo) = Dictyocaryum ptariense 
(Steyerm.) Moore & Steyerm., 1967. 

Desmoncus duidensis Steyerm., Fieldiana Bot. 28(1): 85. 
1951. J. A. Steyermark 57944; 23 Aug. 1944; Ven- 
ezuela, T.F. Amazonas, SE base of Cerro Duida, along 
Base River (Caño Negro); 225 m; F (holo). 

Desmoncus multijugus Steyerm., Fieldiana Bot. 28(1): 
85. 1951. J. A. Steyermark 60968; 18 Dec. 1944; 
Venezuela, Bolivar, Tumeremo; 305 m; F (holo). 

Euterpe ptariana Steyerm., Fieldiana Bot. 28(1): 87. 
1951. J. A. Steyermark 60045; 10-11 Nov. 1944; 
Venezuela, Bolívar, Ptari-tepui; 1,585-1,600 m; F 
(holo). 


Geonoma karuaiana Steyerm., Fieldiana Bot. 28(1): 8 
1951. J. A. Steyermark 60789; 29 Nov. 1944; ‘jk 
ezuela, Bolivar, W of La Laja, base of Sororopan-tepui, 
along Rio Karuai; 1,220 m; F (holo) 


Hyospathe maculata Steyerm., Fieldiana Bot. 28(1): 89. 
1951. J. A. oe 60406 0- - Nov. 1944; 
Venezuela, Boliv. a O- paruam x bervenn San- 


ta Teresita de oran en and Rio Poe. (tributary 

of Rio Mouak); 1,065-1, 220 m; F (holo 

L. var. venezuelana Steyerm., Fieldi- 

t. 28(1): 90. 1951. J. A. Steyermark 57649; 

reed 194 44; Venezuela, Bolivar, between Rio Caroni 
and Ciudad Bolivar; 200 m; F (holo). 

Prestoea humilis Henderson & Steyerm., Brittonia 38: 
311. 1986. 4. Henderson 501; 12 Nov. 1985; Ven 
ezuela, Yaracuy, Cerro la Chapa; 1,200-1,400 m; K, 
MYF, NY, VEN (holo 

Socratea albolineata Steyerm., Fieldiana Bot. 28(1): 9 


Volume 76, Number 3 
1989 


Taylor 659 
Plants Described by 
Julian A. Steyermark 


1951. J. A. Steyermark 60541; 25 Nov. 1944; Ven- 
ezuela, Bolivar, Quebrada O-paru-ma, tributary Rio Pa- 
cairao, below Santa Teresita de Kavanayen; 915-1,065 
m; F (holo 


ARISTOLOCHIACEAE 


—€— mollis Standl. & Steyerm., Field Mus. Nat. 
, Bot. Ser. 23(4): 155. 1944. J. 
51269; 24 Aug. 1942; Guatemala, Dept. Hue 
nango, canyon tributary to Rio biel ag between 
Democracia and canyon of Chamushü; 1,000 m; F 
(holo) = 4. paracleta Pfeiffer, nom. nov 


ASCLEPIADACEAE 


Asclepias woodsoniana Standl. & Steyerm., Field Mus. 
Nat. Hist., ar Ser. 23(5): 224. 1947. J. A. Steyer- 
mark 31781; 15 Nov. 1939; Guatemala, Dept. Jutia- 
pa, between ‘Trapiche Vargas and Asuncion Mita; 500 
m; F (holo). 

Blepharodon pr ape Steyerm., Acta Bot. Ven- 
ez. 2: 319. 1967. J. A. Steyermark 93889; 14 May 
1964; Venezuela, Bolivar. Auyan-tepui; 2,000-2,200 
m; VEN (holo). 

Cynanchum auyantepuiense Steyerm., Acta Bot. Venez. 
2: 320. 1967. J. A. Steyermark 93892; 14 May 1964; 
Venezuela, Bolivar, Auyan-tepui; 2,050 m; VEN (holo). 

Cynanchum stenomeres Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 224. 1947. W. A. Schipp 
674; 5 Feb. s.n.; Belize, All Pines; sea level; F (holo). 

Marsdenia blepharodes Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 226. 1947. J. A. Steyer- 
mark 33615; 6 Jan. 1940; Guatemala, Dept. Quezal- 
tenango, San Juan Patzulin 1,500 m; F (holo). 


ASTERACEAE 


Achnopogon Maguire, Steyerm. & Wurd., Mem. New 
York Bot. Gard. 9(3): 437. 1957, gen. nov. 


. J. Wurdack 681; 9 Feb. 1955; d 
Bolivar, Chimantá Massif, Torono-tepui; 2,090 m 
(holo), NY. 

Ageratum chortianum Standl. & Steyerm., Field Mus 
Nat. Hist., Bot. Ser. 23(2): 98. 1944. J. A. Steyermark 
31269; Nov. 1 iqui 
Montaria Castilla, near Montana Cebollas, along Rio 
Sé Lucia Saso; 1,200-1,500 m; F (holo). 

icone ballotifolia Maguire, Steyerm. rd., Mem. 

ork Bot. Gard. 9(3): 425. 1957. J. A. Steyer- 
du 75520: 2] May 1953; Venezuela, Bolivar, Chi- 
mantá Massif, Chimantá-tepui (Torono-tepui); 1,700 m; 
F (holo), NY. 

Aphanactis ie Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 22(5): 390. 1940. P. C. Standley 58674; 
4 Dec. 1938; Guatemala, Dept. Chimaltenango, Cerro 
Tecpán, Santa Elena; 2,700 m; F (holo). 

Aplopappus gymnocephalus DC. f. albus DE pee & 
Moore, Ann. Missouri Bot. Gard. 20: 797. 1933. J. 
A. Moore & J. A. Steyermark 3488; 17 "july 1931; 
U.S.A., Texas, Culberson Co., Guadalupe Mountains, 
ridge above McKittrick Canyon; MO (holo) = Haplo- 
pappus gymnocephalus DC. 

Archibaccharis blakeana Standl. & Steyerm., Field Mus 


Nat. Hist., Bot. Ser. 22(4): 296. 1940. P. C. Standle ey 


56597; 1938-1 939; Guatemala, Dept. Sacatepéquez, 
near Antigua; 1,500-1,600 m; F (holo). 


Aspilia avilensis Aristeg. & Steyerm., Fl. Venez. 10(2): 

ogl 41; Aug. 1937; Venezuela, Distr. 

Federal, (cam Serranias del Avila; depository not 
indicated. 

Aster anomalus Engelm. f. albidus Steyerm., Rhodora 
51: 119. 1949. J. A. P 65135; 23. Sep. 
1947; U.S.A., Missouri, Barry Natural Bridge 
Hollow, 8-9 mi. SE of — F (holo). 

, Field Mus. Nat. Hist., 
Bot. Ser. 23(3): 141. 1944. C. A. Purpus 12091; May 
1927; Mexico, Chiapas, Hacienda Monserratee; F (holo) 
— Aster moranensis 76. 

Baccharis erectifolia Steyerm., Belda Bot. 28(3): 620. 
195 Steyermark 62583; 6 May 1945; Ven- 
ezuela, Sucre, al Turumiquire; 2,360-2,500 m; F 
(holo). 

Baccharis meridensis Steyerm., Fieldiana Bot. 28(3): 
622. 1953. J. A. Steyermark 56537; 14 May 1944; 
Venezuela, Mérida, Páramo de Los Colorados; 2,745- 

— Psila brachylaenoides DC. var. 
brachylaenoides (DC.) Aristeg., 1964 
Baccharis p Steyerm., Follar Bot. psu ici 
A. Steyermark 59928 d 1944; V 
ezuela, Bolívar, Ptari-tepui; 2, Am 2,450 m; F (holo) 
= Psila brachylaenoides DC. var. oa (DC.) 
Aristeg., 1964. 

Baccharis Visi n Baker subsp. occ PD. Stey- 
rm. & uire, Ácta - Venez. 14(3): 25. 1984. 
B. Marise et al. 65567; 15 Jan. 1981; E 
Dept. Atabapo, Cerra Marahuaca; 2,685 m; VEN (holo), 
NY. 


Baccharis venulosa DC. var. cuspidibracteata Steyerm., 
Fieldiana Bot. 28(3): 624. 1953. 4. Steyermark 
5/298; 15 July 1944; Venezuela, Táchira, base of 
n de Tamá; 2,500-2,895 m; F (holo). 
Cacalia epis Standl. & Steyerm., Field Mus. 
Hist., Bot. 23(2): 99. 1944. J. A. Steyermark 


= Psacalium guatemalensis (Standl. Sá 

Steyerm.) Cuatr., 1955, 1976. 

Cacalia pinetorum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(3): 142. 1944. J. A. Steyermark 


nango, Volcán Santo Tomás; 3,000- 3,200 m; F (holo) 
= Psacalium pinetorum (Standl. & Steyerm.) Cuatr., 
1955, 1976. 


Cacalia pudica Standl. Sí Steyerm., Field Mus. Nat 


Hist., Bot. Ser. 23(5): 255. 1947. J. A. Steyermark 
48344; 7 July 1942; Guatemala, Dept. Huehuete 
nango, Sierra de los Cuchumatanes; 3,400-3,500 m; 


ie 

F (holo) = Senecio nubivagus L. Williams, 1975. 
Calea clematidifolia Steyerm., Fieldiana Bot. 28(3): 627. 
1953. J. A. Steyermark 59072; 1 Oct. 1944; Ven- 
ezuela, Bolivar, Gran Sabana, 1,065-1,220 m; F (holo). 
Calea s Standl. & Steyerm., Field Mus. Nat. 
r. 23(5): 256. 1947. J. A. Steyermark 
Y : gun 1942; Guatemala, Dept. Alta Vera- 
Chinajá; 150-700 m; F (holo) — C. del- 


ist., 
45627; 
paz, Cura 
ophylla Cowan, 1964. 

Calea lucidivenia Gleason & S. F. Blake f. lanceolata 
Steyerm., Fieldiana Bot. 28(3): 628. 1953. J. A. S 
€ 59408; 26 Oct. 1944; Venezuela, Bolivar, 

an Sabana sl, 220 m; F sei 

Calea oliveri Robinson & Gree var. taeniotricha 
Steyerm., Fieldiana Bot. 283) 628. 1953. J. A. Stey- 
ermark 59403; 26 Oct. 1944; Venezuela, Bolivar, near 


660 


Annals of the 
Missouri Botanical Garden 


Rio Karuai, NW of Mission of Santa Teresita de Ka- 
vanayen; 1,220 m; F es ). 


alea savannarum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 257. 1947. J. A. Steyermark 
38658; 3 Apr. 1940; Guatemala, Dept. Izabal, between 


Milla 49.5 and Cristina; 70 m; F (holo) = 
uliginosa Sw., 19 
Calea d d Steyerm, Field Mus. Na 


Spilanthes 


t. Hist., Bot. 
Ser. A 1940. J. A. S erant 37004; 1 
Volcá 


Mar. 1940; Guatemala Dept San Marcos, 
sun 1,300-3,0 F (holo) — ben 
standleyi (Steyerm.) D. Nab 1975. 

-ardonaea Aristeg., Maguire & Steyerm., Mem. New 
York Bot. Gard. 23: 892. 1972, gen. nov. 

Cardonaea j jauaensis Aristeg., Maguire & Steyerm., Mem. 
New York Bot. Gard. 23: 892. 1972. J. A. Steyermark 
97945; 22- 97 Mar. 1967; Venezuela, Bolivar, Meseta 
de Jaua, Cerro Jaua; 1, ,922- 2,100 m; VEN (holo), NY. 

N 


Chimantaea sect. Cinerea Maguire, Steyerm. & Wu 
m. 


rd., 
New York Bot. Gard. 9(3): 429. 1957, sect. 


C e dum Wed rein Steyerm., Ann. Missouri Bot. 
Gard. 71: 335.1 rA; Steyermark et al. 129932; 
14-16 Feb. 1984; pisei Bolivar, Piar, Macizo 
de Chimantá; 1,920 m; VEN (holo). 

nas cinerea (Gleason & S. F. Blake) Maguire, 
Él urd. f. glabra Steyerm., Ann. Missouri 

LOU 7l: 337. . E. Foldats 7100; 15 Aug. 
1968: Venezuela, Bolivar, Auyan-tepui, lado derecho 
del Salto Angel; VEN (holo 

Chimantaea eriocephala Maguire wie & Wurd., 
Mem. New York Bot. Gard. 9(3): 431. 1957. J. J. 
Wurdack 34225; 26 e 1953; Menus Bolivar, 
Chimantá Massif, Churi-tepui (Muru-tepui); NY (holo). 

Chimantaea espeletoidea Api pr o & Wurd., 

m. New York Bot. d. 93): 4 1957. J. A. 
Steyermark & . Wurdack a E Feb. 1955; 
Venezuela, Bolivar, Chimantá Massif; 1,940 m; F (holo), 


QE 


Chimantaea huberi Steyerm., Ann. yo Bot. Gard. 

71: 335. 1984. J. 4. Steyermark et al. 28815; 2- 

5 Feb. 198 83; ala, Bolivar, Piar, y on del 

Chimantá; l, 850 m; VEN (holo), K, MO (3223158), 
N 


Chimantaea ndis Maguire, 2 & Wurd., Vae 
New York 


Bot. Gard. 9(3): 432. 1957. J. A. Stey 
mark & J. J. Wurdack 747; i Feb. 1955; Venezuela, 
Bolivar, Chimantá Massif; 2,120-2,210 N 
(holo), F. 


Chimantea lanocaulis Maguire, Steyerm. & Wurd., ME 
New York 0. 


Chimantá Massif, summit of Apacara-tepui; F (holo), 
NY. 


C d mirabilis Maguire, aro & Wurd., Mem. 
ork Bot. Gard. A3): 432. 1957. J. A. Steyer- 

el & J. J. Wurdack 821; 13 Feb. 1955; Venezuela, 
Bolivar, Chimantá Massif; 2,120 m; F (holo 
Chimantaea rupicola Maguire, Steyerm. & 
k Bot. Gard. 9(3): 431. 1957. J. 4. Steyer 

mark & J. J. Wurdack 748; 11 Feb. 1955; Venezuela, 

_ Bolivar, Chimantá Massif; 2,120-2,210 m; F, NY (holo). 


Z 


mark & J. J. 
Bolivar, Chimantá Massif; 1,925 m; F, NY (holo). 


Wurd., Mem. 


d breweri Steyerm. & Maguire, Acta Bot. 
14(3): 26. 1984 


39. T “Moore s.n. 25 Aug 1939; 
S.A., Missouri, Laclede Co., Mill Creek; E (holo), 
O. 
Cirsium nigriceps Standl. & Steyerm., Field Mus 
Hist., Bot. Ser. 23(5): 258. 1947. J. A. Diss 
50157; 6 Aug. 1942; Guatemala, Dept. Huehuete- 
nango, between Tojquia and Caxin, Sierra de los Cu- 
chumatanes; 3,700 m; F (holo). 
ibadi l L. var. macrophyllum Steyerm., 
Fieldiana Bot. 28(3): 629. 1953. J. A. Steyermark 
62437; 3 May 1945; Venezuela, Silene, Cerro Turu- 
miquire; 1, 700- 1,840 m; F (holo) = C. surinamense 
964. 


nons marahuacensis Steyerm., pars 30: 50. 
1978. S. S. Tillett et al. 752-3 2 Feb. 1975; 
Venezuela, T.F. Amazonas, Cerro Ce e 2,750 
m; VEN (holo). 

Echinacea dw Nutt. f. albida Steyerm., aper 

42. J. E Wr cain be 4 June 1941; 

Ms St. Francois Co., Koester MEA 

nie glade on i of bluffs along Plattin Creek; F 
(holo). 

Echinacea purpurea (L.) Moench. var. arkansana Stey- 
erm., Rhodora 40: 71. 1938. Brinkley 250; 19 July 
1937; U.S.A., Arkansas, Sevier Co., Boggy Springs; F 
(holo). 

Echinacea Eos (L.) Moench. f. liggettii Me laa 
Rhodora 44: 150. 1942. W. Liggett 1; 9 July 
1940; U .S.A., Missouri, Dallas Co., Hwy. 54, W of 
Niangua Riveri F (holo). 

Erigeron Hu Standl. & Steyerm., Field Mus. Nat 

ist., Bot. Ser. 22(4): 301. 1940. P. C. Standley 
65053: 11 Feb. 1939; Guatemala, Dept. Sacatepé- 
quez, Volcán de Agua, above Santa María de Jesús; 
2,250-3,000 m; F (holo). 

Erigeron venezuelensis Steyerm., Fieldiana Bot. 28(3): 

3, nom. nov. = Oritrophium venezuelense 
(Steyerm.) Cuatr., 1964 

Eupatorium antiquorum Standl. & Steyerm., Field Mus. 
Nat. Hiat , Bot. Ser. 22(4): 302. 1940. P. C. a 
60304 18 Dec. 1938; Guatemala, Dept. Sacatepé- 
quez, near Antiqua; 1,500-1,600 m; F (holo). 

Eupatorium benjamin-lincolnii Steyerm., Fieldiana Bot. 
28(3): 635. 1953, nom 

db p ue carmonis Standl. & Steyerm., Field Mus. 
Nat , Bot. Ser. 22(4): 303. 1940. P. C. Standley 
63743; E Jan. 1939; Guatemala, Dept. Sacatepéquez, 
Finca Carmona, hills SE of Antiqua; 1,600-1,800 m; 
F (holo). 

Eupatorium crassicaule Steyerm., Fieldiana Bot. 28(3): 
629. 1953. J. A. Steyermark 57749; 21 Aug. 1944; 

Venezuela, T.F. Amazonas, Cerro Duida; 150 m; F 
(holo). 

Eupatorium croc sa tial Standl. & Steyerm., juo Mus. 
Nat. Hist., r. 23(4): 182. 1944 o 
mark 51498; 29 Aug. 1942; Guatemala, Dept 
huetenango, Ciénaga de Lagartero, below Miramar, 
300 m; F (holo). 

Eupatorium cupressorum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(4): 183. 1944. P. C. Standley 


18 1 Sa 
se NEAR. 2. 220- 3,000 m; F (holo) 


Volume 76, Number 3 
1989 


Taylor 661 
Plants Described by 
Julian A. Steyermark 


"o ge pisi Steyerm., Fieldiana Bot. 
28(3): 636. 1953, no À 
Fupatoriun mm Stand. & Steyerm., Field Mus 
t. Hist., Bot. Ser. 22(4): 303. 1940. P. C. Standley 
71125: 9 Apr. 1939; Guatemala, Dept. Alta Verapaz, 
E of Tactic, on road to Tamahu; 1,500-1,650 m; F 
(holo). 

Eupatorium huehuetecum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 22(4): 304. 1940. P. C. Standley 
65798; 20 Feb. 1939; Guatemala, Dept. Huehuete- 
nango, Rio Pucal; 1,775 m; F (holo 

Eupatorium ibaguense Schultz-Bip. var. dndisoratum Mas 

uire, Steyerm. & Wurd., Mem. New York Bot. Gard. 
9(3): 425. 1957. J. J. Wurdack 34112; 13 Jan. 1953; 
Venezuela, Bolívar, Chimanta Massif, Sarven-tepui; 
1,900-2,050 m; F, NY (holo). 

bi coe Standl. & Steyerm., Field Mus. Nat. 
His . Ser. 23(4): 183. 1944. P. C. Standley 

1940; Guatemala, Dept. Jutiapa, El 
Barrial, E of penes 800 m; F (holo). 

— macram Standl. & Steyerm., Field Mus. Nat. 

a Ser. 23(4): 184. 1944. J. A. Steyermark 
e ; 9 Nov. 1939; Coen. Dept. Chiquimula, 
base a Cerro Colorado along Río Jocotán, near Jocotán; 
400 m; F (holo). 

Fupatorium mendax Standl. & Steyerm., Field Mus. Nat 
Hist., Bot. Ser. 23(4): 185. 1944. J. A. Stovermark 
36341; 21 Feb. 1940; Guatemala, Dept. San Maren 

Rafael and Mexica og 

500- 3,000 m; F (holo) = 

76. 


ir ne mimicum hy ndl. & Steyerm., Field Mus. 

t. Hist m Ser. 23(4): 186. 1944. P. C. Standley 

85 146; 7 Jan. 1941; Guatemala, Dept. Quezalte- 
nango, region of Las Nubes, S of San Martín Chile 
Verde; 2,250 m; F (holo). 

Éupatorium montigenum Standl. & Steyerm., Field Mus. 

at. Hist., Bot. Ser. 23(5): 258. 1947. J. A. Steyer- 
mark 43532; 5 Feb. 1942; Guatemala, Dept. El Pro- 
greso, N of E Piamonte, Volcán Santa Luisa; 2,400- 
3,000 m; F (holo 

py aco ovillum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 22(4): 305. 1940. P. C. Standley 
65862; 20 Feb. 1939; Guatemala, Dept. Totonicapán, 
ue del Aire, on road between Sija and Huehue 

nango; 3,000-3,450 m; F (holo). 

ei S pharcidodes Robinson var. sororopanensis 
Steyerm., Mee Bot. 28(3): 632. 1953. J. A. Stey- 
une ; 13 Nov. 1944; Venezuela, Sororopan- 

ui; dra epee F (holo). 

dean plethadenium Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(4): 186. 1944. P. C. Standley 
61100; 26 Dec. 1938; Guatemala, Dept. Chimalte- 
pango; — of Santa Elena, Cerro Tecpan; 2,400- 
2,700 m; F (holo) = E. semialatum Benth. 

o rojasianum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(4): 188. 1944. P. C. Slandiay 
67221; 1939; Guatemala, Dept. Quezalte- 
nango, near El Muro, below Santa Maria de Jesüs; 
1,200 m; F (holo). 

Bsotorum roupalifolium Robinson f. latifolium Stey- 
erm., Fieldiana Bot. 28(3): 633. 1953. J. A. Steyer- 
npe: 59505; 28 Oct. 1944; Venezuela, Bolivar, Ptari- 
p 1,810 m; F (holo) = E. roupalifolium Robinson 
var. roupalifolium, 1976. 

oru roupalifolium Robinson var. e ut 
lium Steyerm., Fieldiana Bot. 28(3): 636. 1943. J. A 
Steyermark 59865; 4 Nov. 1944; Venezuela, Bolivar. 


Ptari-tepui, 2130 m; da = "x roupalifolium Rob- 
inson var. roupalifolium, 19 

Eupatorium salinum pe & M Field Mus. Nat. 
Hist., Bot. Se , 


Eupatorium saxorum Standl. 
Nat. Hist., Bot. Ser. 23(4): 189. 1944. J. A. Steyer- 
mark 36075; 19 Feb. 1940; Guatemala, Dept. San 
Marcos, between Sibinal and summit of Volcán Tacaná; 
2,500-4,000 m; F (holo). 

Eupatorium stevioides Steyerm., Fieldiana Bot. 28(3): 
637. A. Steyermark 57084; 7 July 1944; 
Venezuela, Táchira, Páramo de la Negra; 2,745-3,045 
m; F (holo). 

Eupatorium PETAT: ie ils 
28(3): 636. 1953, nom 

Eupatorium tepuianum S , Fieldiana Bot. 28(3): 
638. 195 Stey E 59504; 29 2 1944; 
Venezuela, olivas Pari. -tepui; 1,810 m; F (holo). 

Eupatorium vetularum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(4): 190. 1944. P. C. Standley 
84404; 23 Jan. 1941; Guatemala, Dept. Totonicapán, 


Fieldiana Bot. 


Eupatorium zunilanum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot Ser. n 191. 1944. J. A. in 
mark 34744; 22 Jan. 1940; Guatemala, Dept. Que 
zaltenango, Volcán m 2,500-3,800 m; F (holo). 

Gnaphalium caeruleocanum Steyerm., deiude Bot. 
28(3): 639. 1953. J. A. Steyermark 62573; 6 May 

1945; Venezuela, Sucre, Cerro iiid 2,200- 
2,500 m; F (holo). 

Gnaphalium standleyi Steyerm., Field Mus. Nat. Hist., 

Bot. Ser. 23(2): 99. 1944. P. C. Standley 81097; 28 

ec. 1940; Guatemala, Dept. Huehuetenango, region 
of Chemal, Sierra de los Cuchumatanes; 3,300 m; F 
(holo). 


Gong ylolepis ap Maguire, es m. & Wurd., 
Mem. New Y . Gard. 9(3): 437. 1957. J. A. 
Steyermark & ^ T Wurdack 1136; 26 Feb. 1955; 


Venezuela, Bolivar, Chimantá Massif, Agparaman-te- 
pui 1,880-1,955 m; F, NY bis lo). 

ryan M bono Steyerm., Ann. Missouri Bot. 
Gard. 74 TJ Sud B. K. Holst 
130629; al Feb, 1985; Venezuela, T.F. Amazonas, 
Dept. Atabapo, Cerro Marahuaca; 1,220-1,350 m; 
MO (holo), VEN. 

no en Steyerm., Ann. een og Bot. Gard. 
21: 498. E. Wooton s.n.; Aug. 1910; 
U.S.A., No LER ini s Meg p (holo). 

Grindelia aggregata Steyerm., Ann Missouri Bot. Gard. 
21: 566. 1934. J. W. Congdon s.n.; 27 sd 1902; 
Canada, British Columbia, Victoria; MO 

Grindelia woo Steyerm., Ann. Missouri “Bot. Gard. 
21: 229. 4. J. T. Howell 5473; 14 Sep. 1930; 
U.S.A., rd Mendocino Co., Mendocino county 
plain, 4.5 mi. N of Ft. Bragg; CAS (holo), GH, MO, 
UC. 


= 


Grindelia arenicola Steyerm. var. E Stey- 
erm., Ann. Missouri Bot. Gar A 596. 1934. J. T. 
Howell 8106; Sl Sep. 1931; ., Ca TM Men 
ocino Co., 6 mi. S of Point i "CAS (holo), GH, 


Grindelia arenicola Steyerm. f. trichophora Steyerm., 
nn. Missouri Bot. Gard. 21: 596. 1934. W. F. Schmitt 


662 


Annals of the 
Missouri Botanical Garden 


s.n.; 1913; U.S.A., California, San Mateo Co., 
moon Bay; US (holo 

Grindelia arizonica A. Gray var. d Steyerm., Ann. 
Missouri Bot. Gard. 21: 508. ). B. Metcalfe 
744; 19 Sep. 1903; U.S.A., New Marin. Bear Mt., 

near Silver City; GH, MO, MINN, ND-G, NY, RM. 
US (holo). 

Grindelia arizonica A. Gray var. mic d Steyerm., 
nn. Missouri Bot. Gard. 21: 508. 1934. 4. Davidson 
736 (96) ; E iid 1902; U.S.A., Arizona, along San 

Franciso River, 10 mi. N of Clifton; GH (holo), PH, 


Grindelia arizonica A. Gray var. Sty up e. 
Ann. Missouri Bot. Gard. 21: 507. 1934. E. O. Wooton 
2508; 12 Aug. 1904; U.S.A., Colorado, near Bayfield; 

5 (holo). 

Grindelia blakei Steyerm., Ann. Missouri Bot. Gard. 21: 
567. 1934. J Moore & J. A. Steyermark 3686; 
9 Aug. 1931; U.S.A., California, Humboldt Co., Eu- 

; MO (h 

ene var. abbreviata Stey- 

. 21: 228. 1934. Hoff- 

man s.n.; 23 Sep. ; US. A., California, Los An- 

geles Co. DE SBC (holo 

. Greene var. australis Steyerm., 


Lj on 


ifornia, Los Angeles Co., 
RSA (holo). 

Grindelia camporum E. Greene var. interioris (Jepson) 
Steyerm. f. foliacea Steyerm., Ann. Missouri Bot. Gard. 
21: 538. 1934. J. T. Howell 5201; 30 Ma 
U.S.A., California, Solano Co., 1 mi. from Vacaville on 
road to Elmira; CAS (holo), GH, RSA. 

Grindelia camporum E. Greene var. parviflora Steyerm., 
Ann. Missouri Bot. Gard. 21: 534. 1934. J. T. Howell 
5496; 28 Sep. 1930; U.S.A., California, Contra Costa 
County, Liberty School on Marsh Creek Road; CAS 
(holo), MO (1015935). 

Grindelia confusa oou 
Ser. 17(5): 442. 

36; Mexico, Chihuahua, pean T-t plains; F 

Grindelia a dis rm . Missouri Bot. 
21; R. oa 1204; 
Mr Loin. uo US (holo). 

Grindelia greenmanii Steyerm., Ann. Missouri Bot. Gard. 
21: 460. 1934. E. Palmer 471; 10-13 July 1880; 
Mexico, Coahuila, Lerios; GH (holo), PA, US. 

o oa Steyerm., Ann. Missouri Bot. Gard. 21: 

. 1934. L. R. Abrams 3957; 30 June 1903; U.S.A., 
COMER San Diego Co., ene Lake; F, GH, MO 
(130219), NY (holo), RSA 

Grindelia havardii Steyerm., rd. 
21: 474. 1934. J. 4. Moore & J. Steyermark 
3607; 23 July 1931; U.S.A., Texas, Culberson Co., 
near mouth of Mckittrick Canyon, Guadalupe Mts.; 
MO (holo). 

Grindelia hirsutula Hook. & Arn. var. brevisquama 

Missouri Bot. Gard. 21: 572. 1934. 

. ,C alifornia, 

Sonoma Co., Black Point Road; CAS (holo), T 

Grindelia hirsutula Hook. & Arn. var. um 
Steyerm. f. a Steyerm. . Missouri Bot. 
Gard. 21: 573. 1934. J. T. Howell 522 1; 8 June 
1930; U.S.A., Calor: Sonoma Co., 8 mi. W of 
pres on Bodega Road; CAS, GH, MO pon 
1016008), RSA = G. hirsutula Hook. & Arn. var. 


hirsutula. 


3 mi. E of bes Lake: 


Field Mus. Nat. Hist., Bot. 
. H. Le we 1016; 17 Aug. 
(holo). 
Gard. 
1924; Mexico, 


Ann. Missouri Bot. Ga 
A. 


Half- 


Grindelia hirsutula Hook. & Arn. var. brevisquama 
Steyerm. f. REE Serena Ann. Missouri 
Bot. Card, 21: 4. Eastwood s.n.; 

1900; U.S.A., (E dies GH, UC (holo). 
Grindelia hirsutula Hook. & Arn. var. brevisquama 
Steyerm. f. tomentulosa Steyerm., Ann. Missouri Bot. 

W. Suksdorf 279; 23 June 

S.A., California, San Mateo Co., Crystal Springs 

Lakes MO (holo —850778) = G. hirsutula Hook. 


Grindelia icum Hook. & ‘Arn, f. cacumena Steyerm., 
Ann. Missouri Bot. Gard. 21: 571. 1934. H. N. Bo- 
lander 389; 1866; U.S.A., California, Oakland Hills; 

, US (holo). 
Grindelia hirsutula Hook. & Arn. var. calva Steyerm., 
Ann. Missouri Bot. Gard. 21: 575. 1934. F. E. Road- 
house ps Spring 1905; U.S.A., California, San Luis 

Obispo Co., San Luis Obispo; UC (holo). 

ue Tue: Hook. & Arn. var. de ics Stey- 
, Ann. Missouri Bot. Gard. 21: 229. . J. T. 
Howell ed 1 July 1933; U.S.A., di Ri Ven- 
tu mi. E of Ojai; CAS, MO (holo— 1044105). 
Grindelia nelli Steyerm., Ann. Missouri Bot. Gard. 
1934. L. F. Henderson 2791; 10 Aug. 1894; 
U. d Idaho, Kootenai Co., St. Maries River; GH 

(holo), S. 

Grindelia humilis Hook. & Arn. f. n Steyerm 

je Missouri Bot. Gard. 21: 527. A i Howell 
0805; 8 Oct. 1932; U.S.A., Heat, Napa Co. 
TURA Wharf, on Napa River; CAS (holo), MO 
(1028961). 
Grindelia humilis Hook. & Arn. f. d Steyerm., 
Ann. Missouri Bot. n 21: 526. V. L. Greene 
s.n.; 15 Nov. 1 S.A., mim Oakland: de- 
pository not indicated: 

Grindelia inornata E. Greene A angusta Steyerm 
Ann. Missouri Bot. Gard. 21: 934. H. L. Shanks 
597; 1903; U.S.A., i ou [ied Springs; US 


Grindelia integrifolia DC. f. dentata Steyerm., Ann. 
Missouri Bot. Gard. 21: 551. 1934. J. C. Nelson 1670; 
14 July 1917; U.S.A., Oregon, Salem; GH (holo). 

i eie integrifolia DC. var. virgata (Nutt.) T. & G. 

sa Steyerm., Ann. Missouri Bot. Gard. 21: 553. 
1934. E Hall 266; 1871; U.S.A., Oregon; F, GH, 
MO (holo —130132 & 130036), NY, US. 

Grindelia lanceolata Nutt. E tr Steyerm., Ann. 
Missouri Bot. Gard. 21: 51 934. IF. Trelease s.n.; 
11 Sep. 1898; U.S.A., Md Stone Co., Marble 
Cave; MO (holo). 

Grindelia littoralis | e T i Missouri Bot. Gard. 

or s.n.; 20 Sep. 1884; U.S.A., 
Texas, renace leon Bay MO ees 

Grindelia maritima (E. Gre 
Steyerm., Ann. Mi 34. J. 
T. Howell 11658; 15 Sep. 1933; U.S.A., California, 
San Franciso, Laguna Honda; CAS, MO (holo). 

Grindelia microcephala DC. var. adenodonta Steyerm., 
Ann. Missouri Bot. Gard. 21: 467. 1934. F. Lindhei- 
mer 255; July-Aug. 1844; U.S.A., Texas, 39 mi. W 
of San Felipe; CAL, G, MO (holo), P 

Grindelia microcephala DC. var. a Steyerm. 
f. angustior Steyerm., Ann. Missouri Bot. Gard. 21: 
469. 1934. E. J. Palmar 33590; 28 Apr. 1928; U.S.A., 
Texas, Uvalde Co., near Uvalde; MO (holo), NY. 

Grindelia microcephala DC. var. montana Steyerm., 
Ann. Missouri Bot. Gard. 21: 470. 1934. L. C. Smith 


. f. anomala 


Volume 76, Number 3 
1989 


Taylor 663 
Plants Described by 
Julian A. Steyermark 


135; 13 Aug. 1894; Mexico, Oaxaca, San Juan del 
Estado; GH (holo). 

Grindelia microcephala DC. var. pusilla Steyerm., Ann. 
Missouri Bot. Gard. 21: 467. 1934. E. Palmer 469; 
21-28 Jan. 1880; U.S.A., Texas, between the Frio 

and the Agree Rivers, on the road to Laredo; GH 
(holo), MO, S. 

Grindelia nana ic var. "i s Steyerm., Ann. Mis 
souri Bot. Gard. 21: 244. 4. J. P. Tracy 4669; 
12 Sep. 1915; U.S.A., Clon, Humboldt Co., Eel 
River, at mouth of e Creek; UC (holo), US. 

Grindelia nana Nutt. var. altissima Steyerm. f. puberula 

ard. 21: 545. 1934. J. 
s.n. e 1924; U.S.A., Califor- 

» Kelseyville CAS (holo). 

. f. longisquama Steye Ann. 
Missouri Bot. " 21: 543. 1934. Rev. Spalding 
s.n.; n.d.; U.S.A., Idaho, near Clearwater (formerly in 
Oregon); GH (holo), N : 

Grindelia nana Nutt. var. turbinella Steyerm., Ann 
Missouri Bot. Gard. 21: 545. 1934. 4. A. Heller 12579; 
7 Sep. 1917; U.S.A., California, Siskiyou County, Shas- 
ta Valley, 10 mi. S of Gazelle; CAS, GH (holo), MO 
(824996), NY, UC, US = G. nana Nutt. var. nana. 

Grindelia Lir Steyerm., Ann. Missouri Bot. Gard. 
dà bass 4. E. W. Nelson 6864; 8-12 Feb. 1903; 

reden Los Reyes; GH, NY, US (holo). 

Grindelia oxylepis E. Greene var. capitellata Steyerm., 
Ann. Missouri Bot. Gard. 21: 490. 1934. E. Palmer 
520; 21-23 Oct. 1898; Mexico, Durango, Mapimi; 
GH (holo), MO, NY, UC, US. 

Grindelia oxylepis E. Greene var. icr Steyerm., 
Ann. Missouri Bot. Gard. 21: 490. 1934. E. Palmer 
316; Sep. 1898; Mexico, Coahuila, Saltillo; GH (holo), 
MO, NY, UC, US. 

Grindelia palmeri Ste , Ann. Missouri Bot. Gard. 
21: 471. 1934. E. Pano 163; 28 Sep.-3 Oct. 1902; 
Mexico, San Luis Potosi, Alvarez; GH, MO, NY, US 
(holo). 

Grindelia revoluta Steyerm., Ann. Missouri Bot. Gard. 
21: 496. 1934. J. 4. Moore & J. A. Steyermark 
3784; 30 Aug. 1931; U.S.A., Colorado, El Paso Co., 
14 mi. S from Colorado Springs; MO (holo). 

Grindelia robinsonii yu e Ann. Missouri Bot. Gard. 
21: 459. 1934. C. A. Purpus 5722; July 1911; Mex- 
ico, San Luis Potosí, Agua del Medio; GH, MO, NY, 
US (holo). 

Grindelia rubricaulis DC. var. elata Steyerm., Ann. Mis 
souri Bot. Gard. 21: 227. 1934. J. T. Howell 6574; 
29 May 1931; U.S.A., California, Ventura Co., 7 mi. 
SW of Simi; CAS (holo), MO. 

Grindelia rubricaulis DC. var. latifolia (Kell.) us 

:9 


=m 
No) 
ES 


erm., ] ur : 

. A. Eastwood 794; 13 June-3 July 1906; Yd 

RR Casmailia; CAS, GH (holo), US. 

Grindelia rubricaulis DC. var. latifolia (Kell.) Steyerm. 
f. pubescens Steyerm., Ann. Missouri Bot. Gard. 21 
589. 1934. K. Brandegee s.n.; 14 June 1909; 1k SA; 
California, Monterey Co., between Lucia and Little's 
Hot Springs; UC (holo). 

Grindelia rubricaulis DC. var. latifolia (Kell.) Steyerm. 
f. d Steyerm., Ann. Missouri Bot. Gard. 21 0 

. C. P. Smith 1016; 10 July 1905; U.S.A., Cal- 


ifornia, Monterey Co., Pacific Grove, coast above Chi- 


s DC. var. permixta Steyerm., 


. nn 
Missouri Bot. Cani 21: 582. 1934. J. T. Howell 4352; 


22 July 1929; U.S.A., California, Alameda Co., San 
Leandro; CAS (holo 
Grindelia squarrosa (Pursh) Dunal f. iao) Steyerm., 
Ann. Missouri Bot. Gard. 21: 480. 4. J. A. Moore 
A. Steyermark 3768; 23 i 1931; U.S.A., 
Utah, Salt Lake Co., 5 mi. W of Salt Lake City; de- 
pository not designa 
Grindelia squarrosa Pusch) Dunal var. nuda (Wood) A. 
Gray f. Missouri Bot. Gard. 


Sockney on | Quitaque Rosdi CAS, MO (holo), NY. 
Grindelia stricta DC. var. aestuarina Steyerm., Ann. 
Pes Bot. Gard. 21: 560. 1934. J. M. Grant s.n.; 
June 1928; U.S.A., Washington, nua! n RM (holo). 
BU. stricta DC. var. aestuarina Steyerm. f. elon- 
"d T nn. Missouri Bot. Gard. 21: 560. 
EX Powel 7545; 27 July 1931; Canada, 
“sel Columbia, Vancouver Island, Arm of Victoria 
Harbor 3 mi. from Victoria on Island Hwy.; CAS (holo), 

MO (1016655). 

Grindelia stricta DC. var. eeu Steyerm., Ann. 
Missouri Bot. Gard. 21: 559. 1934. J. A. Moore & J. 
A. Steyermark 3687; 9 Aug. 1931; U.S.A., California, 
Humboldt Co., 10 mi. N of Trinidad; MO tha lo). 

Grindelia sublanuginosa Steyerm., Ann. Missouri Bot. 
Gard. 21: 458. 1934. E. W. Nelson 6523; 5 Jan. 
1903; Mexico, Jalisco, near Lake Chapala; NY, US 

lo 


Grindelia tenella Steyerm., Ann. Missouri Bot. Gard. 21: 
934. Berlandier 766 (2186) ; Nov. 1830; Mex- 
ico, Tamaulipas, from Victoria to Tula; GH (holo), NY. 
xir rayo SPEE am. f. flavida Steyerm.; Rhodora 55: 
3. J. A. Steyermark 70126; 21 Aug. 1950; 
n Shelby Co., route 36, 4.4 mi. NW 

of see de limit of Lentner; F (holo). 

Hieracium clivorum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 259. 1947. J. . 
50065; 4 Aug. 1942; Guatemala, 

e San Juan Ixcoy, Sierra i» los Cuchu- 

o). 


ndl. & Steyerm., Field Mus. 
, Bot. Ser. 23(2): 100. 1944. J. 4. Steyer- 

815; 22 va 1940; a Dept. Que- 
dd Volcán Santo Tomás; 3,0 ,200 m; F 
(holo) = H. ponencias Standl. & a. 1970. 

m na DA Wedd. f. tamaense Steyerm., Fieldi- 
ana Bot. 28(3): 641. 1953. J. A. Steyermark 57215; 
14 July. 1944; Mare Táchira, peti between 
Quebrada de Palmar and Quebrada de Paramito, base 
of Páramo qe Tam a 500 m; F (holo). 

Hieracium g Standl. & Steyerm., pu Mus. 
Nat. M Ser. 23(2): 101. 1944. J. . d eur 
mark 34860; 22 Jan. 1940; Guatemala, Dept. Que 
seran, e ens 3,000-3,800 m; F (holo). 

Hi Standl. & Steyerm., Field Mus. Nat. 

, Bot, Ser. 23(2): 102. 1944. J. A. Steyermark 
3276 ; 5 Dec. 1939; Guatemala, Dept. Jalapa, be- 
tween Miramundo and summit of Montana Miramundo; 
2,000-2,500 m; F (holo) = H. irazuense Benth., 1976. 

Hieracium minarum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(2): 103. 1944. J. A. Steyermark 
29722; 11 Oct. 1939; Guatemala, Dept. Zacapa, Sier- 
ra de las Minas; 1,000-1,500 m; F (holo) = H. gronovii 
L., 1976. 

Hieracium ostreophyllum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(2): 104. 1944. J. A. Steyer- 


664 


Annals of the 
Missouri Botanical Garden 


mark 36690; Feb. 1940; Guatemala, Dept. San Mar- 
cos, 6 mi. SW of Tajumulco, NW Volcán dle aac 
2,300-2,800 m; F (holo) = H. a Ep ostreo 
phyllum (Standl. & Steyerm.) Beaman 
Hogan dario Standl. & Ste RE: Field Mus. 
Na Sou 23(2): vn 1944. J. A. Steyer- 
E Feb, 1940; Benter Dept. San 
n Sibinal and su of Volcán Be 
F (holo) = Il. in irazuense Benth., 
Hymenostephium viride Steyerm o Bot. TN 
641. 1953 A. Steyermark 570 : 
Venezuela, Mér da 
Jungs pt Standl. & Ste 
r. 23(3): 142. 1944. H. von Tuerck- 
71903; Guatemala, Dept. Alta Vera- 
paz, Tactic; 1, 800 m; F (holo). 
Jungia reticulata Steyerm., Fieldiana Bot. 28(3): 642. 
Ste »yermark 57310; 15 July 1944; Ven- 
e "Táchira: base of Páramo de Tamá; 2,500 
2 895 m; F (holo). 
Laphamia D eda iU Ann. Missouri Bot. 
ard. 19: 392 2. J. A. Moore & J. A. Steyermark 
3547; 20 July rai U.S.A., Texas iri rson Co. 
Guadalupe Mounta trick Canyon; GH 
(h vo Perityle aa Sigjecn) fa din 
198 


heim 8410; 


isum. s ense Standl. & Steyerm., Field Mus 
Nat. Hist., Bot 23(1): 27. 1943. J. A. oa 
30543; Feb. 1 P Guatemala, Dept. San Marcos, SW 
of P eo NW Volcán Tajumulco; 2,300- 2,500 
m; F (ho 


Liatris cylindracea Michx. f. bartelli Steyerm., Rhodora 
59: 23. . K. E. Bartel s.n.; 13 Aug. 1955; U.S.A., 
Indians, Lake Co., lake sand along Side road, Clark 

, NW of Gary; F (holo). 
Lucilia venezuelensis Steyerm., Fieldiana Bot. 28(3): 
4 5: A. Ste yermark 35637; 13 Mar. 1944; 


Venezuela, rior upper headwaters of Rio Torno. 
below El Aguila and above Tea 3,650-3,965 
m; F (holo 

Mikania auyantepuensis S. F. Blake 
ata Steyerm., Poen Bot. 28(3): 6 
Steyermark 58125; 31 Aug. 1944; 


var. Apo e nt 
4 


T.F. 


cidem 


Amazonas, Cerr Dida: 1,675 m; F (holo). 
ikania Sy sd Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 260. 194 A. Steyermark 


43292; 26 A 1942; Guatemala, Dept. Zacapa, Sier- 
ra de las Minas, Volcán Gemelos; 3,000 m; F (holo) 
Mikania vus nsis ire & VE T od Mus 
Nat. Hist., Bot. Ser. 23(2): 105. 1944 M e 
40; 25 Feb 1905; Guatemala, Dept. sed Puerto 
rrios, sea level; F (holo) = M. tm var. gua- 
temale nsis (Standl. € Steyerm.) L. Williar ns, 1975. 
h ookeriana DC. var. 
. 28(3): 658. 1953. J. 4. j 
; 10- T Nov. 1944; Venezuela, Bolivar, Ptari 
iere 1 585-1,600 m; F (holo 
Mikania huitzensis Standl. i Steye rm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5); 2 1947. J. A. Steyermark 
48662; 14 July 1942; Cede Dept. Huehuete- 
nango, bes de los Cuchüitnatanes: Cerro Huitz; 1,500- 
2,600 F (holo) = M. cordifolia (L.f.) Willd., 1976. 
Mikania lucida F. Blake f. hirticaulis Steyerm., Fieldi- 
peg Bot. 28(3): 659. 1953 . Steyermark 60735; 
8 Nov. 1944; Venezuela, Bolívar, Sororopan-tepul; 
r pe^ m; F (ho 
Mikania eds ensis Steyerm. & Maguire, Acta Bot. 


Venez. 14(3): 28. 1984. B. Maguire et al. 65616; 
15 Jan. 1981; Venezuela, T.F. Amazonas, Dept. Ata 
apo, Cerro Marahuaca; 2,685 m; VEN (holo), NY. 

Mikania mic iro rine Steyerm., Ann. Missouri Bot. 
Gard. 71: 339. 1984. J. 4. Steyermark 129456; 10 
Oct. 1983; Gael, T.F. Amazonas, Cerro Mara- 
huaca; 2,560 m; VEN (holo). 

Mikania es Standl. & Steyerm., Field Mus. Nat. 

ist., Bot. Ser. 23(5); 261. 1947. J. A. siena 
47257; 7 a 1942; Guatemala, Dept. Sololá, Volcán 
San Pedro, Lago de Atilan, above San Pedro; 2,500 
2,800 m; F (holo). 
Mikania phelpsii Maguire & Steyerm., Brittonia 7: di 
A. Steyermark 59527; 30 Oct. 1944; Ver 
venale: Bolívar, Ptari- -tepui; 1,810- RS 100 m; F (holo) 
= M. lucida S. F. Blake, 1939, 1 
Mikania robinsoniana nm Fieldia iana Bot. 28(3): 
, nom. nov. = M. nigropunctata var. cryp- 
dona Robins: 1964. 
Mikania (pd dn d wi Bot. 28(3): 660. 
1953. mark 5 4 Nov. 1944; Ven- 
ezuela, Bolivar: Puri -tepul; os d m; F (holo) = M. 
sprucet Baker, 1964. 
Oliganthes roraimensis Steyerm., ase Bot. 28(3): 
2. 1953. J. del Steyermark 58678; 25 Sep. 1944; 
Venezuela, Bolivar, Mt. Roraima; os m; F d zz 
Pollalesta roraimensis (Steyerm.) Aristeg., : I 

UN marahuacense Fi & Magu im 
Bot. Venez. 14(3): 28. 1984. J. 4. EE et al. 
125925; 31 Jan. Tos Venezuela, T.F. Amazonas, 
Dept. Atabapo, Cerro Marahuaca; 2,580 m; NY, VEN 
(holo). 

Oyedaea blakeana Steyerm., doy Bot. "eh 663. 
1953. J. A. Steyermark 58047; 16 Aug. 1944; Ven- 
ezuela, T. F. Amazonas, ind Duida; 1,095- cd pM m; 
F (holo). 

Oyedaea verbesinoides DC. var. glabrior Steyerm., 
Fieldiana Bot. 28(3): 664. 1953. J. A. Steyermark 
61612; * Mar. 1945; Venezuela, Anzoátegui, Cerro 
Peonia; 1,800- 2, 000 m; F (holo). 

Oyedaea Nui) ^s DC. var. hypomalaca Steyerm., 
Fieldiana Bot. 28(3): 664. 1953. J. A. Steyermark 
62095; 15 Apr. 1945; Venezuela, Monagas, Cerro 
Negro; 1,500-2,180 m; F (holo). 

Pectis flava Standl. & Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 23(2): 106. 1944. J. A. Steyermark 304 15; 
24 Oct. 1939; poster: Dept. Jutiapa, along railroad 
between Agua Blanca and Amatillo; 950-990 m; F 
eeg P. uniaristata DC. var. holostemma A. Gray, 


T venezut , Fieldiana Bot. 28(3): 666. 
1953. J. A. (paid 57528; 31 Ju y 1944; Ven- 
ezuela, Bolivar, 5 km W 

decia menium inamoenum Standl. & Stey 

. Hist., Bot i 
Mae 29737; Oct. 
Sierra de "m MIEL Sa Alejandria; 1, 000- l, 500 
m; F (holo). 

Perymenium jalapanum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(3): 144. 1944. de . Steyer- 
mark 32658; Guatemala, Dept. Jalapa, between Mura- 
mln and su of Montana MEL: 2,000- 

2,300 m; F (holo 2 

Pipmoaipha a Aristeg. S Steyerm., Bol. 
Venez. Ci. Nat. 32: 402. 1976. J. A. ir " 

5; 22-28 Feb. 1974; Venezuela. Bolivar, 

Meseta del Jaua, Cerro Jaua; 1,800 m; F (holo). 


= 


Volume 76, Number 3 
1989 


Taylor 665 
Plants Described by 


Julian A. Steyermark 


Polymnia pou Steyerm., Fieldiana Bot. 28(3): 667. 
1953. J. A. Steyermark 57519; 22 July 1944; Ven- 


06. 1944. J. A. Steyermark 36181; 20 

Feb. 1940; quat Dept. San e sd between La 

Vega Ridge along Rio Vega, Volcán Tacaná; 2,500- 

— Rumfordia pcne] nel 
Standl. & Steyerm., 1947 

Polymnia verapazensis Standl. € Steyerm., Field Mus. 
Nat . Hist., Bot. Ser. 23(3): 144. 1944. H. von Tuerck- 


p = Ru umfordia guate- 

malensis (Coult.) S. F. Blake, 1976. 

chia bracteata Maguire, Steyerm. & Wurd., Mem. 
New York Bot. Gard. 9(3): 435. 1957. J. A. Steyer- 
mark & J. J. Wurdack 683; 9 Feb. 1955; Venezuela, 
Bolivar, Chimantá Massif; 1,970 m; NY (holo), F. 

Quelchia cardonae Steyerm., Fieldiana Bot. 28(3): 670. 

53. F. Cardona 2288; Oct. 1947; Venezuela, Bo- 
copan, Rio Caroni; 2,200 m; F (holo). 


mark & J. J. W 
Bolivar, Chimanta Massif, Torono-tepui; 2,165- 2,180 
pl F (holo), NY. 

Quel iiy x grandifolia Maguire, Steyerm. & Wur 
Mem. New York Bot. Gard. 9(3): 436. 1957. J. V. 
CHI ark & J. J. Wurdack 756; 12 Feb. 1955; 
Venezuela, Bolivar Chimantá Massif; 2,185 m; F (holo). 

Rojasianthe Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 224) 411. 1940, gen. nov. 

Rojasianthe superba Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot Ser. 22(4): 315. 1940. J. A. Steyer- 
mark 35835 5; 16 Feb. 1940; Guatemala, Dept. San 
Marcos, Volcán Tajumulco; 3,300-3,900 m; F (halo): 

Rudbeckia hirta L. f. homochroma Steyerm., Rhodora 
40: 179. 1938. J. A. Steyermark 11345; 3 July 1936; 
U.S.A., Missouri, Oregon Co., 10 mi. W of New Lib- 
erty, Fristoe losehase Ünit, Clark National Forest; MO 
(holo). 


Sabazia urticifolia (H.B.K.) DC. var. A Stey- 
3. J. A. Steyer- 

1944; ost Lara, between 
d Los BN above Humocaro 


ndl. & Steyerm, Field Mus 
, Bot. E 23(3): 145. 1944. P. C. Standley 
8 Dec 8 


= Nov. 1939; Guatemala, Dept . Jutiapa, Laguna Ret 
ana; 600 m; F (holo) = Macianghla “andlon 
(Steyerm. ) King & Robs., 


Schkuhria viscosissima Sta 


dl P QU Wy Field Mus 


a 
No 
nn 
MM 
U 
— 
A 
— 
4A 
— 
w 
2 
- 
ue 
r= 9 
bo 29 
Sao 
[a] 
p 
+ 
c 
ag 
= 
D pP 
J 
c 
(OR 


pt. Quic 

near Sacapulas l 040-1, m; F (holo) = Florestina 
pedata (Cav.) Cassini, ere 

Siegesbeckia nudicaulis Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 262. 1947. J. A. Steyer- 


mark 50114; 5 Aug. 1942; Guatemala, Dept. Hue- 


Silphium perfoliatum L. f. petiolatum Palmer & Stey 
erm., Brittonia 10: 119. 1958. J. A. Steyermark 80474; 


19 Oct. 1955; U.S.A., Missouri, Stone Co., above 


Silphium terebinthinaceum Jacq. var. lucy-brauniae 
Steyerm., Rhodora 53: 134. 1951. E. L. Braun s.n.; 
17 Aug. 1950; U.S.A., Ohio, Adams Co., 2.5 mi. N 
x Ohio route 348, N part of Jefferson Top, SE of 

rub Ridge; Herb. Braun (holo). 

Soliva anthemidifolia R. Br. var. lanuginosa Steyerm. 
Fieldiana Bot. 28(3): 672. 1953. J. A. Steyermark 
55506; 12 Feb. 1944; Venezuela, Lara, between Bue- 
nos Aires to Canyon of El Callado; 2,285-2,740 m; F 
(holo 

Stenopadus affinis Maguire, Steyerm. & Wurd., Mem. 
New York Bot. Gard. 9(3): 427. 1957. J. A. Steyer- 
mark 75110; 18 Apr. 1953; Venezuela, Bolivar, Chi- 
manta Massif, Abacapa-tepui; 850- 1,100 m; F (holo), 
NY. 


Stenopadus chimantensis Maguire, Steyerm. & Wurd., 
Mem. New York Bot. Gard. 9(3): 1957. J. A. Stey- 
ermark & J. J. Wurdack 652; 9 Feb. 1955; a 
Bolivar, bua Massif, Torono-tepui; 2,165-2, 

m; NY (ho 

Stenopadus pe e PA Cuatr. & Steyerm., Bot. Mus. 
Leafl. 17(3): 99. 1955. R. E. Schultes & I. Cabrera 
15031; 23-25 Jan. 1952; Colombia, Comisaria del 
Ma Río Kananari, Cerro Isibukuri, near summit; 


F (holo). 

Stenopadus talaumifolius S. F. Blake var. magnifolius 
Steyerm., Fieldiana Bot. 28(3): 673. 1953. F. Cardona 
2190; er 1947; Venezuela, Bolivar, Cerro Arepuchi; 
600 m; US (holo). 

ac chortiana Standl. & Steyerm., ds Mus. Nat. 

, Bot. Ser. 23(2): 108. 1944. J. 4. Steyermark 
30866; Oct. 1939; Guatemala, Dept. E be- 
tween Finca San Jose and Montana Nube; 1,200- 

— S. hirsuta var. chortiana (Standl. 

m.) Grashoff, 1974 

Stevia E op ie Standl. & probar Field Mus. Nat. 
Hist., Bot. Ser. 23(1): 27. 1943. A. Steyermark 
34757; Jan. 1940; Guatemala, Dept. Quezaltenango, 
Volcán Santo Tomás; 2,500-3,700 m; F (holo) — 

Mb in Schultz-Bip., 1976. 
Stomato chasta colveei Steyerm., Brittonia 32: 21. 1980. 


Tagetes sororia Standl. eyerm., Field Mu 
ist., Bot. Ser gr 146. 1 P. C. Standley 
85228; 29 Jan. 194 dvi Dept. Quezalte- 
nango, N of reed 2,500-2,800 m; F (holo) — 
T. nelsonii Greenm., 197 


l 
Trixis amphimalaca Standl. & Steyerm., Field Mus. Nat. 


Hist., Bot. Ser. 23(3): 147. 1944. P. C. Standley 
82538; 6 Jan. 1941; Guatemala, ee Hue busts 
nango, N of Chiantla; 2,000-2,250 m; F (holo) = 


nelsonii Greenm., 1976. 
Verbesina agricolarum Standl. & Steyerm., Field Mus 
t. Hist., Bot. Ser. 22(4): 319. 1940. J. R. pcdes 
1026; 21 Sep. 1937; Guatemala, Dept. Chimalte- 
nango, Finca La Alameda; F (holo 
me angusta Maguire, Steyerm. urd., Mem. 
ork Bot. Gard. 9(3): 426. 1957. J. A. Steyer- 
nb & J. J. Wurdack 851; 13 Feb. 1955; Venezuela, 
Bolivar, Chimantá Massif; 2,120 m 
Verbesina apleura S. F. Blake var. piens Standl. & 
Steyerm., Field Mus. Nat. Hist., Bot. Ser. 23(2): 109. 


666 


Annals of the 
Missouri Botanical Garden 


1944. P. C. Standley 67540; Mar. 1939; Guatemala, 


eana Steyerm., Fieldiana Bot. 28(3): 674. 
A. Steyermark 56300; 4 May 1944; Ven- 
ezuela, Mérida, a Mucutuy and Mucuchachi; 
1,065-2,430 m; F (ho 
Verbesina — Ai 8 Steyerm., Field Mus. Nat. 
on 23(5): 262. 1947. J. A. Steyermark 
0132; 6 pem 1942; Guatemala, Dept. Huehuete- 
rango, Sierra de los Cuchumatanes; 3,700 m; F ee 
feb na exalata Steyerm., Fieldiana Bot. 28(3): 6 
A. Steyermark 55176; 5 Jan. 1944; Vene- 
ibus Distr. Federal, above amacak Cordillera del Avi- 
la; 1,065-1,520 m; F (holo) = V. turbacensis H.B.K., 


Verbesina guatemalensis Robinson & Greenmann var. 
. & Steyerm., Field Mus. Bot. Ser. 
A. Steyermark 29505; Oct. 
1939; B. Gustemala, Dept. Zacapa, Sierra de las Minas; 
O m; F (holo) = V. guatemalensis H. Robinson 
ann var. guatemalensis, 1976. 
I yes minarum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 263. 1947. J. A. Steyermark 
42480; 12-13 Jan. 1942; Guatemala, Dept. Zacapa, 
Sierra de las Minas, Río Repollal; 2,100-2,400 m; F 
(holo). 


gel dee negrensis Steyerm., ee Bot. 28(3): 675. 

Steyermark 57( 7 July 1944; Vene- 

abe Mérida, between peta and Pirame de la 
Negra; 2,285- 2,430 m; F (holo). 

Verbesina petzalensis Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(3): 147. 1944. P. C. Standley 
82921; 9 Jan. 1941; Guatemala, Dept. Huehuete- 
nango, near crossing of Rio San Juan Ixtán, E of San 
Rafael Petzal;. 1,730 m; F (holo). 

Verbesina tachirensis Steyerm., Fieldiana Bot. 28(3): 
677. 1953. J. A. Srevermatk 57149. 12 July 1944; 
Venezuela, Táchira, between Las Delicias and Para 
guita, bordering Rio Táchira; 1,675-1,890 m; F (holo). 

Henna erra LT f. albiflora Steyerm., Brittonia 
10: E. J. Palmer 61259; 7 Sep. 1955; 
U.S.A., Miso Lawrence Co., along Turnback Creek 
and U.S. Hwy. 166, 3.5 mi. E d Chesapeake; F (holo). 

Vernonia na Stand l. & Steyerm., Field Mus. 
Hist., Bot. Ser. 22(5): 395. 1940. J. A. Steyermark 
36787; 27 Feb. 1940; Guatemala, Dept. San Marcos, 
Volcán Tajumulco; 1,800-2,500 m; F (holo). 

Wenn. medialis Standl. & Steyerm., Field Mus. Pu 
Hist., Bot. Ser. 23(3): 148. 1944. P. C. Standley 
87473; 17 Feb. 1941; d Dept. aie. 
Rio Coyote, along road 4 of Tetalhuleu; 300 
m; F (holo) = V. arborescens is (L) Sw., 1982. 

Vernonia mima Standl. & Steyerm., Field Mus. Nat. 
Hist., r Ser 23(5): 264. 1947. P. C. Standley 
82871; 9 Jan. 1941; Guatemala, Dept. Huehuete- 
ango, near crossing of Rio San Juan Ixtán, E of San 
Rafael Petzal; 1,730 m; F (holo). 

Vernonia oo Raf. f. swinkii Steyerm., Rhodora 
9 à . F. A. Swink 2775; 11 Aug. 1955; 
US. a T Cook Co., prairie S of 95th St. and W 

45, oo Forest Preserve S of Willow 
O10 


i Na t. Hist., Bot. 

26 July 1937; 
Md breed near San Toss: Distr. Bocoyna; 
2,400 r (ho 

Zexmenia eae Standl. & Steyerm., Field Mus. Nat. 


Hist., Bot. Ser. 23(3): 149. 1944. P. C. Standley 


— Wedelia pinetorum (Standl. & Steyerm.) Becker, 
1975 


BALSAMINACEAE 


Impatiens cr Nutt. f. dichroma Steyerm., Rhodora 
8. J. A. Steyermark 25927; 6 Sep. Pads 
U.S.A., Mana Pike Co., Mississippi River, 1.5 
NW of Louisiana; MO (ho la ). 


BERBERIDACEAE 


Berberis ar go Standl. & diee Field Mus. Nat. 
22(3): 140. 1940. J. R. Johnston 
1525; 14 Des 1938; Guatemala, Dept. Sa catepéquez, 
Parramos; F (holo) = Mahonia uem (Standl. & 
Steyerm.) Standl. & Steyerm., 
Mahonia volcan ne v Steyerm., Field Mus. Nat. 
Hist., Bot. 943. P. C. Standley 6. 5221; 
Agua, above Santa 


Podophyllum Erase Lor lmoreanum Steyerm. T 
Rhodora 54: ud V. Border . Steyer- 
mark s.n.; b D 1950; U.S.A., Illinois, Lake Co., 
Biltmore Estates, S of ot n Rd. and W of Kimberly 
Rd., 5 mi. N of Ba iene. F (holo). 


BETULACEAE 


Corylus americana Walt. f. indehiscens Palmer & Stey- 
erm., Ann. Missouri Bot. Gard. 22: 516. 1935. B. F. 
Bush 10332; 5 Sep. 1924; U.S.A., Missouri, Jackson 
Co., Greenwood; GH (holo). 


BIGNONIACEAE 


os standleyi Steyerm., Field Mus. Nat. 
, Bot. Ser. 23(5): 236. 1947. J. A. Steyermark 
33533; 5 Jan. 1940; Guatemala, Dept. Quezaltenango, 
Finca Pirineos and Finca Soledad, Volcán Santa María; 
1,300-1,400 m; F (holo) = Mansoa standleyi (Stey- 
erm.) A. Gentry, 1982. 


BOMBACACEAE 


Catostemma clarkii Steyerm., Ann. Missouri Bot. Gard. 
74: 638. 1987. H. A. Clark 8126; 19 Aug. 1981; 
m NE of San Carlos 


2 Feb. 1949; Venezuela, T.F. Amazo erro Sipapo 
(Paráque), water course above Intermediate Camp; MO 
(holo), NY. 

E Y eri Steyerm., Ann. Missouri Bot. 
Gard. A. Steyermark 74781; 30- 
31 ur pect Venssucl, Bolivar, Chimanta Massif, 
NW slopes > Abacapa, vicinity of camp 1 along Rio 
Abácapa-tepui; F, MO (holo), VEN. 

Catostemma marahuacensis Steyerm., Ann. Missouri Bot. 
Gard. 74: 643. 1987. J. A. Steyermark & B. Holst 
130878; 28 Feb.-1 Mar. 1985; Venezuela, T.F. Ama- 
zonas, Dept. Atabapo, Cerro Marahuaca, “Sima Camp' 
south central portion Caño Negro; 1,140 m; MO (holo — 
3327355). 

Catostemma pubistylum Steyerm., Ann. Missouri Bot. 
Gard. 74: 643. 1987. H. L. Clark & P. Maquirino 
6742; 3 Aug. 1978; Venezuela, T.F. Amazonas, IVIC 


Volume 76, Number 3 
1989 


Taylor 667 
Plants Described by 


Julian A. Steyermark 


main study site, 4.3 km NE of San Carlos de Rio Negro; 
119 m lo). 
itisivmian sancarlosiana Steyerm., Ann. Missouri Bot. 
Gard. 74: 644. 1987. H. L. Clark & K. Clark 8117; 
16 Aug. 1981; Venezuela, T.F. Amazonas, 10.8 km 
NE of San Carlos on Solano Road; 119 m; MO (holo — 
3142831). 
Pochota aracamuniana Steyerm., Ann. Missouri Bot. 
= L. Liesner & G. Carnevali 
; Venezuela, T.F. Amazonas, 
Dept. Rio Negro, Cerro Aracamuni, summit, Proa Camp; 
1,400 m; MO (holo), V 
m ewelii Steyerm., don. Missouri Bot. Gard. 75: 
77. 1988. J. Ewel 135; 23 Apr. 1964; Brazil, Dept. 
ADU near Venezuela frontier, camino al Cerro 
Neblina desade Rio Tucano; 1,250 m; MY (holo). 
Pochota ee Steyerm., Ann. Missouri Bot. Gard. 
75: 1077. L. Liesner 18452; 8-9 Mar. 
1985; sea TF. Amazonas, Dept. Atabapo, Ce- 
rro Marahuaca; 1,100 m; MO (holo— 3483071), VEN. 
Pochota gracilis (Robyns) Steyerm., subsp. bolivarensis 
Missouri Pu Gard. 75: 1078. 1988. 
; 17-29 Mar. 1967; Vene- 
subs. Bolivar, Rio — Rui southern base of Cerro 
Sarisarinama, Meseta de Jaua; 400 m; NY, VEN (holo). 
Pochota liesneri Steyerm., Ann. Missouri Bot. Gard. 75: 
1079. 1988. R. L. Liesner 18637; 14 Mar. 1985; 
Venezuela, T.F. Amazonas, Dept. Atabapo, Salto Yu- 
reba, Cerro Yureba, lower Ventuari; 350 m; MO (holo — 
3483072), VEN. 
Pochota mawarinumae Steyerm., Ann. Missouri Bot. 
Gard. 75: 1079. 1988. R. L. Liesner 16355; 2 Mar. 
1984; Venezuela, T.F. Amazonas, Dept. Rio Negro, 
near Cerro de la Neblina Base Camp, Rio Mawarinuma; 


mp, 
140 m; MO (holo — 3483069, 3483068), VEN. 


19 Feb. 1987; Venena T.F. sonas, Dept. Atures, 
stream 0.5-2 km E of Rio Coro-Coro, W of Serranía 
de Yutaje; 200 m; MO (holo — 3483070), NY, VEN. 

Pochota robynsii Steyerm. & Stevens, Ann. Missouri 
Bot. Gard. 75: 1586, nom. nov. 

Pochota tepuiensis Steyerm., Ann. Missouri Bot. Gard. 
75: 1081 8. J. A. Steyermark et al. 108938; 
11-12 Feb. 1974; Venezuela, Bolivar, Meseta de Jaua, 
Cerro Sarisarinama, summit; VEN (holo). 

Scleronema — Steyerm., Ann. Missouri Bot. Gard. 
74: 645. Maguire et al. 60181; 15 Nov. 
1986; Brazil ndn Serra de Neblina, vicinity Base 
LP Cano Tucano, Rio Cauaburi; 100 m; MO (holo), 


BONNETIACEAE = Theaceae 
BORAGINACEAE 


Lithospermum canescens (Michx.) 


Lehm. f. pallidum 
6. 1955. J. 


Palmer & Stey , Rhodora 57: 31 A. 
Steyermark 73223; E Ey 1952; U.S.A , Missouri, 
"er Co., Hwy. 54, i. SW of jct. with Hwy. 15 

. WSW of Cyl F (holo), MO. 


Me Pais rng e s. f. rosea Steyerm., Rhodora 
62: 131. 1960 um e pneus ark 67235; 17 Apr. 
1949; U.S.A., Miss 1, Cass Co., N side of Big Creek, 

4 mi. SE of RE Hill: F (holo). 


BRASSICACEAE 


Cardamine balnearia Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(4): 157. 1944. P. C. Standley 


83332; 14 Jan. 1941; Guatemala, Dept. Quezalte- 
nango, Aguas Amargas, Volcán Zunil; 2,450 m; F 
olo). 
Cardamine bulbosa (Schreb.) Britt., Sterns & Pogg. f. 
fontinalis png & Steyerm. ges Missouri Bat. Gard. 
25: 771. 1938. J. A. Steyermark 12414; 2 Aug. 
1936; U.S. T Missouri, Shannon Co., Pulltight Spring, 
near Current River; MO (holo—1175572) = ul- 
bosa (Schreb.) Britt., Sterns € Pogg var. a Deea 
1981. 


Cardamine eremita Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(2): 53. 1944. J. A. Steyermark 
50143; 6 Aug. 1942; Guatemala, Dept. Huehuete- 
nango, between Tojquiá and Caxix bluff, summit of 
Sierra de los Cuchumatanes; 3,700 m; F (holo). 

Cardamine jejuna Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(2): 54. 1944. J. A Tag died 
50569; 15 Aug. 1942; Guatemala, Dept. Huehuete- 
nango, Cerro Pixpix, Sierra de los HOMER da 2,800 
m; F (holo). 

Romanschulzia alpina Standl. & Steyerm., Fieldiana 
Bot. 24(4): 377. 1946. J. A. Steyermark 50144; n.d.; 
Guatemala, Huehuetenango, Sierra de los Cuchuma- 
tanes, between Tojquia and Caxin bluff; F (holo) = 
Sisymbrium standleyi Rollins, 1956. 

Selenia oinosepala Steyerm., Field Mus. Nat. Hist., Bot. 
Ser. 17(5): 422. 1938. E. Whitehouse s.n.; 28 Feb. 


1930; U.S.A., Texas, San Benito; F, TEX (holo). 


BROMELIACEAE 


Aechmea siapensis L. B. Smith & Steyerm., Ann. Mis 
souri Bot. Gard.: in press. 1989. R. L. Liesner d ` 
Delascio 21904; 13 Oct. 1987; Venezuela, T.F. A 
zonas, Rio Siapa, 130 km E of San Carlos de Río 
Negro; 180 m; in US (holo), VEN. 

Billbergia manarae Steyerm., Brittonia 30: 39. 1978. 
J. 4 eraman 112181; 22 May a. A MN 


Amazonas, Dept. Ata 
(2911061 & 2911060), NY, US (holo), VEN. 
Brewcaria marahuacae L. B. Smi 
Robinson, Ann. Missouri Bot. 
A. Steyermark et al. 126328; 9- 10 Feb. 1982: 
Venezuela, T.F. Amazonas, Dept. Atabapo, Cerro 
Marahuaca — Atuha —Shiho; 2,480 m; NY, US (holo), 
VEN. 


Brocchinia oliva-estevae Steyerm. & L. B. Smith, Ann. 
Missouri Bot. Gard. 74: 85. 1987. F. Oliva Esteva 


1984; Venezuela, Bolivar, summit of A 


9610; 18 Feb. 1984; Venezuela, Bolívar, Auyan-tepui; 
2,100 m; NY, VEN (holo) = Lindmania smithiana 
(Steyerm. & Luteyn) L. B. Smith, 1986. 

Connellia varadarajanii L. B. Smith & Steyerm., J. 
Bromel. Soc. 35(2): 52. 1985. G. S. Varadarajan 
1205; 12 Jan. 1984; Venezuela, Bolivar, Auyan-tepui; 
1,950-2,000 m; US (holo), VEN. 

Cottendorfia arachnoidea L. B. Smith, p & H. 
Robinson, Acta Bot. Venez. 14(3): 9. 1984. J. 
Steyermark et al. 125996; 1-2 Feb. 1982; Venezuela, 


668 


Annals of the 
Missouri Botanical Garden 


T.F. Amazonas, Dept. Atabapo, Cerro Marahuaca Fhuif; 
2.480-2,500 m; US (holo), NY, VEN = Lindmania 
d gue e a Smith, Steyerm. & H. Robinson) 
L. B. 5m 

Cottendorfa atrorosea L. B. Smith, Steyerm. & H. Rob- 
inson, Brittonia 33: 28. 1981. J. A. Steyermark et al. 
117982; 9 Apr. 1979; Venezuela, Bolivar, cumbre del 
Cerro Guaiquinima; 1,650 m; US (holo), VEN — Lind- 
mania atrorosea (L. B. Smith, Steyerm. & H. Rob- 
inson) L. B. Smith, 1986. 

e aurea L. B. Smith, Steyerm. & H. Robinson, 
ined., mss. = Lindmania aurea L. B. Smith, Steyerm. 

i eg 

Cottendorfia no [cde var. angustior L. B. Smith, 
Steyerm. & nson, Brittonia 33: 28. 1981. J. 
A. Steyermark et a 116180; 28 Feb. 1978; Vene- 
zuela, Bolivar, Auyan-tepui; 1, 950 m; US (holo), VEN 
ses m ania PL var. angustior (L. B. Smith, 

Ste . & H. Robinson) L. B. Smith, 1986. 

ad meas Le De Smith, Steyerm. & H. 

a ined., mss. J. 4. Steyermark & F. Delascio 
1292 EN 0 Mar LA 1983; Venezuela; MO (iso — 


3230 

alo discolor L. B. Smith, Steyerm. & H. Rob- 
Steyerbromelia discolor L. B. 
. Robinson. 


B. Smith, Steyerm : 
Venez. ES ) 15. 1 A. Steyermark et al. 
124526; 17 Feb. 1981; Muela T.F. Amazonas, 
Dept. Atures, Serranía Sipapo; 1,500 m; NY (holo), 
VEN = d nia geniculata var. minor (L. B. Smith, 
Steyerm. & H. xcu L. e oi 1986. 

Cottendorfi huberi L. B. teyerm. & H. Rob 
inson, ined., mss. — nd cle L. B. Smith & 
Steyerm. 

Cottendorfia marahuacae L. B. We ERE & H. 

. 14(3): 9. 1984. B. Ma- 

g 1981; M T.F. 

Amazonas, Cerro Marahuaca; MO (2911057), NY, US 
(holo) = "wu suni wp ae (L. B. Smith, Stey- 
erm. & H. Robinso . Smith, 1986. 

Cottendorfia terramarae L. B. Smith, Steyerm. & H. 
Robinson, ined., mss. = Lindmania terramarae L. B. 
Smith, Steyerm. & H. Robinson, 1986. 

p L B. Smith. & Steyerm., 

. 1968. J. 4. Steyermark & M. 

Lus: 1966; Venezuela, Táchira 

de la Negra on road to Pregonero; 2.600 m; 

VEN. 


Phytologia 
abe 96951; 
, below Páramo 


US (holo), 


iod mania membranacea L. B. Smith & e Acta 
. Venez. 2(5-8): 380. 1967. ermark 
922: 4 Mar. 1966; Venezuela, Paria jode iie ula, 
narrow northern crest between Cerro de Humo and 
Cerro La Pavita; 1,250-1,273 m; US, VEN (holo). 
mania terrestris L. B. Smith & ue ., Ann. Mis- 
uri Bot. Gard. 73: 715. 1 . J. A. der HA 
129596; 10-12 Oct. 1983; R T.F. Amaz 
nas, Dept. Atabapo, Cerro Marahuaca; 2, 560 m; NY, 
US (holo), VEN. 

Lindmania aurea L. B. Smith, nO & H. p 
Ann. Missouri Bot. Gard. 73: 697. 1986. J. A. Ste 
ermark et al. iie 5 Fe 1982; Venezuela, Bo. 
livar, Distr. o del Ca! 1,850 m; MO 
(3228804), NY, US (holo), VEN. 

Lindmania We Steyerm. & L. B. Smith, J. Bromel. 
Soc. 38(2): 51. 1988. B. K. Holst & F. Oliva Esteva 


3539; 22 E 1987; Venezuela, Bolivar, Distr. Piar, 


Murispán-tepui, summit; 2,300 m; F (holo), MO 
(3512232), NY, US, VEN. 

Lindmania huberi L. B. Smith & Steyerm., Ann. Missouri 
Bot. Gard. 7? 

f 983; "eneniela: I: 

zo del Chimantá; 1,850 m; MO (3228812), 

NY, US (holo), VEN. 

Lindmania imitans L. B. Smith, Steyerm. & H. 
inson, Ann. Missouri Bot. Gard. 73: 697. 1986. T "4 
Steyermark et al. 128474; 2-5 Feb. 1983; Venezuela, 

olivar, Distr. Piar, Macizo del Chimantá; 1,850 m; 
MO (3340851), US (holo), VEN. 

Lindmania piresii L. B. Smith, a & Luteyn, Ann. 
Missouri Bot. Gard. 73: 698. 1986. J. Murca Pires 
15010 (33) ; 10 Feb. 1975; Brazil Amazonas, Serra 
Araca; IPEAN, US (holo). 

Lindmania riparia L. B. Smith, Steyerm. & H. Robinson, 
Ann. Missouri Bot. Gard. 73: 695. 1986. J. A. Stey- 
ermark 93895; 15 May 1964; Venezuela, Bolivar, 
Auyan-tepui; 2,050-2,300 m; MAC, NY, US us 

Lindmania saxicola L. B. Smith, AO & H. 
inson, Ann. Missouri Bot. Gard. 73: 693. 1986. h FI 
Steyermark et al. 128945; 7-8 E 1983; Venezuela, 
Bolivar, Distr. Piar, Macizo del Chimantá; 2,450 m; 
US (holo), VEN. 

eU Pag is sessilis L. B. Smith, Steyerm. & H. Robinson, 

nn. Missouri Bot. Gard. 73: 693. 1986. J. A. Stey- 
emi et al. 128104; 26-29 Jan. 1983; buco 
Bolívar, Distr. Piar, Macizo del Chimantá; 2,000 n 
US (holo), MO (128104), VEN. 

Lindmania terramarae L. B. Smith, Steyerm. & H. Rob- 
inson, Ann. Missouri Bot. Gard. 73: 698. 1986. J. A. 
Steyermark & F. Delascio 129067; 26 Mar. 1983; 
Venezuela, T.F. Amazonas, Dept. ' Atabapo, Cerro 
Marahuaca; 2,800 m; MO (3230813), NY, US (holo), 
VEN. 


Navia albiflora L. B. Smith, fee hte & H. Robinson, 
9 


na 
Atabapo, Cur Duida; NY, US (holo), 


zonas; Dept. 
VEN. 


Navia aliciae L. B. Smith, Steyerm. € H. Robinson, 
t 3. 


T.F. Amazonas Dept. et Rio Cunucunuma; 180 
210 m; NY, US (holo), VEN. 
Navia ido L. B. Smith & J. A. Steyerm., Bol. Soc. 
Venez. Ci. Nat. 23: 62. 1962. J. A. Steyermark 89698; 
l Sep. 1961; Venezuela, Bolivar, Sierra de Lema, Sa- 
bana de Triana; 300 m; VEN (holo). 
Navia berrya ana L. B. Smith, Steyerm. Š H. Robinson, 
: A. Stey- 


ermark et al. 130318-A; 1-2 Mar. Ten Venezuela, 
T.F. Amazonas, Dept. Rio Negro, near an of Cerro 
Vinilla; 440-600 m; US (holo), VE 

Navia breweri L. B. Smit Steyerm., Acta Bot. Venez. 
2(5-8) 380. 1967. C. Brewer Jr. 248; 4 Jan. 1965; 
Venezuela, lower part of Cerro Jaua, near Río Cana- 
racuni; 600-700 m; US (holo), VEN. 

Navia carnevalii L. B. Smith & Steyerm., Ann. Missouri 
Bot. Gard.: in press. 1989. R. L. Liesner & G. Car- 
nevali 22322; 22 Oct. 1987; Venezuela, T.F. Ama- 

zonas, Cerro Aracamuni, Quebrada Camp; 600 m; MO, 
US (holo), VEN. 

Navia connata L. B. Smith & Steyerm., Bol. Soc. Venez. 
Ci. Nat. 25: 50. 1963. J. A. emu 90223; 29 
Dec. 1961; Venezuela, Sierra Ichun; 625-725 m; VEN 
(holo). 


Volume 76, Number 3 
1989 


Taylor 669 
Plants Described by 


Julian A. Steyermark 


Navia crassicaulis L. B. Smith, Steyerm. & H. Robinson 
Ann. Missouri Bot. Gard. 73: 708. 1986. R. L. Liesner 


Plateau (arm); 1,750-1,850 m; MO (3340862), US 

(holo). 

Navia culata L. B. Smith, s & H. Robinson, 
Ann. Missouri Bot. Gard. 73: 712. 1986. O. Huber 
6184; 13 June 1981; D T.F. Amazonas, Dept. 
Río Negro, ca. 20 km al SW de Mauaca; 760 m; US 
(holo), VEN. 

Navia delascionis L. B. E Dm & H. pass 
Ann. Missouri Bot. Gard. 709. 1986. J. A. Ste 
ermark et al. 130410; a A Mar. hi Venezuela, 
T.F. Amazonas, Dept. Río Negro, vicinity and no 

Ward from Cerro Vinilla; US (holo), MO (3340645) 

VEN. 


Navia emergens L. B. Smith, Steyerm. & H. Robinson, 
Brittonia T 31. 1981. J. A. Steyermark et al. 117298; 
24 May 1978; Venezuela, Bolivar, ro Guai- 
quinima, : km rio arriba del Salto Szczerbanari; 730- 
900 m; US (holo), VEN 
Navia filifera L. B. Smith, Steyerm. & H. Robinson, 
Ann. Missouri Bot. Gard. 73: 713. 1986. R. L. Liesner 
16025; 16-18 Feb. 1984; Venezuela, TF. Am Co. 
Dept. Río Negro, O Camp III; 1,750-1,850 m 
MÓ (3340858), US (holo). 

mith, a & H. Robinso 

984. J. A. Steyerm ark 
et al. 123846; 11-12, 14 . 1981; Venezuela, Bo- 
livar, pu Heres, Cerro Marutani; 1,200 m; NY, US 
(holo), V 

Navia a Stayer, & Holst, Ernstia 38: 44. 1986. 
J. A. Steyermark et al. 131850; 10 Apr. 1986; Ven- 
ezuela, T.F. Amazonas, base of Cerro Yutaje; MO, US, 
VEN (holo). 

Navia huberiana L. B. Smith, xor & H. Robinson, 

. Gard. 73: 706. 1986. O. Huber 

. 1980: d T.F. Amazonas, Dept. 
Atabapo, 12 km W of Esmeralda, entre el Cerro Duida 

y el Rio Orinoco; 125 2 US (holo), V 

habia igneosicola L. B. Smith, der & H. Robinson, 
Ann. Missouri Bot. Gard. 73: . 1986. J. A. Stey- 
ermark et al. 122478; 14 May im Venezuela, T.F. 
Amazonas, Dept. Atures, 35 km SE of Puerto Aya- 
cucho; 150 m; NY, US (holo), VEN. 

Navia incrassata L. B. Smith & Steyerm., Phytologia 
16(2): 74. 1968. J. A. pia 97881; Venezuela, 
Bolivar, Meseta de Jaua, Cerro Jaua; 1,922-2,100 m; 
US (holo), VEN. 

Navia intermedia L. B. Smith « Steyerm., E 
16(2): 74. 1968. J. A. Steyermark 97865; 22-27 
Mar. 1967; Venezuela, Bolívar, Meseta de a Cerro 
Jaua; 1,922-2,100 x hg (holo), VEN. 

la i i h & Steyerm., Ann. Missouri 

. R. L. Liesner & F. Delascio 

; Venezuela, T.F. Amazonas, 

Cerro Aracamuni, ad Camp; 600 m; MO, US 

(holo), VE 

Navia jauana L. B. Smith, tu & H. prices 

ol. Soc. Venez. Ci. Nat. 287. 1976. J. A. Stey 

ermark et al 108895; io Feb. 1974; Paak d 

Bolivar, Meseta del Jos: Cerro Sarisarinama; 1,400 

m; US (holo), VEN 


Navia lact .B.S mith, Steyerm. & H. Robinson, Acta 
Bot. Venez. 14(3): 14. 1984. B. Maguire et al. 65705; 
15 Jan. enezuela, T.F. Amazonas, Cerro Si- 


papo; NY, = (holo), VEN. 


Navia vide L. B. Smith & Steyerm., Flea gia 
16(2): 75. 1968. J. A. Steyermark 97865a; 22-27 
Mar. i Venezuela, Bolivar, Meseta de Jnd. Cerro 
Jaua; 1,922-2,100 m; US (holo), VEN. 

Navia liesneri L. B. Smith, Steyerm. & H. Robinson, 
Ann. Missouri Bot. Gard. 73: 709. 1986. R. L. Liesner 

> C. Brewer 15866; 11 Feb. 1984; Venezuela, T.F. 
Amazonas, Dept. Río Negro, Cerro de Neblina, Puerto 
Chimo Camp; 150-1,800 m; MO (3340861), US (holo). 

Navia linearis L. B. si i 
Ann. Missouri Bot. Gard. ; 
ermark 129646; 13-14 "t nos Venezuela, T.F. 
Amazonas, Dept. Atabapo, Cerro Marahuaca; 1,560 
m; US (holo N. 

Navia luzuloides L. B. age, hopes & H. pra 

ol. Soc. Venez. Ci. Nat. 289. 1976. J. A. Ste 
ermark et al. 109633; es Feb. 1974; da 
olivar, Meseta del Jaua, Cerro Jaua; 2,000 m; US 
(holo), VEN 

Navia navicularis L. B. Smith & Steyerm., Bol. Soc. 
Venez. Ci. Nat. 25: 52. 1963. J. A. Steyermark 90199; 
27 Dec. 1961; Venezuela, Bolívar, Sierra Ichun; VEN 
(holo). 

Navia ovoidea L. E os Steyerm. & H. Robinson, 
Brittonia 33: 31. 1. J. A. Steyermark et al. 117361; 

ay 1978; Su i Bolivar, Cerro Guaiquinima; 

E 


Smith & Steyerm., Ann. Missouri 

89. R. L. Liesner & F. Delascio 

7; Venezuela, Amazonas, 
Cerro Aracamuni, summit, Popa Camp; 1,550 m; MO, 
US (holo), VEN 

Navia pedemontana L. B. Smith, Steyerm. & H. Rob- 
ins son, Ann. Missouri Bot. Gard. 73: 708. 1986. J. A. 


swa E Rio Cunucunuma; 200-220 m; US 
(holo), V 
Navia Me i B. Smith, orden As I Robinson, Ann. 
issouri Bot. Gard. 6. J. Murca Pires 
; 10 Feb. ion en ihe, Serra 
PEAN, US (holo). 
Nania. alaba L. B. Smith & Steyerm., Ann. Mis- 
uri Bot. Gard.: in press. 1989. R. L. Liesner & E 
Delascio 22062; 17 Oct. 1987; Venezuela, T.F. Am 
zonas, Cerro Aracamuni, summit, Popa Camp; 1.550 
m; MO, US (holo), VEN. 
Navia plowmanii L. B. Smith, Steyerm. & H. Robinson, 
03. 1986. IF. m- 


E > , Le 
km W of Pico Zuloaga; 2,000 m; NY, US (holo), V EN. 

Navia polyglomerata L. B. Smith, a « H. Rob- 

Missouri Bot. Gard. 704. 1986. W. 

] Ys Venezuela, T.F. 

s of Rio Marawinuma, 2-6 km E o 

Base Camp; 160 m; MO (3488107), NY, US (holo), 
VEN. 

Navia "ua dea L. B. Smith, eae A H. Robinson, 
Bol. . Venez. Ci. Nat. 32: 307. 6. J. A. Stey- 
a. n et al. 108880; 10 Feb. 6 Venezuela, 
Bolivar, Meseta del Jaua, Cerro Sarasarinama; 1,410 
m; US (holo), VEN. 

Navia terramarae L. B. Smith & Steyerm., 
Bot. Gard.: in press. 1989. R. L. Liesner & F. Delascio 
21998; 16 Oct. 1987; Venezuela, T.F. Amazonas, 
Cerro Aracamuni, summit, Popa Camp; 1,550 m; MO, 
US (holo), VEN. 


Ann. Missouri 


670 


Annals of the 
Missouri Botanical Garden 


Navia thomasii L. B. Smith, Steyerm. & H. Robinson, 
Ann. Missouri Bot. Gard. 73: 703. 1986. W. W. Thom- 
as 3245; 26 Apr. 1984; Venezuela, T F. Amazonas, 
Dept. Río Negro, Cerro Neblina, ESE facing slope above 
Rio Marawinuma, E of ‘Puerto Chimo’ Camp; 600 m 
NY, US (holo), VEN. 

Pitcairnia venezuelana L. B. Smith & Steyerm., Phy- 
tologia 16(2): 76. 1968. J. 4. Steyermark & = Rabe 
96472; 24 Aug. 1966; Venezuela, Portuguesa, 5 km 
ENE of Agua Blanca, 22 km NE of Felicia E m; 
US (holo), VEN. 

Steyerbromelia dde L. B. Smith, Steyerm. & H. Rob- 
inson, Ann ri Bot. Gard. 73: 700. 1986. J. 4. 
Steyermark s ul 130072; 24- EN m. Poi Ven- 
ezuela, T.F. Amazonas, Dept. Rio o Ara- 
titiyope; 990-1,670 m; MO ird 3237103), 
NY, US (holo), VEN. 

Tillandsia abysmophila L. s Smith & pa ., Ann. 
Missouri Bot. Gard. 73: 7 


Vriesea bland dug 
113. 1982. J. A. Steyermark & V. Carreric 
123808; 1 Deo. 1980; Venezuela, Edo. Yar 
San Felipe, Vuelta de Pavo, 5 mi. 5 C eee l, 200 
m; VEN (holo 

Vriesea La cw (Brongn.) Lemaire = oinochroma 
t , Bu E y es Soc. 17(3): 53. 1967. J. . 
pidan" ; Venezuela, p mbai 


el Rio San E Us VEN (holo). 


BRUNELLIACEAE 


Brunellia cde Szyszyl. var. ptariana Steyerm 
ieldiana Bot. 2J Sieveumatk 
59847; 4 Nov X Venezuela, Bolivar, Ptari-tepui; 
2,130 m; F (holo). 
€—— Tus e n. & Cuatr., Ann. Missouri 
14:635. 1987. J. Ewel 209; 29 Apr. 1964; 
V buses Braxton faded Cerro de la Neblina; 1,500 
m; NY (holo). 
Brunellia paucijuga Steyerm., d E ) 344. 
1964. J. A. Steyermark 53873; 16 Aug. 1943; Ec- 
uador, Prov. El Oro, between tt A Guagra 
Uma; 2,790-2,880 m; F ~ 
Brunellia jeg dee Steyerm., Phytologia Ayi 344. 
1964. J. A. Steyermark 53576 6; 13 July 1943; Ec- 
uador, E Santiago-Zamora, arenillas xil Rio Tin- 
tas, 10 leagues SE of El Pan; 2,195 m; F (holo). 


BURMANNIACEAE 


icr carrenoi Steyerm., Acta Bot. Venez. 6: 89. 
1971. J. A. Steyermark et al. ene 25 Dec. 1970; 
Venezuela, Eu e of K.134 S of El Dorado, 

,350 m; NY, E D VEN (holo) 


19. 1984. 


A Cerra ahuaca; MO (29 
(holo), VEN = B. foliosa Gleason subsp. foliosa, 1986. 
Burmannia ici Steyerm., dern Bot. 28(1): 

165. ; 8 Sep. 1944; 
Venezuela, T. F of Orinoco, near 
Río Sanariapo, vicinity of Sanariapo; 100 m; F (holo), 
US. 


GH, NY, U, 


BURSERACEAE 


Bursera permollis Standl. & cR Field Mus. Nat. 
Hist., xd = 23(4): 166 P. C. Standley 
(5245 -Nov. 1940; E Dept. Jutiapa, 
vitiniby of read) 850 m ios lo). 

Dacryodes i d Meg Stey & Magu 
New York Bot. Gard. 17(1 `: “445. 1967. J. . 
ermark 75358; 14 May 1953; Venezuela, Boiss, 
Chimantá-tepui (Torono-tepui), Chimantá Massif; 1,000 
m; NY (holo). 

Dacryodes costanensis Steyerm., Acta Bot. Venez. E 
235. 1975. L. a ard 3030; Apr. 1958; 

du Guayabitos, arriba de 


a jm 


Baruta; 
Protium prin pans T Fieldiana Bot. 28(2): 27: 
1952. Steyermark 59994; 10-11 Nov. 1944. 
Venen uis. a Ptari-tepui; 1, 585- 1,600 m; F (holo). 
Protium ie ie Steyerm. var. angusitor Steyerm., 
28(2) 276. 1952. A. Steyermark 
59987; 10- "n alag 1944; Venezuela, Bolivar, Ptari- 
tepui; 1,585- 1,600 m; F (holo). 
Protium puberulenta cies erm., Fieldiana Bot. 28(2): 276. 
952. J. A. Steyermark 60460; 23 Nov. 1944; Ven- 
ezuela, Bolivar, 8 mi. NW of Kavanayen; 1,220 m; F 
(holo). 
Trattinickia iod ibid - des Bot. 28(2): d 
952. J. A. Steyermark 60312; 18 Nov. 1944; Ver 
ezuela, Bolivar, ¡eee Santa n de bn 
and base of Ptari-tepui; 1,220 m; F (holo). 
CACTACEAE 
Bonifazia Standl. & d Field Mus. Nat. Hist., 
Bot. Ser. 23(2): 66. 1944, gen. nov 
ce ia ee a ; Standl. & Steyerili;, Field 
, Bot. Ser. 23(2): 66. 1944. P. C. a 
85603; eb. 1941; Guntemals, E Quezaltenango, 
rtin Chile Verde and 
= dm quezal- 
tecum (Steyerm.) L. Williams, 1961. 
Heliocereus heterodoxus Standl. & Steyerm., Field Mus. 


Nat. Hist., Bot. Ser. 23(2): 67. 1944. J. . . Steyermark 
36291; 21 Feb. 1940; Guatemala, Dept. San Marcos, 
Rafael en NE part of 


— H. cin- 
nabarinus (Eichl.) Britt. & Rose, 1961 


CAMPANULACEAE 


Lobelia laxiflora H.B.K. f. lutea Standl. & Steyerm., 
Field Mus. Nat. Hist., Bot. Ser. 23(2): 98. 1944. P. 
C. Standley 67195; 1 Mar. 1939; Guatemala, Dept. 


Lobelia d L. + purpurea Palmer & Steyerm., 
Brittonia 10: 118. 1958. J. 4. Steyermark 79674; 
19 E 1955; ps A. Missouri, Adair Co., along Char- 
iton River, 2.5 mi. S of Youngstown; F (ho lo 

sa a as villetti po Brittonia 30: 50. 1978. 

S. Ti et al. 752-329; 2 Feb. 1975; Venezuela, 
T. E ree Cn an 2,750 m; NY (holo). 


CANELLACEAE 
Cinnamodendron venezuelense Steyerm., Fieldiana Bot. 
28(2): 402. 1952. J. A. Steyermark 61950; 9 Apr. 
1945; Venezuela, Monagas, SW of Caripe, peris Que- 
brada Colorado Grande; 850-1,350 m; F (holo 


Volume 76, Number 3 
1989 


Taylor 671 
Plants Described by 


Julian A. Steyermark 


CAPPARACEAE 


Capparis ae ie rs x Steyerm., Field Mus. Nat. 

23(4): 1 1944. J. A. Steyermark 

45616; l- p r. 1942; T. Dept. Alta Vera- 
paz, Cerro Chinajá; 150-700 m; F (holo). 

ree grandipetala Maguire & Steyerm., in press, 

Mer 1. New York Bot. Gard. N. T. Silva & U. Brazao 

Dec. 1965; Brazil; NY (holo), MO 


Capparis EA Steyerm., Fieldiana Bot. 28(1): 
238. 1951. J. A. Steyermark 6280 0; 18 May 1945; 
Vene de Suere, above Cuchivano, along Rio Guagua, 
tributary to Rio Manzanares; 230-300 m; F (holo). 


CAPRIFOLIACEAE 


Sambucus canadensis L. f. rubra Palmer & Steyerm., 

n. Missouri Bot. Gard. 25: 773, 1938. E. J. Palmer 

18928; 3 Sep. ~ U.S.A., Missouri, Atchison Co., 
Watson; GH (holo). 

Puras brunnescens Standl. & rU Field Mus. 
Natl. Hist., Bot. Ser. 23(2): 89. 1944 A. Steyer- 
mark 49855; 31 July 1942; Cuatemala, Dept. Hue- 
HUP EDS RO; Cruz de Limón, Sierra de los Cuchuma- 


tanes; 2,600-3,000 m; F (holo) = V. jucundum Morton, 
976. 
j iburnum detrac tum Standl. & Steyerm., Field Mus. Nat. 


PAR URB euryphyllum Standl. & Steyerm., Field Mus. 
t. Hist., Bot. Ser. 23(2): 90. 1944. J. A. Steyermark 
dot 8 July 1942; Guatemala, Dept DATE 
, between Tunima and Quisil, Sierra de los Cuc 
matanes; 2,500-3,100 m; F (holo). 

Viburnum mortonianum Standl. & Steyerm., Field Mus. 
Nat. Hist., Pur Ser. 22(4): 294. 1940 A. Steyer- 
mark 31606; 10 Nov. 1939; Cowon: Dept. Chi- 
quimula, Padi Tixixi; 1,500 m; F (holo). 

Viburnum tinoides L.f. var. roraimense (Killip & Smith) 


Steyerm. f. turumiquirense Steyerm.; Field Mus. Nat 
list., Bot. Ser. 28(3): 619. 3. J. A. Steyermark 
62691; 10 Ma 


y 1945; Venezuela, Sucre, Cerro Tu 
rumiquire; 1,300-1,800 m; F (holo). 


CARYOPHYLLACEAE 


Arenaria altorum Standl. & Steyerm., ve Mus. Nat 

Hist., Bot. Ser. 23(2): 49. 1944. J. 4. Steyermark 

33091; 12 Dec. 1939; Guatemala, Dept. Jalapa, near 

Minas de Croma, Potrero Carrillo; 1,500-1,700 m; F 
0). 


Arenaria via ip Standl. & Y Field Mus. Nat. 
Hist., Bot. 23(2): 50. 1944. Ghiesbreght 871; 
1864- rom “Mexico. Chiapas; F (holo). 

— Ts malensis Standl. & Steyerm., Field Mus. 

, Bot. Ser. 23(2): 50. 1944. J. A. Steyermark 
36268; "feb. 1940; Guatemala, Dept. San Marcos, Rio 
Vega, near San Rafael and Guatemala - Mexico border, 


k 
Arenaria patula Michx. f. media Steyerm., Rhodora 43: 
331. 1941. Gattinger s.n.; May; U.S.A., Tennessee, 
rocky glades near Nashville: F (bolo), € : M 
Arenaria m Michx. f. robusta Steyerm., Rhodora 
43: 330. 1941. J. A. Steyermark 26652; 27 May 


1939; U.S.A., Vra Butler Co., along Gillis Bluff, 
section 8 an , 5 mi. SW of Giulin; F (holo), MO 
(1198045) = rient patula var. robusta (Stey- 
erm.) Maguire à 

Cerastium t juniperorum Standl. & Steyerm., Field Mus 
Nat. , Bot. Ser. 23(2): 51. 1944. J. A. CER 
48413; F July 1942; Guatemala, Dept. Huehuetenan- 
go, Tunima, Sierra de los tl 3,400-3,500 
m; F (holo). 

Drymaria firmula Steyerm., Fieldiana Bot. 28(1): 227. 
1951. J. A. Steyermark 56536; 14 May 1944; Ven- 
ezuela, Mérida, between El Molina and San Isidro Alto, 
Páramo de Los Colorados; 2,745-2,955 m; F (holo). 

Drymaria — Standl. & vow Field Mus. Nat. 
Hist., Bot. 23(2) 52. Steyermark 
50243; 8 p 1942; ARR Dept. Huehuetenan- 
go, Chemal, summit of Sierra de los Cuchumatanes; 
3,700 m; F (holo). 

TUN avala Willd. f. a ura Steyerm., Fieldiana 

t. 28(1): 228. 1951. J. A. Steyermark 55217; 3 
oe 1944; Venezuela, = above Humocaro Bajo, 
above Las Sabanetas, Los Aposentos; 2,500-2,530 m; 


F (holo). 

Stellaria venezuelana Steyerm., Fieldiana Bot. 28(1): 
229. 1951. J. A. Steyermark 62739; 11 May 1945; 
Venezuela, Sucre, Cerro de Diablo base; 2,000-2,100 
m; F (holo). 


CECROPIACEAE 


Cecropia sylvicola Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(4): 153. 1944. P. C. 


between Tactic & divide on road to Tamahu; 1,500- 
1,600 m; F (holo). 


CELASTRACEAE 
Maytenus agostinii Steyerm., Ernstia 23: 34. 1984. G. 
Agos ostini & M. Farinas 84; 28 Oct. 1963; Venezuela, 
ragua, Parque Nacional Henri dear V (holo). 
Maytenus S urea — rm., Fie eldian t. 28(2): 334. 
1952. J. A. Steyermark 60385 90-21 Nov. 1944; 
oye Bolívar, pete O- pari ma; 1,005-1,220 
; F (holo). 


onn coriacea Steyerm., Fieldiana Bot. 28(2): 337. 
1952. J. A. Steyermark 56509; 14 May 1944; Ven- 
ezuela, Mérida, above El Molino; 2,010-2135 m; F 
(holo). 

Maytenus E Klotzsch f. crenulata Steyerm.; 
Fieldiana Bot. 28(2): 338. 1952. J. A. Steyermark 
61731; 23 Mar. 1945; Venezuela, Anzoátegui, Fila 
Grande; 900-1,400 m; F (holo). 

e own huberi Steyerm., Ann. Missouri Bot. Gard. 75: 
1062. 8. O. Huber 441; 29 Jan. 1977; Venezuela, 


al sur y Serrania Colmena al Norte; 200-300 m; VEN 
lo 


Maytenus goi Steyerm., Ann. Missouri Bot. Gard. 
75: 1063. 1988. O. Huber 1229; 16 Oct. 1977; 
Venezuela, T.F. Amazonas, Dept. Atures, cuenca del 
Rio Manapiare, los cerros al N del Cerro E 

an Juan de Manapiare; 225 m 


Maytenus jauaensis Steyerm., Bol. Soc. Venez. Ci. Nat. 
32: 346. 1976. J. A. Steyermark et al. 109252; 19- 


672 


Annals of the 
Missouri Botanical Garden 


20 Feb. 1974; Mago a Meseta del Jaua, 
Cerro Sarasarinama; 1,3 ; VEN (holo). 

Maytenus longistipitata ta Ann. Missouri Bot. 
Gard. 75: 1064. 1988. B. Ma aguire et al. 53731; 12 
Sep. 1962; T eNe Bolivar, Cerro Uroi, summit, N 
portion, Río Uroi, Río Chicanán; 700 m; MO, 
(holo). 

Maytenus EA Steyerm., Ann. Missouri Bot 


. T. Plowman & W. Thomas Pid 


15 Apr. Th di T.F. Amazonas, Cerro de 
la Neblina, 26 km ENE íi Base Camp; 2,000 m; F, 
MO (holo), VEN. 


Maytenus dri a bacis rm., Fieldiana Bot. 28(2): 338. 

1952. J. 4. Steyermark 55832; 30 Mar. 1944; Ven- 

ezuela, Mérida, an Pinango and Las Corales; 3,333 
m; F (holo). 


ees pa Steyerm., _Fieldiana Bot. 
338. P! 


28(2): 


Vanseusls: T.F. Am e la Neblina, Camp 
IV, 15 km NNE of Pico Phelps; 780 m; MO (holo), 
EN. 


Microtropis rin Standl. & EUM Field Mus. Nat 
Hist., Bot. eri 170. 1944. J. A. Steyermark 
43284; 26 m 942; eae Dept. Zacapa, $ 
Volcán Gemelos, Ei de las Minas; 2,100-3,200 m; 
F (holo). 

Zinowiewia aymardii Steyerm., Ann. Missouri Bot. Gard. 

; : ». Aymard 4631; 21 Oct. 1986; 
nd Bolivar, Distr. Sifontes, 7 km NW of Caserio 

8 km W of Sta. Elena de Uairén; 850 m; 

MO an eus 


CHRYSOBALANACEAE 


cs sep ensis Standl. & Steyerm., Field Mus 
Nat ., Bot. Ser. 22(5): 335. 1940. 4. Smith F17 79; 
14 Mar r. 1939; Costa Rica, Prov. Alajuela, Villa Que- 
sada, Cantón de San Carlos; 825 m; F (holo). 


CLETHRACEAE 


Clethra oo Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(1): 17. 1943. H. Kuylen 151; July 
1927; Guatemala, Dept. Izabal, s Rios; F (holo 

= C. macrophylla Mart. & Gal., 

Clethra johnstonii Standl, & Ae "Field Mus 
Hist., Bot. Ser. 22(4): 258. 1940. R. Johnston 
1255; 20 Mar 1938; Guatemala, Dept. Guatemala, 
Volcán de Pacaya, above Las Calderas; F (holo). 

Clethra lic Meer s Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(1): 17. 1943. J. 4. Steyermark 
33124; 12 Dec. 1939; Guatemala, Dept. Jalapa, Po- 
trero Carrillo, 13 mi. NE of Jalapa; 1,500-1,700 m; 

holo 


F( 

Clethra pac hec ime 1 Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 22(4): 259. 1940. P. C. Standley 
65123; 11 Feb. 1939. Guatemala, Dept. Sacatepé- 
quas Volcán de Agua, above Santa Maria de Jesüs; 

250-3,000 m; F (holo). 
Clethra no Steyerm., Phytologia 9(6): e 1964. 
sta Solis 5054; 20 July 1943; Ec , Prov. 
Santiago: m Campansa, Cordillera el 2,850 


m; F (ho 
giis skuto hii Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 22(4): 260. 1940. 4. F. Skutch 1453; 


14 Oct. 1934; Guatemala, Dept. Quezaltenango, Pal- 
mar; 1,350 m; F (holo). 


CLUSIACEAE 


Caraipa | ea Steyerm., Fieldiana Bot. 28(2): 384. 
1952. Ll. Williams 14140; 2 Feb. 1942; opaca 
mazonas, Yavita; 128 m; F (holo) = C. pun 

tu lata: Da cke, 1978. 

Caraipa longipedicellata Steyerm., Fieldiana Bot. 28(2): 

A. Steyermark 60708; 28 Nov. 1944; 

Venezuela, Bolivar, between Ptari- tepui and Sororopan- 
tepui; 1,220 m; F (holo). 

chrysochlamys pauciflora Steyerm., Fieldiana Bot. 28(2): 
386. 1952. J. A. Steyermark 60566; 25 Nov. 1944; 
Venezuela, Bolivar, Quebrada O-paru-ma; 915-1,005 
m; F (holo 

Clusia cerroana Steyerm., Fieldiana Bot. 28(2): 386. 
1952. J. 4. Steyermark 59702; | Nov. 1944; Ven- 
ezuela, Bolivar, Ptari-tepui; 1,700-1,800 m; F (holo). 

Clusia hexacarpa Fere var. ptaritepuiana Steyerm. 
iur Bot. 28(2): 387. 1952. J. St Pac nd 

866; 4 Nov. 1944; iR Bolivar, Ptari-tepui; 
dem m; F (holo 

Clusia imbricata Steyerm., Faeidiana Bot. 28(2): 387. 
1952. J. A. Steyermark 59699; | Nov. 1944; Ven- 

ezuela, Bolivar, Ptari- -tepul; 1,700- 1, 800 m; F (holo). 

C lusia lusoria Standl. & Steyerm., Field Mu 5 
Bot. ad . 23(2): 63. 1944. J. 
2 Nov. 1939; Guatemala, Dept. Chiquimula, Montaña 
Norte to El Ju Cerro Brujo SE of Concepción de las 
Minas; 1,700-2,000 m; F (holo). 

Clusia pusilla Steyerm., Pie Bot. rA 2): 390. 1952. 
J. A. Steyermark 59396; 26 Oct. 1944; Venezuela, 
Bolivar, Gran Sabana; 1.220 m; F > lo). 

Clusia reducta Steyerm., Fieldiana Sw AER 391. 1952. 
Ll. Williams 14898; 26 Mar 2; Venezuela, T.F. 
Amazonas, Rio Guianía; 1255 m; r (h olo). 

Clusia roraimana Steyerm., Fieldiana Bot. 28(2): 391. 
1952. J. A. Sem 58606; 25 Sep. 1944; Ven- 
ezuela, Bolivar, Mt. Roraima; 1,030-1,115 m; F (holo). 

Clusia williamsii Steyerm., Fieldiana Bot. 28(2): 392. 
1952. Ll. Williams 14143; 2 Feb. 1942; Venezuela, 
T.F. unas Yavita; 128 m; F (holo). 

Havetia laurifolia .K. var. venezuelana Steyerm., 
Fieldiana Bot. 28(2): 392. 1952. J. A. Steyermark 
50592; 1 ay 1944; Venezuela, Mérida, above Ta- 
bay; 2,285-2,745 m; F (holo). 

Hypericum arbuscula Standl. & uer Field Mus 
s pete Bot. Ser. 23(2): 63. 4. J. R. 345 

; 11l Apr. 1941; DR de Baja Verapaz, 
E F (holo). 

Hypericum calcicola Standl. & Steyerm., Field Mus 
Nat. Hist., Bot. Ser. 23(2): 64. 1944. J. 4. dad 
50160; 6 Aug. 1942; Guatemala, Dept. Huehuetenan- 
go, Sierra de qe Cuchumatanes; 3,700 m; F (holo). 

Hypericum caracasanum SAM. var. oni ri 

+» Fieldiana Bot. 28(2): 393. 1952. J. A. Stey- 

9; 6 May 1945; Venezuela, Sucre, Cerro 
Sud ^ 360-2,500 m; F (holo). 

Hypericum ericifolium pacing Fieldiana Bot. 28(2): 

93. 1952. J. A. Steyerr 21: 
Venezuela, Táchira, Paramito 2,500 m E 

ee Du Steye /"Fieldiana Bot. 28(2): 

A. e 56272; 3 May 1944; 
oí "Mérida. Páramo de Pozo Negro; 2,590- 
3,220 m; F (holo). 


Volume 76, Number 3 
1989 


Taylor 673 
Plants Described by 


Julian A. Steyermark 


Hypericum pseudocaracasanum Steyerm., Fieldiana Bot. 
28(2): 394. 1952. J. A. Steyermark 55346; 6 Feb. 
1944; Venezuela, Trujillo, La Quebrada Cortijo; 2,600- 
2,800 m; F (holo). 

Hypericum pseudomaculatum Mack. & Bush f. flavi- 
dum Steyerm., Rhodora 41: 585. 1939. J. A. Stey- 
ermark 56061; 31 May 1938; U.S.A., Missouri, 
on x; along Mill Creek, 5 mi. SE of Pineville; 

F (holo), M 

Mahurea toco M Steyerm., Fieldiana Bot. 
28(2): 395. 1952. J. A. Steyermark 60129; 14 Nov. 
1944; Venezuela, Bolivar, Sororopan-tepui; 2,255 m; 
F (holo). 

Moronobea d gia cus Steyerm., Fieldiana Bot. 28(2): 
395. 1952. J. A. Steyermark 59985; 10-11 Nov. 

1944; Venezuela, Bolivar, Ptari-tepui; 1,585-1,600 
m; F (holo). 

ii inten ptaritepuiana Steyerm. f. rosea Steyerm. 
Fieldiana Bot. 28(2): 397. 1952. J. A. Steyermark 
60892: 7 ec. 1944; Venezuela, Bolivar, Carrao-tepui; 
2,470-2,500 m; F (holo). 

Oedematopus eed Steyerm., eae Bot. 
28(2): 398. 1952 eyermark 59 ; 1 Nov 

1944; Venezuela, “Bolívar, Pan: -tepui; dro: 1,800 
m; F (holo). 

Rheedia mac ipe Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(2): 65. 1944. P. C. Standley 90524; 
l Apr. 1941; Guatemala, Dept. Alta Verapaz, Rio Frio, 
ca. 8 km below Tactic; 1,400 m; E (holo). 

Tovomita angustata Steyerm., Fieldiana Bot. nv cd 
1952. J. A. Steyermark 60475; 23 Nov. 1944; V 


i Nov 
1944; —_ Bolivar, Ptari-tepui; 1,585-1,600 


i 'omita duidae Steyerm., 2 Bot. 28(2): 400. 
952. J. A. Steyermark 57975; 25-26 Aug. 1944; 
ocu. Amazonas, Caño Negro, Cerro Duida; 


305-1,095 m; F (holo). 


COMBRETACEAE 
Buchenavia ptariensis Steyerm., Fieldiana Bot. 28(2): 
423. 19 J. A. Steyermark 60271; 15-17 Nov. 
1944; Venezuela, Bolivar, between Ptari-tepui E So- 
roropan-tepui; 1,615 m; F (holo). 


COMMELINACEAE 
Aneilema ecuadoriensis Steyerm., Phytologia 9(6): 339. 
1964. J. yermark 52813; 3 June 1943; Ec 


uador, Prov. Azuay, between Chacanceo and Río Blan- 


2 132; A. Steyermark 37 7187; Mar. 
940; Guatemala, Dept. a arcos, above Finca El 
Porvenir, on os Santos Chiquitos, Volcán Taju- 


odos Sa 
mulco; 1,300-1,500 m; F (holo 
Commelina x er andl. & Steyerm., Field Mus. Nat. 
Hist , Bot 23(5): 213. 1947. J. A. Steyermark 
50265; 8 qee 1942; Guatemala, Dept. Huehuetenan- 
^ Sierra de los Cuchumatanes, vicinity of Chemal; 
3,700 m; F (holo). 
Commelina erecta L. var. angustifolia (Michx.) Fern. f. 
cana Standl. & Steyerm., Field Mus. Nat. Hist., Bot. 
Ser. 23(2): 32. 1944. J. A. Steyermark 29136; 5 Oct. 


1939; Guatemala, Dept. Zacapa, along RR between 
La Fragua and Estanzuela; 200 m; F (holo). 

Commelina erecta L. f. candida Standl. & Steyerm., 

Mus. Nat. Hist., Bot. Ser. 23(2): 33. 1944. J. 
A. Steyermark 29267; 7 Oct. 1939; Guatemala, Dept. 
Zacapa, between Zacapa and Santa Marta; 200 m; F 
(holo). 

Commelina standleyi Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 23(2): 33. 1944. J. 4. Steyermark 29644; 


973. J. A. Steyermark 106178; 17-19 
1972; Venezuela, Yaracuy, El Amparo hacia Can- 
de WR 1,220-1,250 m; VEN (holo). 

Tinantia erecta (Jacq.) Sheet, f. puberula Standl. A 

m., Field. Mus. Nat. Hist., Bot. Ser. 23(2): 3 
1944. H. S. Gentry 2525; Aug. 1936; Mexico, Chi 
bas a, Sierra er Rio Mayo; olo). 

E ied leiocalyx . Clarke f. vari Stand & 
Stey , Field. Mus . Nat. Hist., Bot. . 23(2): 34. 
jn. C A. Purpus 16201; Sep. Yo MIC Ve- 

ruz, El Fortin; F (holo). 

Tinantia Va ie inr a Standl. & Steyerm., Field 
Mus , Bot. Ser Ps 35. 1944. Heyde & 
las 6392; Jan. 1894; Guatemala, Dept. Escuintla, 
Santa Lucia; 300 m; olo). 

Tinantia standleyi E Field Mus. Nat. Hist., Bot. 

2): 3 

1940; Guatemala, D 

n Santa María de Jesús and Calahuache; 
1,300-1, 500 m; F (holo). 

ica A aE Standl. & Steyerm., Field Mus 
Nat , Bot. Ser. 23(2): 36. 1944. P. C. Standley 

59358; du. 1938; Guatemala, Dept. d porn 
Vole can de Agua, N of Santa Maria T Jesús; 1,800- 
2,100 m; F (holo) = Aneilema aguensis (Standl. & 
Steyerm.) Standl. & Steyerm., 

Tradescantia disgrega Kunth f. glandulosa Standl. & 

Mus Ser 


m 3 

1944. J. A earner 29751; O ; 
mala, Dept. Zacapa, Sierra de | as, between Rio 
ondo and summit of mountain at Finca Alejandria; 
1,000-1,50 — Tripogandra disgrega 


O m; F (holo) 

Kunth f. poda uh (Standl. & Steyerm.) Standl. & 
Steyerm., 195 

Tradescantia diseragu Kunth f. nos ens ui & 
Steyerm., Field. Mus. Nat. Hist., Bot. 23(2): 37. 
1944. J. Morales R. bep Oct. 192 D, osten: 
Dept. Guatemala; lo) = Tripogandra disgrega 
Kunth f. ioni ae & Steyerm.) Standl. & 
Steyerm., 1952 

Tradescantia guatemalensis ie = Clarke f. alba patie 
& Steyerm., Field Mus Hist., Bot. Ser. 23(2): 
37. 1944. P. C. Standley "0035. Nov. 1938- Feb. 
1939; Guatemala, Dept. Sacatepéquez, near Antigua; 
1,500-1,600 m; F (holo). 

Tradescantia standleyi Steyerm., Field Mus. Nat. Hist., 
Bot. Ser 


Montaña Nonojá, E of 
Tradescantia venezuelensis Steyerm 
28(1): 152. 1951. J. A. Steyermark 62688; 10 May 
1945; Venezuela, Sucre, Cerro Turumiquire; 1,300- 
1,800 m; F (holo). 
Zebrina huehueteca Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 213. 1947. J. A. Steyermark 


674 


Annals of th 
Missouri Bones Garden 


51016; 22 Aug. 1942; Guatemala, Dept. Huehuete- 
nango, along Rio Trapichillo; 1,000-2,100 m; F (holo). 


CONNARACEAE 


Pseudoconnarus krukovii Pr Field Mus. Nat. Hist., 
Bot. Ser. 22(3): 141. 1940. B. 4. Krukoff 6455; 14 
Sep.-11 Oct. 1934; ein Amazonas Mun. Humayta, 
near Tres Casas; A, BM, BR, F (holo), G, K, MO, NY, 

; = P. minii i (Poepp. & Endl.) Radl., 
1983. 

Rourea krukovii Steyerm., Field Mus. Nat. Hist., Bot. 
Ser. 22(3): 142. 1940. B. 4. Krukoff 8556; 11 Sep.- 
26 Oct. 1936; Brazil, Amazonas, Mun. Sào Paulo de 
Nei dis near Palmares; A, BM, BR, F, G, K, MO, 
NY (holo), S, U. 


CONVOLVULACEAE 


Dicranostyles costanensis Steyerm. & Austin, Ann. Mis- 
souri Bot. Gard. 57: 155-157. 1970. J. A. Steyermark 
et al. 100263; n.d.; Venezuela, Yaracuy, Cerro La 
Chapa; 1,200-1,400 m; VEN (holo) 

Dicranostyles imatacensis Steyerm., Acta Bot. Venez, 
3: 190. 1968. J. 4. Steyermark 87129; 31 Oct. 1960; 
Venezuela, Terr. Delta Amacuro, San Victor, Rio Ama- 
curo, Sierra Imataca; 65-80 m; NY (holo) = D. gui- 
anensis A. Mennega, 1982. 

Ipomoea heterodoxa Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(2): 82. 1944. P. Gentle 871; Dec. 
1933; Belize, Maskall; F (holo). 

Ipomoea ophioides Standl. & Steyerm., Field Mus 
Hist., Bot. Ser. 23(2): 82. 1944. P. z Standley 7084; 
Dec. 1940; Guatemala, Dept. Santa Rosa, region of 
La Morenita, NE of Chiquimulilla; 400 m; F (holo). 

Ipomoea santae-rosae Standl. & Steyerm., Field Mus. 

at. Hist., Bot. Ser. 23(2): 81. 1944. P. C. Standley 
79287; Nov.-Dec. 1940; Guatemala, Dept. Santa Rosa, 
near Chiquimulilla; 325 m; F (holo). 

Ipomoea saxorum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(2): 81. 1944. J. 4. Steyermark 
30254; 22 Oct. 1939; Guatemala, Dept. Chiquimula, 
gorge of Rio Chiquimula, between Santa Barbara and 
Petapilla; 350-420 m; F (holo). 

Itzaea Standl. & Steyerm., Field Mus. Nat. Hist., 
Ser. 23(2): 83. 1944, gen. nov. 

Jacquemontia guatemalensis Standl. & Steyerm., Field 
Mus 


Bot. 


4. : i 
Steyermark 30066; Oct. 1939; Guatemala, Dept. Chi- 
quimula, Piro 400 m; F (holo) = J. agrestis 
(Choisy) Mei 
andl. & pou 5 jus Mus. 
re . Steyer 
mark 38533; 1 Apr. 1 ee Dun iba, 
Montaña del Mico; 65- 6. AUN m; F (holo). 
Maripa stellulata Steyerm., Acta Bot. Venez. 3: 209. 
c C. Steyermark 95456; 2 Apr. 1966; 
cene. Caraba: Rio San Gian, Planta Elestics: 
350-550 m; MO, NY, U, US, VEN (holo). 


CRASSULACEAE 


Echeveria bic = (H.B.K.) E. Walther var. turumiqui- 


, Fieldiana Bot. 28(2): 244. 1952. J. 
A. "BRE ES 62491; 5 May 1945; d Sucre, 
Cerro Turumiquire; 1,700- 2, 000 m; F (ho 

Echeveria huehueteca Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(4): 159. 1944. J. A. Steyer- 


mark 50934; 19 Aug. 1942; Guatemala, Dept. H 
huetenango; Cumbre Papal; 1,400-3,000 m; F (holo) 

Echeveria macrantha Stan 
Nat. Hist., Bot. Ser. 23(4). 
mark 32808; 6 Dec. 1939; Guatemala, Dept. Jalapa, 
Montaria Miramundo at Buena Vista; 2,000-2,200 m; 
F (holo). 

Tillaea venezuelensis Steyerm., Fieldiana Bot. 28(4): 914. 

7. J. A. Steyermark 55906; 15 Apr. 1944; Ven- 

ezuela, Mérida, between Chachopo and Los Aparta- 
deros, near El Aquila; 3,030 m; F (holo). 


CRUCIFERAE see Brassicaceae 
CUCURBITACEAE 


Ahzolia Standl. & ees d Mus. Nat. Hist., 
Ser. 23(2): 92. 1944, 

Anguria simplicifolia B en. , Fieldiana Bot. 28(3): 

1953. J. A. Steyermark 60161; 15 Nov. 1944; 
Venezuela, Bolivar, E. 2. 130-2,250 m; 
(holo). 

Corallocarpus ne lege d Standl. & Steyerm., Field 
Mus. Nat. Hist., Bot. Ser. 23(2): 93. 1944. P. A 
Standley one Oct. 1940; keg Dept. 

capa, Zac 200 m; F (holo) = Doyerea ee 
deron ate 1976 

Elaterium macrophyllum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(2): 94. 1944. H. von Tuerck- 
heim II. 1728; Apr. 1907; Guatemala, Dept. Baja Vera- 

az, above Pansal; 1,400 m; F (holo) = Rytidostylis 
macrophyllus (Standl. & Steyerm.) Dieterle, 1976. 
Rytidostylis brevisetosa Steyerm., Bol. Soc. Venez. Ci. 
at. 26: 148. 1965. J. A. Steyermark 91499; 7 June 
1963; Venezuela, Distr. Federal, between Hacienda El 
Limón and Junquito-Colonia Tovar road; 1,750 m; 
VEN (holo) 

Sicydium araguense Steyerm. & Trujillo, Bol. Soc. Ven- 
ez. Ci. Nat. 25: 245. 1964. J. A. Steyermark 89813; 
20 Oct. 1961; Venezuela, Aragua, Parque Nacional 
Henri Pittier; 1,500-1,700 m; US, VEN (holo). 

Sicydium glabrum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. S ) 


Bot. 


go, Montaña Chicharro, Volcán Santa María; 1,400- 
1,500 m; F (holo). 

Sicyos guatemalensis Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(2): 96. 1944. J. A. ane 
32140; 28 Nov. 1939; Guatemala, Dept. Jalapa, vi- 
cinity of Jalapa; 1, 300 m; F (holo). 

ecunumania Standl. & Steyerm., pees Mus. Nat. Hist., 


Bot. Ser. 23(2): 96. 1944, gen. 
Tecunumania quetzalteca re & Steyerm, Field Mus 
, Bot. Ser. 23(2): 97. 1944. P. C. Sta ndley 
Peu. ne Mar. 1939; en Dept. San Marcos, 


Finca Vergel, near Rodeo; 900 m; F (holo). 


CUNONIACEAE 
Weinmannia neblinensis Maguire & Steyerm., Mem 
et al. 42226; 25 Nov. 1957; Venezuela, T.F. Ama- 
zonas, Cerro de la Neblina, Cano Grande; 1,200- 2,200 
m; MO (3291190), NY (holo). 
CUPRESSACEAE 


Juniperus standleyi Steyerm., Field Mus. Nat. Hist., Bot. 
Ser. 23(1): 3. 1944. J. A. Steyermark 36137; Feb. 


Volume 76, Number 3 
1989 


Taylor 675 
Plants Described by 
Julian A. Steyermark 


1940; Guatemala, Dept. San Marcos, Volcán Tacaná; 
4,100-4,400 m; F (holo). 


CYPERACEAE 


ps — Steyerm., ra Bot. 28(1): 

19 A. Steyermark 57629, 2 Aug. 1944; 

Neue Bolivar, between Upata Mie Rio Caroni; 
0). 


Carex huehueteca Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 195. 1947. J. A. Steyermark 
49055; 18 July 1942; Guatemala, Dept. Huehuetenan- 
go, Cananá, Sierra de los Cuchumatanes; 2, 500m;F 
(holo). 

Carex larensis Steyerm., Fieldiana Bot. 28(1): 66. 1951. 
J. A. Steyermark 55470; 11 Feb. 1944; Venezuela, 
Lara, between Buenos Aires and Páramo de las Rosas; 
2.285-3,290 m; F (holo). 

Carex roraimensis Steyerm., Fieldiana a en ): 67. 
1951. J. A. Steyermark 58870 ; 285 - 
ezuela, Bolivar, Mt. Roraima, "teslis E 
Central Rift, Central Swamp; 2,700-2,740 m; F (hol) 

Carex standleyana Steyerm., Ceiba 3: 23. 1952 
Williams 13178; 6 July 1947; Guatemala, Dept. 


Y 


lapa, mountains above Aguacate; 2,150 m; (Loa. 
Carex tachirensis Steyerm., Fieldiana Bot. 28(1): 68 
1951. J. A. Steyermark 57367; 15 July 1944; Ven 


ezuela, near Colombia - Venezuela border Tachira, 
Páramo de Tamá; 3,045-3,475 m; F 
Carex tamana Steyerm., Fieldiana Bot. 280) 70. 1951. 
. Steyermark 57401; 15 July 1944; Venezuela, 
near rv Venezuela border, Táchira, Páramo de 
; 3,045-3,475 m; F (holo). 
nn priua Standl. & Steyerm, Ceiba 4: 64. 
1953. Steyermark 50150; on : d Gua- 
temala, Dept. Huehuetenango, s t Sierra de los 
Cuchumatanes, between Tojquia and d Caxix bluff; 3,700 
m; F (holo). 

Carex toreadora Steyerm., Phytologia a 2 338. 1964. 
J. A. Steyermark 53095; 15 July 1943; Ecuador, 
Prov. Azuay, near Toreador; 3,785-3, oa m; F (holo). 

Carex tunimanenisis Standl. & Ste eyerm., Ceiba 4: 65. 
1953. J. A. Steyermark 48334; 7 Ju uly 1942; Gua- 
temala, Dept. Hu rr E near T 
los qu di dame 3,4 3,500 m; F (holo). 

Gores riw ch Steyr Nen Bot. 28(1): 

. 19 A. Stey pone 6270 ; 10 May 1945; 
duae Sucre, "da Turumiquire, headwaters of 
Rio Hi auc and Rio de Amana; 1,900-2,000 m; 
F (holo). 

Carex venosivaginata Standl. & Steyerm., Ceiba 4: 67- 

8. 19 Steyermark 48554; 14 July 1942; 
Guatemala, Dept. Huehuetenango, Cerro Huitz, Sierra 
de los Cuchumatanes; 2,600 m; F (holo). 

eben Bot. 28(1): 50. 

; 3 Dec. 1944; Ven- 
ezuela, Bolivar, dile panne between Santa 

Teresita de Kavanayen and Rio Tek-Yunsen; 1,375 
; olo). 


m 
Finbriryiis tamaensis Steyerm., Fieldiana Bot. 28(1): 
39. 1951. J. 4. Steyermark 57422; 17 July 1944; 


Colombia-Venezuela border, along Rio Táchira, be- 

tween Delicias and Paraqui e a 675-1,980 m; F (holo). 

Ear tepuianum Stey dera Bot. 28(1): 

1951. J. A. poo 602 15-17 Nov. 

D Venezuela, Bolivar, vicinity ‘Misia Kathy Camp’ 

on mesa M Ra Ptari-tepui and Sororopan-tepui; 1,615 
— Mapania tepuiana (Steyerm.) T. Ko- 


vA 

Mapania maguireana T. Koyama & Steyerm., Mem. 
New York Bot. “Card. di. 63. 1967. B. 
& D. anshawe 32351; Gu 
rupung vat Rises à forest; 1,000 m; NY (holo). 

Oreobolus venezuelensis Steyerm., Bol. Soc. Venez. Ci. 
Nat. 11: 308. 1950. J. A. Steyermark 57198; 14 
July 1944; Venezuela, Táchira, paramito at base of 
Páramo de Tamá; 2,500 m; F (holo). 

d bolivarana Steyerm., Fieldiana Bot. 28(1): 

51. J. A. Steyermark 60206; 15 Nov. 1944; 
sra Bolivar, Salto de | LE -meru, W end 
Sororopan-tepui; 1,500 m; F (holo). 

xb sp ale culmenicola Steyerm., Fieldiana Bot. 

951. J. A. Steyermark 62110; 15 Apr. 

1945; B Monagas, P > de 2,180 m; F 
(holo) = R. dissitiflora Steudel, 
Rhynchospora duidae Steyerm., Fela Bot. 28(1): 
43 J.A. cada mark 68254; 2 Sep. 1944; 
Venezuela, T.F. Amazonas, Cerro Duida: 1,065-1,220 
m; 
Rhynchospora karuaiana Steyerm., Fieldiana Bot. 28(1): 
1951. J. A. Steyermark 60334; 18 Nov. 1944; 
T Bolivar, Rio Karuai, o Teresita de 
Kavanayen and base of di mes 1,200 m; F (holo) 
— R. triflora (Poir.) Vah., 
uy rro paramora Stey n Fieldiana Bot. 28(1): 
1951. J. A. remak 5655 ; 14 May 1944; 
"os Mérida, Páramo de los o just 
El Molino and San Isidro Alto; 2,745-2,955 m; F 
(holo). 

FF o ona ptaritepuiana Steyerm., 
28(1) 45. 1951. J. A. Steyermark 5983 
1944; Too m wo me -tepui; 2, is "i 

— R. lechleri Steudel, 
Qv oe p N Fieldiana Bot. 
28(1): 4 951. J. A. —— 58469; 8 Sep. 
1944; asm T.F. Amazonas, tributary at Orinoco 
River, near Rio Sanariapo, vicinity of Sanariapo; F 
(holo). 

Rhynchospora sororopana Steyerm., Renee: Bot. 28(1): 
46. 1951. Steyermark 60 3 Dec. 1944; 
Venezuela, Bolivar, between n nta RE de Ka- 

along Quebrada Soro- 

a Bockelr 1972. 
Drag Bot. 28(1): 
47. 1951. J. A. eo. 57404; 15 July 1944; 
Venezuela, Táchira, Páramo de Tami, along border; 
3,045-3,475 m; o 

Rhynchospora tepuiana Ste eyer , Fieldiana Bot. 28(1): 
48. 1951. J. A. Steyerm a 58652 ; 25 Sep. 1944; 
lcgi rig Bolivar, ML R aima, near Rondon Camp; 
2,040 m du lo) = js ffesuos C. B. Clarke subsp. 
saaa Steyerm., | 

Rhynchospora per HN owes Bot. 28(1): 

49. 1951. J. A. Steyermark 57391; 15 July 1944; 
Venezuela, Táchira, Páramo de n on border; 
o 


oce Bot. 
; 4 Nov. 
F (holo) 


arke var. venezue- 


lensis Steyerm., Fieldiana Bot. 581) 49. 1951. J. A. 


676 


Annals of the 
Missouri Botanical Garden 


Steyermark 57587; 1 Aug. 1944; Venezuela, Bolivar, 
between Ciudad Bolivar and Rio Caroni; 100 m; F 


zuela, Mérida, El Aquila above Páramo de Mucuchies; 
4,025 m; F (holo). 

Vesicarex Steyerm., Fieldiana Bot. 28(1): 63. 1951 gen. 
nov. 

Vesicarex collumanthus Rud Pisae Bot. 28(1): 
63. 1951. mark 57040; 6 July 1944; 
Venezuela, Mérida, El ake ioe Paramo de Mu- 
cuchies; 4,025 m; F (holo) 


DICHAPETALACEAE 


Dichapetalum bullatum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(4): 169. 1944. J. A. Steyer- 
mark 39874; 7 Dec. 1941; Guatemala, Dept. Izabal, 
along road between Puerto Barrios and Santo Tomas; 


sea level; F (holo). 
DILLENIACEAE 


Doliocarpus esmeraldae Steyerm., Fieldiana Bot. 28(2): 
366. 1952. J. A. peta 57879; 23 Aug. 1944; 
Venezuela, T.F. Amazonas, between Sabana Grande 
and SE base of adn Duida; 200 m; F (holo). 

Doliocarpus ptariensis ibi: Fieldiana Bot. 28(2): 

9 . A. yermark 59972; 10-11 Nov 
1944; Micequels. Bolivar. Ptari-tepui; 1,585- 1.600 
m; F (holo). 


DIOSCOREACEAE 


Dioscorea aby dpi sig ele & ET Mem. New 
York Bot. Gard. 50 ress Maguire et 
al. 42319; 8-9 Dec. 1955. doni i MO (3291186), 
NY (holo). 

Dioscorea bolivarensis Steyerm., Fieldiana Bot. 28(1): 

. 1951. J. A. Steyermark 57551 ; 31 July 1944; 
Venezuela, Bolívar, W of Upata, Río. Upata; 500 m; 
F (holo). 

Dioscorea lasseriana Steyerm., Fieldiana Bot. ae 158. 

951. J. A. Steyermark 61836; 5 Apr. 1949; Ven 
ezuela, Monagas, between La Sa iban a de ws Piedras 
and Cerro Negro, NW of Caripe; 1, 200-1 500 m; F 
holo). 

Dioscorea neblinensis Maguire & Steyerm., Mem. New 
York Bot. Gard. 50: in press. . B. Maguire et 
al. 37328; 16 Jan. 1954; Venexuela: MO (3291183), 
NY (holo). 

Dioscorea sororopana Steyerm., Fieldiana Bot. 28(1): 
159. 1951. J. 4. Steyermark 60124; 14 Nov. 1944; 
Venezuela, Bolivar, Sororopan-tepui; 2,255 m; F (holo). 


DIPSACACEAE 


Dipsacus mom Huds. f. albidus Steyerm, Rhodora 
60: e 1959. K. E. Bartel 1; 25 Aug. 1957; 
lings, Cook Co., Chicago, Mt. Hope Ur ier) 

on iris St.; F (holo). 


DIPTEROCARPACEAE 
Pakaraimaea dipteroc arpacea Maguire & Ashton subsp. 


Steyerm., Mem 


, Bolivar, Cerro Guai- 
EN 


quinima; 750 m; NY (holo). 


DROSERACEAE 


Drosera arenicola Steyerm., Fieldiana Bot. 28(2): 243. 
1952. J. A. Steyermark 60920; 16-17 Sep. 1944; 
Venezuela, Bolivar, vicinity of Santa Teresita de Ka- 
vanayen; 1,220 m; F (holo). 

Drosera felix Steyerm. & L. B. Smith, Rhodora 76: 491. 
1974. J. A. Steyermark et al. 105468; 19 Feb. 1972; 
Venezuela, Bolívar, carretera El Dorado to Santa Elena 


de ci S of El n. ao 1,400 m; MO 


- Fieldiana Bot. 28(2): 
243. 1952. J. A. oma 58472; 8 Sep. 1944; 
Venezuela, T.F. Amazonas, vicinity of Sanariapo; 100 
m; F (holo). 

Drosera tenella Willd. var. esmeraldae Steyerm., Fieldi- 
ana Bot. 28(2): 244. 1952. J. A. Steyermark 57850; 
22 Aug. 1944; Venezuela, T.F. Amazonas, between 
i eit savanna and SE base of Cerro Duida; 200 
m; F (holo) = D. esmeraldae (Steyerm.) Maguire & 
Wurd., 1957. 


EBENACEAE 


Diospyr teguiensi Fieldiana Bot. 28(3): 
489. 1953. J. A. Steyermark 61513; 16 Mar. 1945; 
Venezuela, Anzoátegui, between Bergantín and San José; 
300-400 m; F (holo). 

des johnstoniana Standl. & Steyerm., A 
Bot. 22(3): 165. 1940. P. C. Standley 59934; Gua- 


temala, Dept. Sacatepéquez, along Rio Du dee. near 
m; F e lo iem 


Hor Bot. Ser. 22(4): 263. 1940. ra A. o 
ra de las Minas; 250-400 m; F (holo) = D. yatesiana 
Standl., 1967. 

ELAEOCARPACEAE 


Pittieria 7: 13. 1978. J. A. 


Sloanea autanae Steyerm., 


Steyermark 105222; 21-22 Sep. 1971; Venezuela, 
T.F. Amazonas, Cerro Autana; 1,230-1,270 N 
(holo) = S. steyermarkii C. E. Smith subsp. Mitina 


(Steyerm.) Steyerm., 1988 
Sloanea bolivarensis Beyer. .; Ann. Missouri Bot. Mad. 
75: = 1988. R. L. Liesner & A. González 11479 
. 1981; bar zuela, Bolivar, 7 km NE of Ciudad 
Piar; 350- 500 m; VEN (holo), M 
Sloanea breviseta Severn , Fieldiana ei 28(2): 357. 
1952. J. A. Stey id 622 31 . 1945; Ven- 
ezuela, Monagas, Montaña de cu  600- 900 m; 
F (holo). 
Sloanea carrenoi Steyerm., Bol. Soc. Venez. Ci. Nat. 32: 
349. 1976. J. A. Steyermark et al. 109699; 28 Feb.- 
1,2,5 Mar. 1974; Venezuela, Bolivar, Meseta del Jaua, 
Cerro Jaua; VEN (holo). 


Sloanea cataniapensis Steyerm., Ann. Missouri Bot. Gard. 


Venezuela, T.F. Amazonas, Dept. 
Cataniapo, 48 km SE of Puerto Ayacucho; 200-300 
m; MO, VEN (holo). 
Sloanea caudata Steyerm., Fieldiana Bot. 28(2): 357. 
1952. J. A. Steyermark 60544; 25 Nov. 1944; Ven- 


Volume 76, Number 3 
1989 


Taylor 677 
Plants Described by 
Julian A. Steyermark 


ezuela, Bolivar, Quebrada O-paru-ma; 915-1,065 m; 
F (holo). 

Sloanea cavicola Steyerm., Bol. Soc. Venez. Ci. Nat 
351. 1976. J. A. Steyermark et al. AAT » Feb. 
1974;. Venezuela, Bolivar, Meseta del Jau ro Sa- 
rasarinama; 1,320 m; MO (2582381), VEN yum 

Sloanea davidsei Steyerm., Ann. Missouri Bot. Gard. 75: 
1575. 1988. G. Davidse 27733; 23-25 July 1984; 
Venezuela, T.F. Amazonas, Dept. Rio Negro, Rio Pa 
cimoni, between its mouth and its jct. with the Rio Baria 
and Rio Yatua; 80 m; MO, VEN (holo). 

n . Venez. Ci. Nat 

als ud. Steyermark et al. 109841; 28 
Feb.-1, 2. 5 Mar. 1974; Venezuela, Bolivar, Meseta js 
Jaua, Cerro Jaua; 1,810-1,880 m; VEN (holo) — 
steyermarkii C. E. Smith subsp. Jauaensis HERE 
Steyerm., 1988. 

Sloanea jauaensis Steyerm. var. minor Steyerm., Bol. 
Soc. Venez. Ci. Nat. 32: 354, 1976. J. A. Steyermark 
et al. 109695; 28 Feb. 1976; Venezuela, Bolivar, 
Meseta del Jaua, Cerro Jaua; 1,810-1,880 m; VEN 
(holo) = S. steyermarkii C. E. Smith subsp. jauaensis 
(Steyerm.) Steyerm. , 1988. 

Sloanea larensis Sexe. , Acta Bot. Venez. 10: 239. 
1975. J. A Steyermark et al. 110148; 6 July 1974; 
Venezuela, Lara. Distrito Jiménez, Cubiro; 1,850-1,950 
m; VEN (holo). 

Sloanea urb Steyerm., Ann. Missouri Bot. Gard. 

15: 1577. 1988. R. Liesner 18455; 8-9 Mar. 1985; 
C T.F. Amazonas, Dept. Atabapo, Cerro Ma- 
rahuaca, 1-2 km of Sima Camp; 1,100 m; MO, 
VEN (holo). 

Sloanea maroana Steyerm., Pittieria 7: 14. 1978. J. 4. 
Steyermark & G. 5. Bunting 102799; d. T.F. 
Amazonas, 1 km al este de EET VEN (holo) = S. 
Y ler Spruce ex Benth., 

nea megacarpa PR 8 |. Berti, B 
ie Venez. Ci. Nat. 26: 467. 1966. L. Marcano Berti 

418; 29 Nov.-18 n 1964; Venezuela, T.F. Delta 


= 


Amacuro; E of Río Grande, NE of El Palmar; VEN 
(holo). 

Sloanea a dioe > Steyerm., Ann. Missouri 
Bot. Gard. 75: 15 . F. anb 1051 (also 


numbered 7053, i Ms eae uela, Bolivar, Alto 
Rio Guana (Merevari) near Brazil lentes F (holo), 
US, VEN. 

Sloanea onotillo Steyerm., Fieldiana Bot. 28(2): 358. 

4. Steyermark 61016; 20 Feb. 1945; Ven- 

ezuela, Anzoátegui, along Rio León by Quebrada Danta; 
500 m; F (holo). 

qus onotillo Steyerm. var major Steyerm., Padina 
Bot. 28(2): 359. 1952. J. A. Steyermark 61726; 23 
Mar. 1945; Venezuela, Anzoátegui, Río Marville, Fila 
Grande; 900-1,400 m; F (holo 


Sloanea cde ec Steyerm., Ann. Missouri PX Gard. 
U. Br 


75: 1579. 1988. N. T. Silva € U. Brazáo 60865; 
23 Jan. 1966; Brazil (near Venezuelan kg Serra 
de Neblina, Río Negro, Rio Cauaburi, Río Maturaca, 


between Missáo Salesiana and Serra Pirapucú; 800- 
1,000 m; MO (holo), NY. 

Sloanea petenensis Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(4): 172. 1944. J. A. Steyermark 
45309; 25 Mar. 1942; Guatemala, Dept. Petén, forest 
between i Yalpe Rio San Diego, and 
San Diego m; F (holo). 

Sloanea pittieriana Sieve: rm., nee Bot. 28(2): 359. 

52. J. A. Steyermark 6026 15-17 Nov. 1944; 


Venezuela, Bolivar, between Ptari-tepui and Sororopan- 
tepui; 1,615 m holo 

Sloanea ptariana Steyerm , Fieldiana Bot. 28(2): 300. 
1952. J. A. Steyermark 60668 ; 28 Nov. 1944; Ven- 
ezuela, Bolivar, Ptari-tepui; 1, 220 m; F (holo). 

Sloanea sipapoana Steyerm., Ann. Missouri Bot. Gard. 
75: 1580-1581. 1988. B. Maguire & L. Politi 27674; 
15 Dec. 1948; Venezuela, T.F. Amazonas, Cerro 
papo (Paráque), Camp Savanna; t 500 m; MO (holo), 
NY 


Sloanea subpsilocarpa Steyerm., Ann. Missouri Bot. Gard. 
15: 1583. 1988. L. Marcano Berti 447; 29 Nov.- 
18 Dec. 1964; Venezuela, T.F. Delta Amacuro, E de 
Rio Grande, ENE a Palmar, near limits of Bolivar; 
VEN (holo), MO, 

Sloanea venezuelana 20 Fieldiana Bot. 28(2): 361. 
1952. J. A. Steyermark 62216; 19 Apr. 1945: Ven- 
ezuela, Monagas, Montaña de Aguacate; 600-900 m; 
F (holo 

Sloanea Pa kii Steyerm., Ann. Missouri Bot. Gard. 
75: 1583. 1988. J. Wurdack € J. Monachino 41017; 
30 Dec. 1955; Venezuela, Bolivar, Rio Paraguaza, just 
below Raudal x (ca. 110 km above river mouth); 
115 m; MO (holo), N 

Sloanea d Steyerm.. Pittieria 7: 978. J. 
A. Steyermark & G. S . Bunting Tu seuils 
T.F. pb Cerro Y apacana; 1,000-1,200 m; VEN 
(holo) = S. steyermarkii C. E. Smith subsp. jauaensis 
(Steyerm.) Steyerm., 1988. 


ELATINACEAE 
Elatine ds ce Steyerm., Fieldiana d 28(2): 400. 
ermark 55905; 16 | 944; Ven- 


ela je near El Aguila; 3,960 m; F (holo), MO 
(1621390). 


ERICACEAE 


Be faria neblinensis Maguire, ee ae & Luteyn, Mem. 
ork Bot. Gard. 29(1): 1 1978. B. Maguire 

& 1 poe Pires 60477; 2 B. 1 965; Venezuela, 
T.F. Amazonas, Cerro de la Neblina; 2,500-2,800 m; 
NY (holo), VEN. Note: Bejaria i is the correct spelling. 


et al. 37254; Venezuela, T.F. Amazonas, Cerro de la 
Neblina; 1,500-1,700 m; NY (holo), US. 
Cavendishia salicifolia Maguire, Steyerm. 
fem. ork Bot. Gard. 29(1): 167. 
ded: & D. B. Fanshawe 32430; 29 Oct.-4 Nov 
1951; Guyana, Membaru-Kurupung trail; NY e 
US. 


Ceratostema glandulifera Maguire, Steyerm. $ Luteyn, 
Mem. New York Bot. ard. 29(1): 160. 1978. S. 5. 
Tillett et al. 45057; 8 Aug. 1960; Guyana, upper 
Mazaruni River, Mt. Ayanganna; 1,370-1,525 m; NY 
(holo). 

Ledothamnus atroadenus Maguire, Steyerm. & Luteyn, 
Mem. New York Bot. Gard. 29(1): 249. 1978. J. 4. 
Steyermark 73886; 13 Apr. 1953; Venezuela, Bolivar, 
Chimantá Massif, Abacapa-tepui; 2,125-2,300 m; F, 
NY (holo), V 

Ledothamnus dec ila Maguire, e & Luteyn, 
Mem. New York Bot. Ll 29(1): 143. 1978. J. 4. 
Steyermark & J. J. Wurdack 7. - 11 
Venezuela, Bolivar, Chimantá Massif; 2,200 m; F, NY 
(holo), VEN 


678 


Annals of the 
Missouri Botanical Garden 


Ledothamnus jauaensis Maguire, Steyerm. & Luteyn, 
Mem. New York Bot. Gard. 29(1): 146. 1978. J. A. 
Steyermark 97918; 22-27 Mar. 1967; Venezuela, 


Bolivar, Meseta de Jaua, Cerro Sarasarinama; 2,000 


m; NY (holo), VEN. 

Ledothamnus luteus Maguire, Steyerm. & Luteyn, Mem. 
New York Bot. Gard 29(1): 146. 1978 J. A. Steyer- 
mark & J. J. Wurdack 953; 20 Feb. 1955; Venezuela, 
Bolivar, Chimantá Massif; 1,895- 1,910 m; F, NY (holo), 
VEN. 


Ledothamnus stenopetalus iiri Steyerm. & Luteyn, 
Mem. New York Bot. Gard. 29(1): pe 1978. J. A. 
Steyermark 97944; 22-27 Mar. 1967; Venezuela, 
Bolívar, Meseta de E Im Sarasarinama; 1,922- 
2,100 m; NY (holo), V 

Ledothamnus stey polo x C. Smith subsp. longisetus 
Maguire, Steyerm. & Luteyn, Mem. New York Bot. 
Gard. 29(1): 152. 1978. J. A. Steyermark 97878; 
22-27 Mar. 1967; Venezuela, Bolivar, Meseta de Jaua, 
Cerro Sarasarinama; 1,922-2,100 m; NY (holo), VEN. 

Mycerinus chimantensis Maguire, Steyerm. & Luteyn, 
Mem. New York Bot. ide 29): 177. 1978. J. A. 
Steyermark & J. J. Wurdack 825; 13 Feb. 1955; 
Venezuela, Bolivar, Chimantá Massif; 2,210 m; F, NY 
(holo), VEN. 

Mycerinus O Steyerm. & Maguire, Bol. Soc. 
Ci. Nat. 32: 373. 1976. J. A. Steyermark et al. 109181; 
16-18 Feb. 1974; Venezuela, Bolivar, Meseta del Jaua, 
Cerro Sarisarinama; 1,400 m; VEN (holo). 

ui usc merumensis Maguire, Steyerm. & Luteyn, Mem. 

ork Bot. Gard. 29(1): 168. 1978. s. S. Tillett 
et Al 44867. , 12 July 1960; Guyana, Merume Mts., 


e, Steyerm. & Luteyn, Mem. 
vss rk Bot. Gard. 29(1): 171. 1978. R. S. Cowan 
& J. J. Wurdack 31061; 31 Jan. 1951; Venezuela, 
T.F. Amazonas, Serranía Parú, Rio Parú, Cano Asisa, 
along W Rim; 2,000 m; NY (holo). 

Orthaea pubifolia Maguire, Steyerm. & Luteyn, Mem. 

York Gard. 29(1): 171. 1978. B. Maguire & D. 
Fanshawe 32339; 29 Oct.-4 Nov. 1951; Guyana, 
Pakaraima Mts., Membaru-Kurupung trail; 1,000 m 
NY (holo). 

Orthaea thibaudioides Maguire, Steyerm. & Luteyn, 
Mem. New York Bot. Gard. 29(1): 173. 1978. B. 
Maguire et al. 30945; 15 Jan. 1951; Venezuela, T.F. 
Amazonas, Cerro Moriche, Rio Ventuari; 1,250 m; NY 
hol 


olo). 

Orthaea venamensis Maguire, Steyerm. & Luteyn, ius 
New York Bot. Gard. 29(1): 174. 1978. J. A. St 
ermark et al. 92423; 29-30 Dec. 1963; aloe 
Bolivar, Cerro Venamo; 950- 1,150 m; NY, VEN (holo). 


1957; Venezuela, T.F. Ama 
zonas, add de la Neblina, Rio Yatua; 1,900 m; NY 
(holo). 

Pernettia saxicola Standl. & RPM ve Mus. Nat 

ist., Bot. Ser. 23(3): 139. 1944. J. A. Steyermark 
36110; 19 Feb. 1940; — Dept. San Marcos, 
Volcan: Tack 4,400 m; F (holo). 

Psammisia breweri Eom. & Maguire, Bol. Soc. Ven- 
ez. Ci. Nat. 32: 377. 1976. J. A. Steyermark et al. 
109076; 12- 15 Feb. 19745 S ene Bolivar, Me- 
seta del Jaua, Cerro Sarasarinama; 1,000 ^E VEN 
N Psammisia urichiana (Britt.) A. C. Smith, 


1978 
Whadadendron roseum (Loisel.) Rehder f. albidum Stey- 


erm., Rhodora 62: 131. 1960. J. A. Steyermark 8522; 
= Ma ay d .S.A., Missouri, Ste. Genevieve Coun- 
y, E of Chimney Murus along River Aux Vases, 5 mi. 

E of Pickle; "MO. ree (holo). 
s intermedia Steyerm., Acta Bot. Venez. 2: 303. 
1967. J. A. Stey Nen iun 93980; 15 May 1964; Ven- 
ezuela, Bolívar, Auyan-tepui; 2,050-2,300 m; VEN 

lo 


(ho 
Tepuia multiglandulosa Steyerm. & Maguire, s Bot. 
z 1967 A. Steyermark 75733; 19 
June 1953; Val ra Auyan-tepui; 2, Do m; 
VEN (holo). 

RU vs tatei Camp var. — Steyerm. & Maguire, 

t. Venez, 2: 308. 1967. J. A. Steyermark & 

> 4 Feb. 1955; Venezuela, Bolivar, 

Auyan- -tepui; l, 925- 1,940 m; VEN (holo). 

"e paresciól Steyerm., Acta Bot. Venez. 2: 306. 
1967 Vareschi & E. Foldats 4966; Apr. 1956; 
e Bolivar, Auyan-tepui; 2,300 m; VEN (holo). 

Thibaudia breweri mn rm. & Maguire, mn Pu Venez. 
Ci. Nat. 32: 378.1 : t al. 109315; 
22-28 Feb. 1974; ld Bolivar, n del Jaua, 
Cerro Jaua; 1,750-1,800 m; VEN (holo 

Thibaudia carrenoi Steyerm. & Maguire, Bol. Soc. Ven- 
ez. Ci. Nat. 32: 380. 1976. J. A. Steyermark et al. 
109298; 22-28 Feb. 1974; Venezuela, Bolivar, Me- 
seta del Jaua, Cerro Jaua; 1,750-1,800 m; VEN = 

Thibaudia dolichandra Maguire, Steyerm. & Lute 
Mem. New York Bot. Gard. 29(1): 180. 1978. B. & 
C. K. Maguire 35149; 11 Feb. 1953; Venezuela, T.F. 
Amazonas, Cerro Yutaje, Serrania Yutaje; 1,400 m; 
NY (holo 

PBibaudia Jaunenils Steyerm. & Maguire, Bol. Soc. Ven- 
ez. Ci. Nat. 32: 383. 1976. J. A. Steyermark et al. 
109584; 27 Feb. 1974; Venezuela, Bolivar, Meseta 
del Jaua, Cerro Jaua; 2,228-2,250 m; VEN (holo). 

m. & Luteyn, Mem 


et al. 29564; 20 Nov. 1950; Venera. T.F. 
zonas, Cerro Duida; NY (holo). 


Thibaudia smithiana Maguire, Steyerm. & Luteyn, Mem. 


Amazonas, Cerro uos (Paraque); ¡A 815 m; NY (holo). 

Vaccinium chimantense Maguire, Steyerm. & Lute 

em. New York Bot. Gard. 29(1): 189. 1978. J. 4. 

Steyermark & J. J. Wurdack 669; 9 Feb. 1955; 
Venezuela, eie Vg es Mee Torono-tepui; 
2,165-2,180 , NY (ho 

Vaccinium E Sundl PY Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(3): 139. 1944. J. A. SHE 
mark 48572; 14 July 1942; Guatemala, Dept. Hue 
huetenango, Cerro Huitz; 1,500-2,600 m; F (holo). 

Vaccinium minarum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23: 219. 1947. J. A. Steyermark 
43295; 26 Jan. 1942; Guatemala, Dept. Zacapa, Vol- 
cán eem Sierra de las Minas; 2,100-3,200 m; F 


Vi accinium puberulum Klotzsch var. jauaensis Maguire, 
Mem. New York Bot. Gard. 29(1): 
. A. Steyermark 98054; 22-27 Mar. 
1967; Venezuela, Bolivar, Meseta de Jaua, Cerro Sar- 


asarinama; 1,922-2,100 m; NY, VEN (holo). 


ERIOCAULACEAE 


Paepalanthus holstii Steyerm., Ann. Missouri Bot. Gard. 


75: 311. 1988. B. K. Holst 3523; 22 May 1987; 


Volume 76, Number 3 
1989 


Taylor 679 
Plants Described by 


Julian A. Steyermark 


Venezuela, Bolivar, Distr. Piar, Vesp tepui, sum- 
mit; 2,300 m; MO (holo-3446413), V 
Paepalanthus meseticola Mold. & M Bol. Soc 
Venez. Ci. Nat. 32: 281. 1976. J. A. Steyermark et 
al. d 22- 28 Feb. 1974; Venezuela, Bolivar, 
Cerro Jau old. Herbarium (holo). VEN. 
Paepalanthus veneran Mold. & de ~ Soc 


mark et 
liva 


Meseta de Jaua, p Jaua; 1,750-1,800 m; Mold 
Ean (holo), VE 


ERTHROXYLACEAE 


Arto lon venez pod Steyerm., Fieldiana Bot. 28(2): 

95 4. Steyermark 60371; 20-21 Nov. 

ay Tamal, Bolívar, Quebrada O-paru-ma, be- 

tween Santa Teresita de Kavanayen and Rio Pacairao; 

1,065- " E m; F (holo) = Erthroxylum mucronatum 
Benth., 


EUPHORBIACEAE 


Amanoa pubescens Steyerm., Fieldiana Bot. 28(2): 304. 
1952. Ll. Williams 14439: 20 Feb. 1942; Venezuela, 
T.F. Amazonas, Rio Guainia; 128 m; F (holo). 

Bernardia venezuelana al Fieldiana Bot. 28(2): 
306. 1952. J. A. Steyermark 556 27-28 Feb. 
1944; Venezuela, Distr. Federal, arcis del Avila; 
F (holo). 

Chaetocarpus williamsii Steyerm., Dro Bot. 28(2): 
06. 1952. Ll. Williams 14312; 12 1942; Ven- 


ezuela, T.F. Amazonas, Rio Guainía, rubei 127 m; 


(holo). 
Conceveiba krukoffii Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 17(5): 414. 1938. B. A. Krukoff 8396; 26 


ct.-11 Dec. 1936; n Amazonas, Mun. Sào Paulo 
de Olivenca, basin of Creek Belem, basin of Rio Soli 
moes; d uM "NY (holo) = C. guianensis 


Aublet 

centia magnifica Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 17(5): 414. 1938. B. A. Krukoff 8698; 26 
Oct.-11 Dae, 1936; Brazil, Amazonas, Mun. Sao Paulo 
de Olivenca, basin of Creek Belem, basin of Rio Soli- 
moes; MO (1250474), NU (holo). 

Conceveiba simulata Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 17(5): 416. 1938. B. A. Krukoff 8616; 26 
Oct.-11 Dec. 1936; Brazil, Amazonas, Mun. Sao Paulo 

Belem, basin of Rio Soli- 

— C. guianensis 


Conceveibastrum ptarianum Steyerm., Fieldiana Bot. 
28(2): 308. 1952. J. A. oriol 60021; er 11 
Nov. 1944; Venezuela, Bolívar, Ptari-tepui; 1,585- 

inr ird ptariana (Steyerm.) 


Croi zatia a Paru Bot. 28(2): 1952, gen p 

erm., Nin 30(1) 4 

: 1973; o 

Distr. Federal, Cerro Naiguata: ^ 000 m; DAV, VEN 

holo). 

Croizatia neotropica Steyerm., Powers Bot. 28(2): 309. 
19 . Steyermark 61523; 18 Mar. 1945; Ven 
ezuela, pa do a E of Bergantín; F 
(holo). 

Croton chamanus Steyerm., ws Bot. 28(2): 312. 
1952. J. A. Steyermark 56232; 2 May 1944; Ven- 
ezuela, Mérida, Río Chama, mum Los González; 1,220- 
1,820 m; F (holo). 


Croton croizatii Steyerm., Fieldiana Bot. 28(2): 312. 
1952. J. A. Steyermark 56124; 26 Apr. 1944; Ven- 
canela, Mérida, near Hacienda Agua Blanca; 975 m; 
F (holo). 

Croton deserticolus Steyerm., re Bot. 28(2): 313. 
1952. J. A. Steyermark 56821; 28 May 1944; Ven- 


By zn between Carora e Barquisimeto; 500 


G e Dini S Steyerm., Los Angeles Co. Mus. Contr. 
Sci. 21: 4. 1958. E. Y 
Goiás, S Serra Dourada; 

Croton ma imae ed Brittonia 32: 21. 1980; 
J. A. Steyermark et al. 117473; 26 May 1978; Ven- 

ezuela, Bolivar. da id VEN (holo). 

ind iwi podes brc 30: 41. 1978. O, 
Hu & A. Cleef 5 4 Mar. 1977; Venezuela, 
Distr. "Federal, P ond El Avila; 2,000 m; 
VEN (holo). 

Croton inaequilobus Steyerm.; Los Angeles Co. Mus. 
C cl 9 . Y. Daw son. 14685; 25 


ur kavanayensis Steyerm., Fieldiana Bot. a 313. 

52. J. A. Steyermark 60386; 20- 1944; 

nos Bolfvas. Quebrada O-paru-ma; 1, 065- 1,220 
m; lo 

Croton renal Steyerm., Fieldiana Bot. 28(2): 314. 
1952. J. A. Steyermark 56806; 28 May 1944; Ven- 
ezuela, Lara, between *Teujllo-Lara boundary; 305 m; 

(holo). 

Croton roraimensis Croizat var. subintegrus Steyerm. 

Fieldiana Bot. 28(2): 315. 1952. J. A. Steyermark 
59925; 7 Nov. 1944; Venezuela, Bolivar, Ptari-tepui; 
2,410-2,450 m; F (holo) = C. roraimensis Croizat 
var. roraimensis. 

Croton standleyi Steyerm., Field Mus. Nat. Hist., Bot. 
Ser. 22(3): 151. 1940. M. E. Davidson 892; 12 July 
1938; Panamá, Chiriqui, Distr. Boquete, Volcán Chi- 
riquí; 2,100 m; F (holo), MO (1172438) = C. pungens 
Jacq., 7. 

Croton subincanus Muell. Arg. var. —€— 

yermark et al. 113330; 


quinima, a lo largo del Rio Szczerbanari (Rio Carapo); 
MO (iso— 2774296) 

Croton sucrensis Steyerm. Pec Bot. 28(2): 3 
1952. J. A. Steyermark 62 ; 23 May 1945; e 
np. Sucre, of Cumana, near Quetepe, between 
km and 20; 30 m; F (holo). 

Tide je dois Steyerm., Fieldiana Bot. 28(2): 
315. 1952. J. A. Steyermark 62532; 5 May 1945; 
Venezuela, Sucre, above La Trinidad SW of Cocollar; 
2,100-2,200 m; F (holo) 

Drypetes amazonica ia ., Field Mus. 
Bot. Ser. 17(5): 420. 193 


Nat. Hist., 


. Nat. . . Ser. : : . J. A. 
Steyermark 51646; 30 Aug. 1942; Guatemala, Dept. 
Huehuetenango, Sierra de los Cuchumatanes; 800- 
1,200 m; F (holo). 

ag wo [red wer Standl. & Steyerm., F ield Mus 
Nat , Bot. Ser. 23(3): 119. 1944. P. `. Standley 
80907: E Dec. 1940; Guatemala, Dept. Chimaltenan- 
go, between Chimaltenango and San Martin eiiim 


1,500-1,700 m; F (holo) 


680 


Annals of the 
Missouri Botanical Garden 


Euphorbia latazi H.B.K. var. glabra Steyerm., Fieldiana 
= 28(2): 316. 1952. J. A. Steyermark 55924; 15 
Feii Venezuela, Mérida, Moconoque; 2,510 m; 

F (ho 
Sia haha machrisiae Steyerm., Los Angeles Co. Mus. 
Contr. Sci. 21: 11. 1958. E. Y. Dawson 14594; 24 


Euphorbia pu Standl. & Steyerm., Field Mus 
Nat. Hist., Bot. Ser. 23(3): 120. 1944. W. A. Keller. 
man 5175; 10 Mar. 1905; Guatemala, Dept. Baja 
Verapaz, Sierra de las Minas, opposite El Rancho; F 
(holo). 

ag mi a & Steyerm., Field Mus. Nat. 
Hist., 23(3): 120. 1944. J. A. Steyermark 
50739; o yo 1942; Guatemala, Dept. Huehuete- 
nango, between San Ildefonso Ixtahuacán and Cuilco; 
1,350-1,600 m; F (holo). 

Euphorbia aes sa Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(3): 121. 1944. C. L. Wilson 
272; 4 Ma 1939; Guatemala, Dept. Alta Verapaz, 
Rio Chiacte; 480 m; F (holo). 

Gymnanthes a Standl. & Steyerm., Field 
Mus. Nat. Hist., Bot. Ser. 23(3): 122. 1944. J. A. 
Steyermark 33501; 5 Jan. 1940; Guatemala, Dept. 
Quezaltenango, S. Volcán Santa Maria; 1,300-1,400 
m; F (holo). 

“Hieronyma” croizatii Steyerm., Fieldiana Bot. 28(2): 
317. 1952. J. A. Steyermark 57005; 25 June 1944; 
Venezuela, Distr. Federal, E of El Junquito; 1,980- 
2,130 m; F (holo). Note: Hyeronima is the correct 
spelling. 

“Hieronyma” oblonga (Tul.) Muell. Arg. var. genuina 
Muell. Arg. f. glabra Steyerm., Fieldiana Bot. 28(4): 
951. 1957. J. A. Steyermark 60686; 28 Nov. 1944; 
Venezuela, Bolivar, Ptari-tepui; 1,220 m; F (holo) — 
ei oblonga (Tul.) Muell. Arg. var. oblonga, 

967. 


— 


iy standleyi Pew Field Mus. Nat. Hist., Bot 
Ser. 22(3): 152. Matuda 2260; July 1936; 
Mexico, Oaxaca, diee F (holo). 

Mabea elata Steyerm., Field Mus. Nat. Hist., Bot. Ser. 


17(5): 418. 1958. G. Klug 3206; Aug.-Sep. 1933; 
Perú, Loreto, between Balsapuerto and Moyobamba; 
600-1,200 m; F (holo). 

Mabea excelsa Standl. & s a Field Mus. Nat. Hist., 
Bot. Ser. 23(3): 123. 1944. 4. F. Skutch 2008; 27 
Dec. 1934; Guatemala, Dept. Quezaltenango, Color 
bia; 850 m; F (holo). 

Mabea klugii Steyerm., Field Mus. Nat. Hist., Bot Ser 

1938. G. Klug 1969; Jan.-Feb. 1931; 


17(5): 416 
Colombia, Conisañk del Putumayo, Umbria; F (holo). 
Mabea pirioides Steyerm., Field Mus. Nat. Hist., Bot. 


ae 17(5): 419. 1938. B. A. Krukoff 1983; 24 Oct. 
32 A io Campo de Boa Esperanca, Maranhao, 
Ma: ume River region; F (holo). 

M es prse i Steyerm., Field Mus. Nat. Hist., Bot. 
Ser. 17(5): 417. 1938. G. Klug 2064; Mar. -Apr. 
1931; Perú, Dept. Loreto, Florida, Rio Putumayo, at 
mouth of Rio Zubineta; 200 m; F (holo). 

Omphalea elaeophoroides Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 22(3): 152. 1940. B. A. Krukoff 6297; 
14 Sep.-11 Oct. 1934; Brazil, Amazonas, Piraninhon, 
Mun. Humayta, near Tres Casas, basin of Río Madeira; 

, NY (holo). 

Pedilanthus camporum Standl. & Steyerm., Field Mus. 

Nat. Hist., Bot. Ser. 23(3): 124. 1944. P. C. Standley 


87781; 18 Feb. 1941; Guatemala, Dept. Retalhuleu, 
between Nueva Linda and S E UN, 120 m; F (holo) 
— P. tithymaloides (L.) P 
Phyllanthus bolivarensis Aden , Fieldiana Bot. 28(2): 
2. J. A. Steyermark 57688; 31 July-1 Aug. 
ioi eins Bolivar, 1-10 is NW of Upata; 
700 m; F (holo) 

Phyllanthus carrenoi Steyerm., Bol. Soc. Venez. Ci. Nat. 
32: 343. 197 Steyermark et al. 109596; 27 
Feb. 1974; Veneris. Bolivar, Cerro Jaua, Cumbre; 
2,228-2,250 m; VEN (holo). 

Phyllanthus croizatii Steyerm., 
317. 1952. J. A. Steyermark 56854 
Venezuela, Yaracuy, near Taria; m: ( clo). 

Phyllanthus dawsonii Steyerm., Los Angeles Co. Mus. 

contr. Sci. 21: 15. , nom. nov. 

Phyllanthus HE Steyerm. & Luteyn, Ann. 
Missouri Bot. Gard. 71: 317. 1984. J. A. Steyermark 
129816; 8 Feb. 1984, enla T.F. Amazonas, Río 
Negro, Cerro de la Neblina; 1,880 m; MO (3223155), 
NY, VEN (holo). 

Phyllanthus d hide » Mose I 28(2): 318. 

1952. mark 56 28 1944; Ven- 
me Lara Tren Trojilo- den onda. and Ca- 
; 305 (holo). 

Phrlianihus longipes, Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 22(3): . 1940. P. H. Gentle 2619; 13 
May 1938; ad D Cayo Distr., Vaca; F (holo), 


Fieldiana oe 28(2): 
: y 1944; 


M 
Phyllanthus major Steyerm., Fieldiana Bot. en 318. 
952. J. A. Steyermark 59482; 29 Oct. 1944; Ven- 


Phyllanthus orinocensis Steyer 
321. 1952. J. 2 md 57891; 23 Aug. 1944; 
Veiene; T. * Ait 1azonas, SE base of Cerro Duida; 
210 m; F (ho 

Phyllanthus bus ola ao 

75. M. Farinas et 
UE TF. Amazonas, 
VEN (holo 

Phyllanthus vacciniifolius ( (Muell. Arg.) Muell. Arg. Se 
aa Steyerm., Ann. Missouri Bot. Gard. 

: 4. J. A. Steyermark et al. 130328; iir 
1984; Nansen: T.F. Amazonas, Serrania Vinila, N 
of Cerro Aratitiyope; 440 m; VEN (holo). 

Phyllanthus PAP a Los Angeles Co. 
Mus. Contr : 8. E. Y. Dawson 14918; 
17 May 1956; Brazil, em d Dourado, 20 km 
E of Formoso; F, G, LAM, R (holo). 

Phyllanthus zanthoxyloides Steyerm., Fieldiana Bot. 
28(2): 321. 1952. J. A. Steyermark 62189; 19 Apr. 

1945; A Monagas, Montaña de Aguacate; 
600-900 m; F (holo). 

Richeria submembranacea Steyerm., Field Mus. Nat. 
Hist., Bot. Ser 17(5): 419. 1938. B. A. Krukoff 8513; 
11 Sep.-26 Oct. 1936; Brazil, Amazonas, Mun. Sào. 
Paulo de Olivenca, near Palmares, basin of Rio Soli- 
moes; NY (holo) = ea racemosa (Poepp. & Endl.) Pax 
& K. Hoffn 

UAE. Bm Bot. 

, gen. 

senderos aiii n., Bot. Mus. Leafl. 15(2): 

l. J. A. Steyermark 60849; 4-5 Dec. 1944; 


Acta Bot. Venez. 10: 
l. 433; Jan.-Feb. 1969; 
pes Duida; 1,000 m; 


Mus. Leafl. 15(2): 45. 


Volume 76, Number 3 
1989 


Taylor 681 
Plants Described by 


Julian A. Steyermark 


Venezuela, Bolivar, Carrao-tepui, SE base; 1,460-1,615 
m; F (holo). 
Stillingia cruenta Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(3): 125. 1944. P. C. Standley 
91207; 4 ae Ma Guatemala, Dept. Baja Verapaz, 
above Sa ; 1,500 m; F (holo). 
Terorchidium brevifolium Standl. & Steyerm., Field Mus 
r. 23(3): 126. 1944. C. L. Wibon 
He 11 Feb. 1039; Guatemala, Dept. Alta Verapaz, 
Rubelpec; F (holo). 


FABACEAE 


Aeschynomene d Standl. & Steyerm., Field Mus 
Nat. Hist., iae er. 23(1): 9. 1943. J. A. Steyermark 

1939; Guatemala, Zacapa, Santa Ro- 
salia, 2 m LS x Zacapa; 200 m; F (holo). 

P PUE tricholoma Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(1): 10. 1943. P. C. Standley 
73714; Oct. 1940; Guatemala, Dept. Chiquimula, near 
divide on road from Zacapa to Chiquimula; 660 m 

l 


(holo). 
Aldina berryi Cowan & Steyerm., Ann. Missouri Bot. 
ard. 71: 312. 1984. J. A. Steyermark et al. 117468; 
26 May 1978; Venezuela, Bolívar, Cerro Guaiquinima; 
950 m; MO (3242295, 3234984), US, VEN (holo). 


Apios americana Medic. f. pilosa Steyerm., Rhodora 
“40: 179. 1938. J. A. Steyermark 11390; 7 July 1936; 
U.S.A., Missouri, Butler Co., Clark 


ational Forest, 

Wappapello Purchase Unit, Mud Creek, 2 mi. NW 
Rombauer; olo). 

eS carcerea Standl. & RS Field Mus. Nat. 

, Bot. Ser. 23(4): 161. 1944 . Steyermark 
ds 20 dun. 1942; Guatemala, Dept. El Progreso, 
between Calera and Volcán Siglo; 2,000-2,200 m; 
(holo). 

Canavalia munda Standl. & Steyerm., e Mus. Nat. 

ist., Bot. Ser. 23(1): 10. 1943. J. 4. Steyermark 
33179; Dec. 1939; Guatemala, Dept. Quezaltenango, 
Finca Pirineos, Volcán Santa María; 1,300-1,500 m; 
F (holo). 

Cassia prre- Michx. f. aia ur & Steyerm., 
Ann. Missouri Bot. Gard. 22: 574 5. L. P. Jensen 
sna 20 Aug 1928; U.S.A., eh Franklin Co. 

ray Summit; MO (holo). 

Cladrastis en e f.) Koch. f. tomentosa Steyerm., 

hodora 40: 487. 1938. E. J. Palmer 35387 
29; U.S.A., Alabama, Tuscaloosa Co., along rock 
_ bluffs of Bla ck Warrior 


Z 
E 
= 


Crudia lacus Standl. & Steyerm., : 
Bot. Ser. 22(5): 339. 1940. J. A. dal 39611; 


Dalbergia pacifica Standl. 8 Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 22(4): 236 0. P. C. Standley 
62109; 5 Jan. 1939. Guatemala, Dept. Suchitepéquez, 
Cocales; 215 m; F (holo 

Dipteryx EEV Steyerm., Ann. Missouri Bot. Gard. 
11: 313. 1984. J. 4. Steyermark et al. 113200; 20- 

25 Jan. 1977; Venezuela, Bolivar, Cerro Guaiquinima; 
MO (2769905), VEN (holo) = Spirotropis angusti- 
folia (DC.) Baill 


Eee a acto (Walp.) O. F. Cook f. redmondii 
Ste r, Phytologia 48(4): 286. 1981. P. 
Retina a n.; : 9 Mar. 1 1981; Venezuela, Miranda, Los 
Chorros, Avenida principal, Caracas; MO (3234983), 
VEN (holo). 

Galactia acuminata Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 22(3): 144. 1940. E. Matuda 1741; 12 Aug. 
1937; Mexico, Chiapas, Finca Juárez; F (holo). 


a . 

Galactia sparsiflora Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(4): 162. 1944. J. A. Steyermark 
43837; 11 Feb. 1942; Guatemala, Dept. El Si Sa 
E of Finca Piamonte, Sierra de las Minas; 2,500 m 


(holo). 

a a a modestus Standl. & Steyerm., Field Mus 
Nat og Ser. 23(2): 56. 1944. J. A. Seana 
39238; 1940; Guatemala, Dept. Izabal, Bay of 
Santo RUN between Escobas and Santo Tomás; sea 
level; F (holo). 

Lonchocarpus og rid llus — & Steyerm., Field 
Mus. Nat. Bot. Ser. 23(2): 56. 1944. P. C 
Standley 74569; Oct. 1940; Guatemala, Dept. Chi- 
quimula, between Ramirez & Cumbre de Chiquimula; 
400-600 m; F (holo 

to fruticerrum Standl. & Steyerm., Field Mus. 

at , Bot. 22(4): 240. 1940. J. A. Steyer- 
mah TAR. a "Oct. 1939; Guatemala, Dept. Chi- 
qmm = Chiquimula and La Laguna; 500- 
1,000 m; F (holo). 
Mimosa canahuensis Standl. & vx Field Mus. 
r. 23(4): 163. 1944. J. A. Steyer- 
. 1942; Aib un Dept. El 
Progreso, Montan Canahui; 1,600- 2,300 m; F (holo). 


29018; Oct. 1939; Dostenals. Dept. Zacapa, near 
Santa Rosalia; 250-350 m; F (holo). 
— Standl. rs = Mus. Nat. Hist., Bot. 
23(1): 12. 1943, 
ritos p cerrar Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(1): 13. 1943. P. C. Standley 
75307; Oct. 1940; Guatemala, Dept. Jutiapa, near 
Jutiapa; 850 m; F (holo). 
doe saxosum Standl. & Steyerm., Field Mus 
t. Hist., Bot. Ser. 23(4): 163. 1944. P. C. Standley 
74367; 14 Oct. 1940; Guatemala, Dept. Chiquimula, 
Quebrada Shusho, above Chiquimula; 480 m; F (holo). 
Pithecolobium zollerianum Standl. € Steyerm., Field 
Mus. Nat. Hist., Bot. Ser. 22(5): 343. 1940. J. A 
Steyermark 37440; 9 Mar. 1940; Guatemala, Dept. 


dera Shac, Volcán Tajumulco; 1,300-1,500 m; F (holo). 
Psoralea io Pursh f. alba Steyerm., eg 41: 

585. 193 . Steyermark 5779; ne 1938; 

U.S.A., van Benton o., along Osage River, 25 
mi. W of Warsaw; MO (holo— 1174669), F. 
Rhynchosia jalapensis Standl. & Steyerm., Field Mus 


Lm 
pz 
> 
e 
3 
- 
Ou 
= 
N 
pS 
bo 
== 
RE 
=] 
mM 
No) 
A 
N 
v 
at 
Lo” 
S 
> 
A 
m 
3 
a 
* 
15 


label) 1977; 


Vorena, Bolívar, Cerro Gonilna, 


682 


Annals of the 
Missouri Botanical Garden 


Caparo); MO (2775619); US, VEN (holo). 


FAGACEAE 
Quercus n Michx. f. angustior Palmer & Steyerm., 
Ann Missouri Bot. Gard. 25: 771. 1 A. Stey- 
ermark 12318; 27 July 1936; U.S 


Bot. Gard. 22: 5 ] 
1919; U.S.A., Missouri, Butler Co., Poplar os 
GH (holo). (Q. phellos x Q Siu 
ercus X mutabilis Palmer Peek , Ann. Missouri 
Bot. Gard. 22: 521. 1935. EJ Pie 26069; 10 
4; U.S.A., Missouri, Bates Co., Montieth Junc- 


tion; GH (holo). (Q. palustris X Q. shumardii var. 
schneckii 

Quercus Xvaga Palmer F Steyerm., 2 Missouri Bot. 
Gard. 22: 521. 1935. E. J. Palmer 25421; 13 Jun 
1924; U.S.A., Missouri, Nodaway va Maryville GH 
(holo). (Q. palustris x Q. velutina) 


O 


FLACOURTIACEAE 


Banara larensis Steyerm., Fieldiana Bot. 28(2): 404. 
1952. J. A. Steyermark 55424; 10 Feb. 1944; Ven- 
ezuela, Lara, between Las E Los Aposentos 


and Buenos Aires; 1,675-1,430 m; F (holo), K, NY, 
US. 


Banara paucinervosa Steyerm., Fieldiana Bot. 28(2): 
406. 1952. J. A. Steyermark 55824; 27 Mar. 1944; 
Venezuela, Mérida, above y Cuadras "pe Quebrada 
Molino; 1,820-2,255 m; F (holo), NY, Xy- 
losma paucinervosa (Steyerm.) Sleumer, id 

Bartholomaea Standl. & Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 22(4): 251. 1940, gen. nov. 

Bartholomaea mollis Standl. & Steyerm., Field Mus 
Nat. Hist., Bot. Ser. 22(4): 252. 1940. J. 4. Steyer- 
mark 39452; 27 Mar. 1939; Guatemala, Dept. Izabal, 
Río Dulce, near Livingston; F (holo). 

Casearia tachirensis Steyerm., Fieldiana Bot. 28(2): 406. 
1952. J. A. Steyermark 57276; 14 July 1944; Ven- 
ezuela, Táchira, above Betania, at base of Páramo de 


Tamá; 2,285-2,430 m; F (holo), N 


97851; 22-27 Mar. 1967; Venezuela, Bolivar, Meseta 
de Jaua, Cerro Jaua; 1,922-2,100 m; L, NY, 
VEN (holo). 

Hasseltia monagensis Steyerm., Fieldiana Bot. 28(2): 
407. 1952. J. A. Steyermark dd 7 Apr. 1945; 
Venezuela, Monagas, Cerro de la Cueva de Dona Anita; 
1,100-1,200 m; F (holo), NY, US, VEN = H. flori- 
bunda H.B.K., 


Homalium anzoateguiensis Steyerm 


" WT Bot. 


m; F (holo), L, NY, US, VEN = H. 
racemosum Jct i 
irri casiquiar, s Steyerm., Fieldiana Bot. 28(2): 
1952. Ll. Vi liams 15564; 25 May 1942; Ven- 
uara, Alto e 
— R. specio 
.) Monatcbina, 1980. 
Xylosma sessile Standl. m Field Mus Nat. Hist., 
Bot. Ser. 23(4): 177. 1944. H. von Tuerckheim Il. 1617; 


& 


Jan. 1907; Guatemala, Dept. Alta Verapaz, Cobán; F 
(holo), US = X. amense Turcz 

Xylosma trinervium Standl.& Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(4): 178. 1944. J. 


nango, along Rio Cuilco, between Cuilco 
1,200-1,300 m; F (holo) — Nonlin. spa 
(Hemsley) S. Watson, 1980. 


GARRYACEAE 


Garrya corvorum Standl. & Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 23(1): 16. 1943. P. C. Standley 81651; 
Dec. 194 0; Guatemala, Dept. Huehuetenango, region 
of Chemal, Sierra de los Cuchumatanes at km 36; 3,300 
m; F (holo). 


GENTIANACEAE 


cen bella Maguire & Steyerm., Mem. New York 

t. Gard, 32: 383. 1981. B. Maguire et al. 42096; 

19 Nas l1 Venezuela, T.F. Amazonas, Cerro de 
la Neblina; 1,800 m; NY (holo), US, VEN. 

Cent taurium rosans Standl. & Steyerm., Field Mus. Nat 


= C. pauci- 
orum (Mart. & Gal.) Rob., 1976 

Chorisepalum acuminatum Steyerm., Bol. Soc. Venez. 
Ci. Nat. 23: 75. 1962. J. A. Steyermark & S. Nilsson 
450; 21 Apr. 1960; Venezuela, Bolivar, Cerro Vena- 
mo; 1,140 m; UP, VEN (holo) = C. psychotroides 

w ar. acuminatum (Steyerm.) Maguire, 1981. 

Chorisepalum breweri Steyerm. & M 
Venez. Ci. Nat. 32: 39 
al. 109255; 19-20 Feb. 
Meseta del Jaua, Cerro Sarasariñama; 1,350 m 
(holo). 

Gentiana guatemalensis Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(2): 75. 1944. P. C. Standley 
81113; Dec. 1940; Guatemala, Dept. Huehuetenango, 
region Chemal, Sierra de los Cuchumatanes; 3,300 
m; F (holo). 

Gentiana lewisiae Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(2): 76. 1944. P.C. Standley 84556; 
Guatemala, Dept. Totonicapan, Pacaja, region of El 
Desconsuelo; 3,100-3,200 m; F (holo). 

Gentiana pumilio Standl. & Steyerm., Field Mus. Nat. 


; VEN 


"i Volcán Taju- 


, Bol. Soc. Venez. Ci. 

Nat. 26: 439. 1966. J. A. ear & S. Nilsson 
732; 25 Apr. 1960; Venezuela, Bolíva ao 
at km 146; 1,220 m; VEN (holo) = bahia cardonae 
ipis) es 1981. Note: Lisianthius is the cor- 
rect sp 

Lisianthli]us alioli Standl. & Steyerm., Field Mus. 

ist., Bot. Ser. 22(4): 267. 1940. C. L. Wilson 


ark & . W 
E Bolivar. e Massif, Summit Camp; 1940 


Volume 76, Number 3 
1989 


Taylor 683 
Plants Described by 


Julian A. Steyermark 


m; NY (holo) = Celiantha chimantensis (Steyerm. & 
Maguire) Maguire, 1981. 


€— diia Standl. & Steyerm., Field Mus. Nat 
Hist., S s 67. 1940. J. A. teyermark 
js abal, between 


940; yep Dept. Iz 

Milla a : p ridge 6 mi. from Izabal, Montana del 
Mico; 65-600 m; E (holo), 

Lisianth{ilus jauaensis Steyerm. & Maguire, Mem. New 
York Bot. Gard. 23: 879. 1972. J. A. Steyermark 
98068; 22-27 Mar. 1967; Venezuela, Bolivar, Cerro 
Jaua; 1,922-2,100 m; VEN (holo 

Lisianth[ ilus petenensis Standl. & Steyerm., Bull. Torrey 
Bot. Club 84(1): 46. 1957. C. H. Lundell 3153; 4 

1933; Guatemala, Dept. Peten, Nicton, Lake Pe- 
ten; US (holo). 

SE ha scabridulus Steyerm., beo Bot. 28(3): 
496. teyermark 5 ; 25-26 Aug. 
= E T.F. Amazonas, eu Duida; 1,200 

F (holo) = Irlbachia tatei (Gleason) Maguire, 1981. 
en aa neblinae "rer & down Mem. New 

. B. Maguire et al. 
37103; 6 Jan. 1954; ae a Amazonas, Cerro 
de la Neblina; 2,000 m; NY (holo) 

Macrocarpaea rugosa Steyerm., Bol. Soc. Venez. Ci. 
Nat. 25: 83. 1963. J. A. Steyermark, & J. J. Wurdack 
861; 13 Feb. 1955; Venezuela, Bolivar, Chimantá Mas- 
sif; 2,185-2,210 m; C, NY, VEN (holo). 

Schultesia guianensis (Aublet) Malme f. lutescens Standl. 
& Steyerm., Field Mus. Nat. Hist., Bot. S 
77. 1944. P. C. Standley 76561 Nov. 1940 Gua- 


Dept. Rio Negro, Cerro Aracamuni, summit, Popa Camp; 
1,550 m; MO (holo), VEN. 

Symbolanthus huachamacariensis Steyerm., Ann. Mis- 
souri Bot. Gard. 75: 1083. 1988. B. Maguire et al. 
29859; 4 Dec. 1950; Venezuela, T.F. Amazonas, Cerro 


Cae iare lensis Steyerm:, Ann. Missouri Bot. 
.F. A nas, Dept. Atures, sum- 
Yavi; 100 m; a: MO (holo) VEN. 
Tapeinostemo eweri Steyermark Maguire, FE 
Bot. Venez. 143) 42. 1984. B. pens et al. 655 
19 Jan. 1981; Macer T.F. Amazonas, Cerro ii, 
huaca; 2,685; m; K, MG, MO, NY, US, VEN (holo). 
Tapeinostemon jauaensis "eia & Maguire, Bol. Soc. 
Ven . Nat. 32: 394. 1976. J. A. Steyermark et 
al. 109535; 26 Feb. 1974; Venezuela, Bolivar, Meseta 
del Jaua, Cerro Jaua; 1,800 m; VEN (holo). 
d longiflorum Maguire & Steyerm., Mem 
ork Bot. Gard. 32: 359. 1981. B. Maguire et 
al. 42134; 17 Nov. 1957; Venezuela, T.F. Amazonas, 
Cerro de la Neblina; 1,800 m; K, NY (holo), US, VEN. 


Gard. 32: 362. 1981. J. A. Steyermark 103937; 16- 
17 Oct. 1970; Venezuela, Cerro de la Neblina NY 
(holo), VEN. 

Tapeinostemon ptariense Steyerm., Lloydia 14(1): = 
1951. J. A. Steyermark 59861; 4 Nov. 1944; 


ezuela, Bolivar, Ptari-tepui, E. of ‘Cave Rock’; 213 m; 
F (holo) = T. pena Benth., 1981. 
Tapeinostemon r m Maguir e & Steye Mem 
New York Bot. Cad. "32: 366. 1981. J. 4 am 
& J. J. Wurdack 1227; 1 Mar. 1955; Venezuela, 
Bolivar, Chimantá Massif, Torono-tepui; NY (holo), 
VEN. 


rca dd zamoranum Steyerm., Lloydia 14(1): 63. 
1951. J. A. Steyermark 54560; 10 Oct. 1943; Ec- 
uador, Prov. Santiago- amora, vicinity of Rancho 
puer along Quebrada Honda; 2,500-2,700 m; 
F (holo). 
br "via Steyerm., Ann. Missouri Bot. Gard. 28: 
1941. P. H. Allen 2240; Panama, Coclé, N of 
A a k Antón; F (holo) = V. acata (Standl.) 
Standl. & Steyerm., 1969. 


GESNERIACEAE 


Besleria mortoniana mere i dies 22: 303. 1970. 
A yermark 94962; 966; Veneta, 
Sucre, Peninsula de Paria, yee PE Humo; 1,060 
1,273 m; US (holo), VEN. 

Solenophora abietorum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 236. 1947. J. A. Steyer- 
mark 48643; 14 July 1942; Guatemala, Dept. Hue 
huetenango, Cerro Huitz, Sierra de los Cuchumatanes; 


1,500-2,600 m; F (holo) 


GRAMINEAE see Poaceae 
GROSSULARIACEAE 


Phyllonoma cacuminis Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 22(5): 335. 1940. J. A. Steyer- 
mark 29870; 13 Oct. 1939; Guatemala, Dept. Zacapa, 
Quebrada Alejandria, summit of Sierra de las Minas, 
vicinity of Finca Alejandria; 2,500 m; F (holo). 


GUNNERACEAE 


Gunnera pittieriana Badillo & Steyerm., Acta B pes 
: = 311. 1973. V. M. Badillo 4866; 18 June 972; 
uela, Aragua, 2 Nacional Henri DI Alto 
o). 


ue 
ha Choroni: MY (hol 
GUTTIFERAE = CLUSIACEAE 


HELICONIACEAE 


"on schneeana Steyerm., Fieldiana Bot. 28(1): 161. 
l. J. A. Steyermark 56083; 25 Apr. 1944; Ven- 
ien Mérida, above La Azulita, Hacienda Agua Blan- 

ca; 1,310-1,340 m; F (holo). 


HUMIRIACEAE 


“Houmiria” po Steyerm., Fieldiana Bot. 28(2): 270. 
952. J. A. Steyermark 60289; 15-17 Nov. 1944; 
Venezuela, Bolívar, between Ptari-tepui and Sororopan- 
tepui; 1,615 m; F (holo). Note: Humiria is the correct 
spelling. 


HYDROPHYLLACEAE 


sph saa i appendiculatum Michx. f. album Stey- 
. Rhodora 54: 257. 1952. J. A. Steyermark 67519; 

19 ) May 1949; U.S.A., Missouri, Johnson Co., Knob- 
noster State Park, 3-5 mi. SW of Knobnoster; F (holo). 
Phacelia hirsuta Nutt. f. albiflora Palmer & Steyerm., 


— 


684 


Annals of the 
Missouri Botanical Garden 


Brittonia 10: 116. 1958. E. J. Palmer 65070; 15 
May 1957; U.S.A., Missouri, Newton Co., 1 mi. S of 
Granby; F (halo ). 


HYPERICACEAE = Clusiaceae 


ICACINACEAE 


Emmotum ptarianum Steyerm., Fieldiana Bot. 28(2): 
341. 1952. J. A. Steyermark 59974; 10-11 Nov. 
1944; Venezuela, Bolivar, Ptari-tepui; 1,585-1,600 
m; F (holo) = E. glabrum Benth. ex Miers, 1978. 

Oecopetalum greenmanii Standl. & Steyerm.; Field. Mus. 
Nat. Hist., Bot. Ser. 22(3): 154. 1940. J. A. Steyer- 
mark 39516; 16 Apr. 1940; Guatemala, Dept. Izabal, 
Rio Dulce, 2-4 mi. W of Livingston; sea level; F (holo). 


IRIDACEAE 
Orthrosanthus chimboracensis (H.B.K.) Baker var. 
o (H.B.K.) Steyerm. f. albus Steyerm., Lloy- 


11(1): 19. 1948. J. A. Steyermark 55339a; n.d.; 
Venezuela, Trujillo, La Quebrada Cortijo above Hu- 
mocaro Bajo; F (holo) = O. acorifolius (Kunth) Ra- 


na, 
Orthrosanthus chinborne ensis (H.B.K.) Baker var. cen- 
l 


A. Steyermark 31913; n.d.; Guatemala, Jutiapa, Vol 
cán Suchitán, NW of ción Mita; 0). 

Orthrosanthus chimboracensis (H.B.K.) Baker var 
exsertus Foster f. albus Steyerm., Lloydia 11(1): 17 
1948. Runyon 875; 8 hu 1926; Mexico, Tamauli 
pas, Santa Rita, Ranch Tamaulipas; 1,500 m; US (holo) 
= O. exsertus (Foster) Ravenna, 1987. 

Orthrosanthus chimboracensis (H.B .K.) Baker var. in- 
termedius om Lloydia 11(1): 19. 1948. Type 
not indica 


Trimezia ios AN Steyerm., Ann. Missouri Bot. Gard. 
71: 301. 1984. J. A. Steyermark et al. 128071; 26- 
29 Jan. 1983; Venezuela, Bolivar, Distr. Piar, Macizo 
de Chimantá, sector centro-noreste del mae -tepui; 
2,000 m; VEN (holo). 

Trimezia y avc beide rm. Fieldiana Bot. 28(1): 160. 
1951. Steyermark 60834; 4 Dec. 1944; Ven 
ezuela, "oa Rio Tek-Yunsen between Santa Te- 
resita de Kavanayen and Carrao-tepui; 1,375 m; F 

0) 


IXONANTHACEAE 


Ochthocosmus attenuatus Steyerm. & Luteyn, Brittonia 
32: 142. 1980. J. A. Steyermark et al. 117570; 15- 
16 Nov. 1978; Venezuela, Bolivar, near San Rafael de 
Camoiran, km 175 S of El Dorado; 1,090 m; NY (holo), 


Oc jhocosmus berryi Steyerm., Ann. Missouri Bot. Gard. 
71: 313. 1984. J. E Steyermark et al. 130228; 26 
Feb. 1984; Venezue mazonas, Cerro Aratiti- 
yope; MO Bum "VEN (holo). 

Ochthocosmus ates aes Steyerm. 
Ci. Nat. 26: 421. 1966. J. A. Ste 

Dec. 1963; an Bolivar, Cerro Venemo; 
950-1,150 m; VEN = a O. roraimae var. gran- 
difolius (Steye Lut 

Ochthocosmus longipe dic in Steyerm: & Luteyn, 
Brittonia 32: 141. 1980. J. A. Stey 
117569; 15-18 Nov. 1978; AR 
175 S of El Dorado, near San Rafael de Camoiran; 
1,090 m; NY, VEN (holo). 


ol. . Venez. 
yermark et 2 92379; 


Ochthocosmus micranthus Steyerm., Ann. Missouri Bot. 
Gard 75: 1988. L. Hernández 348 
1986; Venezuela, vade, ori 10 i SW of 
i m; MO, VEN (ho 


Amazonas, NW base Cerro Yapacana, Savanna No. 3; 
125 m; NY, US, VEN (holo). 
Ochthocosmus multiflorus Ducke var. 
Steyerm. & Luteyn, dicare 32: 141. P 
Steyermark et al. 113837; 2 Mar. 1977; Vende 
T.F. Amazonas, edu lado sur del Rio Ventuari; 


125 m; F, NY, VEN (holo) 


canaripoensis 
1980. J. A. 


JUGLANDACEAE 


Carya buckleyi Sarg. var. arkansana Sarg. f. glabra 
Palmer & Steyerm., Ann. Missouri Bot. Gard. 25: 770. 
1938. J. A. mail 11461; 8 July 1936; U.S.A., 
Missouri, Butler Co., near Cane Creek, off N For 
Hollow, 10 mi. SE of Ellsinore; MO (holo— 1130651) 

— C. texana Buckl. var. texana, 1981. 


LAMIACEAE 


Sc utellaria nervosa Pursh f. alba Steyerm., Rhodora 54: 
^" . A. Steyermark 68024; 21 May 1949; 
Miss ssouri, Chariton Co., along Yellow Creek, 

near i ert F (holo). 


LAURACEAE 


Benzoin aestivale (L.) Nees var. pubescens Palmer & 
| 45. 1935. 
; U.S.A., Missouri, 
Jasper Co., Alba; GH (holo) = Lindera benzoin (L.) 
Blume, 1981. 
d laetevirens Standl. & Steyerm., Field. Mus 
, Bot. Ser. 23(3): 114. 1944. J. A. an 
49189; July 1942; Guatemala, Dept. Huehuetenango, 
Cerro Chiblac; 1,200-2,000 m; F (holo). 
Ocotea vara pasensis Standl. & Steyerm., Field. s. 
., Bot. Ser. 23(3): 114. 1944. P. C. Standley 
1939; Guatemala, Dept. Alta Verapaz, 
1,480 m; F a 


Bot. Sar, 2: 


ist., Bot. Ser. 23(2): 116. 1944. / 
36207; 20 Feb. 1940; Guatemala, Dept. San Marcos, 
Volcán Tacaná; 2,500-3,000 m; l 
Phoebe iugi Mis co Standl. A Steyerm., Fi eld. Mus 
t. Hist., Bot. Ser. 23(3): 1 "1944. P. C. NON 
70009; Mar. 1939; RDUM. Dept. Alta Verapaz 


SE of Hsu 1,500 m 
Allen, 
Phoebe on Standl. & Steyerm., Field. Mus. Nat. 


; F (holo) = Ocotea standleyi 


Hist., Bot. Ser. 23(3): 117. 1944. A. F. Skutch 1367; 
Oct. 1934; Guatemala, Dept. Quezaltenango, Colomba; 
575m LE (h olo). 


Phoebe savannarum Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(3): 118. 1944. J. A. Steyermark 
45712; Apr. 1942; Guatemala, Dept. Alta Verapaz, 
Cerro Chinajá at Sachaj and Sacacao; 150-180 m; F 
(holo). 


Volume 76, Number 3 
1989 


Taylor 685 
Plants Described by 
Julian A. Steyermark 


LECYTHIDACEAE 


L "s karuaiensis Steyerm., F E add ur 422. 

2. J. A. Steyermark 6076 944; Ven- 

Lb Bolivar, base of S on iE l ae m; F 

(holo), oe aa E alutacea (A. C. Smith) 
Mori, 


LEGUMINOSAE = Fabaceae 
LENNOACEAE 
Lennoa madreporoides Llave & Lex subsp. australis 
Steyerm., Acta Bot. Venez. 3: 230. 1968. J. A. Stey 
ermark & A. Braun 94518; 23 Jan. p Von 
Falcón, Peninsula de Paraguaná M 
(1897510), NY, US, VEN pa 


Lexarza var. madreporoides, 19 


oro; 
eL des 


LENTIBULARIACEAE 


Genlisea esmeraldae Steyerm., Fieldiana Bot. E k 534. 
1953. J. A. Steyermark 57835a; 22 Aug. 1944; Ven- 
ela, T.F. Amazonas; bus base of Cerro D 200 

m, F pes — G. pygmaea St. Hiliare, 1967 

isea nigrocaulis met , Bull. Torcy Bot. Club 
75(6): 657. 1948. J. A. Ste mark 59644; 1 Nov. 

1944; Venezuela, Bolívar, Ptari-tepui; 1,600 m; F (holo) 
= ygmaea St. Hiliare, 1967. 

anlisen icd a Steyerm., ae Bot. 28(3): 
534. 1093 . J. A. Steyermark 58445; 8 Sep. 1944; 

T.F. Amazonas, below meok of Rio Sa- 

nariapo, a alo ong Orinoco River; 100 m; F (holo). 

Pinguicula clivorum Standl. & Steyerm., Field. Mus. 
Nat. Hist., Bot. Ser. 23(4): 179. 1944. J. A. Steyer- 
mark 50061; 4 Aug. 1942; Guatemala, Dept. Hue- 
huetenango, above San Se du Sierra de los Cu- 
eme 2,400 m; F (holo) = P. lilacina Schlecht 
& Cham 4. 

l tricularia aig zonasana Steyerm., Fieldiana Bot. 28(3): 
535. 1953. J. A. Steyermark 58505; 8 Sep. 1944; 
iota T. F. Amazonas, vicinity " Sanariapo 100 
m; F (holo) = U. hydrocarpa Vahl, 


Utricularia arenicola Tutin var. b it Steyerm. 


Fieldiana Bot. 28(3): 535. 1953. J. A. Steyermark 
59335; 26 Oct. 1944; Venezuela, Bolivar, Gran Sa- 
bana; He m; F (holo) = U. nana St. Hiliare & 
Girard, 


Mob us Em Steyerm., Fieldiana Bot. 28(3): 
1953 A. Steyermark 59351; 26 Oct. 1944; 
Venezuela, Bolivar: Gran Sabana; 1.220 m; F (holo) 
— U. adpressa Salzmann, 1967. 
Utricularia Dd Steyerm., i Bot. 28(3): 
36. 1953. , teyermark 5 4 Nov. 1944; 
Venezuela, Bolivar, Ptari-tepui; ^ x m; F (holo). 
— arce Steyerm., Fieldiana Bot. 28(3): 
3. J. A. Steyermark 58520; 11 Sep. 1944; 
diei Y F. Amazonas, Puerto Ayacucho; 200 m; 
F (holo) = U. cucullata St. Hiliare & Girard, 1967. 
Utricularia baldwinii Steyerm., Bull. Torrey Bot. Club 
79(4): 310. 1952. J. T. Baldwin, E rit 7:3 Lu. 


Yel Bot. 28(3): 


U tric ularia boliv 'arana Steyerm., 


A. oboe md 3 Oct. 1944; 
Venezuela, Bolivät, So aima, Gran Sabana; 
l, sai m; F (holo) = U. la St. Hiliare & 


ard, 
beam campbelliana Oliver var. minor Steyerm., 


Fieldiana Bot. 28(3): 539. 1953. J. A. Steyermark 
59753; 2 Nov. 1944; Venezuela, Bolivar, Ptari-tepui; 
2,000-2,200 m; F (holo) = U. 
var. campbelliana, 1967. 

Utricularia cearana Steyerm., Bull. Torrey Bot. Club 
79(4): 310. 1952. H. C. Cutler 8365; 10 June 1952; 
Brazil, Ceara, 4 km N i Soure (Caucaia); 

Utricularia acce Stey 
1953. J. A aa 59404; 26 Oct. 1944; Ven- 

bana; 1,220 m; F (holo) = U. 


campbelliana Oliv. 


ezuela, Bolivar; er Sa 
fimbriata Kunth, Ts 

Utricularia a Steyerm. f. diminutiva Steyerm., 
m Bot. 28(3): 540. 1953. J. A. Steyermark 

; 3 Oct. 1944; Venezuela, Bolivar, S of Mt 
ata Gran Sabana; 1,065 m; F (holo). 

eee a wu Steyerm., Fieldiana Bot. 28(3): 

3. J. A. Steyermark 60119; 14 Nov. 1944; 
Visi Bolivar, Sororopan-tepui; 2,255 m; F (holo) 
— U. calycifida Benjamin, 1967. 
Utricularia canlen Steyerm., Bull. Torrey Bot. Club 79(4): 
1952. H. C. Cutler 8363; 10 June 1945; Brazil, 
Ceara, 4 ion N of Soure (Caucaia); F (holo) = U. viscosa 
uce ex Oliver, 1967. 
lbricularia dawsonii Steyerm., Los Angeles Co. Mus. 
Sci. 21: 20. 1958. E. Y. Dawson 14770; 1 
May 1956; Brazil, Golás, Chapada dos Veadeiros, 10 
km from Veadeiros on Cavalcante road; F (holo), LAM, 
R = U. amethystina St. Wem are & Girard, 1967. 

Utricularia heterochroma Ste erm., Veram Bot. 28(3): 
544. 1953 Vbi d 59930; 7 Nov. 1944; 
Venezuela, Bolivar: Ptari-tepui; ION 2,450 m; F 
holo) 

Utricularia humboldtii Schomb. f. cuneata. Steyerm., 
Fieldiana B 
58229; 2 Sep. 1944; dran T.F. 
Duida; 1,025-1,200 m; F (holo) = 
H. Schomb. var. —€—Ó . Mudo 

d kaieteurensis Stey Bull. Torrey Bot. 

75(6): 658. 1948. B. Maguire de D. B. Fanshawe 

av 4 May 1944; Guyana, Kaieteur Plateau; F 

TE NY = U. amethystina St. Hiliare & Girard, 


Am 
U. humboldtii R. 


Utricularia maguirei Steyerm., - Torrey Bot. Club 
i t D. anshawe 
ana, ee ret F 
holo), NY = U. calycifida Benjamin, 1967 
Utricularia www Steyerm., Fieldiana Bot. 28(3): 
3. J. A. Steyermark 58446; 8 Sep. 1944; 
Venezuela, T.F. Amazonas, below meti of Rio Sa- 
nariapo, along Orinoco River; 100 m; F (holo). 
Utricularia d Steyerm. f. parva Steyerm., Field- 
28(3): 546. 1953. J. A. Steyermark 59333; 
26 Oct. ge Venezuela, Bolivar, Gran Sabana; 1,220 
m; F (holo) = U. oliveriana Steyerm. var. oliveriana, 
1967 


-~J 


Utricularia > Steyerm., Fieldiana Bot. 28(3): 
548. A. Steyermark 58488; 8 Sep. 1944; 
es H T. F. Amazonas, Sanariapo; 100 m,F (holo) 

= U. fimbriata Kunth, 1967. 
Utricularia subpeltata Steyerm., Fieldiana Bot. 28(3): 
. J. A. Steyermark 60477; 23 Nov. 1944; 
Vénisauels. Bolivar, between Santa Teresita de Kava- 
nayen and ca. 8 km NW of Kavanayen; 1,220 m; F 
(holo) = U. pubescens Smith in Rees, 1967. 

Utricularia tepuiana Steyerm., Fieldiana Bot. 28(3): 550. 
1953. J. A. Steyermark 59517; 29 Oct. 1944; Ven- 
ezuela, Bolivar. Ptari-tepui; 1,810 m; F ias )= U. 
amethystina St. Hiliare & Girard, 1967 


686 


Annals of the 
Missouri Botanical Garden 


Utricularia turumiquirensis Steyerm., Fieldiana Bot. 
. J. A. Steyermark 62752 


Diablo and Cerro de Neveri, along headwaters o 
de Amana; 2,000 m; F (holo) = U. amethystina St. 
Hiliare & Girard, 1967. 
Utricularia venezuelana Steyerm., Fieldiana Bot. 28(3): 
. A. Steyermark 59625a; 1 Nov. 1944; 
Venezuela, Bo lívar, Ptari-tepui; 1,600 m; F (holo) = 
U. pubescens Smith in Rees, 1967. 
Utricularia williamsii Steyerm., Ceiba 1: 126. 1950. 
Ll. Williams 10639; 17 Oct. 1946; Honduras, Dept. 


Morazán, near Hoya Grande, drainage of Rio Yeguare; 


1,500 m; F (holo) = U. amethystina St. Hiliare & 
Girard, 1967. 
LILIACEAE 
vni scilloides e Cory f. digan vp ale 
Rhodor 40: 178. 1938. J. A. Steyermark 22499; 23 
pr. S.A., M St. Louis Co., ip Hills, 
MO A 11746 11), 
ee RR (Raf.) Cory f. variegata Steyerm., 
Rhodor : 116. 1949. J. A. Steyermark 28324; 25 


V n U.S.A., Missouri, Christian Co., Pedelo 

Creek, 4.5. mi. NE of Sparta; F (holo). 

Excremis coarctata (Ruiz & Pavón) Baker f. alba Stey- 
erm., Fieldiana Bot. 28(1): 153. 1951. J. A. Steyer- 
a ry 16-17 June 1945; Venezuela, Male 
a s Chorros, Pico de Naiguata; 2,200- 2,765 
m; F (hol 0). 

Vietneria paniculata Steyerm., doses Bot. 28(1): 153. 
1951. Steyermark 59362; 26 Oct. 1944; Ven- 
ezuela, Bolivar, Gran Sa Sed L 220 m; F (holo). 

Smilacina crassipes Standl. & Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 23(5): 214. 1947. J. A. Steyermark 
51003; 20 Aug. 1942; Guatemala, Dept. Huehuete- 
nango, Cerro Pueblo Viejo; 1,900 m; F (holo) = Maian- 
themum gigas var. crassipes (Standl. & Steyerm.) 
LaFrankie, 1986. 

m duidae Steyerm., Fieldiana Bot. 28(1): 156. 

EY A Steyermark 58716; 1 Sep. 1944; Ven- 
is T.F. Amazonas, Cerro Duida, Broddbiuis Hills; 
1,700-1,980 m; F (holo). 

Tofieldin sessiliflora Hook. var. robustior Steyerm., 
Fieldiana Bot. 28(1): 157. 1951. J Steyermark 

1944; Venezuela, Lara, Las Sabanetas, 
above Los Aposentos, W of Humocaro Bajo; 2,530 m; 
F (holo). 

Trillium ozarkanum Palmer & Steyerm., Ann. T 
Bot. Gard. 22: 504. 1935. J. A. Steyermark 1862 


of Cassville; 

ozarkanum (Palmer & Steyerm.) Stey , 1960. 
Trillium recurvatum Beck f. foliosum HM Rhodora 

61: 124. 1959. W. Adams s.n.; 6 Apr 


. pet etaloideum Stayer: is da 
Adams s.n.; © Apr. 
U.S.A., Missouri, Cape Cirardeau Co., Houck ta 
B2 Road; SE Missouri State Univ. (holo). 
Trillium recurvatum Beck f. shayi Palmer & Steyerm., 
Ann. Missouri Bot. Gard. 22: 504, 1935, nom. nov. 


TUN recurvatum Bec 
: 124. 1959 


LIMNOCHARITACEAE 


“Hydrocleis” standleyi Steyerm., Field. Mus. Nat. Hist., 
1944. J. A. Steyermark 30416; 


Bot. Ser. 23(2): 31. 


Oct. 1939; Guatemala, Dept. Jutiapa, between Agua 
Blanca and Amatillo; 950 m; F (holo). Note: Hydro- 
cleys is the correct spelling 

LINACEAE 


Ochthocosmus see under Ixonanthaceae 


LISSOCARPACEAE 


, Ann. Missouri Bot. Gard. 


. Holst 


Lissocarpa stenocarpa Steyerm. 
74: 104. 1987. J. 4. Steyermark & B. 
i 28 Feb.-1 Mar. 1 
zonas, Dept. Atabapo, Cerro Mad 1,140 m; MO 
(holo — 3327360), VEN. 


LOASACEAE 


"Cajaphora" larensis Steyerm., Fieldiana Bot. 28(2): 
412. 1952. J. A. Steyermark 55528; 12 Feb. 1944; 
Venezuela, Lara, between Buenos Aires to Canyon of 
El Callado, above Humocaro Alto; 2,285-2,740 m; F 
(holo). Note: Caiophora is the correct spelling. 

“Cajaphora” venez uelensis Steyerm., Fieldiana Bot. 

. J. A. Steyermark 56594; 18 May 
1944; Venezuela, Mérida, La Isla, above Tabay; 2,285- 
2,745 m ). 

Fucnide guatemalensis $ Standl. & Steyerm., Field. Mus. 

at. Hist., Bot. Ser. 23(4): 178. 1944. J. A. Steyer- 
mark 50818; 17 du 1942; Guatemala, Dept. Hue 
huetenango, NW of Cuilco, Cerro Chiquihui, bote 
Carrizal; 2,000 m; F (holo). 

Schismocarpus matudai Steyerm., Bull. Torrey Bot. Club 
80(2): 138. 1953. E. Matuda 5330; Mexico, Chiapas, 
along Río Cintalapa, between Jilquero and Pinuela, Mun. 
Escuintla; 500 m; Matuda Herbarium (holo). 


LOGANIACEAE 


E eds cd a mea & Eu ova Field. 
fus Pope Bot. 23(2): 44. P. C 

anilla 5479; 17 Feb. 1939; a Dept. 
ui Ln El ree near Zunil; F (holo) = Bud- 
dleja RPE des A. Gray, 1969. Note: Buddleja is 
the correct spelling 

"Buddleia" au Sa Standl. & Steyerm., Field. Mus 
Nat. Hist., Bot. Ser. 23(2): 72. 1944. J. A. B ud 
37276; 8 Mar. 1940; Guatemala, Dept. San Marcos, 
Loma de la Paloma, above Finca El Porvenir, Volcán 
Tajamulco; 1, pria 700 m; F (holo) = Buddleja cor- 
data H.B.K., 

“Buddleia” "ih Standl. z Uy ., Field. Mus. 
Nat. Hist., Bot. Ser. 23(5): 223. 1947 A. Steyer- 
mark 43628; 7 Feb. 1942; ae ne De ept. El Pro- 
greso, Sierra i: las Minas, Joya Pacayal; 2,500-3,000 
m; F (holo 

" Buddleia" lo ade Standl. & Steyerm., Field. Mus 
Nat. Hist., Bot. Ser. 23(2): 71. 1944. J. A. Pecan 


go, in Rio Samalá, below Zunil; 2,500 m; F (holo) = 
Buddleja crotonoides A. Gray, 1969. 
i dl. & Steyerm., Field. Mus. Nat 


48692; 15 July 1942; Guatemala, Dept. H 

go, between Yulhuitz and Maxbal, Sierra de los 

chumatanes; 1,400-1,500 m; F (holo). 
LORANTHACEAE 


Phthirusa hippocrateoides Steyerm. & Maguire, Mem. 
New York Bot. Gard. 17(1): 442. 1967. J. A. Stey- 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


687 


ermark 75059; 17 Apr. 1953; Venezuela, Bolivar, 
Abacapa-tepui, Chimanta Massif; 1,312-1,500 m; NY 
(holo) = Struthanthus syringifolius Mart., 1982. 
Phthirusa pM Steyerm., neers Bot. 28(1): 225. 
951. J. A. Steyermark 59549; 30 Oct. 1944; Ven- 
ezuela, Bolivar, Ptari-tepui; 2, pa 2. 285 2 F (holo) 
= Struthanthus Bo era Mart., 

Phthirusa n nsis Steyerm., a M 28(1): 
224. 1951. J. Fi Steyermark 58943; 30 Sep. 1944; 
Venezuela, Bolivar, Mt. Roraima; 2,040-2,255 m; F 
(holo). 

Psittacanthus costanensis Steyerm., Acta Bot. Venez. 3: 
189. 1968. J. A. & C. Steyermark 95215; 28 dun 
1966; Venezuela, Carabobo, Rio San Gian, S Borb 
rata, La Toma; 750-1,000 m; NY, US, VEN (holo). 

Psittacanthus mayanus Standl. & Steyerm., Field. Mus. 

ist., Bot. Ser. 23(2): 41. 1944. P. Gentle 116; 

1933; Belize, Santa Rita; F (holo). 

Struthanthus brachybotrys dr & Steyerm., Field. 
Mus. Nat. Hist., Bot. Ser. 23(2): 42. 1944. P. - 
Siandley 82430; Jan. 1941; Guatemala, Dept. 
huetenango, Rio Pucal ca. 14 km S of ie ee: 
1,790 m; F (holo). 

Struthanthus chimantensis Steyerm. & Maguire, Mem. 
New York Bot. Gard. 17(1): 443. 1967. J. A. Stey- 
ermark 74816; 5 tale rd Mera Bolivar, Aba- 

pa-tepui, Chimantá ssif; 0-1,100 m; 
(201171), VEN um a 2 d (Gleason) Steyerm. 
Maguire, 1967. 

Struthanthus johnstonii Standl. & Steyerm., Field. Mus 
Nat. Hist., Bot. Ser. 23(2): 43. 1944. J. R. Jolason 
1887; 2 [on 1941; Cuatemala. Dept. Huehuetenango, 
W of Aguacatán; F (holo). 

Struthanthus mucronatus Steyerm., Bol. Soc. Venez. Ci. 
Nat. 26: 418. 1966. J. A. Steyermark et al. 92336; 
29-30 Dec. 1963; Venezuela, Bolivar, Cerro Venamo; 
950-1,150 m; VEN (holo) = S. icta ec 


a & Maguire var. mucronatu 


a 
E 

Uu 

e 


Srushanthus EA Standl. & Steyerm., Field. Mus. 
, Bot. . 23(4): 154. 1944. J. A. ag d 
mark 50672; rae 1942; Bep in nid o 
huetenango, above San Ildefonso Ixtahuacàn; 1, 600- 
1,700 m; F (ho ai LT hinge (Standl. & 
Steyerm.) Kuijt, 
ie cour "re ns: Standl. & Steyerm., Field. Mus 
Nat , Bot. Ser. 23(2): 43. 1944. H. von Turek: 
heim IL 11240: June 1906; Guatemala, Dept. Alta Vera- 
paz, Cobán; 1,350 m; F (holo). 
d xpo ie Vii ae Standl. & Er val , Field. Mus 
2s nu 23(4): 155 4. J. A. Steyer- 
den 5153 ; 29 e 1942; od Dept. Hue 
i. Ciénaga de Lagartero, below Miramar: 
300 m; F (holo). 


LYTHRACEAE 


a MAL Mee H.B.K. var. tachirensis Steyerm 
Fie a Bot. 28(2): 420. 1952. J. A Steyermark 
57162: 12 July 1944; Venezuela, Táchira, between 
Villapaez and Betania along Rio Táchira; 2,130-2,285 
m; F (holo). 


MAGNOLIACEAE 


Magnolia chimantensis Steyerm. & Maguire, Mem. New 


ork Bot. Gard. 17(1): 443. 1967. J. A. Steyermark 


75840; 20 June 1988; Herve Bolivar, Apacara- 

m; VEN (holo). 

Magnolia P des Sew - 
233. 1951. J. A. Steyerm dry 59547 ; 30 Oct. 1944; 
Venezuela, Bolivar, alie 2. 100- 2,285 m; F 
(holo). 

o erae roraimae Steyerm., Fieldiana Bot. 28(1): 234. 

l. J. A. Steyermark 59000; 30 Sep. 1944; Ven- 

inn Bolivar r, Mt. Roraima; 2, 155.3 m; F (holo). 


MALPIGHIACEAE 


Banisteria NO Steyerm., ioe Bot. 28(2): 
280. 1952. eyermark 61914; 7 Apr. 1945; 
Venezuela, m agas, "Cerro de la ed de Dona Anita; 
1,100-1,200 m; c: lo). 

cipes cretacea (Gleason) Steyerm. var. com- 
posita Ste - Fieldiana Bot. 28(2): 281. 1952. J. 
A. a 58191; 1 Sep. 1944; Venezuela, T.F. 
Amazonas, Cerro Duida; 1,700- i E m; F (holo) = 

. hypoleuca (Benth.) d 

Blepharandra ptariana Steyerr "Ficldiana Bot. 28(2): 
282. 1952. J. A. SUM 60310 ; 18 Nov. did 
Venezuela, Bolivar, Ptari- n A 2201 m; F (holo) = 
hypoleuca (Benth.) Griseb., 

Burdachia duckei Steyerm., ae Bot. 28(2): 283. 
1952. A. Ducke 522; 14 July 1937; Brazil, Manaus, 
Igarape de Cachoeira Grande; F (holo), MO (1157729). 

Burdachia williamsii Steyerm., Fieldiana Bot. 28(2): 
285. 1952. Ll. Williams 14321; 1942; Venezuela, T. 
F. Amazonas, Rio Guainia, Maroa; 127 m; F (holo). 

Byrsonima adr Steyerm. pe Bot. 28(2): 

A. Steyermark 60272; 15-17 Nov. 
1944; silii Bolivar, between y Sororopan: tepui and 
Ptari-tepui; 1,615 m; F (holo) — B. chalcophylla var. 
carraoana (Steyerm.) Anders., 1981. 

Brysonima carraoana Steyerm Pree Bot. 28(2): 
287. 1952. J. A. Steyermark 60883; 7 Dec. 1944; 
Venezuela, Tokens Carrao-tepui; 2, ve 500 m; F 
(holo) = B. chalcophylla var. carraoana (Steyerm.) 
Anders., 1981. 

Byrsonima leutz pmi Steyerm., Fieldiana Bot. 28(2): 
288. 1952, no 

md villiamsii ra mM Piden Bot. 28(2): 288. 

lL. Williams 14154; eb. 1942; Venezuela, 
X: y oes Yavita; s m; F (holo). 

“Heteropterys” quetepensis Steyerm., Fieldiana Bot. 
28(2): 292. 1952. J. A. Steyermark 62880; 23 May 
1945; Venezuela, Sucre, E of Cumana, near Quetepe, 
between km 18-20; 30 m; iow Note: Heteropteris 
is the correct laonseetid) spellin 

Hiraea tepuiensis Steyerm., Fieldana Bot. 28(2): 293. 

1952. A. S. Pinkus 90; 28 1939; Moe 
Bolivar, vicinity Arabupu, Mt. Roraima; 1,500 m; F 
(holo). 

Mascagnia ua Standl. & Steyerm., Field. Mus. Nat. 

23(4): 168. 1944. J. A. Steyermark 

56; 14 . 1940; Guatemala, Dept. Izabal, Rio 

Dulce; sea level F (holo). 

“Tetrapterys” phylladenophora E & Steyerm 

pr . New York Bot. Gard. 9: 4 


. ZA. JE 


- & 
Venezuela Bolív ar, Uaipán-tepui; NY (holo) = 
is rhodoptero Oliver, 1981. Note: Tetrapteris is the 
pe spelling 
“Tetrapterys” all Steyerm., Fieldiana Bot. 28(2): 
294. 1952. J. A. Steyermark 59391; 26 Oct. 1944; 
Venezuela, Balivar. Gran Sabana; 1,220 m; F (holo). 


688 


Annals of the 
Missouri Botanical Garden 


MALVACEAE 


Abutilon hig sor P rH S Ps Mus. Nat. 
Hist., Bot. Ser. 23(4): 3 P. ead 
(4420; 15 Oct. 1940; Ed odd on 
road Pique Agua Blanca & Cumbre de e 
350-500 r 0). 

Abutilon pachecoanum Standl. & PW Field. Mus 

. Hist., Bot. Ser. 23(2): 61. 1944. P. C. ondas 
83528; Jan. 1941; Guatemala, Dept Quezaltenango, 
near Las Nubes, S of San Martin Chile Verde; 2,250 
m; F (holo). 

Abutilon prono Steyerm., Fieldiana Bot. 
28(2): 36 A. Steyermark 61519; 16 Mar. 
1945; enla Anzoátegui, between San José & 
Cucuta; 600-700 m; F (holo). 

Abutilon hag pe & Steyerm., Field. Mus. Nat. 
Hist., Bot. 23(4): 173. 1944. ad 
43439; 4 Feb, 1942; Guatemala, Dept. El Progreso, 
hills between Finca Piamonte & slopes SE, Sierra de 
las Minas; 2,400 m; F (holo). 

Abutilon turumiquirense Steyerm., dg, Bot. 28(2): 

3. 1952. J. A. Steyermark 62689; 10 May 1945; 

Venezuela, Sucre, Cerro iia 1,300-1,800 
m; F (holo 

eti guatemalense por & 2 ., Field. 

Nat. Hist., Bot. Ser. 23(4): 174. 1944. 
Standley (69. 34; 20-27 Nov. 1940; ue Dept. 
Santa Rosa, near Cuilapa; 900 m; F (holo). 

Pavonia imatacensis Steyerm., Acta Bot. Venez. 3: 181. 

A. Steyermark 88149; 17 Dec. 1960; Ven- 
nieli, Bolivar: Sierra Imataca, Rio Toro (Rio Grande); 
200-250 m; VEN (holo). 

Pseudabutilon inornatum Standl. " uy Field. Mus 
Nat. Hist., Bot. Ser. 23(1): 15. . P. C. Standley 
61425; Dec. 1938; Guatemala, E Guatemala, near 
Amatitlán; 1,170 m; E (holo). 


MARANTACEAE 


Econ acuminata Steyerm., yea Por 28(1): 161. 
1951. Ll. Williams 15532; 24 Ma 2; Venezuela, 
T. u pá Alto Casiquiare, C ouis 120 m; 


Calathea coccinea Standl. E Steyerm., Field. Mus. Nat 
Hist., Bot. Ser. 23(2): 39. 1944. . Steyermark 

1940; A Dept. San Marcos, 

shave Finca El Porvenir, along Rio Cabüs, Volcán 
Tajumulco; 1,300-1,500 m; F Gholo). 

Calathea duidae Steyerm., Fieldiana Bot. 28(1): 162. 
1951. . Steyermark 57888; 23 Aug. 1944; Ven- 
ezuela, T. F. Amazonas, SE base of Cerro Duida; 210 
m; F (holo). 

Calathea lasseriana Steyerm., Fieldiana Bot. 28(1): 163. 
1951. J. 4. Steyermark 57920; 23 Aug. 1944; Ven- 
ezuela, T.F. Amazonas, SE base of Cerro Duida; 215 
m; F (holo). 

Ischnosiphon lasseriana Steyerm., Fieldiana Eon 28(1): 

4. l. . Steyermark 60557; 15 Nov. 1944; 
Venezuela, Bolívar, 
nayen; F (holo). 

Monotagma duidae Steyerm., Aera d 28(1): 164. 

A. Steyermark 578 22 Aug. 1944; Ven- 
ezuela, T. F. Amazonas, SE Ls of ad Duida; 200 
m; F (holo 

d yapacadenss up rs & Bunting, Acta 
B . 8: 1973 A. Steyermark & € 
Bunting 103062; 3 May 1970; Venezuela, T.F. Am 
zonas, Cerro Yapacana; 125-400 m; VEN (holo). 


below Santa Teresita ds Kava- 


Thymocarpus Joe, Steyerm. & Sivad., Brittonia 33: 
22. 1981, Gen. nov 
Thymocarpus iade Nicolson, Steyerm. & Sivad., 
Brittonia 33: 24. 1981; B. Maguire et al. 53608; 5 
. 1962; Venezuela, Bolivar: SE end of Cerro Piton, 
Cordillera Epicara, alta Rio Cuyuni; 40 m; MO 
(2892301), NY, US, VEN (holo). 


MARCGRAVIACEAE 


Marcgravia sororopaniana Steyerm., eiae Bot. 
28(2): 378. 1952. Steyermark 60127; 14 Nov. 
1944; Venezuela, Bolivar, Sororopan- Pr 2,255 m; 


F (holo). 


Ruyschia longistylis Standl. & Steyerm., Field. Mus. 
Nat. Hist., Bot. Ser. 23(4): 175. 1944. J. A. — 
mark 49545; 26 July 1942; Guatemala, Dept. H 


ue 

huetenango, Cafetal of Finca Soledad, Seri de los 
ei tanes; 1,150 m; F (holo) = R. enervia Lun- 
dell, 

oo nS Aublet var. tomentella Steyerm., 
Price Bot. 28(2): 378. . Steyermark 
60652; 27 Nov. 1944; Venezuela, Bolan, Ptari-tepui; 
dum m; F (holo). 

Souroubea micrantha Standl. & Steyerm., Field. Mus. 

t. Hist., Bot. Ser. 23(2): 62. 1944. P. C. Standley 
72793; Apr.-May 1939, Guatemala, Dept. Izabal, near 
Puerto Barrios; sea level; F (holo) = 5. triandra Lun- 
dell, 1961. 

Souroubea puberula Standl. & Steyerm., Field. Mus. Nat 

ist., Bot. Ser. 23(4): 175. 1944. J. A. Steyermark 
46802; 2 June 1942; Guatemala, Dept. Suchitepéquez, 
Loma Grande, as Finca El Naranjo, Volcán de Santa 
Clara; 2,000 m; F (holo) = S. exauriculata Delpino, 
1961 


MELASTOMATACEAE 


C oe as Standl. & Steyerm., Field. Mus. 
at , Bot. Ser. 22(5): 361. 1940. P. C. Standley 
70317; 2 Apr. 1939; Guatemala, Dept. Alta Verapaz, 
Cocolá, NE of Carchá; 1,200 m; F (holo) = C. hirtella 
Cogn. 63. 
Conostegia volcanalis Standl. & Steyerm., Field. Mus 
t. Hist., ine er. 23(3): 136. 1944. P. C. ula 
onis: ;8 939; Guatemala, Dept. Quezaltenan- 
o, Chiquihu ite: ji 410 m; F (holo). 


Dolana calciphila Standl. & Steyerm., Field. Mus. Nat 
H d h 


- Qui 

— Triolena eaiciphla (Standl. & 
Steyerm.) S . & L. Williams, 196: 

Diolena roseiflora Standl. ar eyerm., Field. Mus 3. Nat. 
Hist., Bot. Ser. 23(3): 1944. f: A. Stey beds 
41819; 24 Dec. 1941; oe si are alor 
Rio Tameja, Cerro San Gil; 50 m; first cited s specime sj 
type not designated. = a coser (Standl. & 
Steyerm.) Standl. & L. William 

Diolena stenophylla Standl. & dem. 

. Hist., Bot. Ser. 23(3): 135. 1944 Steyer- 

mark 48728; 15-16 July 1942; Guatemala, Dept. 
Huehuetenango, between Maxbal and lake to the SE; 
1,500 m; F (holo) = Triolena dd (Standl. & 
Steyerm). Standl. & L. William 

Miconia ec io Standl. & abad ., Field. Must. 
Nat. Hist., Ser. 23(4): 180. 1944. J. A. Steyer- 
mark 48831: : "is 16 July 1942; Guatemala, Dept. 
Hue huetenango, vicinity of Maxbal, Sierra de los Cu 


chumatanes; 1,500 m; F (holo). 


Field. Mus. 


Volume 76, Number 3 
1989 


Taylor 689 
Plants Described by 


Julian A. Steyermark 


Miconia wp oie Standl. & Steyerm., Field. Mus. Nat. 
p 9r e (4): 181. 1944. J. A. Steyermark 
-10 Nov. 1939; Cuatemala, Dept. Chiquimula, 

e. Pix 1,500 m; F (holo). 
Monochaetum guatemalense Standl & aim Field. 
Mus. Nat. Hist., Bot. Ser. 23(3): 136. 1944. J. 4. 
Steyermark 48512; 10 July 1942; Guatemala, Dept. 


along Quebrada Suchiate, near SE portion of Volcán 
Tacaná; 2,000-3,000 m; F (holo). 

Triolena izabalensis Standl. '& Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 23(3): 138. 1944. C. L. Wilson 175; 
7 Feb. 1939; Guatemala, Dept. Izabal, near Rio Lám 
para; F (holo). 


MELIACEAE 


c Té ug o Standl. & Steyerm., Field Mus. 

, Bot. Ser. 23(4): 167. 1944. J. A. Steyer- 

pon 46355 ; 10 May 1942; Guatemala, Dept. Alta 

Verapaz, along Route n between Chirreacté & Sem- 

ococh; 500-900 m; F (holo) = P chirriactensis 
(Standl. & Steyerm. ) Penn., 1981. 

Trichilia [o Standl. & i Field. Mus. Nat. 
Hist., Bot. Ser. 23(4): 167. 1944. J. 4. Steyermark 
45827; 21 A r. 1942; Guatemala, Dept. Petén, along 
Río Santa Isabel, between mouth of Rio Sebol and El 
Porvenir; s m F (holo) = T. breviflora S. F. Blake 
& Standl., 

“Trichilia Ee Steyerm., mo E 28(2): 278. 
. Steyermark 5997 ] Nov. 1944; 

Serenus Bolivar: Ptari-tepui; 1, re E ion m; F (holo) 
— Matayba oligantha Sandw., 1981. Note: The Tri- 
chilia name was a homonym, non A. C. Smith, 1933. 
Trichilia subsimplex Steyerm., Fieldiana Bot. 28(2): 279. 
Steyermark 58061 

Venezuela, T.F. A 
(holo), NY, US, VEN - T. pallida Sw., 1981. 


MENDONCIACEAE 


Mendoncia guatemalensis Standl. & Steyerm., Field. 
Mus. Nat. Hist., Bot. Ser. 23(5): 245. 1947. J. A. 
Steyermark 49270; 23 July 1942; Guatemala, Dept. 
Huehuetenango, between Ixcán & Rio Ixcán; 150- 
200 m; F (holo). 

Mendoncia mellobarretoana Steyerm., Field. Mus. Nat 
Hist., Bot. Ser. 17(5): 421. 1938. H. L. Mello ek 
226; 1 Dec. 1934; Brazil, Suyo, Jardim Botan 
Mun. Belo Horizonte, Minas Gerais; F (holo). 


MENISPERMACEAE 


dpi salvadorensis Steyerm., Ceiba 4: 301. 1955. 
J. i ucker 979; 23 Feb. 1942; El Salvador, Dept. 
San Miguel, Volcán San Miguel, 0.8 km SW of buildings 
of hacienda of Max Haltmeyer; F (holo). 

"s vulcania Standl. & Steyerm., Field. Mus 

Hist., Bot. Ser. 23(1): 8. 1943. P. C. Standley 
"o 8; bur 1939; Guatemala, Dept. Escuintla, near 
Barranco Hondo, above Las Lajas; 1,200 m; F (holo). 


MONIMIACEAE 


Mollinedia glabricaulis Maguire & Steyerm., Mem. New 
York Bot. Gard. 50: in press. 1989. 


Mollinedia neblinensis Maguire & Steyerm., Mem. New 
York Bot. Gard. 50: in press. 1989. V. T. Silva & U. 
Brazao 60678; Brazil, Serra da dg Rio Negro, 
Rio Cauaburi, Rio Maturacá, between and Palmito 


camps; 600-1,300 m; MO eres NY (holo). 
uu" ptariensis Steyerm., Fieldiana Bot. 28(1): 
35. 1951; J. A. Cid qe 60014; 10-11 Nov. 


1984; Venezuela Bolívar, Ptari-tepui; 1,585-1,600 


Mollinedia 1 rue Steyerm., Fieldiana Bot. 28(1): 
236. 1951. J. A. Steyermark 59004; ; 30 Sep. 1944; 
Venezuela, Bolivar, Mt. Roraima; 2 .040- 2.255 m; F 
(holo). 


Siparuna manarae Steyerm., Bol. . Venez. Ci. Nat. 
26: 453. 1966. B. J. Manara a 27 June 1964; 
Venezuela, Distr. Federal, Cerro Naiguitá, Quebrada 
Río Chiquito; VEN (holo) = $. pede | var. 


marnarae "Misa ) Steyerm., 1978 

Siparuna rugosa Steyerm., Phytologia 9(6): 342. 1964. 
M. Acosta pue 7008; 18 Jan. 1944; Ecuador, Prov. 

hincha a de Lache e Condor NT Cor- 

dillera Occidental: 2,700-3,100 m; F (holo 

Siparuna verrucosa Steyerm., Lp y 9(6): 343. i did 
M. Acosta Solis 8233; 19 June 1944; Ecuador, Pro 
Sube: Urcusique, faldas occidental del ael 


3,300 m; F (holo) 


MORACEAE 


Ficus cabusana Standl. & Steyerm., Field. Mus. Nat 
Hist., Bot. Ser. 22(4): 226. 1940. J. A. Steyermark 
7583; 12 Mar. 1940; Guatemala, Dept. San Marcos, 


Pseudolmedia simiarum Standl. & Steyerm., Field. Mus. 
Nat. Hist., Bot. Ser. 23(4): 154. 1944. J. A. Steyer- 
mark 48741; 15-16 July 1942; Guatemala, Dept. 
Huehuetenango, Maxbal; 1,500 m; F (holo). 


MYRICACEAE 


Myrica rotundata Steyerm. & Maguire, Mem. New York 
Bot. Gard. 17(1): 441. 1967. J. A. Steyermark 75804; 
20 June 1953; Venezuela, Bolivar, Apacara-tepui, Chi- 
manta Massif; 2,000-2,150 m; VEN (holo). 


MYRSINACEAE 


Ardisia ipn e oo ay Field. Mus. Nat. Hist., 
219. 194 A. Steyermark 41923; 
25 Dec. iod Gane. Dept. Izabal, Cerro San Gil; 
300-900 m; F (holo). 


Ardisia ca rlsonae Steyerm., 


Ceiba 4: 301. 1955. M. C. 


Om;F duo) = Deis Nanthes 
carlsonae (Steyerm. ) Lundell 983. 

Ardisia meridensis Steyerm., iu Bot. 28(3): 454. 
1953. J. A. Steyermark 56325; 6 May 1944; Ven- 

Mérida, between Mucuchachi e Canagua; 


Hist., Bot. Ser. 23(5): 2 
48558; 14 July 1942; dE Dept. Huehuetenan- 
go, Cerro Huiz, Cerro de los Cuchumatanes; 1,500 
2, 600 m; F (holo) = Gentlea vatteri (Standl. & Stey- 
erm.) Lundell, 1964 

Conomorpha apiculata Steyerm., Fieldiana Bot. 28(3): 
456. 1953. J. A. Steyermark 57986; 25-26 Aug. 


690 


Annals of the 
Missouri Botanical Garden 


1,095 m 
Conom morpha densipapillata Steyerm., Fieldiana Bot. 
28(3): 457. 1953. J. A. Steyermark 56539; 14 May 
1944; Venezuela, Merida, Páramo de Los Colorados, 
between El Molino & San Isidro Alto; 2,745-2,955 
m; F (holo). 
erue de depressa Steyerm., Fieldiana Bot. 28(3): 
458. 1953. J. A. Steyermark 60898; 7 Dec. 1944; 
Venezuela, Bolivar Carrao-tepui; 2,470-2,500 m; F 
(holo). 


1944; Venezuela, T.F. Amazonas, Cerro Duida; 305- 
; F (holo). 


Conomorpha frigidicola Steyerm., Fieldiana Bot. 28(3): 
5 A. Steyermark 56516; 14 May 1944; 
enezuela, Mérida, between El Molino & ridge above 

San Isidro Alto; 2,430-2,895 m; F end — Cybian- 
thus frigidicola (Steyerm.) Agost., 

Conomorpha hieronymoides roig Fieldiana Bot. 


rezuela, Lar netas; 2,530 m; F 
(holo ra - Cybia anthus laurifolius (Mez) Agost., 1972. 
, Fieldiana Bot. 


Venezuela, Mérida, above Jaji, Los Quebraditos; 2,590 
m; F (holo 
Conomorpha larensis Steyerm., Fieldiana Bot. 28(3): 
A. Steyermark 55392; 8 Feb. 1944; 
Venezuela, Lara, above Humocaro Baja, S of Las Sa- 
banetas; 2,430-2,745 m; F (holo). 

Conomorpha lepidota Gleason f. acutata Steyerm., Field- 
iana Bot. 28(3): 465. 1953. J. A. once 58265; 
2 Sep. 1944; Venezuela, T.F. Amazonas, Cerro Duida; 
1,025-1,200 iu F P c Cybianthus lepidotus 
(Gleason) Agost., 

Conomorpha job d Fieldiana Bot. 28(3): 465. 
1953. Steyermark 59136; 2 Oct. 1944; Ven- 
ezuela, Bolivar, S of Mt. Roraima, Gran Sabana; 1,065 
m; 

mala remi Steyerm., Fieldiana Bot. 28(3): 
466. 1953. J. A. Steyermark 57456; 18 July 1944; 
Venezuela, Táchira, Alto de Lirio; 1,890-2,285 m; F 
(holo). 


Conomorpha ptariensis Steyerm., A Bot. 28(3): 

66. 19 A. Steyermark 59632; | Nov. 1944; 

Venezuela, Bolivar Ptari-tepui; 1, cud m; F (holo), MO 
(1621331). 


C — o roraimae Steyerm., Fieldiana Bot. 28(3): 


3 
1944; Venezuela, Táchira Quebra a de Palmar at foot 
of Páramo de Tamá; 2,500 m; F (holo). 

Conomorpha sulcata ie Fieldiana Bot. 28(3): 473 

3. J. A. Steyermark 57435; 17 July 1944; Ven- 
ezuela, Pachi above Betania below Páramo de Tamá 
2,530 m; F (holo). 

Conomorpha tamana Steyerm., ery Bot. 28(3): 475. 
1953. J. A. Steyermark 5735 ; 15 July 1944; Ven- 
ezuela, Táchira, base of Él de Tamá: 2,500- 
2,895 m; F (holo). 

M ptariensis Steyerm., ge. Bot. 28(3): 

. 1953. J. A. Steyermark 59497; 29 Oct. 1944; 
e Bolivar, dei Wm l, 810 m ; F (holo). 
, Fieldiana Bot. 28(3): 477. 


ezuela, Bolivar, Mt. Roraima; 2, 620- 2, 740 pic E (bolo), 


Eugenia Ts dnd Steyerm., Fieldiana Bot. 
nom. nov. 


Eug 
Nat. Hist., 


Rapanea reticulata Steyerm., Fieldiana Bot. 28(3): 477. 
1953. J. A. Steyermark 58983; 30 Sep. 1944; Ven- 
ezuela, Bolivar, Mt. Roraima; 2, 040- 2,255 m; F (holo). 

Stylogyne turumiquirensis Steyerm., Fieldiana Bot. 28(3): 

A. Steyermark 62539; 5 May 1945; 
Venezuela, Sutra: Cerro Turumiquire; 2,100-2,200 
m; F (holo). 


MYRTACEAE 


Aulomyrcia amshoffiana Steyerm., s Bot. 28(4): 
1004. 1957. J. A. Steyermark 60354; 20-21 Nov. 
1944; Venezuela, Bolivar, Quebrada O- im ma; 1,065- 
1,220 m; F (holo) = Myrcia albldotomantosu (Am- 
shoff) McVaugh, 1969. 

Aulomyrcia bolivariensis Steyerm., Fieldiana Bot. 28(4): 
1004. 1957. J. A. Steyermark 59534; 30 Oct. 1944; 
Venezuela, Bolivar, Ptari-tepui; 1,800-2,100 m; F (holo) 
— Myrcia bolivarensis (Steyerm.) McVaugh, 1969. 

Aulomyrcia karuaiensis Steyerm., Fieldiana Bot. 28(4): 
1005. 1957. J. A. Steyermark 60702; 28 Nov. 1944; 
Venezuela, Bolivar, Ptari-tepui; 1,200 m; F (holo) — 
Marlierea karuaiensis (Steyerm.) McVaugh, 1958. 

d parvifolia Steyerm., Fieldiana Bot. 28(4): 

006. 1957. J. A. Steyermark 60285; 15-17 Nov. 
1944 Venezuela, Bolivar, between Ptari-te 5 
ui; 1,615 m; F (holo) = Myrcia ondal 


a vigas Steyerm., Fieldiana Bot. 28(4): 
1006. 7. J. A. Steyermark 59724; 1 Nov. 1944; 
MEAN A Bolivar, Ptari-tepui; 1,700-1,800 m; F (holo) 
= Myrcia ptariensis (Steyerm.) McVaugh, 1969. 

dica salticola Steyerm., Fieldiana Bot. 28(4): 
1007. 1957. J. A. Steyermark 60213; 15 Nov. 1944; 
Venezuela, Bolívar, Salto de Iwaracaru-meru, W end 
of Sororopan-tepui; 1,615 m; F (holo) = Myrcia sal- 
ticola (Steyerm. ) McVaugh, 1969. 

d nes Steyerm., Fieldiana Bot. 28(4): 1008. 

1957. J. A. Steyermark 62375; 28 Apr. 1945; Ven- 
ezuela, Suum. Cocollar Valley; 820 m; F (holo). 

Calycolpus calophyllus (H.B.K.) Berg var. di e dg 
Steyerm., Ann. Missouri Bot. Gard. 71 J. 
A. Steyermark & F. Delascio 129391; 5 eni 198%; 
Venezuela, T.F. Amazonas, Dept. Atabapo, Salto Ma- 
tushi, Río Cununcunuma; VEN (holo). 

erem ce Steyerm., Fieldiana Bot. 28(4): 
1009. . S. Pinkus 92; 30 Dec. 1938; Ven- 
ezuela, Bolivar, Distr. Mt. Roraima, near Arabupu; 
1,300 m 

Calypiranthes aquilarii ed & ue dn Field. Mus. 
Nat. Hist., Bot. Ser. 23(3): 128. 1944. J. del 

: antenna, i Quiche; F (holo) = 
Wonders Standl., 1963 
Calyptranthes macrantha Standl. & Steyerm., jus 


us. Nat. Hist., Bot. Ser. 23(3): 127. 1944. P. 
Standley 90532; 1 Apr. 1941; Guatemala, Dept. A is 
'erapaz, Rio Frio, ca. 8 km below Tactic; 1,400 m; 
F (holo). 


Calyptranthes meridensis Steyerm., Fieldiana Bot. 28(4): 
1009. 1957. J. A. Steyermark 56589; 18 May 1944; 


Venezuela, Mérida, above Tabay; 2,285-2,745 m; F 
(holo). 


28(4): 
m 
nia cacuminum 


Sta & Steyerm., Field. Mus 
Bot. Ser. DE 357. 1940. J. A. ien 


Volume 76, Number 3 
1989 


Taylor 691 
Plants Described by 
Julian A. Steyermark 


mark 30903; 31 Oct. 1939; Guatemala, Dept. Chi- 
ntaña Volcáncitos), be- 
E of Concepción 


, o). 

Eugenia caurensis Steyerm., Fieldiana Bot. 28(4): 1010. 
1957. Ll. Williams 11505; 12 Mar. 1939; Venezuela, 
Bolivar, Medio Caura, en la playa da Ce de Para; 
170 m; F (holo) = E. biflora (L.) DC., 

Eugenia cervina Standl. & Steyerm., Field "Mus 
Hist., Ser. 23(3): 128. 1943. J. A. Pini 
44691; : Mar. 1942; Guatemala, Dept. Alta Verapaz, 
near Rio Icvolay, near Hacienda ica 5 mi. NW 
of Cubilquitz; 250-350 m; F (ho 

Eugenia chiquimulana Standl. & n om Field. Mus. 
Nat. Hist., Bot. Ser. 22(5): 358. 1940. J. A. Steyer- 
m 31453; 8 Nov. 1939; Guatemala, Dept. Chi- 
quimula, near summit of Volcán de Quezaltepeque; 

Sm., 


ey a m; F (holo) = E. pachychlamys Donn. 


Eugenia chinajensis Standl. & Steyerm., Field. Mus 
. Hist., Bot. Ser. 23(3): 129. 1944. J. A. Steyer- 
d 45142; 20 Mar. 1942; kr ua DR Alta 
Verapaz, S onm Chinajá; 150-180 m; F (holo). 
Suns crebrifolia Steyerm., vage Bot. 28(4): 1011. 
57. J. A. Steyermark 57204; 14 July 1944; Ven- 
spiel. Táchira, 2 km above id & 7 km above 
Villapaez, in Paramito between Quebrada de Palmas & 
Quebrada de Paramito; 2,500 m; F (holo). 
i Standl. & Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 23(3): 129. 1944. J. R. Johnston 
1480; 21 Dos 1938; Guatemala, Dept. Jutiapa, Ju- 
tiapa; F (holo). 
Eugenia llewelynii Steyerm., Fieldiana Bot. 28(4): 1011. 
1957. Ll. Williams 14962; 15 Apr. 1942; Venezuela, 


y : 
Distr. Federal, hills of Bot. Gard., Caracas; 920 m 
MICH, VEN (holo). 

a meridensis Steyerm., ear ie 28(4): 1012. 

1957. J. A. Steyermark 5625 944; Ven- 
ch. Mérida, between La ^el San José & Mu- 
cutuy; 1,820-2,590 m; F (holo). 

Eugenia musarum Standl. & Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 22(5): 358. 1940. J. 4. Steyermark 
39165; 8 Apr. 1940; Guatemala, Dept Iza io 
Juyama, SE of Cheyenne, ca. 15 mi. S 
50-100 m; F (holo) = Psidium musarum (Standl. 
Steyerm.) McVaug 

us e eges Standl. & Steyerm., eu Mus. Nat. 

E 130. 1944. J. A. Steyermark 
42; Guatemala, Dept. Huehuete- 

o, Cur es Papal 1,800-3,000 m; F (holo). 

Xn ia patalensis Standl. & Steyerm., Field. Mus. Nat. 

Hist., Bot. Ser. 23(3): 130. 1944. P. C. Standley 

69568; 30 Mar. 1939; Guatemala, Dept. Baja in 
paz, region of Patal; 1, 7 holo) = E. gua 
malensis Donn. Sm., 

e ptariensis Steyerm., Filan Bot. 28(4): 1013. 

1957. J. A. Steyermark 60599; 27 Nov. 1944; Ven- 
d Bolívar, above Rio "e Ptari-tepui; 1,220- 
1,375 m; F (holo) = E. anastomosans DC., 1969 

Eugenia rheedioides Standl. & Steyerm., Field. Mus 
Nat. Hist., Bot. Ser. 23: 131. 1944. J. I. Aguilar 
1757; 1942; Guatemala, D ept. err without local- 
ity; F (holo) = E. myrtifolia Sims, 196 

Eugenia rondonensis Steyerm., Fidia Bot. 28(4): 


1013. 1957. J. A. Steyermark 58956; 30 Sep. 1944; 
kr Bolívar, Mt. Roraima; 2,040-2,255 m; F 
(holo) = Myrcianthes species ?, 1969 fide McVaugh. 


31327; 6 Nov. 1939; Guatemala, Chi uimula, Mon- 
taña Castilla, vicin Ny of Montana Cebollas, along Rio 
Santa Lucia Saso; i Fi 500 m; F (holo). 
d savannarum S . & Steyerm., Field. Mus 
, Bot. Ser. E 132. 1944. J. A. Steyer- 
ius 457 >10; 6 Apr. 1942; Guatemala, Dept. Alta 
Verapaz, base of Cerro Chinaja; 150-180 m; F (holo). 
Eugenia simiarum Standl 
Hist., Bot. Ser : . 
38274; 28 Mar. 1940; Ciistemala. Dept. Izabal, be- 
tween Bananera & La Presa, Montaña del Mico; 40- 
300 m; F (holo) = E. choapamensis Standl., 1963. 
Eugenia tachirensis Steyerm., inan Bot. 28(4): 1014. 
57. J. A. Steyermark 57170; 12 July 1944; Ven- 
, Táchira, base of ‘pies de Tama; 2,300- 
o). 


Eugenia ppg Steyerm., det Bot. 28(4): 1015. 
1957. J. A. Steyermark 57172a; 13 July 1944; Ven- 
ss int Táchira, Montana of Ris Táchira at base of 
Páramo de Tamá; 2,430 m; F (holo). 

elie tepuiensis Steyerm., ora Bot. 28(4): 1015. 

1957. J. A. Steyermark 60248; 15-17 Nov. 1944; 
Venezuela, Bolívar, m m -tepui & Sororopan- 
tepui; 1,615 m; F (ho 
ugenia an e c" Fieldiana Bot. 28(4): 
1015. 1957. J. A. Steyermark 62548; 5 May 1945; 
Venezuela, Sucre, SW Cocollar, Cerro Turumiquire; 
2,100-2,200 m; F (ho 

Gomidesia bonnetiasy ae Steyerm., Fieldiana Bot. 
28(4): 1016. 1957. J. A. 
Venezuela, Bolivar, 

; F (holo) = Myrcia bonnetiasylvestris (Steyerm.) 
diu 1984. 

E sanisidrensis Steyerm., Fieldiana Bot. 28(4): 

1018. 1957. J. A. Steyermark 56551; 14 May 1944; 
Venezuela, Mérida, above San Isidro Alto; 1,820 m; F 
(holo). 


Myrcia sororopanensis Steyerm., Fieldiana Bot. 28(4): 
1019. 1957. J. A. Steyermark 60102; 14 Nov. 1944; 
Venezuela, Bolivar: Sororopan-tepui; 2, 255 m; F (holo). 

Myrcia tepuiensis Steyerm., e bur 28(4): 1019. 
1957. J. A. Steyermark 60284; 15-17 Nov. 1944; 
Venezuela, Bolivar, m Ptari- bali, & Sororopan- 
PUR l, 615 m; F (ho lo). 

rciaria ieldiana Bot. 28(4): 1020. 
1957. Ll. Williams 11691; 3 Mar. 1939; Venezuela, 
Bolivar, Medio Caura, La Prisión; 120 m; F (holo) = 
M. dubia (H.B.K.) McVaugh, 1969. 

Myrtus myricoides H.B.K. var. Pre cab rs Stey- 
erm., Fieldiana Bot. 28(4): 1022. 1957. J. Ds Stey- 
ermark 62628; 6 May d Venezuela, ce e, Cerro 
P un i 2,200 m; F (holo) = Ugni ois det 
(H.B.K.) Berg. 

T fruticosa Steyerm., Fieldiana er a 1023. 
19 ? 1945; Ven- 


Y) 
St 
> 
c 
x 
3 
& 
m 
m 
a 
pa 
a 


holo) = S. de e 
distin 1969. (= S. dussii (Berg) | wil ined.). 


692 


Annals of the 
Missouri Botanical Garden 


NYCTAGINACEAE 


Guapira eg rosa Steyerm., Ann. Missouri Bot. Gard. 
14: 615 7. L. Marcano Berti 551; 12 Jan. 1965; 
nu [oed E of Rio Grande, ENE of El Palmar, 
near limits of T.F. Amacuro; MO (2670763, 2775683), 
NY, VEN (holo). 

Guapira ayacuc hae Steyerm., Ann. Missouri Bot. Gard. 

1987. J. A. Steyermark & O. Huber 113856; 
4 May 1977; ea T.F. Amazonas, Tobogan de 
la Selva, 35 km S of Puerto Ayacucho; 85 m; MO 
(2775745), VEN (hol 0). 

Gua ping bolivare n. Missouri Bot. Gard. 

6. 1987. R. T oe B. K. Holst 20800; 
11 May 1986; Venezuela, Bolivar, Dept. Piar, summit 
of Amaruay-tepui, S side, E half; 950-1,100 m; MO 
(holo), V 

Guapira davidsei Steyerm., Ann. Missouri Bot. Gard. 

l . G. Davidse & A. González 16382; 
28 Mar. -2 i 1979; Venezuela, T.F. Amazonas, 
Dept. Tucupita, 5-14 km ESE of Los Castillos de 
Guayana; 50-2,000 m; MO, VEN (holo). 

Guapira aee od Steyerm., Ann. Missouri pu Gard. 
74 . Marcano Berti & P. S 


š t 
io Negro, between San Carios t El Solano; MER 
(holo), MO (3423162). 

Guapira Wreg i "rti Steyerm., Ann. Missouri Bot. 

: 987. L. iate 'ano Berti 209; 26 May 
1964; E T.F. curo, E of Río Grande, 
ENE of El Palmar, near limits of Edo. Bolivar; MO 
(2701699, 2680864), VEN (holo). 

Guapira neblinensis Maguire & Steyerm., Mem. New 

50: in press. 1989. B. Maguire et 
al. 42068; 14-16 Nov. 1957; Venezuela; M 
(3291201), NY (holo). 

Guapira sancarlosiana Steyerm., Ann. Missouri Bot. 
Gard. 74: 618. 1987. H. L. Clark & P. Maquirino 
7794; 8 Jan. 1981; Venezuela, T.F. Amazonas, 4.3 
km NNE from San Carlos on Solano Road, IVIC study 
site; 119 m; MO (holo— 2991814). 

Guapira P Steyerm., Ann. Missouri Bot. Gard. 
74: 620. 1 . C. Blanco 1158; May 1971; Vene- 
zuela, T.F. cee Reserva Forestal Sipapo, left 
margin of Rio Sipapo, Bloque 1; VEN (holo). 

ea E hella Standl. & Steyerm. . Field. Mus. Nat. 

e Ser. 23(1): 5. 1943. P. C. Standley 73841; 
Oct d 40; Guatemala, Dept. Zacapa, road between 
et and Chuquimula; 500-660 m; F (holo). 

Neea amaruayensis Steyerm., Ann. Missouri Bot. G 

987. R. L. Liesner & B. K. Holst 20514. 

1986; Venezuela, Bolivar, Amaruay- ae W 

— 3395522), VE 
Missouri Bot. | d. 


ezuela, Bolivar, de Santa d 
800-900 m; MER (holo), MO (2406055), NY. 

Neea bracteosa Steyerm., Ann. Missouri Bot. Gard. 74: 
624. 1987. R. 
9 Apr. 1981; Venezuela, epres 
km NE of Ciudad Piar; 250- 300 | m; MO (310009 3), 
VEN (holo 

Neea brevipedunculata Steyerm., Ann. Missouri Bot. 
Gard. 74 7. J. A. Steyermark et al. 122366; 
12 May DU Venezuela, T.F. sige ar between 
Paso El Diablo and Cano de Culebra, 30 km SE 
of Puerto Ayacucho; 100 m; MO, on a 


Neea cedensis Steyerm., Ann. Missouri Bot. Gard. 74 


Neea sebastianii Steyerm 


Neea subglabrata Steyerm., Ann 


625. 1987. B. Stergios et al. 8611; 16 Apr. 1984; 
Venezuela, Bolivar, Cedeno, bosque de galeria 
del Cano Chaviripa, Carretera Caicara — El Burro; MO 
(holo — 3395524), PORT. 

Neea ius Steyerm., Ann. Missouri Bot. Gard. 74: dd 
1987. H. L. Clark 6980; n.d.; Venezuela, T.F. A 
zonas, A; 3 km NNE on Solano Road; 119m; NY (holo). 

Neea wi Steyerm., Ann. Missouri Bot. Gard. 74: 
621. . G. Davidse & A. € izülez 16558; 4-6 
Apr. 1919; gere T.F. Amacuro, Dept. Tucupita, 
ca. 13 km by road ESE of nine pe Sierra Imata; 400 

o). 


up s o da Neblina; 1,350-2,000 m; M 

(3291199), NY (holo). 

Neea guaiquinimae Steyerm., Ann. Missouri er Gard. 
74: 627. A. Steyermark et al. 117971; 9 
Apr. 1979; Mone ds Bolívar Cae 


summit, NE sector; 1,650 m; MO (2774805), VEN 
(holo). 


Neea huachamacarae Steyerm., Ann. Missouri Bot. Gard. 
74: 628. 1987. L. Liesner 18403; 7 Mar. 1985; 
Venezuela, T.F. Amazonas, Rio Cunucunuma, base of 
trail to Cerro Huachamacari; 220 m; MO (3395518), 
VEN (holo). 

Neea a Steyerm., Ann. Missouri Bot. Gard. 74 
628. 1987. O. Huber & J. Cerda 1456; 26 Jan. 1978; 
Venezuela, . Amazonas, 11 km N of Puerto Aya- 
cucho, toward El Burro; 90 m; VEN (holo 

Neea liesneri Steyerm., Ann. Missouri Bot. | 
629. 1987. 
Amazonas, iin o Negro, Cerro de la N 

m of b ius on Rio Mawarinuma; 140 m; 
(3395523), VEN (holo). 

Neea ponp e Steyerm., 

74: 630. 1987. R. L. Li ie 


n Missouri Bot. Gard. 

096; | Dec. 1977; 
Venezuela, . Amazonas km E of San Car- 
los; 120 m; MO (3020010), SIN (holo). 

Neea marahuacae Steyerm., Ann. Missouri Bot. Gard. 
74: 631. 1987. R. » y e 18428; 8-9 Mar. 1985; 
Venezuela, T.F. Amazonas, Cerro Marahuaca, 1-2 km 
N of Sima Camp; 1 100 m; MO (3395521), VEN 


0}. 

Neea neblinensis Maguire & Steyerm., Mem. New York 

a Gard. 

2072; 14- 16 Nov 
pili de la Neblina, Rio Yatua, just S of Camp 3; 700 
m; MO (3291200), ue o). 

Neea Man Steyerm., Ann. Missouri ET Gard. 74: 
631. A. Ea 106982; 18 Apr.-23 
May Ten ena T.F. Amazonas, Sierra Parima, 
Simarawochi, Rio Matacuni, 6-7 km W of Venezuelan- 
Brazilian border; 795-830 m; MO (2669239), NY, 
VEN (holo). 

Neea ey Steyerm., Ann. Missouri Bot. Gard. 74: 
632. . O. Huber 1693; 14-28 Feb. 1978; Ven- 
ita us Amazonas, third savanna at W base of 
Cerro Yapacana; 100 m; VEN (holo). 

., Ànn. Missouri Bot. Gard. 74: 

633. 1987. R. L. Liesner & H. Clark 8959; 31 Jan. 

1980; Venezuela, T.F. Amazonas, Isla Sebastian, Rio 

Casiquiare above Chapezón, between Boca and Solano; 

120 m; MO (3395520), VEN (holo 

Missouri Bot. Gard. 


74: 633. 1987. J. A. Steyermark et al. 117615; 15 


Volume 76, Number 3 
1989 


Taylor 693 
Plants Described by 


Julian A. Steyermark 


Dec. 1978; Mas qum Dade. between Betania and 

Santa Elena de Uai O km W of Santa Elena; 900 

m; MO (2774 451), NE VEN (holo). 

eea irae Steye Ann. Missouri Bot. Gard. 74: 

634. 7. J.A. Siren. & J. J. Wurdack 1104; 

23 a 1955; d Bolivar, Chimantá Massif, 

Torono tepui, su -facing forested slopes above 
valley of South Caño? 1,955-2,090 m; F, MO, NY, 

N (holo). 

Salpianthus ne Field. Mus. Nat. Hist., 
Bot. Ser. 22(3): 1 d Send 83; 11 Nov 
1937; exico, EN pd a del Consuelo, MA 


Madre del Sur, N of Rio Balsas. "Distr, Adama; F (holo). 


OCHNACEAE 


Adenanthe Maguire, Steyerm. & Wurd., — New York 
Bot. Gard. 10(4): 19. 1961, gen. no 

Adenanthe bicarpellata ei. Steyerm: & Wur 
Mem. New York Bot. Gard. 10(4 ): 


Venezuela, Bolivar, Chimantá Massif; 1,940 m; MO 
(2011680), NY (holo). 

Adenanthe bicarpellata Maguire, Steyerm. & Wurd. 
var. chimantensis Steyerm., ined.?. J. A. e al 
et al. 128637; 2-5 Feb. 1983; Veuesuela: MO (iso 


kid asisae Maguir re & Steyerm., Mem. New York 
t. Gard. 50: in press. 1989. J. Hoyos & G. Morillo 

90; 7 May 1973; Venezuela, T.F. Amazonas, Serrania 
Parú, Cerro Asisa (La Momia); l, dun m; VE 

Ouratea ayacuchae Maguire & 
York Bot. Gard. 50: E press. 1989. O. Huber ~ g 
Cerda 1447; 16 Jan. 1978; Venezuela, T.F. 
nas, Dept. Atures, Puerto Ayacucho; 90 m; VEN (holo. 

Ouratea brevicalyx Maguire & Steyerm., Mem. New 

89. R. Cowan & J. 

Amazonas, base of Cerro Parú, Serranía Parú; 100 m; 
NY (holo). 

Ouratea brevipedicellata Maguire & Steyerm., m. 
New York Bot. Gard. a 9. B. p i 
et al. 41909; 15 Oct. . Am 
zonas, Río Guainía E Maroa and Giunti Blanco. 
130 m; NY (holo). 

Ouratea croiz atii Maguire & Steyerm., Mem. New York 

50: in press. 1989. L. Croizat 624; 15 
enezuela, T.F. Amazonas, Raudal Boba- 
dilla, alto Orinoco; NY (holo 

Ouratea culmenicola Maguire & Steyerm., Mem. New 
York Bot. Gard. 50: in press. 1989. J. : Steyermark 
et al. 117393; 23-24 May 1978; Ve ezuela, Cerro 
Guaiquinima, 1 km Rio del Salto O 730- 
750 m; MO (holo), VEN. 

Ouratea deminuta Maguire & Steyerm., Mem. New York 
Bot. Gard. 50: in press. 1989. O. Huber 2504; 23 
Aug. 1978; Venezuela, T.F. Amazonas, Dept. Atabapo, 
Sabana de Minicio; 90 m; VEN (holo). 

Ouratea la rió Ducke subsp. pervenulosa Maguire 

Steyerm., Mem. New York Bot. Gard. 50: in press. 
1989. "n pens et al. 36657; 27 Nov. 1953; Ven- 
ezuela, 50 km above mouth of Pacimoni; 120 m; NY 


(holo). 

Ouratea duidae Steyerm., Fieldiana Bot. 28(2) 369. 
1952. J. A. Steyermark 58284; 2 Sep. 1944; Ven- 
ezuela, T. F. Amazonas, Cerro Duida; 1,025-1,200 m; 
F (holo), NY. 

Ouratea evoluta Maguire & Steyerm., Mem. New York 


Bot. Gard. 50: in press. 1989. B. Maguire et al. 
29253; 17 Oct. 1950; Venezuela, T.F. Amazonas, 15 
km above San Fernando de Atabapo; 100 m; NY (holo). 

Ouratea fasciculata Maguire & Steyerm., Mem. New 
York Bot. Gard. 50: in press. 1989. Forest Dept, 
Record = 7227 (Field No. G251); 28 Sep. 1952; 
Guyana, 6 mi. E of Onoro Creek; NY (holo). 

Ouratea o (Dwyer) Sandw. & Maguire subsp. 
pachypoda Maguire & Steyerm., Mem. New Yor 
Bot. Gard. 50: in press. 1989. B. Maguire & D. B. 
Fanshawe 32629; 16 Nov. 1951; Guyana, Sagarai- 
madoi Savanna, upper Mazaruni River, Pakaraima 
Mountains; NY (holo 

Ouratea guaiquinimae Maguire & Steyerm., ined. mss. 
= O. guaiquinimensis Sastre, 1988 

Durates huberi Maguire & Steyerm., Mem. New York 
Bot. Gard. 50: in press. 1989. O. Huber 1904; 30 
May 1978; Venezuela, T.F. Amazonas, Dept. Atabapo, 
Rio Ventuari; 98 m; VEN (holo). 

Ouratea darte Maguire e& de , Mem. New 
Yo t. Gard. 50: in press. 1989. F. la iris 
25 Nov. 1965; beoe T.F. Amazon 2 


er near confluenc 1 NY (holo) 
N. 

Ouratea ee Maguire & Steyerm., Mem. New 

York Gard. 50: in press. 1989. O. Huber ra e 


Medina p 763; 6 Feb. 1981; Venezuela, T. 
zonas, Dept. Rio Negro, Río Siapo; 125 m; NY (holo), 
VEN. 


Ouratea macurucoensis Maguire & Steyerm., Mem. New 
Yor Gard. 50: in press. 1989. P. E. Berry 2152; 
25 Hs 1975; Venezuela, T.F. Amazonas, Piedra Blan- 
ca, 1 km Río arriba de Macuruco; 130 m; VEN (holo). 

uri marahuacensis Maguire & Seien. ., Mem. New 

k Bot. Gard. 50: in press. 1989. J. A. Steyermark 
a > K. Holst 130907; 1 Mar. 1985; Venezuela, T.F. 
Amazonas, Cerro Marahuaca, along Pan of Cano 
Negro; 1,140-1,150 m; MO (holo), NY, ] 


Ouratea medinae Maguire & Steyerm., AP New 
Atabapo, San Formand i Atabapo; VEN (holo). 


Ouratea ornata Maguire & (M Mem. New York 
ard. 50: in press. 1989. B. Maguire & L. Politi 
27 185: 1-3 Dec. 1948; Venezuela, mazonas, 
Cerro Sipapo (Paraque); 605-910 m; NY (holo). 
og papillata Maguire & Steyerm., Mem. New York 
. Gard. 50: in press. 1989. J. A. 5 
s. uae 103271; 7 May 197 
ao along Rio Yatua, base of Cerro Yapacana; 
; NY, VEN (holo). 
us paruensis Maguire & a ., Mem. New York 
. R. Cowan & J. 


et al. 
Bolivar, Meseta del tea. Cerro Jani 1,800 m; VEN 
(holo). 
Ouratea ii )hila Maguire & Steyerm., ined. mss. 
= O. steyermarkii Sastre, 1988. 

Ouratea pseudotatei Maguire & Steyerm., Mem. New 
York Bot. Gard. 50: in press. 1989. 4. 5. Pinkus 277; 
Jan. 1939; Guyana, Arabaru River (Kako tributary), 
upper Mazaruni drainage; NY (holo). 


694 


Annals of the 
Missouri Botanical Garden 


Ouratea ptaritepuiensis Steyerm., rae Bot. 28(2): 
n 1952. J. A. Steyermark 59642; 1 Nov. 1944; 
zuela, Bo livar, Pari iind h 000 m; F (holo). 
Shee pum ila Maguire & ne , Mem. New Yor 
Bot. Gard. 50: in press. 19 9. 0. "Huber & S. S. Tillett 
5481, A July 1980; eu. T.F. Amazonas, Dept. 
Atabapo, 5 mi. S of Laguna Pcia m; VEN (holo). 
d Poen Maguire & Steye 
Mag i Lair ., Mem. New : 
in E. O. Huber & S. S. Tillett 5536; 27 
July TUM aiam T.F. Amazonas, Dept. Azures, 
2 km N of Rio Guayapo Medio; VEN (holo). 
ad ramosissima Maguire & Steyerm., Mem 
York Bot. Gard. 50: in press. 1989. F. Cardona 2204; 
Oct. 1947; Venezuela, Bolivar, Cari de Acopán (Ma- 
(holo). 


cizo de Chimanta): 900 m; NY, VEN holo 


no, N of Cerro Yapacana; 130 m; NY (holo). 
Ouratea sipapoensis Maguire > 
Bot. Gard. 50: in press. . Mag 
L. Politi 27631; 12 Dec. 1948; Vonenels. T.F. Ama 
zonas, Cerro Sipapo; 1,700 m; NY (holo). 

Ouratea subamplexic aulis Maguir e & Steyerm., Mem. 

e 50: in press. 1989. F. Papa 
2784; 1 Jan. 1957; Venezuela, Bolivar, 105 
El Dorado hacia m Elena; NY (holo). 
Ouratea venezuelensis Stey , Fieldiana E Aoi 
370. 1952. Ll. Williams. 14152; 2 Feb. ; Ven- 
uela, T.F. Amazonas, del Cano Temi, Misa 128 
m; F (olo. 
Ouratea siglas kiana Magui ire & Steyerm., Mem. New 
ard. 50: in press. 1989. J. J. Wurdack & 
L. S. didi. 42719; | June 1959; Venezuela, T.F. 
Amazonas, Rio Orinoco just above mouth of Río Ata- 
bapo; 125 m; NY (holo). 
Ouratea yapacanae Maguire & Steyerm., ined. mss. = 
. yapacanae Sastre, 1988. 

Perissocarpa, Steyerm. p Papui; Ann. Missouri Bot. 
Gard. 71: 319. 1984, gen. nov. 

Perissocarpa oiii. s (Maguire) Steyerm. & Ma 
guire subsp. tachirensis E aguir 
Missouri Bot. Gard. 71: . 1984. J. A. al 
& B. Manara i 2i June 1981; Venezuela, 
Táchira, along Río San Buena, area of Presa Las Cue- 
vas, ca. 10 km E of La Fundación: 550-600 m; VEN 
(holo). 

Perissocarpa umbellifera RE & Maguire, Ann. 
Missouri Bot. Gard. 71: . 1984. B. Maguire et al. 
29529; 18 Nov. co ME T.F. Amazonas, 
Cerro Duida, Río Cunucunuma; 1,400 m; NY, VEN 

lo 


(holo). 

Poecilandra pe ENT eie Bot. 28(2): 371. 
1952. J. A. 1944; Ven- 

F (holo). 

din Steyerm., Fieldiana Bot. 

: 376. 1952. se 

Sauvages duidae Sa Fita [e 28(2): 372. 

1952. J. A. Steyermark 5802 . 1944; Ven- 
ezuela, T: F. Amazonas, near a Ne 1,400 m; F 
(holo). 

Sauvagesia guianensis (Eichl.) Sastre oe aracamu- 
niensis Steyerm., Ann. Missouri Bot. Gard.: in pre 
1989. R. L. Liesner & F. Delascio 22075; a 

E, Ana azonas, Dept. Río Negro, Cerro Aracamuni, 


summit, Popa Camp; 1, 550 m; MO (holo), VEN. 


Schoepfia tepuiensis Steyerm 


Sauvagesia guianensis (Eichl.) Sastre subsp. guaiquin- 
imensis Steyerm., Ann. Missouri Bot. Gard. 74: 101. 
987. J 17498; acres 

. Amazonas, “Cerro Guaiquinima; 1,540 m; MO 

(holo), V 

Sauvagesia guianensis (Eichl.) Sastre subsp. Sipapoen- 
sis Steyerm., Ann. Missouri Bot. Gard. 74: 101. 1987. 
J.. nun et al. 124530; encre: T.F. Ama- 
zonas, Serranía Sipapo; 1,500 m; MO (holo), VEN. 

Sauvagesia imthurniana (Oliver) Dwyer subsp. chiman- 
ensis Maguire, Steyerm. urd., Mem. New York 
Bot. Gard. 10(4): 17. 1961. J. J. Wurdack 34201; 
26 Jan. 1953; Venezuela, Bolivar, Churi-tepui (Muru- 
tepui); 2,250-2,300 m; F, NY (holo). 

Sauvagesia longipes Steyerm., jose Bot. 28(2): 374. 
195 A. Steye ad 6021: ; 15 Nov. 1944; Ven- 
melo. Bolivat: Salto de pic meru, Sororopan- 
tepui; 1,615 m; F (holo), MO (1313407). 

Sauvagesia marahuacensis Steyerm., Ann. Missouri Bot. 
Gard. 74: 98. 1987. R. L. Liesner 1767 7a; Venezuela, 
T.F. Amazonas, Dept. Atabapo, Cerro Marahuaca, Rio 
Yameduaka arriba; MO (holo), VEN 

Sauvagesia marahuacensis eae subsp. neblinensis 
Steyerm., Ann. Missouri . Gard.: 87. 

. L. Liesner 16809; Venezuela. T. 
Rio Cs Cerro de la Neblina, Camp IV; 780 m; MO 
(holo), 

P an miniata Steyerm., tapeo: Bot. 28(2): 376. 
1952. J. A. Steyermark 58467; 8 Se ep. 1944; Ven- 
ezuela, T.F. Amazonas, vicinity of Sanariapo; 100 m; 
F (holo), MO (324780). 

Tyleria breweriana Steyerm., Ann. Missouri Bot. Gard. 
71: 322. 1984. J. 4. Steyermark et al. 124326; 14 
Feb. 1981; Venezuela, Bolivar, Dept. Sucre, Meseta 

. olo). 


n a ., Mem. New 
972. J. A. Steyermark 

98049; 22-27 Mar. 1967; cet d Bolivar: Meseta 

de Jaua, Cerro Jaua; 1,922-2,100 m; NY (holo), VEN. 


OLACACEAE 


Dulacia redmondii Steyerm., Phytologia 38(3): 217. 
9 A. Steyermark & P. Redmond 112829; 28 


Heisteria Vd Steyerm., Bol. Soc 
A. Steyermark 92077 
1963; E Distr. Federal, Cerro Naig 
1,000-1,300 m; NY, S, US, VEN (holo) = H. latifolia 
Standl., 1984 
Liriosma tepuiensis Steyerm., Bol. Soc. Venez. Ci. Nat. 
32: 328. 1976. J. A. Steyermark et al. 109041; 13 
Feb. 1974; Nena: Bolivar, Meseta del Jaua, Cerro 
Sarisarinama; 1,320 m; F, K, L, NY, VEN (holo) — 


Dulacia tepuiensis (Steyerm.) Sleumer & Steyerm., 
1984. 


i5- 19 Nov. 


Schoepfia clarkii Steyerm., Ann. Missouri Bot. Gard. 75: 
1061. 1988. H. L. Clark 8111; 16 Aug. 1987; Ven- 
ezuela, T.F. Amazonas, 10.8 km NE of San Carlos on 
Solano road; 119 m; MO (holo). 

., Bol. Soc. Venez. Ci. Nat. 

J. A. Steyermark 93413; 5 May 1964; 

Venezuela, Bolivar, Auyan-tepuy; 1,850 m; K, NY, P, 

S, U, US, VEN (holo) 


Volume 76, Number 3 
1989 


Taylor 695 
Plants Described by 
Julian A. Steyermark 


OLEACEAE 


Fraxinus bicolor Standl. & Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 23(2): 73. 1944. P. C. Standley 80408; 
Dec. 1940; Guatemala, Dept. Guatemala, near Fiscal; 
1,100 m; F (holo). 

Fraxinus cavekiana Standl. & Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 23(2): 74. 1944. A. F. Skutch 1660; 
Nov ee Guatemala, Dept. Quiche, Nebaj; 1,860 


m; 

Fraxinus om Standl. & Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 23(2): 74. 1944. P. C. Standley 76162; 
Oct.-Nov n Guatemala, Dept. Jutiapa, near Ju- 
tiapa; 850 m; F (holo). 

Linociera avilensis Steyerm., Brittonia 30: 45. 1978. E. 
Delgado 233; 9 May 1939; Venezuela, Distr. Federal, 
Cerro Avila; VEN (holo). 

Menodora coulteri A. Gray var. minima Steyerm., Ann. 
Missouri Bot. Gard. 19: 121. 1932. J. E. Kirkwood 
20; June 1908; Mexico, Zacatecas, Cedros, near Ar- 
royos; F, MO (holo). 

Menodora decemfida (Gill.) A. Gray var. longifolia Stey- 

. Missouri Bot. Gard. 19: 143. 1932. E. 

Palmer 429; Apr., Aug. 1896; Mexico, Durango, San- 
tiago, Papasquiara; MO (holo). 

Menodora helianthemoides Humb. & pd var. en- 
ue Steyerm., Ann. Missouri Bot. Gard. 19: 119. 
1932. J. Gregg 44; 7 May 1948; Mexico, Coahuilla, 
near Saltillo; GH, MO (holo). 

Menodora helianthemoides Humb. & Bonpl. var. hu- 


> 
d 
d 


1878; Mexico, 

San Luis Potosi, region of San Luis Potosí 1,849- 
2,439 m; F, G, MO (holo). 

Menodora helianthemoides Humb. & Bonpl. var. mag- 
niflora Steyerm., Ann. Missouri Bot. Gard. 19: 118. 
1932. C. C. Parry & E. Palmer 570pp; 1878; Mexico, 
San Luis Potosi, region of San Luis Potosi; 1,829- 


2,439 m; F, GH, MO (holo 
Menodora heterophylla Moric. ex DC. var. australis 
Steyerm., Ann. Missouri Bot. Gard. 19: 127. 1932. 


Pegler 950; Aug. 1903; South Africa, Transvaal, near 
Rustenburg; 1,220 m; B (holo). 

Menodora intricata Brandg. var. purpusii Steyerm., Ann. 
Missouri Bot. Gard. 19: 3 . Purpus 
1318; June 1905; Mexico, Puebla, macia F, GH, 
MO (holo), NY. 

Menodora pinnatisecta Steyerm., Ann. Missouri Bot. 
Gard. 19: 133. 1932. Jorgensen 2643; 9 Nov. 1917; 
Argentina, Terr. del Chaco, Las Palmas; GH (holo). 

Menodora scabra Engelm. ex A. Gray var. longituba 
Steyerm., Ann. Missouri Bot. Gard. 19: 141. 1932. 
Smart 2 ; U.S.A., Arizona, Massatzal 


. ex A. Gray var. ramosissima 

Steyerm., Missouri Bot. Gard. 19 1932 

E. J. Palmer 31083; 19 June 1926; U.S.A., Texas, 
). 


. Gray var. dee Steyerm. 
2. S. B. Pa zi 
9795; 23 Apr. 1915; U.S.A., Cn San Bernar- 
dino Co., Mojave Desert, 14 mi. NE of Barstow; CAS 
(holo). 


ONAGRACEAE 


Fuchsia adpressipilis Steyerm., Fieldiana Bot. 28(2): 
438. 1952. J. A. Steyermark 55381; 8 Feb. 1944; 


ezuela, Lara, between Santo Domingo and Los Que- 


Ven 
braditos, S of Las Sabanetas; 2,430-2,475 m; F (holo). 


. A. Steyermark 56162 
exitela- Mérida. between La Azulita ipid: La Trampa; 
1,280-2,225 m; F (holo). 


ar. tamaensis Steyerm 


57175; 13 July 1944; Venezuela, Táchira, base of 
áramo de Tamá, between Betania and Tama; 2,430 

; F (holo 
“Ludvigia” alternifolia L. var. mek Palmer & 


1933; , Miss 
MO ias = Tea alternifolia L. var. alternifolia, 


196 
Oenothera pilosella Raf. f. laevigata Palmer & Stey- 
erm., Brittonia 10: 116. 1958. J. A. Steyermark 78703; 
25 Jun ne 1955; U.S.A., Missouri, Howell Co., 4 mi. S 
of West Po F (holo) — O. pilosella Raf. var. pi- 
losella, 198 


ORCHIDACEAE 


Scaphyglottis michelangeliorum Carnevali & Steyerm., 
Phytologia 55(5) 289. 1984. J. E n ermar 
: 1983; Venezuela, T.F. 
nas, faldas del Cerro karaliui 1,550 m; VEN donet 


OXALIDACEAE 
E calcicola Standl. & Steyerm., Field. Mus. Nat 
, Bot. Ser. 23(3): 118. 1944. J. A. Steyermark 


50238; 8 Aug. 1942; Guatemala, Dept. H 
go, vicinity Chemal, Sierra de los Cuchumatanes; 3,700 


m; F (holo). 
Oxalis minarum Standl. & Steyerm., Field. Mus. Nat 
Hist., Bot. Ser. 23(2): 58. 1944 A Steyermark 


29718; Oct. 1939; Guatemala, De 
de las Minas, Finca Alejandria; 1,000-1,500 m; F 
(holo). 


PASSIFLORACEAE 


Dilkea magnifica Steyerm., Acta Bot. Venez. 3: 186. 
1968. J. A. Steyermark 87606; 18 Nov. 1960; Ven- 
ezuela, Terr. Delta Amacuro, Cerro La Paloma, Sierra 


Imataca; 100-200 m; NY (holo). 
Passiflora bomareifolia Steyerm. & Maguire, Mem. New 
York Bot. Gard. 17(1): 455. 1967. J. A. Steyermark 


J. Wurdack 1370; 11 Mar. 1955; Venezuela, 
Bolivar, Amuri-tepui, Chimantá Massif; 850- 1,275 m; 
MO (2011386), VEN (holo) 

Passiflora nuriensis Steyerm., Acta Bot. Venez. 3: 188. 
1968. J. A. Steyermark 88425; 14 Jan. 1961; Ven- 
ezuela, Bolivar, Altiplanicie de Nuria, Agua Linda; 400 
m; MO (1803670, 180670), NY (holo). 


PIPERACEAE 


Peperomia acuminata Ruíz & Pavón f. rubra Steyerm., 
Fl. s 2(2): 33. 1984. L. Ruíz Terán 6461; 29 
Sep. 1971; Venezuela, Mérida, Dept. Rangel, entre 
a del Padre y El Baho, unos 10 km al E de la 
Sierra Nevada de Santo Domingo; 2,300-3,340 m; 
MERE, VEN (holo). 

Peperomia aristeguietae Steyerm., Acta Bot. Venez. 6: 


83. 1971. J. A. Steyermark 103733; 17 Aug. 1970; 


696 


Annals of the 
Missouri Botanical Garden 


Venezuela, Distr. Federal, S rd Carvao, Rio Aguas Ca- 


lientes; e 100 m; VEN (ho 
Peperomi a E Venez. 2(1): 50. 1984. 
L. peretas & F. Pannier 1924; 6-7 July 1953; 
Venezuela, Yaracuy, Sierra de Aroa; VEN (holo). 
Peperomia atabapoensis Steyerm., Fl. Venez. 2(1): 52. 
9 s 


84. S. S. Tillett et al. 743-230; 23-26 Mar. 1974; 
Venezuela, mazonas, Dept. A tabapo, Nn Pa- 
von, margen derecha del Rio Orinoco, arri e San 


Fernando de Atabapo; 130 m; VEN (holo). 
Peperomia berryi Steyerm., Fl. Venez. 2(1): 53. 1984. 
P. E. Berry et al. 3923; 19 June 1982; Maie 
Falcón, Urapagual, ‘Los Haitoncitos,' ca. 2 km S 
Curimagua, ladera sur de la Sierra de San Luis 1,200- 
1,300 m; VEN (holo). 
Peperomia boomii Steyerm., Brittonia 38: 220. 1986. 
Boom & M. Grillo 6417; 26 Oct. 1985; Venenuela: 
Bolivar. Distr. Cedeño, vicinity of Panare, village of 
Corozal, 6 km from Mer oo toward Caicara; 200 
m; MYR, NY (holo), V 
Peperomia borburatensis aa NO Venez. 2(1): 61. 
. Steyermark 953: ; 30 Mar. 1966; 
Venezuela, C arabobo, selva sempreverde a ip largo de 


oe iri alia Acta Bot. Venez. 8: 
973. Steyermark & G. S. Bunting 105. 306; 
a Nov. 1971 l; Nansnusis. Yaracuy, Rio Guayabito; 
200-250 m; VEN (holo). 
Peperomia caraboboensis Steyerm. & Bunting, Acta Bot. 
Ve L1 . Steyermark et al. 102431; 
4 Jan. 1970; Venezuela, Carabobo, Rio e 150 
m; VEN (holo). 
de va carnevalii Steyerm., Fl. Ven P 2(1): 67. 
19 G. Carnevali & H. Pivat 640; 2 May 1982; 
V enecaels: Aragua, cumbre de la und C RE Cuy- 
agua; 400 m; MY, VEN (holo). 

Peperomia chapensis Steyerm., Fl. Venez. 2(1): 70. 1984. 
J. A. Steyermark & G. S. Bunting 97664; Venezuela, 
Yaracuy, Cerro La Chapa, N de Nirgua; 1,200-1,360 
m; VEN (holo 

Peperomia c horoniana Trel. & Yuncker var. heterodoxa 
Steyerm., Fl. Venez. 2(1): 74. 1984. S. Bunting 
4461; 7 Jul 1971; Venezuela, amen uy, Cerro La 
Chapa, ca. 5 km N de Nirgua; MY (holo). 

Peperomia ig Hn Standl. & Steyerm., deca Bot. 
24(3) 240. 2. J. A. Steyermark 4 ; Guate- 

d between Tum. ani Sebol; 

00-500 m; F (holo). 
Peperomia croizatiana AR Acta Bot. Venez. 6: 
1971. J. A mark & G. Wessels-Boer 
10037 la; Venezuela, ee Cerro Negro, Sierra 
oa; 100 m; VEN (holo). 
Peperomia c Ma Ay Miq. 7 ia Steyerm., Fl. 
ez. 2 19 ermark et al. 122730; 

98 Deo Venezuela, Distr. Mara, Caño 
bdg vip la Haci Azul y la base del Cerro 
Yolanda, 15 km a poses 5 Rancho 505; 200- 
250 m; es (holo), MO. 

Peperomia delascioi Steyerm., Brittonia 40: 294. 1988. 
F. Delascio 13161; 25-26 Mar. 1987; Venezuela, 
Bolivar, Distr. Piar, Kamarkaibaray-tepui, al este del 
Auyan-tepui; 2,400-2,500 m; VEN (holo), MO. 

Peperomia HERES Standl. & Steyerm., Fieldiana 
Bot. 24(3): 244. 1952. J. A. Steyermark 49410; n.d.; 
Guatemala, Huehuetenango, between Ixcan e Finca 


San Rafael; 200-800 m; F (holo). 


Peperomia dr onensis T rm., ae Venez. 2(1): 95. 
1984. R. Lies t al. 7729; 21 June 1979; 
Venezuela, Fale on, (dem Nacional Quebrada de la 
Cueva del Toro; 600 m; VEN (holo), MO. 

Pom is edd ata Steyerm., Fl. Venez. 2(1): 99. 1984. 

Maguire et al. 36889; 24 Dec. 1953; Venezuela, 
T.F. Amazonas, Cerro de la Neblina, Río Yatua, 3 km 
al sur del Campamento 3; 900 m; NY (holo). 

Peperomia ee Steyerm., Fl. Venez. 2(1): 
101. 1984. R. L. Liesner & A. Conssi 10555; 14 
Mar. 1981; Venez pe Táchira, 18 km al suroeste de 
La Fundación; 950 m; VEN (holo e 

Peperomia pi ig ., Ann. Miss isso 
74: 87. 1987. R. L. Liesner & B. Stannard 16901: 
21-24 Mar. 1984; Venezuela, T.F. Amazonas, Cerro 
de la Neblina, pred V, valley N base of Pico Cardona; 
MO (holo): 34620 

d LIN ie (Sw. ) A. Dietr. var. obtusa Steyerm., 
Fl. Venez. 2(1): 112. 1984. J. A. Bautista 5037; 8 

971; Venezuela, Barinas, Distr. Ezequiel Za- 

Andres Eloy Blanco, Reserva Forestal de 
Caparo; 100 m; VEN (holo i MER. 

Peperomia pages Standl. & Steyerm., versns 
Bot. 24(3): 251. 1952. J. 4. puc 51340; n.d.; 
Guatemala, Hushuet te ace Sierra de los Cuchuma- 
tanes; 800- 900 m; F (ho 

Peperomia honigii Steyerm., e Venez. 2(1): 125. 1984. 
S. S. Tillett & K. Honig 746-798; 27 June-5 July 
1974; Venezuela, Zuli 
Motilones, "Sierra de Perija; 3,000 m; V 

Peperomia ure Standl. & Steyerm., jer Bot. 
24(3) 253 J. A. Steyermark 48613; n.d.; 
Guatemala, MA etd RM Cerro Huitz; 1, pd 32. 000 
m; F (holo). 

dba ic piii Steyerm., Fl. Venez. 2(1): 131. 1984. 
J. A. Ste inei Ns L. Liesner 118651; Venezuela, 
“r Que a ua Azul, al sur de El Reposo, 14 
km E de b 2,150-2,300 m; VEN (holo), 
MO. 


Peperomia manarae Steyerm, ., Fl. Venez. 2(1): 
1984. Manara s.n., 20 Mar. 1980; Venezuela, 
Aragua, justamente por debajo de El Portachuelo, a lo 
largo de la ra que desciende hacia Ocumare de 
la Costa, ade ‘Nacional Henri Pittier; 950 m; VEN 

lo). 


Peperomia marahuacensis Steyerm., Ann. Missouri Bot. 
Gard. 74: 87. 1987. J. A. Steyermark & B. K. Holst 
130742a; 26-27 Feb. 1985; Venezuela, T.F. Ama- 
zonas, Dept. Atabapo, Cerro rr summit, 'Si- 
mas En xà of summit camp; 2,520-2,620 m; MO 
(holo), V 

Pisae mic icroreticulata Fg dae Bol. Soc. Venez. 

Ps Nat. 26: 411. 1966. J. A. Steyermark et al. 92807; 

7 Jan. 1964; ee Bolivar, Cerro Venamo; 
5 220- 1,275 m; VEN (holo) = P. alata Ruíz & Pavón, 
1984. 

nun pc Standl. & Steyerm., Fieldiana Bor. 
24(3): 4. Steyermark 43747 
PE El Progreso, Sierra de las Minas; F (holo) 


Peperomia minarum Steyerm., Fl. Venez. 158. 

. Steyermark et al. 119958; 11 

1979; Venezuela. Táchira, cumbre Vibe de 
Cerro Azul, Cerro las Minas, 18 km SE de Santa Ana; 
1,200-1,380 m; MO, VEN (holo). 

Peperomia ouabiane sanluisensis Steyerm., 


Fl. Venez. 2(1): 173. 1984. H van der Werff et al. 


A 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


697 


237; 18 Jan. 1979; Venezuela, Falcón, bosque nu- 
blado, arriba de La Chapa, Sierra de San Luis; 1,100 
m; VEN (holo). 

mo pariensis Steyerm., Fl. Venez. 2(1): 176. 

1984. . Steyermark & M. Rabe 96216; 9-12 
Aug. 1966; Annie Sucre, Peninsula de Paria, fila 
de Cuchilla de Cueva de Tigre, de Rio Nuevo, Oeste 
de Cerro de Humo; 750-850 m; VEN (holo), US. 

Peperomia portuguesensis Steyerm., Fl. Venez. 2(1): 
195. 1984. J. A. Steyermark et al. 126610; 29 Oct. 
1982; Venezuela, Portuguesa, 15 km al Este de Cha- 
basquen, 67 km al Noroeste de Guanare; 1,450-1,520 
m; MER, MO, PORT, VEN (holo). 

— nd Steyerm., Fl. Ven aa 

198. 1984. L. Aristeguieta & F. Pannier 1948; = 
7 July Tun Venezuela, Yaracuy, Cafetales, Sierra js 
La Aroa; VEN (holo), NY. 

Peperomia pseudojamesoniana Steyerm., Fl. Venez. 2(1): 
199. 1984. J. A. Steyermark & R. L. Liesner 118338; 
22-23 July 1979; Venezuela, Táchira, Quebrada Agua 
Azul, Sur de El Reposo, 14 km SE de Delicias; 2,150- 
2,300 m; VEN (holo), MO. 

Peperomia mu Steyerm., ^ Venez. 2 257. 1984, 
m ah ca al. 761-83; Jan. ; Venezuela, 


Merida, Sucre, a i ae o de la Toma de Agua 
arriba de La Rancheria, entre dA es y La Victoris 
acia La Rancheria y Las Palmas; 1,400 m; VEN 
(holo). 

Peperomia Paus Standl. & Steyerm., Pe Bot. 
24(3): 273. 1952. J. A. Steyermark 46. uate- 


mala, cated 14 May 1942; F (holo) = =P pres 
C. DC., 1966. 


Peperomia Ate Steyerm., Fl. Venez. 2(1): 261. 1984. 
B. Trujillo & E. del Castillo 7919; 30 Mar. 1967; 
Vanes. Merida, El Valle-La Culata; 2,650 m; MY 
(holo) 


Peperomia yatuensis Steyerm., I Venez. 2(1): 275. 
1984. B. Maguire et al. 36891; 24 Dec. 1953; Ven- 
ezuela, T.F. Amazonas, Cerro FR m Neblina, Rio Yatua, 
3 km al Sur del Campamento 3; 900 m; VEN (holo). 

Peperomia yutajensis Steyerm m., Fl Venez. 2(1) 276, 

984. B. & C. Maguire 35304; 17-19 Feb. 1953; 
Venezuela, T.F. Amazonas, peg Yutaje, Rio Mana- 
piare; 1,800 m; NY (holo), V 

Piper agostiniorum Steyerm., Fl. pum Al): 317. 1984, 
G. & T. Agostini 1120; 3 May 1972; Venezuela, 
Falcón, Dept. Democracia, Cerro Monte (Montana); 
900-1,100 m; VEN (holo 

Piper arieianum C. DC. var. yaracuyense (Yuncker) 
Steye f. barbulaense Steyerm., Fl. Venez. 2(1): 
343, 1984. C. Joly 5630; 20 Sep. 1968; Venezuela, 
Carabobo, La Florida, Sierra Barbula; 1,100 m; CAR 
(holo). 

Piper arieianum C. DC 
Steyerm. f. falconense Steyerm., Fl. Vene 

84. van der Werff et al. 548; 7 May 1979; 
Venezuela, Falcón, Montaña de Paraguariba, Sierra de 
San Luis, 3 km al ENE del Cerro Galicia: 1,400 m; 
VEN (holo). 

Piper. arieianum C. DC. var. Ho (Yuncker) 

m. f. nubicolum Steyerm., Fl. Venez. 2(1): 342. 

1984. p H. G. Alston 6167; 8 Jan. 1975. Veda, 

Carabobo, arriba de la Hacienda Cura, entre Valencia 
cay; 1,600 m; às ee), US. 

Piper arieianum D yaracuyense (Yuncker) 
a f. puberulun D. Fl. Venez. 2(1): 341. 

984. R. L. Liesner & A. González 10494; 10-13 


var. yaracuyense (Yuncker) 


z. 2(1): 341 


Mar. 
dación, alrededor de la 
m; VEN (holo), MO. 

Piper Lo Steyerm., Fl. Venez. 2(1): 354. 1984. P. 
E. Ber D. Robinson 3825; 10 Apr. 1982; 
dale poen Parque National Guatopo, ca. 1.5 
km E de la autopista Los Alpes-Altagracia, en el punto 
sur de la curva; MO (3438655), VEN (holo). 

Piper bredemeyeri Jacq. f. escabridum Steyerm., Fl. 
Venez. 2(1): 360. 1984. E. Conicit 2663; 11 Nov 
1977; Venezuela, Falcón, Dept. Bolivar, San José, N 
de San Luis, Sierra de San Luis; VEN (holo). 

Piper canovillosum Steyerm., Fl. Venez. (1): 362. 1984. 
J. A. Steyermark £ M. Rabe 97257; 3-4 Sep. 1966; 
V enezuela, Trujillo, carretera vieja entre Trujillo y Bo- 
cono, entre Urbina y San Rafael, 30 km de Bocono; 
2,300-2,500 m; US (holo), V 

Piper cernuum Vell. f. papillatum Steyerm., Fl. Venez. 
2(1): 369. 1984. J. A. Steyermark 105390; 23 Jan. 
1972; Venezuela, Yaracuy, Quebrada Honda, 17.3 km 
del pueblo de Aroa; 1,100 m; VEN (holo). 

r. araguense (Trel. & Yuncker) 


1981; Venezuela, Táchira, 10 km E de la Fun- 
Represa Dorada; 600-1,000 


Venezuela, Aragua, Parque Nacional Henri Pittier, Por- 
tachuelo; 860 m; VEN (holo 

Piper cerronianum Steyerm., Fl. Venez. 2(1): 375. 1984. 
R. L. Liesner et al. 8176; 27 June 1979; Venezuela, 
Falcón, entre el Edo. Lara y el Edo. Falcón, Cerro 
Cerron; 1,800-2,000 m; VEN (holo), MO. 

Piper ii n "ie rim Fl. Venez. 2(1): 380. 1984. 
R González 9767; 31 Mar. 1980; 
Verexdela. iud Sierra de Aroa, Cerro Tigre, 10 
km al E de Aroa; saad Ba 200 m; VEN (holo 

Piper crenulatum Ste , Fl. Venez. 2(1): 385. 1984. 
J. A. Ste yermark et al 10 0884; 13 Jan. 1968; Ven- 


imari, 20 km a 
35 km al sur de Alquitrana, Suroeste de Santa Ana; 
2,500 m; US (holo), VEN. 

Piper cuyunianum Steyerm., Fl. Venez. 2(1): 391. 1984. 
J. A. Steyermark 89565; 28 Aug. 1961; Venezuela, 
Bolivar, cabeceras del Rio Chicanan, 80 km 
de El Dorado; 500 m; VEN (holo). 

Piper ori Steyerm., Fl. bong "nid 394. 1984. 
G. S. Bunting 2332; 28 Sep. 1967; Vene h- 
ira, — Rubio-Las 
de Aldea de Toronjal, sitio 19 pm arriba de Diamante; 
MY (holo), VEN. 

Pipar diffamatum Trel. & Yuncker var. angustius Stey- 

, Fl. Venez. 2(1): 402. 1984. R. Wingfield 5495; 
27 May 1978; Venezuela, Falcón, Sierra de San Luis, 
arriba de Santa María; 1,240 m; VEN (holo 

ich. f. longipilum Steyerm., E 

A. Steyermark & V. 
Espinosa 105826; 20 Apr. «i May 1972; uM 
Aragua, Parque Nacional Pittier, cabeceras del Rio 
Grande del Medio entre la Fila Alta de Choroni y la 
Quebrada Río Hondo al Sur de Tremaria; 1,000-1,500 
m; NY, US, VEN (holo). 

"D dunstervilleorum Steyerm., Fl. Venez. 2(1): 418. 

1984 . Steyermark et al. 104454; 25 Dec. 1970; 


al Suroeste 


n 134, al Sur de El Dorado; 1,300-1,350 
m; VEN (holo). 
Piper frioense Standl. & Steyerm., Fieldiana Bot. 24(3): 


698 


Annals of the 
Missouri Botanical Garden 


299, 1952. P. C. Standley 90328; n.d.; Guatemala, 
Alta Verapaz, near Tactic; 1,500 m; 


Sn 
1981; Venesuels: Táchira, 10 km E de la Fundación. 
a 23 km por carretera, alrededor de la Represa Dora- 

00-900 m; MO, VEN (holo). 

Piper ie Steyerm., Fl. Venez. 2(2): 427. 1984. 
J. A. Steyermark 9895 6; 16 July 1967; Venezuela, 
Falcon, Sierra de San Luis, Selva Nublada, vecindad 
del ge Parador, al Sur de La Tabla; 1,450 m; US, 
VEN (holo). 

Piper ire en Steye 

z. 2(2): 429. 

1979; Vane Falcón, selva nu- 

a Chapa; 1,200 

; VEN (holo). Note: P. gf. ne a qu is the 
d spelling. 

Piper gentryi Steyerm., Ann. Missouri Bot. Gard. 74: 
9. A. Gentry & B. Stein 46887; 23 Apr. 
1984; Venenisla; T.F. Amazonas, Cerro Neblina, trail 
S from Base Camp; 140 m; MO (holo— 3462077). 

Piper heterobracteum Steyerm., Fl. Venez. 2(2): 439. 
A. Steyermark 99933; 31 Aug. 1967; Ven 

i. Zulia, Sierra de Perija, vecinidad de la velada 

Misión de Los pue de 


rm. f. oe. Stey- 


Piper hippocrepiforme Steyerm., A: enez. A2 440. 
1984. J. A. Steyermark et al. ; 3 Jan. 1964; 
Venezuela, Bolivar, Cerro — es 400- 1,450 m; 

VEN (holo). 

Piper ixocubvainense Standl. & Steyerm., jeu Bot. 
24(3): 305. 1952. J. A. Steyermark 44987; n.d.; 
Guatemala, Alta Verapaz, Montaña Eu F (holo). 

Piper s Steyerm . Soc. Venez. Ci. Nat. 32: 

976. J. A. Steyermark et al. 109689; 28 Feb.- 


319. 
1. 2; x Mar. 1974; Venezuela, Bolivar, Meseta del 
Jaua, Cerru Jaua; 1,810-1,880 m; VEN (holo 


Piper yla Standl. & Steyerm., PRERE Bot. 24(3 ): 
: A. Steyermark 4 ; Guatemala, 
n Volcán de Santa uu l, 300-2,600 
m; F (holo). 

Piper liesneri Steyerm., Fl. Venez. 2(2): 467. 1984. R. 
L. Liesner 6405; 9 Apr. 1979; Venezuela, T.F. Ama- 
zonas, IVIC Estación de Investigación a 4 km NE de 
San Carlos de Río Negro, ca. 20 km S de la confluencia 
del Río Negro y Brazo Casiquiare; 120 m; MO, VEN 

lo 


(holo). 

Piper linguliforme Hp e Fl. Venez. 2(2): 468. 1984. 
R. onzález 10022; Venezuela, Yara- 
cuy, Sierra de Fab 9 i 
aerea), 0-3 km NE por carretera entre Cocorote 
Mons 15 km NE de Cocorote y 1 km SE de los Cru- 

100-1,500 m; MO, VEN (holo). 

] Ste Piper T To Stey- 

E J. A. Steyermark 

oe Serra de 
us extremo SE, fila entre a ríos Tejar y Cocoritico, 
14-15 km NE de Urachiche, a 3 km NE del Caserio 
eed Aires; 1,350-1,500 m; VEN (holo). 

Piper i at re Steyerm., Fl. Venez. 2(2): 
469. 1984. R. L. Liesner & A. González 10891; 20- 
21 Mar. 1981; ae Tachira, ca. 35 km SSE de 
San Cristóbal, La Buenana, 6- E A E de la Quebrada 
Colorado; 600-1, 2: m; MO, VEN (holo). 

i col heirs icum C. DC. f. ona Steye rm., Fl. 
Venez. 2(2): 474. 1984. J. A. Steyermark 104978; 


ud 


984. H. van der Werff 


1 Mar. 1971; Venezuela, Trujillo, entre La Pena 
Agua de Obispo, 22-28 km de Carache; 2,400- 2,500 
m; VEN (holo). 

Piper marturetense Trel. & Yuncker f. pilosum Stey- 
erm., Fl. Venez. 2(2): 485. 1984. J. A. Steyermark 
et al 124826; 1 Mar. 1981; Venezuela, Yaracu 
Serrania de Aroa, extremo SE, selva nublada, fila pur 


Tejar y Cocoritico, 14-15 km al NNE de 


; l 

Piper eiu Standl. & Steyerm., e Bot. 24(3): 
309. 1952. J. A. Steyermark 42667; n.d.; Guatemala, 
And Lr de las Minas; 2,200- a, 400 m; F (holo). 

Piper morilloi Steyerm., Fl. Venez. 2(2): 489. 1984. G. 
Morillo 5510; 8 Feb. 1977; Venezuela, T.F. Amaz 
nas, 1-4 km S de Solano, entre el Cano Chola y Solano; 
120 m; VEN (holo). 

Piper mosaicum Steyerm., Ann. Missouri Bot. Gard.: in 
press. 1989. R. L. Liesner & F. Delascio 21968; 16 
Oct. 1987; Venezuela, T.F. Amazonas, Dept. Rio Ne- 
gro, Cerro Aracamuni, summit, Popa Camp; 1,550 m; 
MO (holo), VEN. 


Piper nubigenum Kunth var. Wir oed Steyerm., Fl. 


Venez. 2(2): 497. 1984. J. A. Steyermark 104989; 
l Mar. 1971; Venezuela, Traj entre La Peña 
Agua de ee 22-28 km de Carache; 2,400-2,500 
m; VEN (ho 


Piper otto- eris Steyerm., Fl. Venez. 2(2): 503. pe 
O. Huber 1848; 29 May 1978; Venezuela, T.F. Am 
zonas, Dept. Atabapo, E del campamento, a 
de Canaripo; 98 m; VEN (holo). 

Piper otto-huberi Steyerm. var. eciliatum Steyerm., Fl. 
Venez. 2(2): 504. 1984. P. E. Berry 694; 22 May 

1975; Venezuela, T.F. Amazonas, al lado de la pista 
de Santa Barbara de Orinoco; VEN (holo 

Piper papilliferum Steyerm., Fl. Venez. 2(2): 507. 1984. 
G. Morillo 4136; 28 Apr. 1978; Venezuela, T.F. Ama- 
zonas, alrededores de San Simon Cocuy; VEN (holo). 

Piper pavasense Steyerm., Fl. Venez. 2(2): 510. 1984. 

A. Steyermark et al. 417 209; 22 May 1978; Ven- 
anol: Bolivar. Las Pavas arriba del Salto Para, Río 
Caura; 250-260 m; VEN (holo). 
Piper perijaense Steyerm., Fl. Venez. 2(2): 517. 1984. 
angoux 10193; 15 Sep. 1974; Venezuela, Zulia, 
Sierra de Perija, al pie de la montana, Serranía de 
Abusanqui; 250 m; VEN (holo). 

Piper perstipulare Steyerm., Fl. Venez. 2(2): 519. 1984. 
S. S. & C. L. Tillett 45501; 23 Sep. 1960; Guyana, 
Cuenca del Alto Río Mazaruni, Río Kako, Campamento 
3, 0.4 km Lr de las cataratas del Kako; 500 m; 
NY, VEN (ho 

TEA Standl. & Steyerm., Fieldiana Bot. 

d k 41928; 


huete Victoria, near Barillas, Sierra de iln 
Cian EE 1,800-2,000 m; F (holo). 

Piper politii Yuncker subsp. toronoense Steyerm., Fl. 
Venez. 2(2): 530. 1984. J. A. Steyermark & J. J. 
Wurdack 1197; 27 Feb. 1955; Venezuela, Bolivar, 
Macizo de Chimantá, Torono-tepui; 1,880-1,970 m; 
F, NY, VEN (holo). 

Piper politii Yuncker subsp. sipapoense Steyerm., Fl. 
Venez. 2(2): 532. 1984. B. RAE & L. Politi 28516; 
21 Jan. 1949; Venezuela, T.F. Amazonas, Cerro Sipapo 
(Paraque), arriba del Cano Grande: 1,500 m; NY (holo). 

Piper pseudoacreanum Steyerm., Fl. Venez. 2(2): 5: 


Volume 76, Number 3 


Taylor 699 
Plants Described by 


Julian A. Steyermark 


1984. R. L. Liesner 6222; 5 Apr. 1979; Venezuela, 
0.5 km S de San Carlos de Rio Negro; 120 m; M 
(2734623), VEN (holo 

Bi pseudoaequale Ste eye 2(2): 540. 

1984. J. A 1966; Ven- 
ezuela, Sucre, Peninsula de Paria, Cerro de Humo, 
entre la Laguna y Roma, NE de Irapa; 900-1,060 m; 
VEN (holo). 

dee pseudobredemeyeri Steyerm., Fl. e s 542. 

84. teyermark 94897; 2 Mar. ; Ven- 
eH ‘Sucre Peninsula de ah ad A pus 

1,273 , US, VEN (holo). 

"uu pedals ui oum Steyerm., Fl. Venez. 2(2): 543. 

984. R did & H. van der Werff 6571; 25 
i. 1978; Venezuela, Falcón, Sierra de S i 
nos nublada anha del Hotel Parador: 1,500 m; VEN 
(holo). 

de pseudohastularum Steyerm., Fl. Venez. 2(2): 548. 

1984. J. A. Steyermark et al. 11997 1; 11 Nov. 1979; 


rm., Fl. T 


m ae eed Steyerm., Fl. Venez. 2(2): 

984. B. Ma n.; 27 Mar. 1971; Venezuela, 

T. F. Amazonas, 1 pos de aeropuerto de San Carlos de 
Rio Negro; VEN (holo 

Piper nm Standl. & Steyerm., Fieldiana Bot. 
24(3) 320. 1952. J. A. Steyermark 43622; n.d.; 
rue Dept. El Progreso, Sierra de las Minas: 

2,600 m; F (holo). 

s ronald Steyerm., Fl. Venez. 2(2): 556. 1984. R. 
L. Liesner & M. Guariglia 11873; 6 May 1981; 
la Táchira, Cerro Minas; 1,200-1,380 m; 
MO, VEN (holo). 

Piper santae-clarae Standl. & Steyerm., Fieldiana Bot. 
24(3) 321. 1952. J. A. Steyermark 46624; n.d.; 
Guatemala, Dept. Suchitepéquez, Volcán de Santa Clara; 
1,250-2,600 m; F (holo). 

Piper sierra-aroense Steyerm., Fl. Venez. 2(2): 563 
1984. R. L. Liesner & A. bits 9699; 30 Mar. 

1980; Venezuela, eo Cerro Tigre; 800-1,000 
m; MO, VEN (holo). 

Piper subsessilifolium C. DC. var. morii Steyerm., Fl. 

V «LL. 


cumbre Gamelotal en El Amparo, 7.5 km N de Salom; 

1,250 m; MO (3438654), VEN (holo). 

Piper tacariguense Steyerm., Fl. Venez. 2(2): 570. 1984. 
Steyermark et al. 121697; 23 Feb. 1980; Ven- 


ezuela, Sucre, Dept. Arismendi, Peninsula de Paria, 


del Cerro de Humo; 750 m; MO, VEN (holo). 
Piper tamayoanum Steyerm., Fl. Venez. 2(2): 572. 1984. 
Steyermark & R. L. Liesner 127445; 1 Dec 
198 2: Venezuela, Bolivar, Distr. Roscio, near base al 
Cerro Ceita; 950 m; MO, VEN (holo). 
Piper y rend Steyerm., Fl. Venez. 2(2): 575. 1984. 
A mark & J. J. Wurdack 1346; 10 Mar. 
1955; TEA Bolívar, Macizo de Chimantá, Amuri- 
tepuy; 1,365 m; VEN (holo). 

Piper toronotepuiense Steyerm., Fl. Venez. 2(2): 576. 
1984. teyermark 75476 ; 19-20 May 1953; 
Venezuela, Bolivar, Macizo de Chinamtá, Chimantá- 
tepui (Torono-tepui); 1,700 m; VEN (holo 

Piper ero Standl. & Steyerm., Fieldiana Bot. 24(3): 

. P. H. Gentle Il n.d.; Belize, El Cayo 
Biss; - p F (holo), MIC 
Piper venamoense Steyerm., » Neues 2(2): 589. 1984. 


J. A. Steyermark et al. 92631; 3 Jan. 1964; Vene- 
zuela, Bolivar, Cerro Venamo; 1,400-1,450 m; VEN 
lo 


(ho 

Piper veraguens : var. venezuelense Steyerm., 
Fl. Venez. 2(2) 5 90. 1984. F. died 4082; 11 Aug. 
1964; Venezuela, Trujillo, 16 e Bucono, a lo largo 
de la carretera a Biscucuy; 1, 850 m; MER, VEN (holo). 

Piper vitaceum Yuncker var. venesuelense Sopan, Fl. 
Venez. 2(2): 594. 1984. J. A. Ste, S. Nilsson 
176; 15-17 Apr. 1960; uds aha la Fila de 
La Danta, entre Luepa y Cerro Venamo; 1,200 m 
VEN (holo). 

Piper d ME Ar Venez. 2(2): 595. 1984. 

«Lb. er et al. 8 27 June 1979; Venezuela, 

Falcón. area E gi pe el Edo. Lara y el Edo. 
Falcón, Cerro Cerron; 1,800-2,000 m; VEN (holo). 

Sarcorhachis venezuelana Steyerm., dea 3: 33. 
1971. J. A. & C. Steyermark 95152; 27-28 Mar. 
1966; Venezuela, Carabobo, del Rio E Gian, al Sur 
de Borburata, arriba de la planta electrica, entre Los 

N (h 


Tanques y La Toma; 750 m; US, VEN (holo). 


POACEAE 


Muhlenbergia brachyphylla Bush f. aristata Palmer & 
Steyerm., Brittonia 10: 110. 1958. J. A. e lal 
82921; 7 Oct. 1956; U.S. A., Missouri, Pike Co., vi- 
cinity of Stillhouse Cave, along City Fork of Pen 
Creek; F (holo), US. 


POLEMONIACEAE 


Phlox divaricata zh var. laphamii Woodson f. candida 
almer Steyerm., Rhodora 57: 316. 1955. E. J. 
€ 55411. 30 pu 1953; U.S.A., Miss ou Bar- 
o. l mi. W of Nashville; Palmer Herbarium 

(holo), F. 


POLYGALACEAE 


Polygala appressa Benth. var. kavanayena Steyerm., 
pere Bot. 28(2): 298. 1952. J. A. Steyermark 
59359; 26 Oct. 1944; Venezuela, Bolivar, Gran Sa- 
bana; T 220 m; F (holo). 

"une blakeana Steyerm., Fieldiana Bot. 28(2): 298. 

A. Steyermark 59065; 10 Oct. 1944; Ven- 
era Bolívar, Gran Sabana; 1, 065- l, 200 m; F (holo). 

Polygala rimulicola Steyerm., Ann. Missouri Bot. Gard. 

19: A. Moore & J. A. Steyermark 
3515; 20 July 1931; U.S.A., Texas, Culberson Co., 
Smith Canyon, Guadalupe Mts.; 1,900 m; GH (holo), 
MO (1011635). 

Pa sanariapoana Steyerm., Fieldiana Bot. aus 
301. 1952. J. A. Steyermark 59452; 8 Sep. 19 
Venezuela, T.F. Amazonas, Sanariapo; 100 m; F ein 

Polygala santanderensis Killip prier Fieldiana 
Bot. 28(2): 301. 1952. Killip & Smith 16585; 5-6 
Jan. 1927; Colombia, Dept. Santander, Rio one val- 
ley; 2,000-2,300 m; F, US (holo). 


PORTULACACEAE 


Montia calcicola Standl. & Steyerm., Field. Mus. Nat. 
Hist., Bot. Ser. 23(2): 48. 1944. J. A. Steyermark 
50308; 8 Aug. 1942; Guatemala, Dept. Huehuetenan- 
go, Cerro Chemal; 3,700-3,800 m; F (holo 

eyerm., Ann. Missouri . Gard. 

& G. al 8200; 

Bolivar, carretera Ciudad 


26 Feb. 1980; Venezüela, 


700 


Annals of the 
Missouri Botanical Garden 


Bolivar- Maripa, 100 km W of Ciudad Bolívar; 100- 
200 m; VEN (holo). 

Portulaca pygmaea Steyerm., Ann. Missouri Bot. Gard. 
15: 1058. 1988. B. Maguire et al. 36188; 12 Nov. 
1953; Venezuela, T.F. Amazonas, 1 km E of Hotel 
Amazonas, Puerto Ayacucho; 100 m; NY (holo). 


PRIMULACEAE 
Lysimachia quadriflora Sims f. albescens Steyerm., Rho- 
dora 54: 257. 1952. J. 4. Steyermark 68397; 4 July 
1949; U.S.A., Missouri, Carter Co., along route 60 
Pune spring branch tributary to right fork of Carter 


reek, 6.7 mi. NE of Van Buren; F (holo). 
PROTEACEAE 
Euplassa chimantensis Steyerm., Bol. Soc. Venez. Ci. 
Nat. 25: 77. I ES Steyermark & e J. riii 


1103; 23 Feb. ar, Chimant 
Massif, Torono- Pi 1, 955-2,090 m; VEN (holo), F, 
NY. 


28(1): 
. 1944; 
anta Teresita de Kava- 


Euplassa venezuelana Steyerm., Fieldiana Bot. 
217. 1951. Steyermark 60824; 30 Nov 
Venezuela, Bolívar, NW of S 


. Missouri Bot. Gard. 

1987. B. Maguire & L. Politi 28409; 17 

Jan. 1949; cae T.F. Amazonas, Rio Cuao, Rio 
Orinoco; NY (holo). 

Panopsis pue ia Steyerm., Bol. . Venez. Ci. 
Nat. 25: 79. A. dae a J. Wurdack 
1036; 21 SN S Venezuela, Bolivar, Chimantá 
Massif; 2,030-2,150 m; > NY, VEN (holo). 

Panopsis parimensis Steyert , Ann. M issouri Bot. Gard. 
14: 612. 1987. O. Huber 6136; 12 June 1981; Ven- 
ezuela, T.F. Amazonas, Dept. Atabapo, Sierra Parima, 
25 km E de Parima ‘B, cabeceras del Rio rama: 
MO (3295700), NY, VEN (holo). 

decias — s Steyerm., ined.? J. A. Steyermark 

al. 1 ; 27 Mar. 1972; std dedi Sierra 
de Pei Suroeste de Pishikakao E Iría hacia la Misión 
de Suc 1,500-1,800 m; MO (iso — 2680841). 

Panopsis ana d Fieldiana Bot. 28(1): 218. 
1951. J. A. Steyermark 59677; 1 Nov. 1944; Ven- 

ezuela, Bolivar, Ptari- nia 1,600 m; F (holo). 

jane tepuiana Pile 23 o Bot. 28(1): 219. 

A. Steyermark 5 -11 Nov. 
Venezuela Bolivar, Ptari- fate rt 1,600 m; F 

Nonnus chimantensis Steyerm., Bol. . Venez. Ci 
Nat. 25: 

1105; 23 Feb. ! Bolivar, Chimantá 
Massif; 2,090 m; VEN (holo), F, MO (2011335), NY. 
Roupala griotii Steyerm., pese 44(5): 321. 1979. 
Steyermark et al. ; 22 Feb. 1979; Ven- 
sanels. T.F. Amazonas, nein : Rio Coro-Coro 

y del aeropuerto de Yutaje; VEN (holo). 

Roupala minima Steyerm., Fieldiana Bot. 28(1): 220. 
1951. J. 4. CHR dua 60234; 15-17 Nov. 1944; 


Venezuela, Bolivar, between Pini: -tepui and Sororopan- 


.K. var. ecuadorensis Steyerm., 
Phytologia 9(6): 342. 1964. J. 4. Steyermark 54825; 
20 Oct. 1943; Ecuador, Prov. Loja, along Rio Cachi- 
yacu at Tambo Cachiyacu; 7 000 m; F (holo). 

Roupala paruensis Steyerm . Missouri Bot. Gard. 
74: 613. 1987. R. S. Cosa" ry J. Wurdack 31378; 


10 Feb. 1951; Venezuela, T.F. Amazonas, Cerro Parú, 
S-SE to edge descent to tributary of Caño Asisa, Rio 
Ventuari; 2,000 m; NY hour ). 
i og sororopana Ste doceri a Bot. 28(1): 220. 
l. J. A. ión 6008 -13 ‘Nov. 1944; Ven- 
a Bolivar, Sororopan- s 2,225-2,255 m; F 
(holo). 


QUIINIACEAE 


Froesia venezuelensis Steyerm. & Bunting, Brittonia 27: 
175. 1975. J. A. Steyermark & G. S. Bunting 105237; 
12 Oct. 1971; Venezuela, Yaracuy, entre Nirgua y 
Las Marias; 1,000-1,200 m; NY, US, VEN (holo). 


RANUNCULACEAE 


Anemone virginiana L. f. plena Palmer & Steyerm., 
Brittonia 10: 113. 1958. G. E. Moore s.n.; 
U.S.A., Missouri, St. Louis Co., Rockwoods Reserva: 
tion; c (holo 
ge caleoides Sondi, & Steyerm., Field Mus. Nat 
, Bot. Ser. 23(2): 52. 1944. P. C. Standley 587 32; 
TS apis Guatemala, Dept. Chimaltenango, Cerro 
ecp gion of Santa Elena; 2,700 m; F (holo). 
Clematis M COMME Small ex Rydb. f. pubescens Stey- 
erm., Rhodora 40: 71. 1938. Brinkley 268; 24 July 
1937; U.S.A., Arkansas, Sevier Co., Boggy Springs; F 


(holo). 
Ranunculus oe H.B.K. f. subintegrus Steyerm., 
Fieldiana Bot. 28(1): 232. 1951. eyermark 
y 1944; Venezuela, Mérida, above Los 
Apartaderos, pros. Laguna Mucubaji towards Lagu 


na 
Negra; 3,625-3,655 m; F (holo) = R. ym 


Thalictrum johnstonii Standl. & Steyerm., Field Mus. 

Bes Hist., Bot. Ser. 22(4): 229. 1940. J. R. Johnston 

43; 20 June 1940; Guatemala, Dept. Totonicapán, 
Du F (holo). 

Thalictrum standleyi Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 22(4): 229. 1940. J. A. Steyermark 36258; 
20 Feb. 1940; Guatemala, Dept. San Marcos, Río 
Vega, near San Rafael and Guatemala-Mexican bound- 
ary Volcán Tacaná; 2,500-3,000 m; F (holo). 


RAPATEACEAE 
Kunbardrin radiata Maguire & Steyerm., Acta Ama- 
: 207. 1979. J. A. Steyermark & O. Huber 


13851; 4 May 1977; Venezuela, T.F. Amazonas, 35 
Ya S of Puerto Ayacucho at Tobogán; 85 m; NY (holo), 
VEN. 


Rapatea aracamuniana Steyerm., Ann. Missouri Bot 
. R. L. ius & F. Boon 
- 1987; Venezuela, T.F. Amazonas, 
SE Rio Negro, p Aracamuni; 600 m; MO (holo), 
VEN. 


s chimantensis Steyerm., Ann. Missouri Bot. Gard. 
i 98 A. St cad 75584; 24 May 
1953; Venezuela, "Bolivar, Chimantá Massif, lower 
southwestern slopes of Chimantá- iur 1,000 m; MO, 
NY, VEN (holo). 
i dels albiflora Steyerm., x bins Missouri Bot. Gard. 
ermark et al. 108873; 
10 Feb. 1974; Vesali, Bolivar. Meseta del Jaua, 
Cerro Sarisariama, NE part; 1,410 m; VEN (holo), NY. 
Stegolepis allenii Steyerm., Field Mus. Nat. Hist., Bot. 
Ser. 22(5): 325. 1940. P. H. Allen 2153; 23 June 


Volume 76, Number 3 
1989 


Taylor 701 
Plants Described by 
Julian A. Steyermark 


1940; Panama, Prov. Cocle, hills N of El Valle de 
Anton; 1,000 m; F (holo), MO (1227389) = Epidryos 
allenii (Steyerm.) Maguire, iil 

B os ap ue Steyerm., Fieldiana Bot. 28(1): 
130. 1951. J. A. St M. 58040; 26 Aug. 1944; 


eet TF. Amazonas, Cerro Duida; 1,095-1,520 


m; F (holo). 
Stegolepis a Steyerm., Ann. Missouri Bot. Gard. 
74: 610. . O. Huber 9467; 28 Apr. 1984; Ven- 


ezuela, Bolivar Kukenán-tepui; 2,500 m; MO, VEN 
lo 


(ho 

Stegolepis humilis Steyerm., Ann. Missouri Bot. Gard. 
14: 611. 1987. J. 4. Steyermark et al. 132006; 22- 
24 May 1986; Venezuela, Bolivar, Camarcaibari-tepui, 
SW -facing lm 1,825 m; MO (holo— 3514022, 
3393686), V 

Stegolepis T Steyerm., Ann. Missouri Bot. 
Gard. 71: 300. 1984. J. A. Steyermark et al. 129962; 
14 Feb. 1984; Venezuela, Bolivar, Chimantá Massif, 
Acopan- tepui; 1,950 m; VEN (holo), NY. 

Stegolepis minor Steyerm., Ann. Missouri Bot. Gard. 75: 
315. 1988. J. A. Steyermark 89577; 28 Aug. 1961; 
Venezuela, Bolivar, Sierra de Lema, cabeceras del Rio 
Chicanán; NY, VEN (holo) 

-— parv irs Steyerm., Fieldiana Bot. 28(1): 

4. Steyermark 59514; i is 1944; 
ea Politan, Ptari-tepui; 1,810 F (holo). 

e ptaritepuiensis Steyerm., ¿Felina Bot. 28(1): 

1951. J. A. Steyermark 5 ; 1 Nov. 1944; 
Vasa Bolivar, Ptari-tepui; E 600 m; F (holo). 

Stegolepis era Steyerm., Ann. Missouri Bot. 

; 4. J. A. Steyermark & F. Delascio 

r. 1983; Venezuela, T.F. Ama 

2,600 m; VEN (holo). 


129197; 30 “re Ap 
zonas, Cerro Marahuaca; 2,580- 


RHAMNACEAE 
Fieldiana Bot. 28(4): 


Colubrina ica Steyerm., 
71. 1 , nom. nov. 
Fieldiana Bot. 28(2): 

'enezuela, Distr. 
l, 600-1,800 m; F 


Colubrina venez szuelensis Steyerm.; 
355. 1 T. González s.n.; n. 
Voderal. didis: de Caracas; 


olo). 
Gouania e aeo kii Steyerm., Ann. Missouri Bot. Gard. 
75: 1065. 1988. J. J. Wurdack & J. V. Monachino 
39810; 12 oe 1955; Venezuela, Bolivar, Distr. Ce- 
deno, Cerro San Borja; 100-300 m; MO (holo), NY, 
VEN. 


Rhamnus acuminata Maguire & ge Mem. New 
York Bot. Gard. 50: in press. 1989, B. Maguire et 
al. 42496; 25 Dec. 1957; abe T.F. Amazonas, 
Cerro de la Neblina, summit, Canon Grande; 1,050- 
1,150 m; MO (3291195), NY (holo). 

Rhamnus chimantensis oe & Maguire, Mem. New 
York eee Ms 17(1) 4 . J. A. Steyermark 
75946; 21-22 June 1953; Venezuela: Bolivar, Apa- 
cara- ra Chimanta Massif; 2,450-2,500 m; NY 
(holo). 

Rhamnus longipes Steyerm., Ann. Missouri Bot. Gard. 
75: 1066. 1988. R. S. Cowan & J. J. Wurdack 
31388; 10 Feb. 1951; Venezuela, T.F. Amazonas, 
Serranía Parú; 2,000 m; NY (holo). 

Rhamnus marahuacensis Steyerm. & Maguire, Acta Bot. 
Venez. de 22 A. Steyermark et al. 

2 Feb. 1982; Vadi, T.F. Amazonas, 

Atabapo, Cerro Marahuaca; 2,480-2,500 m 

NY, VEN (holo). 


Rhamnus microreticulata Maguire & Steyerm., ined. 
mss. = R. neblinensis Maguire & Steyerm., 1989 
Rhamnus neblinensis Maguire & Steyerm., Mem. New 

York Bot. G 

1954; Venezuela, T.F. Amazonas, 

Cerro de la Neblina; summit; 1,800 m; MO (3291187), 
(holo). 

Rhamnus psilocarpa Maguire & Steyerm., Mem. New 
York Bot. Gard. 50: in press. 1989. B. Maguire et 
al. 37230; 10 Jan. 1954; Venezuela, T.F. Amazonas, 
Cerro de la Neblina, summit; 1,700-2,000 m; MO 
(3291189), NY (holo) 

Rhamnus sipap rm., Ann. Missouri Bot. Gard. 
75: 1066. 1987. B. Maguire Fs L Politi 28656; 26- 
28 Jan. 1949; Venezuela, T.F. Amazonas, Cerro Sipapo 
(Paraque) rim head of S basin; 1970 m; NY (holo), 


Rhamnus subpedunculata Maguire & Steyerm., ined. 
mss. = R. acuminata Maguire & Steyerm., 1989. 


RHIZOPHORACEAE 


Hypogyneae "pe & iun, 
Gard. 70: 1 71983, tribe n 
Po pol sl Kuhlmann en Ba lsamocaulon Stey- 
erm. & Liesner, Ann. Missouri Bot. Gard. 70: 190. 
983, subg. nov. 
Sterigmapetalum exappendiculatum Steyerm. & Lies- 
, š i . Gard. 88. 1983. B. 
Maguire et al. 46876; 20-21 Aug. 1962; Venezuela, 
Bolívar, La Escalera, Alto Río Cuyuni, Rio Uiri-Yuk; 
1,000 m; NY, VEN (holo 


Sterigmapetalum laa Steyerm 


Ann. Missouri Bot. 


m., Fieldiana Bot. 


. A. Steyermark 90428; 29 Dec. 1961; Ven- 

, Bolivar, Sierra Ichun; 500-625 m; NY, US, 
VEN (holo). 

Sterigmapetalum ene one pU & Liesner, Ann. 
Missouri Bot. Gar 1983. J. A. Steyermark 
99409; 21 July os Vena Falcón, Sierra de 


1,050 m; US (holo). 
E tachirense Steyerm. & Liesner, Ann. 
ri Bot. Gard. 70: 190. 1983. J. A. Steyerm ark 
et P 120032. 11 Nov. 1979; Venezuela, 
erro Azul, at Cerro Las Minas, 18 km S 
Ana; 1,200-1,380 m; MO (3466058), VEN (holo). 


ROSACEAE 
Amelanchier canadensis (L.) Medic. f. nuda ups & 
Missouri us Gar d. 25; + 1998, 


. S of Linesville: 


Rosa setigera Michx. f. alba m Rhodora 54: 

254. 1952. J. A. Steyermark 7 7 July 1951; 

uA. ie Reynolds Co., n mi. “SE of Bunker, 
along Bes Fork; F (holo), GH, MO. 


Rosa setigera Michx. f. inermis Palmer & Steyerm., 


702 


Annals of the 
Missouri Botanical Garden 


Ann. Missouri Bot. Gard. 22: 569. 1935. E. J. Palmer 
39039; 26 Apr. 1931; U.S.A., Missouri, Bollinger, 
Patton; GH (holo). 
Rosa setigera Mickx. var. serena Palmer & Steyerm., 
Ann. Missouri Bot. Gard. 22: 569. 1935. E. J. Palmer 
A. Steyermark 41642; 2 July 1933; U.S.A., 
Missour ri, Ripley Co., Ponder; GH (holo). 
Rubus flagellaris Willd. var. WU Bailey f. roseus 
Steyerm., Rhodora 62: 130. O. J. A. Steyermark 
84212; 8 May 1957; U.S.A., Mant Bollinger Co., 
in valley of creek tributary to Castor River, NW o 
Buchanan Post Office; UMO (holo 
Rubus flagellaris Willd. f. roseo- plen nus Palmer & Stey- 
ayy Brittonia 10: 114. 1958. E. J. Palmer 59655; 
3 May 1 1955; U.S. E Missouri, Barton Co., 0.5 mi. 
SE of Libera; F (holo) = R. flagellaris var. occidualis 
eei f. roseo-plenus (Palmer & Steyerm.) Steyerm., 
1960 
Rubus hadroc E Standl. & Steyerm., Fieldiana Bot. 
. P. C. Standley 86270; 6 Feb. 
dioi Dept. San Marcos, Barranco Emi- 
nencia; 2,500-2,700 m; F (holo). 
Rubus hadrocarpus Standl. & Steyerm. f. adenophorus 
5 & Steyerm., — Bot. 24(4): 478. 1946. 
J. A. Steyermark 49859; 31 July 1942; eges 
Dept. Huehuetenango, no de Limón; 2, 600-3 
lo 


Rubus pensilo anicus Poir. f. albinus Palmer « Steyerm., 
Brittonia 10: 114. . J. A. ese ale 78868; 
23 July 1955; U.S.A., Missouri, Howe , 6 mi. NE 
of West Plains, 5 mi. SSE of White o F (holo), 
GH, MO, US 


RUBIACEAE 


Alibe d acuminata (Benth.) Sandw. var. obtusiuscula 
Steyer em. New York Bot. Gard. 12(3): 223 
Alios 5264; 30 Jan. 1950; Venezuela, 

Bolivar, Ciudad Bolivar; VEN (holo 
Alibertia davidsei Steyerm., Ann. Missi Bot. Gard. 
9. G. Davidse & A. González 12910; 
8- 9 May 1977; Venezuela, Apure, Distr. Pedro Ca- 
mejo, 11 km E of Paso de San Pablo, 2 km ENE of 
Fundo Picachón, along Rio Capanapano; 45 m; MO 

(268781), VEN (holo). 

Alibertia latifolia Benth. var. parague age bes gas 3 
Mem. New York Bot. Gard. 12(3): 225 5. J. J. 
Wurdack & J. Monachino 39784; 10 Des 1955; 
Venezuela, Bolivar, Rio Paragueni; 90 m; NY (holo). 

Alibertia myrciifolia (Spruce) K. apis var. tepuiensis 
ARCHI. Mem. New York Gard. 12(3) 226. 

65. B. & Maguire us 22 Nov. 1954; 

B Terr. Rio Brando: Serra Tepequem; 800 m; NY 

(holo). 


Alibertia Vias Steyerm., Mem. New York Bot. Gard. 
12(3): 227. 1965. S. S. Tilleti et al. 45180; 7 Aug. 
0; EN upper Mazaruni River basin, Mt. Ayan- 


19 
ganna; 900-1,100 m; NY (holo). 


Alseis microcarpa Stan dl. & Steyerm., Fieldiana Bot. 
28(3): 565. 1953. J. A. Steyermark 62232; 1 
1945; Venezuela, Monagas, Montana de Aguacate; along 


Quebrada de Pajarral; 600-900 m; F (holo). 
& Steyerm., Fieldiana Bot. 


elix; F (holo) — 9, labatiodes 
Karst 


. ex Schumann, 1974 


Amaioua brevidentata Steyerm., 


E 
Gard. 17(1) 


“Arachnothrix” calycophy lla Steyerm., Mem 


Mem. New York Bot. 
Gard. 12(3): 216. 1965. J. A. Steyermark 89802; 20 
Oct. 1961; Venezuela, Aragua, Parque Nacional Henri 


Amaioua guianensis Aublet var. macrantha Steyerm., 
Mem. ork Bot. Gard. 12(3): 216. 1965. B. 
Maguire et al. 42025; 11 Nov. 1957; Venezuela, T.F. 
Amazonas, Cerro de la Neblina, Camp 3; 700 m; NY 
(holo). 

am sya ET Steyerm., Mem. New York Bot. 
Gard. 23(1): 3 2. F. R. Fosberg & N. C. Fassett 
21825; 20 hg 3544 Colombia, Dept. Santander, E 
of Cimatara, 17 km WNW of Landazuri; 230-330 m; 
US (holo), N 

Amphidasya neblinae Steyerm., Mem. New York Bot. 
Gard. 23(1): 320. 1972. B. Maguire et al. 60295; 1 
Dec. 1965; Brazil; Amazonas, Rio Negro, Rio Cauaburi, 
Rio oce =S of Serra de Neblina; MO (2029484), 
NY (holo), V 

Aphanocarpus RM erm., ione New York Bot. Gard. 
12(3): 263. 1965, gen. 


Aphanocarpus steyermarkii Stand ) eed > elon- 


m., Ann. Missouri Bot. Gard. 71: 331. 


Jan. 1983; Venezuela, Bolivar, Piar, Macia de Chi- 
mantá; 2,000 m; VEN (holo). 

Aphanocarpus steyermarkii (Standl.) puede gla- 
brior Steyerm., Ann. Missouri Bot. Gard. 71: 330. 
1984. J. A. Steyermark etal. 115968; 25 Feb, 1978; 
Venezuela, Bolívar, cumbre de Aprada-tepui; 2,460- 
2,500 m; VEN (holo 


Appunia e Steyerm. ., Mem. New York Bot. Gard. 


17(1): 35 7. A. C. d 1-4 Nov. 1937; 
Guyana, s of Rupunu ar mouth of Char- 
wair Creek; NY (holo) — Morinda da (Steyerm.) 
Steyerm., 1972. 


Appunia e Steyerm., Mem. New York. 


Gard. 17(1): 3 1967. Fores Dept. Field No. CAP 

41; 16 Feb. 1946 Guyana, Mazaruni Station; NY 

(holo) = Morinda Pius (Steyerm.) Steyerm., 
2. 


Appunia tenuiflora (Benth.) Jacks. & Hook. var. leio- 


phyla Steyerm., Mem. New York Bot. Gard. 17(1): 
1967. Forest Dept. Field No. WB 500; Nov. 
ee Guyana, Iramaipang, Kanuku Mountains; 150 
m; NY (holo) = Morinda tenuiflora (Benth.) Steyerm. 
var. leiophylla (Steyerm.) Steyerm., 1972 
unia venezuelensis Steyerm., Mem. New York Bot. 
356. 1967. J. A. Steyermark 88277; 10 
Jan. 1961; Venezuela, Bolívar, Altiplanicie de Nuria; 
400 m; VEN (holo) = Morinda venezuelensis (Stey- 
erm.) Steyerm., 


. New York 


ra 
raquita, Rio Tá chira; L 675-1,980 m; VEN 
Note: Arachnothyrx is the correct pe 
costanensis Steyerm em w York 
Bot. Gard. 17(1): 259. 1967. J. P Sueyenmark 61424. 
7 Mar. 1945; Venezuela, Anzoátegui, Cerro Peonia 
(oan Coroy); 1,000-1,450 m; VEN (holo). 
“Arachnothrix” fosbergii Steyerm., Mem. New York 
Bot. Gard. 17(1): 255. 1967. F. R. Fosberg & M. A. 
Giler E 10 Feb. 1945; Ecuador, Loja, Rio Salado; 
NY (holo 5 
2 lojensis Steyerm., Mem. New York Bot. 


Volume 76, Number 3 
1989 


Taylor 703 
Plants Described by 
Julian A. Steyermark 


Gard. 17(1): 255. 1967. F. R. Fosberg & M. A. Giler 

23141; Ecuador, Loja, Rio San Francisco, Cordillera 
de Zamora (El Condor); NY (holo). 

“Arachnothrix” i (Benth.) Planch. var. breviloba 
Steyerm., Mem. New York Bot. Gard. 17(1): 253. 
1967. F. R. Fosberg 21418; 14 Dec. 1943; Colombia, 
Norte de Santander, La Camarona; 1,700-1,850 m; 

NY (holo). 
** Arachnothrix’ ed (Benth.) Planch. var. calycina 
York na Gard. 17(1): 255. 

mero a 24332; 
22 Sep. 1959; Colombia, Ma gdalena, Sierra Nevada 
de Santa Marta, Hoya del Rio Donachui; 1,350-1,500 
m; VEN (holo ). 

“Arachnothrix” irn (Benth.) Planch. var. glandu- 
lifera Steyerm., ew York. Bot. Gard. 17(1): 
253. 1967. J. A. eodd 61317. 1-2 Mar. 1945; 
Venezuela, Anzo Stegul, Lys. Rio Zumbador, near base 
of Piedra Blanca; 800-1,000 m; VEN (holo). 

" Arachnothrix” reflexa ( (Benth, ) Planch. var. meridensis 

. New York Bot. Gard. 17(1): 253. 

ited: L. Bernardi 985; 21 Sep. 1953; 

Venezuela, jid Santo Domingo; 1,600 m; depos- 

itory not desi signa 

" Arachnothrix" er (Standl.) Steyerm. var. tachi- 
rensis Steyerm., Acta Bot. Venez. 6: 110. 1971. L. 
Cardenas de Guavara 805; 29 Dec. 1968; Venezuela, 

chira, Distr. Lobatera, Parque Rio Conejo-Borota; 


m3 
un 
= 
[e] 


“Arachnothrix” venezuelensis Steyerm., Mem. New York 
Bot. Gard. 17(1): 256. 1967. C. E. Chardon 179; 26 
Oct. 1940; Venezuela, Aragua, Carretera a Choroni; 
1,300 m; VEN (holo). 

Arcytophyllum nitidum (H.B.K.) Schlecht. subsp. ca- 
racasanum 
ving Acta p Venez. 6: 1 

4509; Sep. 1961; Venezuela, Mérida, ‘teleferico de 
Mérida; VEN. (holo). 

Arcytophyllum nitidum (H.B.K.) Schlecht. f. hispidum 
Steyerm., Acta Bot. Venez. 6: 117. 1971. J. A. Stey- 
ermark et al. 98558; 20-23 May 1967; Venezuela, 
Táchira, Páramo de Tamá; 2,750- 2,950 m; depository 
not designated. 

Arcytophyllum Cp uen Steyerm., de Bot. Venez. 6: 
113. 1971. Y. schi & K. Magdefrau 6850; 3 
Apr. 1958; Modus. Mérida, Pico Bolivar, Laguna 
Anteojos; 4,100 m; VEN (holo). 

Arcytophyllum Pg dae Steyerm., Acta Bot. Ven- 
ez. 6: 120. 1971. J. A. Steyermark & T. Koyama 
102360; 17 i 1969; Venezuela, Mérida, estacion 
La Aguada, between Merida and Pico Espejo; 3,475 
m; VEN (holo). 

Bertiera aequaliramosa Steyerm., Mem. New York Bot. 
Gard. 17(1): 319. P. Cooper 586; Jan 

r. 1928; Panama, Prov. Bocas del Toro, eet 
Buena Vista Camp on Chiriqui trail 380 m; NY (holo). 

Bertiera pe Tir le w York Bot. 
Gard. 17(1): 319. . H. S ae et al 55299; 1 
Sep. 1963; riii T Rivier, 4 km S Juliana 
Top, Wilhelmina Gebergte; 700 m; NY (holo). 

Bertiera guianensis Aublet var. glabriflora Steyerm., 
M New York Bot. Gard. 17(1): 318. 1967. R. 
Spruce 4109; 1855-1856; Perú, Prope Tarapoto; NY 
(holo). 

Bertiera guianensis Aublet var. leiophylla Steyerm., 

lem. New York Bot. Gard. 17(1): 318. 1967. H. 


Pittier 5178; Jan. 1912; Panamá, Prov. Chiriqui, vi- 
cinity of San Felix; 0-120 s NY (holo). 
ertiera guianensis Aublet subsp. pubiflora Steyerm., 
Mem. New York Bot. Gard. n 318. 1967. G. Klug 
2164; May-July 1931; Perú, Loreto, Rio Zubineta 
mouth, Florida, Rio Putpumayo; 180 m; NY (holo). 

Borojoa ee Steyerm., Bol. Soc. Venez. Ci. Nat. 26: 
47 A. Ducke 1141; 3 Oct. 1942; Brazil, 
eet po (Boca de Javary); NY, US er. 


ezuela, Miranda, Parque Naciónal de Guatopo; doo m; 
VEN (holo). 

Borojoa venez NC Steyerm., Bol. Soc. Venez. Ci. 
Nat. 26: 475. 1966. J. Garcia 128; 8 Aug. 1951; 
Venezuela, eats Rancho Grande; 1,130 m; VEN 
holo 


olo 
Borreria aristeguietaeana Steyerm., Bol. Soc. Venez. Ci. 
Nat. 29: 1971. E. Casano 46; 4 Sep. 1965; 
Venezuela, nica Estación Biologica de D Tis anos, 

Calabozo: VEN (holo 

Borreria bolivarensis Steyerm., Mem. New York Bot. 
Gard. 23: 819. 1972. J. A. Steyermark 87964; 12 
Dec. 1960; Tiai, Bolivar, Sierra Imataca, Río 
Toro (Río Grande); 200-250 m; VEN 

prieky capitata (Ruíz & Pavón) 

em. New York Bot. Gard. 23: 823. 
lia 4250; July 1960; Venezuela, ee 0, 
Sabanas, Estación Biologica de Los Llanos; VEN (holo). 

Borreria cataractarum Steyerm. i ork Bot. 
Gard. 23: 813. 1972. B. Maguire et al. 4612; 7 Sep. 
1961; Guyana, S Pakaraima Mountains, base of Chim- 
pau Falls; depository not designated. 

Borreria p i ient Steyerm., Mem. New York Bot 

28. 1972. J. J. Wurdack &L. = Adderley 
42701; 30 May 1959; Venezuela, T.F. Am s, Rio 
Orinoco 2 km below mouth of Rio Atabapo, "Saban ta 
Morocto; 125-150 m; NY (holo), VEN. 

Borreria intricata Steyerm., Mem. New York Bot. Gard. 
23: 828. 1972. J. J. Wurdack & L. S. Adderley 
42988; 12 June 1 959; Colombia, os tage Manacal 
on Rio Atabapo; 125-140 m; NY (holo), V 

lia Sah Steyerm., Fl. Venez. 9(3): 165. 1974. 

x 10157; 20 Apr. 1974; Venezuela, T.F. 

m E of Cano Parucito, E of Hato 
Yavi, N of Cano Massa, S of Cerro Corobita; 150 
m; VEN (holo). 

Borreria bis (Lam.) e var. edentata Steyerm., 
Acta Bot. Venez. 6: 196. 1971. H. Pittier 9829; 18 
Sep. 1921; Venezuela, ie Cotiza, around Caracas; 
800-1,000 m; NY (holo), VEN. 

Borreria macrocephala Standl. & Steyerm., Fieldiana 
Bot. 28(3): 566. 1953. Ll. Williams 13853; 19 Jan. 
1943; Venezuela, T.F. Amazonas, banks of Cano Rana, 
Rio Atabapo; 280 m; F (holo). 

Borreria oligodonta Steyerm., Mem. New York Bot. 
Gard. 23: 826. 1972. G. H. H. Tate 1147; Dec. 1937- 
Jan. 1938; Venezuela, Bolivar, Auyan-tepui; NY (holo). 

Borreria pygmaea Spruce var. robusta Steyerm., Mem. 
New York Bot. Gard. 93: 812. 1972. B. Maguire et 
al. 29242a; 24 Oct. 1950; Venezuela, T. mazonas, 
Puerto Ayacucho, E of Hotel Amazonas; 150 m; 

holo 


(MEIN dem Steyerm., Mem. New York Bot. Gard. 
23: 812. 1972. J. J. Wurdack & L. Adderley 42915; 
10 June Ts Venezuela, T.F. Amazonas, Rio Ata- 


704 


Annals of the 
Missouri Botanical Garden 


bapo, Yavita- Pimichin trail near Yavita; 125-140 m; 
NY (holo), VEN. 

Borreria PE Standl. & Steyerm., Field Mus 
Hist., Bot. Ser. 23(1): 21. 1943. P. C. Standley 7600: 
Nov. 1940; Guatemala, Dept du between ae 
and La Burrera; 800 m; F (holo 

Botryarrhena venezuelensis Steyerm., Ann. Missouri Bot. 
Gard. 70(1): 207. 1983. G. Davide " al. 17427; 8 
May 1979; Venezuela, T.F. Amazonas, Dept. Atabapo, 
SE a a n part of Cano Yagua at Cucurbital 
de Yagua; 1l ; MO (2769061), VEN (holo). 

odes vede M A d. a "gs Mus. Nat. 

t., Bot. Ser. 22(5): 3 940. J. 4. Steyermark 
31901; 18 Nov. 1939; een Dept. din Vol- 
cán Suchitán, NW of Asunción ~ ; 2,050 m; F (holo) 
= B. laevis Mart. & Gal., 

Bouvardia pachecoana Standl. & Ste eyerm., 
Nat. Hist., Bot. Ser. 23(1): 22. 1943. P. ts 
86226; Feb. 1941; Guatemala, Dept. 
Barranco a road between San a and 
San Rafael Pie de la Cuesta; 2,500-2,700 m; F (holo) 
= B. dictyoneura Standl., 1975. 

Bouvardia salvadorensis Steyerm., C pred 
J re 


Pain El $ 


Field Mus 


. 1955. 


Calycophyllum inns Pm. El M York 
Bot. 1963. S. S. Tillett & C. L. 
Tillett 415367: 30 S en 1960; Guyana, Karow- -tipu, 
upper cd river basin, Kako River; 950 m; N 
(holo), VE 

oe iae pu Mem. New York 
Bot. Gard. 23(1): 2 2. S. S. Tillett et al. 44837; 

uly 1960; in je River, SE ridge of 
Merume Mt.; 1,140 m; NY (holo). 

Calycophyllum spectabile Steyerm., New York 
Bot. Gard. 10(5): 188. 1963. B. pa & D. B. 
Fanshawe 32219; 23 Oct. 1951; Guyana, between 
es ee and Mt. Yamanaik, upper Maza- 
runi ; NY (holo), VEN. 

Calycophyllum v venez ca ense ure Mem. New York 

uw 1963. J. A. Steyermark & L. 

962; naa, Bolívar, Ce- 


w York Bot. 
JJW ick e L. S. Ad- 
derley 43383; 10 July 1959: Venezuela T.F. Ama- 
zonas, near left ban no Duquiapo, Río Cassi- 
quiare, 2 km above Salimos “100 m; NY (holo), VEN. 
oe a 'arensis Standl. & Steyerm., ea Bot. 
28(3): 5 A. me yermark 6018. a Nov. 
944; ‘eS Bol: Sororopan-te ui 
2,250 m; F (holo) = Psychotria bolivarensis cene 
& Steyerm.) Steyerm 
Cephaelis cacuminis Td P Acta Bot. Venez. 2: 335. 
1967. J. A. cl 93564; 7 May 1 
ezuela, Bolivar, Auyan-tepui; 1,800 m; VEN (holo) = 
Psychotria cacuminis (Steyerm.) Steyerm., 
Cephaelis deis A Mem. New York Bot. 
Gard. 17(1): 428 . B. Maguire et al. 53562; 3 
= ep. 1962; de E along Río Franela, Ce- 
o Piton, Cordillera Epicara, Alto Rio Cuyuni, Rio Chi 
canan; 400 m; VEN (holo) = Psychotria sandwithiana 
(Steyerm.) Steyerm., 1972. 
Cephaelis tepuiensis Steyerm., 


Gard. 17(1): 430. 1967. B. 


Mem. New York Bot 
Maguire et al. 46864: 


20-21 Aug. 1962; Venezuela, Bolivar, La Escalera, 
Rio Uiriyuk, Alto Rio Cuyuni; 1,000 m; MO (2389390), 
VEN (holo) = Psychotria tepuiensis (Steyerm.) Stey- 
erm 72 

Cephaelis tinctoria Standl. & Steyerm., Fieldiana Bot. 
28(3): 568 53. J. A. Steyermark 55256; 4 Feb. 
1944; Venezuela, Lara, Palojosco above Los Aposentos, 
above Humocaro Bajo; 2,530-2,680 m; F (holo) = 
Psychotria eciliata Steyerm., nom. nov., 

Cephalodendron AR Mem. New York Bot. Gard. 

en. nov. 

Cephalodendron aracamuniensis Steyerm., Ann. Mis- 
souri Bot. Gard.: in press. 1989. R. L. Liesner & F. 
Delascio 22064; 17 Oct. 1987; Venezuela, T.F. Ama- 
zonas, Dept. Rio Negro, Cerro Aracamuni, summit, 
Popa Camp; 1,550 m; MO (holo), VEN. 

Wi er nier globosum Steyerm., Mem. New York 
Bot. Gard. 23(1): 228. 1972. B. Maguire et al. 42467; 
24 Dec. 1957; Venezuela, T.F. Amazonas, Cerro de 
la Neblina; 1,100 m; NY (holo), VEN. 

Chalepophyllum guianense Hook. f. var. cuneatum 
Steyerm., Mem. New York Bot. Gard. 10(5): 193. 
1963. B. Maguire 33700; 30 Mar. 1952; Venezuela, 
Bolivar, near Kavanayen; 1,300 m; NY (holo 

Chalepophyllum ocium Steyerm., Mem. New York 
Bot. Gard. 10(5): 193. 1963. B. Maguire et al. 40637; 
7-8 Feb. 1955; Guyana, between Ayanganna and Chi- 
nowieng, Mt. Yi Pakaraima Mountains; 100- 
1,200 m; NY (holo) 

Chalepophyllum nao Standl. & Steyerm., devis 

568. 1953. Ll. Williams 12970; 18 May 
1940; Venezuela, T» Amazonas, Puerto one 
100 m; F (holo) = Acanthella sprucei pont f., 1973. 


Chimarrhis bathysoides Steyerm., Mem w York Bot. 
Gard. 12(3): 181. 1965. B. Maguire y L. Politi 
: 49; Venezuela, mazonas, 


Cerro Sipapo, near Base Camp; 125 m; NY (holo). 

Chimarrhis cubensis Steyerm., jii m. New York Bot. 
Gard. 12(3): L ov. 

Chimarrhis decur "Ceiba 3: 18. 1952. P. 
H. Allen 3566; 28 ne 1946, Pa anama, Prov. Coclé, 

{ = Allenanthus eryth- 
rocarpus Standl., 

Chimarrhis microcarpa Standl. var. speciosa Steyerm., 
Mem i y York Bot. Gard. 12(3): 183. 1965. T. 
Gar etg 47; 18 Sep. 1951; Venezuela, Aragua, Ran- 
cho Lalas VEN (holo). 

Camaras O Steyerm., Acta Bot. Venez. 8: 

A. inmane et al. 105658; 25 Mar. 

197 2; Ide Zulia, Sierra Perija, Río Omira-Kuna 

(Tumuriasa); 1,470-1,560 m; MO (2250725), VEN 

(holo). Note: No. 105655 is a paratype, collected at 
the same locality. 

Chimarrhis venezuelensis Standl. & Steyerm., Pure. 
Bot. 28(3): 570. 1953 A. Steyermark 55408; 8 
Feb. 1944; Venezuela, Lara, mountain en Santo 
Domingo and Los Bigs S of Las Sabanetas; 
2,430-2,475 m; F (holo) = Dioic oe dioicum 
(K. Schum. & K. Krause) Steyern 4. 

Chiococca alba (L. : A. Hitchc. subsp. din ifolia (W ullsch. 
ex Griseb.) Steyerm. var. micrantha (Johnston) ina 
erm. f. pilosa oa. Acta Bot. Venez. 6: 141 


Volume 76, Number 3 
1989 


Taylor 705 
Plants Described by 


Julian A. Steyermark 


1971. J. Saer 4; 1923; Venezuela, Lara, cercanias de 
Barquisimento; VEN (holo). 

Chiococca auyantepuiensis Steyerm., Mem. New York 
Bot. Gard. 23(1): 374. 1972. J. A. gc eR: 93652; 
10 May 1964; Venezuela, Bolivar, Auyan-tepui; 1,800 
m; VEN (holo). 

Chiococca lucens Standl. & Steyerm., Fieldiana Bot. 
28(3): 571. 1953. J. A. Steyermark 60184; 15 Nov. 
1944; bri ee Bolivar, Sororopan-tepui; 2,130- 
2,250 m; F (ho 

Chiococca fos NM Steyerm., Mem. New York 
Bot. Gard. 23(1): 373. 1972. B. A. Whitton 120; 20 
Aug. 1959; Guyana, Potaro Distr., Mt. Kanaima; 485 
m; NY (holo). 

Chiococca eia con Steyerm., Ll Bot. Venez. 6: 
135. 1971. J. A. Parade ada 75; 8 Oct. 1900; 
Ven ata Distr. Federal, Cerro een 1,400 m 
MO (2270191), die ded ob ). 

Chiococca nitida nth. r. chimantensis Steyerm., 
Mem. New York E Gard. 2301): 376. 1972. J. A. 
al alah 75406; 15 May 1953; Venezuela, Bolivar, 

anta Massif, Chimanta-tepui (Torono-tepui); MO 
(2012043, 2011333), NY, US, VEN (holo). 

Chiococca semipilosa Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 22(4): 278. 1940. J. A. Steyer- 
mark 314 06; 7 Nov. 1939; ca Dept. Chi- 
quimula, Caracol Mt., N of Quezaltepeque; 
1,200-1,400 m; F helo: 

Chione guatemalensis Standl. & Steyerm., Field Mus 

at. Hist., Bot. Ser. ee. 


Chiriqui, Cerro Horqueta, Boquete region; 2,167 m; F 
(holo), MO. 

Chomelia delascioi Steyerm., Mem. Soc. Ci. Nat. La 
Salle 40(113): 83. 1980. F. Delascio et al. 8471; 
Aug. 1979; Venezuela, Bolivar, Distr. Roscio, San Mar- 

e de Tumeremo, Rio Cuyuni; 
VEN (holo), LA SALLE 

Chomelia gte x Steyerm., Ann. Missouri Bot. Gard 
. G. Morillo Ere May 1982; Vene- 

Ro Caura, Ks m S of Las Pavas 


is 657. 1 


, Bolivar 
. New York Bot. 


Feb. 1968; French Guiana, Grand Tamouri, Riviere 
Camopi; P (holo) 
Chomelia monachinoi Steyerm., York Bot 
33 


. New 
Gard. 17(1): 336. 1967. J. J. B & J. Mone 


, Fl. Venez. 9(2): 854. 1974. 
a & R. Monte: 5254; 22 May 1973; Ven- 
ezuela, "Ee cercanias a Mantecal, alto Apure; 80 
m; VEN (holo). 
Chom elia schomburgkii a eum Mem. New York Bot. 
: 339. 1967. R. Schomburgk 924; 1841; 


74: 105. 1987. B. Stergios & G. iiber 2804; 20 


puesto 
Anacoco; Pari VEN (holo) 


Chomelia venezuelensis Steyerm., Mem. New York Bot. 


Gard. 17(1): 336. 1967. E. Rohl 103; 1939; Vene- 
zuela, Distr. Federal, Los Castillitos, near Caracas; VEN 
(holo). 

Chondrococcus Steyerm., Mem. New York Bot. Gard. 
23(1) 403. 1972, gen. nov. = Coccochondra Rau- 
schert, 1982. 

Chondrococcus laevis Steyerm., Mem. New York Bot. 
Gard. 23(1): 405. 1972. R. S. Cowan & J. J. Wurdack 
31130; 2 Feb. 1951; Venezuela, T.F. Amazonas, Rio 
Ventuari, Caño Asisa, Rio Parú, Serranía Parú; 2,000 
m; NY (holo), VEN = Coccochondra laevis (Steyerm.) 
Rauschert, 1982. 

Cinchona henleana Karst. var. trichophylla Steyerm., 


Acta Bot. Venez. 8: 247. 1973. J. A. & C. Steyermark 


Gian, E Los Tanques, S Borburata; 750- 1,100 m; VEN 
(holo). 

C nm rim Steyerm., Ann. Missouri Bot. 
Gard. 398. 1987. G. Davidse & A. C. González 
15825; 27 Feb. 1979; Venezuela, Apure, Dept. Pedro 
Camejo, Cano El Cabello, between the Rio n and 
Rio Cinaruco; 75 m; (holo— 2746721), V 

Coccocypselum brevipetiolatum Steyerm., 

k Bot. Gard. 17(1): 304. 1967. F. W. Pannal 
1676; 4 Sep. 1917; Colombia, Cundinamarca, Bue- 
navista to Pipiral, SE of Quetame; 1,000-1,200 m; 
NY (holo). 

Coccocypselum croatii Steyerm., Ann. Missouri Bot. Gard. 
74: 106. 1987. T. B. Croat 54112; 25 July 1982; 
Venezuela, Bolivar, vicinity of Icabarú; 600 m; 
(holo — 3327537). 

Coccocypselum guianense (Aublet) Schumann var. pa- 
tens Steyerm dps A York Bot. Gard. 1 7(1): 303. 

967. T. Lasser 25 Apr. 1946; Venezuela, 
Bolivar, Alto udis VEN (holo). 

C LOW Cra ved teyerm., Ann — Bot. Gard. 
74: 657. . O. Huber 9123; 1 Mar. 1984; Ven- 
ezuela, Dd Distr. Roscio, an = Caco, 25 
km NW of San Ignacio de Yuruani 2.5 km ESE of 
Wanarü; 1,150-1,200 m; MO (3321365), VEN (holo). 

Coccocypselum trinitense Steyerm., Mem. New Yor 
Bot. aa 17(1): 300. 1967. J. A. Steyermark 90919; 
3 July 1962; Trinidad, Aripa Savanna, Long Stretch 
Forest rine EN (holo). 

eren um New York Bot. Gard. 


Coussarea amapaensis CAM Mem. New York Bot. 
Gard. 17(1): 366. 1967. J. Murca Pires et al. 50285; 
31 July 1961; Brazil, Amapa, Serra do Navio, on Rio 
Amapari; NY (holo 

Coussarea bernardii Sidi. Mem. New York Bot. 
Gard. 17(1): 371. 1967. L. Bernardi 2325; June 
1955; Venezuela, Mérida, Pueblos del Sur; 1,600- 
2,200 m; NY (holo). 

cancel pice nsis Steyerm., Ceiba 3: 20. 1952. P. 

576; 9-11 June 1947; Panamá, Prov. Da 
rién, ae Rio Chich, vicinity of Yaviza; 33 m; F (holo), 
MO 


Coussaria evoluta Steyerm., Ann. Missouri Bot. Gard. 
15: 334. 1988. R. L. Liesner & D. Bell 17493; 4 


Dec. 1984; Venezuela, T.F. Amazonas, Cerro de la 


Neblina; 140-340 m; MO (holo— 3524658), VEN. 
Coussarea a piel pe ur Mem. New York Bot. 
Gard. 17(1): 365. . B. Maguire & D. B. Fan- 


shawe 23072; 28 D 1944; Guyana, Potaro River 
Gorge; NY (holo). 


706 


Annals of the 
Missouri Botanical Garden 


Coussarea hallei Steyerm., Mem. New York Bot. Gard. 
23(1) 387. 1972. F. Halle 1018; 30 Jan. 1965; 
French Guiana, Route du lac des Americains; P (holo). 

Coussarea hirticalyx Standl. var. glabrior Steyerm., Mem. 

ork Bot. Gard. 17(1): 363. 1967. R. E. Schultes 

& c. 4 Black 8156; 11-18 Sep. 1946; Colombia, 

Amazonas, near Sergipe, Putumayo (Ica) River; 100 
m; US (holo). 

oce klugii Steyerm., Mem. New York Bot. Gard. 

17(1): 363. 1967. G. Klug 2362; Oct.-Nov. 1931; 


Peru, Loreto, e of Rio Zubineta Rio Putumayo, 


Florida; 180 m; F (holo). 

Modi "rd Steyerm., Mem. New York Bot. Gard. 
17(1) 3 1967. T. ur 1682; 27 May 1946; 
Venezue ii Kl Rio Uairen; VEN (holo). 


n 

Coussarea liesneri Steyerm., Ann. Missouri Bot. Gard.: 
in press. 1989, R. L. Liesner & F. Delascio 22262; 
21 Oct. 1987; Venezuela, T.F. Amazonas, Dept. Rio 
Negro, dub Aracamuni, Quebrada Camp; 600 m; 
MO, VEN (holo). 

Coussarea mediocris Standl. & Re Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 248. . A. i ise 
48732; 15 July 1942; EE Dept. Huehue 
nango, id Maxbal, Sierra de los ee 
1,500 m; F (holo). 

Coussarea pittieri Steyerm., Bol. Soc. Ven i. 
25(106): 70. 1963. J. A. Steyermark ae 5 Pu 
1963; Venezuela, Áragua, Parque Nacional Pittier, halt. 
way up Fila de Paraiso; 1,400 m; (holo). 

Coussarea revoluta Steyerm., Mem. New 
17(1): 369. 1967. 1 
43078; 20 June 1959; Venezuela, T.F. Ama 
Orinoco, just below mouth of Río Cunucunuma; 
m; NY (holo 

Coussarea terepaimensis Steyerm., Acta Bot. Venez. 8: 

Smith V-6649; 14 Apr. 1971; 
Venezuela, Lata, i Fila, 6 km SE de Terepairma; 


VEN (holo). 
Cuatrecasasiodendron Standl. & Steyerm., Acta Bot. 
Venez. 4(1): 29. 1964, gen. nov. 
Cuatrecasasiodendron oun Standl. & Stey- 
erm., Acta Bot. Venez. 4(1): 3 964. J. Cuatrecasas 
13694; 19 Dec. 1942; E Dept. del Valle, 
Cordillera Occidental, Hoya del Rio Anchicaya, Que- 
brada del Retiro; 300 m; F (holo). 
Cuatrecasasiodendron spectabile Steyerm., -o D 
. . Cuatrecasas 1716 
30 Apr. 1944; Colombia, Dept. del Valle, Co p" 
Pacifico, Rio Cajambre, Barco; 5-80 m; US (holo), 


Declieuxia dasyphylla Schumann ex Steyerm. f. ciliata 
Steyerm., Lo Contr. Sci. 21: 26. 


eye Angeles Co. Mug 
1958. E. Y. Dawson 14163a; 13 Apr. 1956; Brazil, 
Goiás, Chapada dos ACIE, F, R (holo) = D. das- 


yphylla Schumann ex Steyerm. var. dasyphylla, 1976. 
d irr" Steyerm., Mem. New York Bot. 
Gard. 10(5): 196. 1963. B. Maguire et al. 37592; 7 
Feb. 1954; Till) T.F. Amazonas, right bank of 
Rio Pacimoni; 100-140 m; NY (holo 
Dendrosipanea wurdackii Steyerm., Mem. New York 
Bot. Gard. 10(5): 198. 1963. B. Maguire & J. J. 
“urdack 3558; 14 Apr. 1953; Venezuela, T.F. Ama- 
zonas, Sabana El Venado on left bank of Caro Pimichin, 
above Pimichin, Río Guainia; 140 m; NY (holo). 
eppea amaranthina Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 22(4): 281. 1940. J. A. Steyer- 
mark 30926; 1 Nov. 1939; Guatemala, Dept. Chi- 


air Cerro Brujo, vicinity of Rio Negro, below 


Montaña Montenegro, near Brujo; 1,500-2,000 m; F 
(holo). 

Deppea E Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 22(5): 385. 1940. J. A UE 


Diodia d iium Los Angeles Co. Mus. Contr. 
Sci. 21: 26. 1958. E. Y. Dawson 14672; 24 Apr. 
1956; Baa Coiás, Chapada dos Veadeiros, 14 km S 
of Veadeiros; F, LAM, R (holo), U 

Diodia hyssopifolia ( (Willd.) Cham. & Schl. var. leio- 
carpa Steyerm., Mem. New York Bot. Gard. 23: 797. 
1972, J. 4. Steyermark & C. K. Maguire 60125; 7 
Nov. 1905; dins Amazonas, de Negro, along Rio 
Cauaburi; deposi qe not designa 

Diodia hysopifolia. (W illd.) Cham & Schlecht. var. 
linearis (Willd.) Steyerm. f. glabriuscula 2 - 

em. New York Bot. Gard. 23: 796. 1972. H. H. 

. 1904; Guyana, Bu River, 
Tinamu Falls; NY (holo). 

Diodia hyssopifolia ( (wil : Cham. & Schlecht. f. psilo- 
clada Steyerm., Mem 


e ; olo). 
Diodia radula (Willd. & Hoffmanns.) Cham. & Schlecht. 
subsp. Vp d dw Steyerm., Acta Bot. Venez. 6: 
181. 1971. F. J. Breteler 4095; 12 Aug. 1964; Ven- 
ezuela, Trujillo 22 km from Boconó along road to 
Trujillo; 1,900 m; VEN (holo). 

Diodia teres Walter var. dos ponis (A. Gray) Vd 
f. latior Steyerm., Acta Bot. Venez. 6: 184. » H. 
H. Smith 2628; 1.898- 1/899 m; abia Du 
Magdalena, Santa Marta; NY (holo). 

Diodia teres Walter subsp. prostrata bab Steyerm. 
f. latifolia Steyerm., Acta Bot. Venez. 6: 186. 1971 

. Blydenstein 250; 16 le 1961; la Guár- 
ico, Alto Llano, Hato El Carmen, Estación Biologica de 
Los Llanos; VEN (holo). 

Diodia teres Walter subsp. prostrata Gadde Steyerm. 
f. leiocarpa Steyerm., Acta Bot. Venez. 6: 187. 1971 
F. J. Breteler 4204; 4 Sep. 1964; Venezuela Barinas, 
l4 km from Barinas along road to Guanare; 200 m; 
VEN (holo). 

Dioicodendron iod 

4. 1 v. 


Bol. Soc. Venez. Ci. Nat. 25: 


Dioicodendron cuatrecasasii Steyerm., Bol. Soc. Venez. 
Ci. Nat e 21. J. Cuatrecasas 15192; 19- 
28 Aug. 1943; Colombia, Dept. del Valle, Cordillera 
Prud Hoja del Rio Digua, Piedra de Moler; 900- 
1,180 m; F, US, VEN (holo) 

Duroia amapana Steyerm., Mem. New York Bot. Gard. 
12(3) 209. 1965. J. Murca Pires et al. 51601; 9 
Oct. 1961; Brazil, Terr. Amapa, Camp 13, Rio Ara- 
guari; NY (holo). 

Duroia bolivarensis Steyerm., Mem. New York Bot. Gard. 
12(3): 206. 1965. F. Cardona 819; 2-15 Aug. 1943; 
Venezuela, Bolivar, Alto Rio Paragua, orillas del Rio 
Tonoro; F, VEN (holo) 

Duroia eriopila L.f. var. brevidentata Steyerm., Mem. 

k Bot. Gard. 12(3): 203. 1965. A. Ducke 
1332; 31 Aug. 1943; Brazil, Rio Branco, Serra Grande; 


NY (holo). 
Duroia eriopila L.f. f. glabra Steyerm., Mem. New York 


Volume 76, Number 3 
1989 


Taylor 707 
Plants Described by 


Julian A. Steyermark 


Bot. Gard. 12(3): 203. 1965. 4. C. Smith 2792; 15- 
24 Dec. 1937; Guyana, near mouth of Onoro Creek, 
basin of Essequibo River; NY (holo). 

Duroia eriopila L.f. var. tafelbergensis Steyerm., Mem. 
New York Bot. Gard. 12(3): 203. 1965. B. Maguire 
24393; 16 Aug. 1944; Suriname, Tafelberg, between 
savannas 4 and 5; 535 m; NY (holo). 

Duroia Qna iae Standl. & Steyerm., Field Mus. Nat 
Hist., Bot. 22(3): 186. 1940. J. A. Steyermark 
39382; 14 jud 1940; Guatemala, Dept. Izabal, Rio 
Dulce, near Livingston; sea level; F (holo). 

Duroia gransabanensis Steyerm., Mem. New York Bot. 
Gard. 12(3): 205. 1965. F. Tamayo 3133; 15 Mar. 
1946; Venezuela, Bolivar, Gran Sabana, Uari; VEN 


(holo). 

Duroia ias Steyerm., Brittonia 33: 385-400. 
G. nce et al. 7606; 27 Sep. 1968; Brazil, Acre, 
Mun. Sena Madueira, 4 km E of, Saá Madueira; NY, 
VEN (holo). 

Duroia kotchubaeoides Steyerm., Mem. New York Bot. 
Gard. 12(3): 201. 1965. J. Silverio Level 147; 12 
Sep. 1954; Venezuela, T.F. Amazonas, Rio Orinoco, 
Caño Tama-Tama; 150 m; NY (holo). 

Duroia maguirei Steyerm., Mem. New York Bot. Gard. 
23(1): 343. 1972. B. Maguire et al. 60071; 4 Nov. 
1965; Brazil, Amazonas, Río Cauaburi, base of Cacho- 
eria Caranguejo; NY (holo), VEN. 

Duroia maguirei Steyerm. var. patentinervia Steyerm., 

m. New York Bot. Card. 23(1): 345. 1972. J. A. 
Steyermark $: G. S. Bunting 102939; 22 Apr. 1970; 

Hiis uela, . Amazonas, 7-9 km from Yavita on 

oad to Pimichin: NY, VEN (holo). 
roia merumensis Steyerm., Mem. New York Bot. Gard. 
12(3): 207. 1965. S. S. Tillett et al. 44835; 6 July 
1960; Guyana, SE ridge of Merume Mt., upper Ma- 
zaruni river basin, Partang River, Merume Mountains; 
1,140 m; NY (holo 

Duroia nitida Steyerm., Mem. New York Bot. Gard. 
12(3): 208. 1965. A. Ducke 799; 28 Sep. 1941; 
Brazil, Amazonas, Cachoeira Iracema, Rio Irubu; NY, 
US (holo). 

Duroia paruensis Steyerm., Mem. New York Bot. Gard. 
12(3): 206. 1965. R. Cowan & J. J. Wurdack 31447; 
13 Feb. 1951; Venezuela, T.F. Amazonas, Serranía 
Parú; NY (holo). 

Duroia prancei Steyerm., Mem. New York Bot. Gard. 
23(1): 345. 1972. G. T. Prance et al. 59234; 2 Oct. 
1964; Brazil, Mato Grosso, 3 km from Garapu; NY 
(holo). 

Duroia retrorsipila Steyerm., Mem. New York Bot. Gard. 
l 208. 1965. J. A. Steyermark & S. Nilsson 
382; 20-22 Apr. 1960; Venezuela, Bolivar, vicinity 
Cerro Uei, vicinity roa campañent o 125, between 


Luepa & Cerro Venamo; 1, ee m; VEN (holo). 
ies e Steyerm., Mer Ne w York Bot. Gard. 
12(3): 204. 1965. B. nar et al. 53676; 9-11 


Sep. 1962 Venezuela, Bolivar, Rio Chicanan, Cordi- 
llera Epicara, Cerro Piton; 400 m; VEN (holo 
Elaeagia alterniramosa Steyerm., Acta Bot. Vaisi 4(1): 
39. 1964. J. Cuatrecasas 23870, 27 Mar. 1947; 
Colombia, Dept. del Valle, ao Occidental, Hoya 
del Río Digua, Rio San Jua do E ), VEN 
y ] w York Bot. Gard. 
. J. M. Su 1559; n.d.; Colombia, 
Dept. del Valle, Cordillera Occidental, Cali; 1,800 m; 
US (holo). 


Elaeagia cuatrecasasii Steyerm., Acta Bot. Venez. 4(1): 


8. 1964. J. Cuatrecasas 21682; 24 July 1946; Co- 
lombia, Dept. del Valle, Cordillera Occidental, Hoya del 
Rio Cali; 2,500 m; US (holo), V 

Elaeagia ies e Steyerm., Bol. Soc. Venez. Ci. 

. 1960. J. A. Steyermark 54305; 10 
Sep. 1943; Ecuador, Prov. Santiago-Zamora, on trail 
between Pailas and El Pan; 30-50 ft.; F (holo) 

Elaeagia a Standl. & Steyerm., Fieldiana Bot. 
28(3): 574. 1953. J. A. Steyermark 61373; 5 Mar. 
1945; Veuesuele, Anzoátegui, NE of Bergatin, along 
headwaters of tributary to Rio Neveri; 800-1,200 m; 
F (holo). 

Elaeagia up a Steyerm., Bol. Soc. Venez. Ci. Nat. 
96: 240. qe L. Aristeguieta 1989; Aug. 1953; 
Venezue D ua, Parque Nacional Henri Pittier, 
Rancho Grande: VEN (holo). 

Elaeagia maguirei Standl. var. pubens Steyerm., Mem. 
New York Bot. Gard. 12(3): 190. 1965. J. A. Stey- 

L. Aristeguieta 102; 6-11 Mar. 1962; 

Venezuela, Bolivar, NE of Luepa, vicinity of km 143 

Dorado, Rio Aponguao drainage; 800-1,200 
m; VEN (holo). 

Elaeagia microcarpa Steyerm., Mem. New York Bot. 
Gard. 12(3): 189. 1955. B. A. Krukoff 10650; 27- 
31 July 1939; Bolivia, Dept. La Paz, Prov. 5 Yungas, 
Asunta (near Evenay), Basin of Rio Bopi; NY, US (holo). 


Colombia, Comisario del Putumaya; 2,200-2,400 m; 
US (holo). 

Elaeagia ruiz-teranii uis Pittieria 4: 15. 1972. 
L. Ruíz Terán & M. Lopez Figueiras 441; n.d.; 
Venezuela, Mérida, Los E s Distr. Sucre, Mun. Es 


1960; Venezuela, Bolivar, between Luepa and 
m; NY (holo), VEN. Note: Em- 
e correct pics À 

umbellata (Spreng.) Schumann LT 


& M. Saldiva 555; 20 Mar. 1951; Venezuela, Aragua 


Trayecto Guamitas-La Mesa; VEN (holo 
rines anisocalyx Poepp. var. . schwackei 
(Schumann & Krause) Steyerm. f dentata Steyerm., 
em. New ork Bot. Gard : 375. T. O. 


San Carlos bei abb Me 600 ; (holo). 
Faramea berryi Steyerm., Pittieria 9: 5. 1981. P. E. 

Berry L. Chesney 2096; 26 Feb.-2 Mar. 1976; 

Venezuela, T.F. Amazonas, Estación Exp. de Sta. Bar- 


bara del Orinoco, 1-2 km 

m; MO, VEN (holo). 

Faramea boomii Steyerm., Ann. Missouri Bot. Gard. 75: 
335. 1988. B. Boom & A. Weitzman 5278; 4 Dec. 
1984; Venezuela, : F. Amazonas, Cerro de la Neblina, 
Base Camp; ; AAU, R, COL, GH, INPA, 

K, MG, MO (holo 3393678), NY, P, PORT, U, US, 
N. 


al Sur de Trapichote; 130 


Faramea ier Steyerm., Mem. New York Bot. Gard. 
17(1): 381. 1967. Forest Dept. Field No. Fanshawe 
877; 10 July 1942; Guyana, Keriti Creek; K (holo). 


Faramea cardonae Standl. & Steyerm., Fieldiana Bot. 


708 


Annals of the 
Missouri Botanical Garden 


28(3): 574. 1953. F. Cardona P. 1115; 10 July 1944; 
Venezuela, Bolivar, Cerro Guaiquinima, Alto Rio Para- 
gua; 1,200 m; F (holo), US. 

Baras amea anulare ensis Steyerm., Ann. Missouri Bot. Gar 
1987. H. van der Werff & R. Ortiz 5457; 

y 1983; Venezuela, Táchira, Dept. Lobertera, 
La uide 2,000 m; MO (holo), VEN. 

Faramea costata Steyerm. ., Mem. New York Bot. Gard. 
17(1): Pu 1967. J. & P. A. Florschutz 1180; 5 
Fe 51; Suriname, Saramacca River, upstream of 
Asus Fall: N olo 

gs iei c pres Steyerm., Mem. New York Bot. Gard. 
17(1): 380. 1967. B. Maguire & D. B. Fanshawe 
32329; os Oct. 1951; Guyana, Kamarang Station, 
upper TANT River; 500 m; NY (holo). 

., Bol. Soc. Venez. Ci. Nat. 

63. J. Garcia 80; 21 June 1951; 

Venezuela, Aragua, Rancho Grande; 1,100 m; VEN 

(holo). 


Faramea ip Steyerm., Mem. New York Bot. Gard. 
17(1): 3 967. Forest Dept. Field No. G569; 9 
Nov. T (send Sierra Irikoume, Acari Mountains; 
NY (holo). 

Faramea ENE Steyerm., Mem. New York Bot. Gard. 
17(1): 3 1967. B. Trujillo 2763; 22 Au 
Ven Led. Tus Morterito, Estación Experimental de 
Sanare; 1,600 m; MY (holo). 
aramea lourteigiana Steyerm., 
1972, nom. no 

Faramea maguire Steyerm, Mem. New York Bot. Gard. 
17(1): 3 . B. Maguire & D. B. eie end 
32357; oe es z Nov. 1951; Guyana, Membar 
Kurupang trail; 1,000 m; NY (holo). 

Faramea morilloi Steyerm., Ann. Missouri Bot. Gard. 
75: 337. 1988. G. Morillo 6716; 17 Nov. 1977; 
Venezuela, T.F. Amazonas, Parcela Fénologica, 16 km 
e Cruce con la carretera a Puerta Ayacucho-Sanar- 
iapo; 100 m; VEN (holo 

Faramea multiflora A. Rich. var. epedunculata Stey- 

New York Bot. Gard. Sa 392. 1967. 

, Zulia, Selvas de 


Bradea 1(16): 147. 


Faramea neblinae Steyerm. m. New York Bot. Gard. 
17(1): 395. 1967. B. OMEN et al. 42515; 25 Dec. 
1957; Venezuela, T.F. Amazonas, Cerro de la Neblina, 
Cano Grande; 1,050-1,100 m; MO (1811401), NY 
holo). 

Faramea occidentalis (L.) A. Rich. var. brachycalyx 
d m., Mem. New York Bot. Gard. 17(1): 385. 

. H. Curran s.n.; Apr.-May 1916; Colombia, 
Sm Island of Mompos, gera of Loba; US (holo). 

Faramea occidentalis (L.) A. Rich. subsp. lonchocalyx 
teyerm., Mem. New York Bot. Gard. 17(1): 385 
1967. O. Haught 3068; 6 Feb. 1940; Ecuador, Ma- 
nabi, near Guale; 250 m; US (holo). 

Faramea races (L.) A. Rich. var. eeu 
Steyerm., m. w York Bot. Gard. 17(1): 
1967. H. Pittier 12192; 16 Aug. 1926; ede 
Carabobo, Canaobito near Urama; VEN (holo). 

Faramea pres ola Steyerm., Ann. Missouri Bot. Gard. 
75: 338. 1988. R. L. Liesner 16960; 26 Mar. 1984; 
Venezuela, TE ie azonas, Dept. io Negro, upper 

N. 

rollina ctea Steyerm., “Mam: New York Bot. 

Gard. 17 70) 1967. J. Silverio Level 145; 12 

Sep. 1954; Venezuela, Amazonas, Cano Tama-Tama, 


Rio Orinoco; 150 m; NY (holo). 


Faramea pseudospathacea Steyerm., Mem. New York 
Bot. Gard. 17(1): 383. 1967. L. Schlim 531; May 
1846-1852; Colombia, Prov. Ocana, Pericos; 1,210 
m; F (holo). 

Faraon silvae Steyerm., Mem. New York Bot. Gard. 

23(1): 389. 1972. N. T. Silva & U. Brazao 60700; 
31 Dec. 1965; Brazil, dd Serra de Neblina; 
400-600 m; NY (holo), VEN. 


Faramea yavitensis Steyerm 


Ann. Missouri Bot. Gard. 


Faramea yutajensis Steyerm., A 
75: 339. 1988. R. L. Liesner & B. K. Holst 21640; 
4 Mar. 1987; Venezuela, T.F. Amazonas, 8 km NW 
of Yutajé, W of Serranía de Yutajé; 1,500-1,760 m; 
MO (holo — 3482984), VEN. 

Ferdinandusa boomii Steyerm., Ann. Missouri Bot. Gard. 
75: 339. 1988. B. Boom & A. Weitzman 5274; 8 
Dec. 1984; Venezuela, T.F. Amazonas, Cerro de la 
Neblina, Base Camp; BR, COL, GH, INPA, K, MO 
(holo — 3393679), NY, PORT, U, VEN. 

Ferdinandusa duckei Steyerm., Mem. New York Bot. 

23(1) 284. 1972. A. Ducke 1167; 19 Jan. 
1943; Brazil, Amazonas, Rio Taruma-miry, Manaos; 
NY (holo), US. 

Ferdinandusa goudotiana Schumann var. eciliata Stey- 

erm., Mem. New York Bot. Gard. 23(1): 281. 1972. 
. Schomburgk 955; n.d.; Venezuela, Bolivar, Rorai- 
ma; K (holo 

Ferdinandusa sia Mauer var. psilocarpa 
teyerm., Mem. New . Gard. 23(1): 281. 

1972. S. S. Tillett et al. 45184; T Aug 1960; Guyana, 
Mt. Ayanganna; 900-1,000 m; NY (holo). 

Ferdinandusa neblinensis Steyerm., Mem. New York 
Bot. Gard. 23(1): 283. 1972. B. Maguire et al. 36800; 
22 Dec. 1953; Venezuela, T.F. Amazonas, Rio Yatua, 
Cerro de la Neblina; 200 m; NY (holo), VEN. 

Ferdinandusa schultesii ug Mem. New York Bot. 
Gard. 23(1): 279. 1972. R. E. Schultes & I. Cabrera 
14475; 28 Sep. 1951; * Colombia. Amazonas- Vaupés 
Río Apaporis, Río Kananan, Cerro Isibukuri; US (holo). 

Galium canescens H.B.K. f. alstonii Steyerm., Acta Bot. 
Venez. 8: 253. 1973. 4. H. G. Alston 6533; 23 Jan. 
1939; Venezuela, Mérida, between Timotes & Chacho- 
po; 2,150-2,400 m; NY (holo). 

Genipa americana L. var. caruto (H.B.K.) K. Schum. 
f. E Steyerm. Mem. New York Bot. Gard. 
23(1): 350. 1972. P. Jorgensen 3694; July 1931; 
Paraguay, Villarrica; NY (holo 

Genipa ERA Steyerm., Mem. New York Bot. Gard. 
23(1): 353. 1972. J. Murca Pires & P. B. Cavalcante 
mers Bio Aug. 1962; Brazil, Amapá, Terr. Amapa, 

o Cujubim; NY (holo). 
Geophila hid due Miq. var. rape Steyerm., Mem 
Bot. Gard. 23(1): 393. 1972. E. P. Killip 
& 4 C. Smith 26780; 20-21 July 1925: Perú, Dept. 
oe Cahuapanas, on Rio Pichis; 340 m; NY (holo), 


Te m dps es 35(6): 401. 1977. 

A i Cro ; 3 Oct 8; Panama Canal Zone, 
el Tud Zenek ail MO Aas lo). 

ia harleyi Steyerm., in . Harley et al. 
15006; 10 Jan. 1974; Brazil, Bala Itabuna, in the 
eh of Centro de Pesquisas do Cacau; MO (iso 
2417 

Ge pida EM (Muell. Arg.) Steyerm. var. nebli- 
nae Steyerm., Mem. New York Bot. Gard. 23(1): 398. 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


709 


1972. B. Maguire et al. 36975; 29 Dec. 1953; Ven 
ezuela, T.F. Amazonas, Hio Yao, Cerro de la Neblina; 
650 m; NY (holo), VEN. 

Geophila Pa aa , Mem. New York Bot. Gard. 
quad .G. T. Juande et al. 8985; 29 Nov. 

968; Ae n. Rondonia, Serro do Tres Irmàos; 
a (holo), VEN. 

Gleasonia duidana Standl. var. latifolia Steyerm., Mem. 
New York Bot. Gard. 17(1): 239. s B. Maguire 
& C. K. Maguire 35344; 27 Feb. 1953; venecia: 
T.F. Amazonas, exo Yutaje, Camp o 1,300 m 
GH, NY (holo), U, VEN. 

Gleasonia duidana May var. oblanceolata Steyerm., 
] York Bot. Gard. 17(1) 239. 1967. F. 
Cardona 1610; 26 June 1946; Venezuela, Bolivar 

Cerro Apacara, Caroni; 800-900 m; NY, VEN (holo) 
— G. duidana Standl. var. duidana, 1984. 
decode ciliata Steyerm., Acta Bot. Venez. 6: 123. 
971. J. Saer 635; Aug. 1930; Venezuela, Lara, Distr. 
ae Serranias de Terepaima; 800-1,000 m; VEN 
(holo). 

Gonzalagunia dicocca Cham. & Schlecht. var. gui- 

anensis Steyerm. . New York Bot. Gard. 23(1): 

1972. B. Maguire & D. B. Fanshawe 32456; 
8 Nov. 1951; Guyana, Paruima Falls to Paruima Mis- 
sion; 600 m; NY (holo), VEN. 


Gonzalagunia dicocca Cham. ia Schlecht. subsp. ve 
zuelensis Steyerm., Mem. mig, ba Gard. 23(1): 
2. A. Allart 225: Nov. Tener. 


Miranda, Quebrada de las Te a. 

(holo). 

Guettarda bernardii Steyerm., Acta Bot. Venez. 6: 128. 
1971. Bernardi 2011; 21. Feb. 1955; Venezuela, 
Barinas, Mun. Pedraza, Alto de Aguada-Los Piedrones; 
1,200 m; NY (holo). 

Guettarda crispiflora Vahl be id e Steyerm., 
Act s. Venez. 6: 129. . H. Pittier 9967; 25 
Dec. 1921; Venezuela, ¡eta Colmi Tovar; 1,900- 
2, m m; VEN (holo). 

Guettarda fanshawei Steyerm., Mem. New York Bot. 
Gard. 23(1): 359. 1972. D. B. Fanshawe 1255 (Rec. 
3991); 21 Apr. 1943; Guyana, Groete Creek, Lower 
Essequibo River; NY (holo). 

Guettarda re d Steyerm., Ann ouri Bot. Gar 
71: 1175. 1984. J. A. Qu et oL 1223168; de 
May 1980; Venezuela, T.F. Amazonas, along road be- 
eee Paso El Diablo end Cano de Culebra, 20-30 km 
SE of Puerto Ayachcho; 100 m; MO, VEN (holo). 

Guettarda steyermarkii Standl. f. n Standl. & atey: 
erm., p Bot. 28(3): 578 


cienda E Jou of ee Ric 


Guettarda eremi Sah f. pilosior Steyerm., Acta 
Bot. Venez. 6: . 1971. E. L. Little 15372; 22 
July d Venezuela, Mir ida, 40 km ESE of Tabay, 
5 km E of San Juan Bautista; VEN (holo). 

a 222 Spruce ex Benth. var. subcuneata 

ew Yo - Bot. Gard. 10(5): 207. R. 

ce we Rio Negro, 

— H. nitida Spruce 


Henriquezia € Spru B Heu i Hook; var, 
apiculata Stey New York Bot. Gard. 10(5 
204. 1963. 4. Duc ke 8l; 19 Dec. 1935; Brazil, es 


as, Manaos; A, F, MO, NY, R, US (holo) = H. 
verticillata Spruce var. verticillata, 1984. 

Hillia bonoi Steyerm., Ernstia 42: 5. 1987. J. Bono 
3797; 2 Apr. 1984; Venezuela, Tachira, Dept. Junin, 
Rubio y Las Delicias; 2,000 m do (holo). 

Hillia costanensis uad Mem. 
23(1): 2 2. J. A. ima 99274; 20 July 
1967; ean Falcon, Sierra de San Luis; 800- 
900 m; VEN (holo 

Hillia us Stoperi New York Bot. Gard. 

972. E. Jonas 7192; 17 Aug. 1968; 


29( 1-2 
d E Auyantepuy, Valle de Orquideas; 
1,200 m; VEN (holo). 

Hillia irwinii Steyerm., Mem. New York Bot. Gard. 
23(1): 287. 1972 W. A. Egler & H. 5. Irwin 46683; 
26 Aug. 1961; Brazil, Terr. Amapá, near Cachoeira 
Mac , Rio Jari; 200 m; NY (ho 

Hillia macrocarpa Standl. & Steyerm., Field Mus. Nat. 
Hist., zm r api 23. nr J Md diea 


Serrania Yutaje, Bio Manapiare; 1,750 m; NY (holo). 
Hillia marcano-bertii di ie pus. 9: 6. 1986. L. 
9-979; 5 Dec. 1979; 

Venezuela, Táchira, M Db. Parque 'Caza- 

dero', carretera Palo Grande- Minas de Barbon de Loba- 

tera; MER, VEN (holo). 

Hillia microcarpa Steyerm., Mem. New York Bot. Gard. 
23(1): 292. 1972. J. A. Steyermark 56591; 18 May 
1944; Venezuela, Mérida, above Tabay; 2,285- 2,745 
m; F (holo), VEN 

Hillia psammophila d Mem. New York Bot. 
Gard. 10(5): 211. 1963. J. . 4. Steyermark & S. Nils- 
son 392; 20 Apr. 1960; Venezuela: Bolivar, vicinity 
of camp 125 at km 125 between Luepa and Cerro 
Venamo, vicinity of Cerro Uei; 1,100 m; NY, VEN 
(holo). 

Hillia sc ipe Steyerm., Mem w York Bot. Gard. 
23(1): 2 1972. R. E. Séhulies € hi Cabrera 13075; 
18 July 1951: een ne onas-V p mouth of 

oris; 250 m; US | 


1964; Perú, Dept. pia Prov 
end of Pomacocha (lake) on 


to Rioja; 2,000 m; VEN (holo 
Hillia ge Steyerm Miiri Bot. Gard. 7 
1176. 1984. J. A. Steyermark et al. 122876; 29 e 


1980; Venezuela, Zulia, Mara, vicinity of Rio — 
between Rancho 505 and Cerro Yolanda; 200-270 m; 
MO, VEN (holo 
Hindsia s ex Lindl, sect. Brevicalyx Steyerm., Brit- 
tonia 30: 34. 1978, sect. nov 
Hindsia irwinii E li Brittonia 30: 34. 1978. H. S. 
Irwin et al. 23537; 19 Feb. 1969; Brazil, Minas Gerais, 
ca. 15 km W of Grão Mogol, Serra do Espinhaco; 950 
m; MO (2817604), NY, UB (holo), VEN. 
ur cn Steyerm., Acta Bot. Venez. 1(1): 
5. O. Haught 5400; 28 Dec. 1946; uer 
Dept ud Valle de San Juan de Micay; 100 m 
(holo). 
mo lasseri Steyerm., Acta Bot. Venez. 1(1): 8 
65. H. Pittier 14103; 8, 29 Aug. 1937; "wid 


710 


Annals of the 
Missouri Botanical Garden 


Aragua, selvas nubladas de valle de Ocumare, cerca 
e Rancho Grande; 1,000 m; VEN (holo). 
Hippo latifolia Steyerm., Acta Bot. Venez. 1(1): 9 
de Wurdack 1817; 3-4 Sep. 1962; i 
. Amazonas, Prov. de Bagua, Valle del Rio Ma- 
ranon Sh de las Cascadas de Mayasi cerca del Cam- 
pament e Montenegro; 450 m; US (holo). 
wr subelongara po Acta Bot. Venez. 1(1): 
me 3084; May 1933; Peru, Dept. 
Lor o 220 m; US (holo). 
Hippos wurdackii Steyerm., Acta Bot. Venez. 1(1): 9 
965 J. Wurdack 2089; 4-7 Oct. 1962; Pan 
Dent Loreto, Prov. de Alto Amazonas, del Rio Maranon 


Pe 
< 


Hoffmannia aroensis Stey erm., ad: 
k & G. Wessels- Boer 100400; 13 NS 


67; Venezuela, el Cerro Negro, Sierra de 


Aroa, arriba de San Felipe; 1,100-1,250 m; VEN 
(holo). 
i draps Sn ar yerm., Pittieria 4: 20. 1972. 


12 May 1 950; Venezuela, Mé- 
e Portachuelo a d bos del Cacique; 950-1,100 
m; NY (holo). 
bodie bernardii tao m. var. aracalensis Stey- 
, Pittieria 4: 20. 2. B. Trujillo & G. S. Bunting 
, 7591; 30 May en Morel: Yaracuy, Aracal; 
600-800 m; NY (holo). 

Hoffmannia costensis Standl. & Steyerm., peer. Bot. 
28(3): 578. 1953. E H A 62542; 5 May 
1945; Venezuela, S abov rinidad; 2,100- 
2,200 m; F (holo) = H Torote. Standl. & ran 
s costensis (Standl. & Steyerm. Fosse. 1972. 

Hoffmannia huehueteca Standl. & $ , Field um 

Ta J. 


Nat. Pot i: 


de los Cuchumatanes), Cerro Negro; 
1,600- 2.000 m; F (holo). 
Holman larensis Steyerm., Pittieria 4: 20. 1972. J. 
'rmark et al. 103322; 3 Aug. 1970; Vene- 
Hn p n Distr. Palavecino, Terepaima, 20 km S de 
Cabudare; 1,200-1,300 m; VEN ard 
M deis a ifolia Steyerm., Bol. Venez. Ci. 
. 25(107): 231. 1964. J. J. Wurdack 911; 19 
n 1962; Peru, Dept. Amaz zonas, Prov. Bongara, NW 
Pon muchas 2,300-2,700 m; US, VEN (holo). 
Hoffinannia pauciflora Standl. venezuelensis 
Steyerm., Pittieria 4: 25. 1972. F. Breteler 3463; 20 
Dec. 1963; een Mérida, La Mucuy, 15 mi. NE 
of Mérida; 2,350 m; VEN (holo 
Hoffmannia quadrifolia Standl. & Steye rm., Field Mus. 
at. Hist., Bot. Ser. y pe 1947. Steyer- 
mark 42600; 12-13 Jar 1942; Guat Ela, Dept. 
Zacapa, Sierra de las Minas, Monte Virgen; 2,400 m; 
F (holo). 
dl css rac emife ra Standl. & pelea Field Mus. 
, Bot. Ser. 23(5): 249. . J. A. Steyer- 
ipn 41910; 25 Dec. 1941; cease Dept. Izabal. 
Cerro San Gil; 300-900 m; F (holo). 


;. Wessels- ues 100402: 2: I: 3 
Nov. 1967 /; Venezuela, Yaracuy, Sierra de Aroa, C erro 
Negro, arriba de San Felipe; 1,100-1,250 m; depos- 
itory not designated. 

Hoffmannia terepaimensis Steyerm., Pittieria 4: 24. 
1972. J. A. Steyermark et al. 103352; 14 run "bn 
Venezuela, Laro, La To ma, Loma Redonda, 
epaima, 23 km S of Babudare; 1,100 m; an vind 


Hoffmannia tuerckheimii Donn. Sm. var. glabra Standl. 
ed Field Mus. Nat. , Bot. Ser. 23(5): 
250. . J. A. zii deni Peer 10 Feb. 1942; 
Guate nita Dept. El Progresso, Sierra de las Minas, 
Montana Canahui; 1,600-2 300 m; F (holo). 
Hoffmannia anicola Standl. & Steyerm., Field Mus. 
at. Hist., Ser. 23(5): 251. 1947. J. A. Steyer 

mark 47931; 30 June 1942. Guatemala, Dept. Sololá, 
S Volcán Atitlán, above Finca Moca; 1,000-1,250 m; 

F (holo). 
Holstianthus Steyerm., 
6, gen. nov. 
Holstianthus Vp coa Steyerm., Ann. Missouri Bot. 
A. Steyermark & B. K. Holst 
1300. 37; 23- 24 Ps 1985; Venezuela, T.F. Amazo- 
nas, Dept. Atabapo, above Caño Negro, Cerro Mara- 

huaca; 1,220-1,350 m; MO (holo), VEN. 
Houstonia pusilla Sc ur f. rosea Steyerm., Rhodora 
9. 1 A. Steyermark 4599a; 17 Apr. 
, along road D, 4 


Ann. Missouri Bot. Gard. 73 


1937; U.S.A., Mis ssouri, Pulaski Co. 
mi. W of Taromsi F (holo 
Isertia haenkeana DC. var. edi nsis Steyerm., Mem. 
New York Bot. Gard. 17(1): 298. 1967. J. A. Stey 
7 . 1960; Venezuela, Miranda; 
e Nacional de Guatopo; 500 m; VEN digi 
robes poor Vahl var. hirta Steyer . New 
York Bot. Gard. 17(1): 2 1967. Fore si n Field 
No. W B388: Oct. 1948; Gayana, Wabuwak, Kanuku 
Mountains; 605 m; NY (holo). 
Isertia peas a Mem. 
. H. S. Irwin et al. 


New York Bot. 
54949; 


22 Aug. 1963; ed E ey Gebergete, Fred- 
erik Top, 3 km SE of Juliana Top; 1,050 m; NY (holo). 


Ixora agostiniana Steyerm., 

A. Steyermark & G. Agostini 91120; 19 

July 1962; Venezuela, Sucre, Cerro Patao; 800-825 
m; VEN (holo). 

Ixora aluminic ve Steyerm., Mem. New York Bot. Gard. 
17(1): 3 967. B. Maguire 40799; 12 Mar. 1955; 
Suriname, Nasen Mountains, Marowijne River, Pla- 
e ; 400-550 m; NY (holo). 

Ixora DEA SA Mem. New York Bot. Gard. 

je d E d Murca Pires et al. 51210; 25 
A cs Amapá, Rio Araguari, Serra do 
Navio; un (holo). 

Ixora intropilosa Steyerm., er New York Bot. Gard. 

S . Tillett et al. 45085; 10 
Aug. 1960; Guyana, Bu. a ip to Thompson Camp, 
Mt. Ayanganna; 760-1,4 ES m; NY (holo). 
Ixora nr Steyerm., Mem. New York Bot. Gard. 17(1): 
384. 1967. J. Murca (n 500. = l Sep. 1961; 
Brazil, ine Rio Araguari; NY (ho 

Ixora prin Steyerm., Mem. New cu Bot. Gard. 
17(1): 348 . D. B. Fanshawe 1147 (Field. No.): 
26 Jan. 194 i Guan Kakaralli-Clump Eagle Mt., 
Potaro River; K (holo). 

Ixora rias Steyerm., Mem. New York Bot. Gard. 
17(1): 344. 1967. R. Spruce E June 1850; Brazil, 
Pará, Tapajoz, ee K (holo). 

Ixora Apr d Mem. New York Bot. Gard. 
23(1): 3 2. J. A. Steyermark & G. S. la 
Du 21 ris 1970; Venezuela, along road between 
Yavita and Pimichin, 1 km from Yavita; 125 m; NY 
VEN (holo). 

Joosia subg. NN Steyerm., 

1975, subg. nov. 
Joosia oda Steyerm., 


4. LIVI. 2. 1). 


Brittonia 27: 252. 


Brittonia 27: 257. 1975. 


Volume 76, Number 3 
1989 


Taylor 711 
Plants Described by 
Julian A. Steyermark 


J. A. Steyermark 54506; 9 Oct. 1943; Ecuador, San- 
US 


tiago-Zamora, Quebrada Achupallas; F (holo), 
Joosia macrocalyx Stan Steyerm., Brittonia 27 
259. 1975. Ownbey 2739; 25 Mar. cuador, 


Napo-Pastaza, ridge S of Rio Pano, Cordillera de Gua- 
camayos, Archidona-Tena region; 1,605 m; F (holo), 


US. 

Joosia nn ee Brittonia 27: 257. 1975 
J. A. 58; 10 Oct. 1943; Ecuador, 
Santiago- Zamora, ne a Honda, vicinity Rancho 
Achupallas; 2,500-2,700 m; F (rotoj, NY, US. 

Joosia ulei Serai Brittonia 27: 255. 1975. E. Ule 
6795; Mar. 1903; dii quam Pampas de Ponasa; 
1,200 m; 6, HBG (ho 

Joosia p E a var. caucana Steyerm., Brit- 
tonia 27: 254. 1944. Core 1367; 27 Sep. 1944; Co- 
ju ibia, Cauca, Rio Pinche, near headwaters, NW of 

gelia; 1,060 m; NY, US (holo). 

ne umbellifera i. to macarensis Steyerm., 
Brittonia 27: 254. 5. Philipson et al. 2072; 10 
Jan. 1950; Co bus E eta, N Ridge, Central 
Mountains, Serranía de la Macarena; 1,500 m; F, US 
(holo). 

Joosia a o p occidentalis Steyerm., 
Brittonia 27: 254. . J. Cuatrecasas 14827; 4 
Aug. 1943; Colombia, vals Cordillera Occidental, Hoya 
del Río Anchicaya, El Prado; US (holo). 

“Kotchubaea” duckei Steyerm., Mem. New York Bot. 
Gard. 10(5): 216. 1963. 4. Ducke 1557; 5 Apr. 1944; 
Brazil, Amazonas, Sáo Paulo de Olivenca; F, NY, US 


boen A d Steyerm., Mem. N 
Gard. 10(5): 218. 1963. J. 4. Steyermark 75544; 24 
May 1953; Venezuela, Bolivar, near Rio Tirica, lower 
slopes Chimantá-tepui (Torono-tepui), Chimanta Massif; 
1,000 m; F, NY, VEN Moi). 

“Kotchubaea” micrantha Steyerm., Mem. New York 
Bot. Gard. 10(5): 215. 1963. E Foldats 3758; Sep. 

1960; Venezuela, T.F. Amazonas, de Alto Rio Atacavi; 
NY, VEN (holo). 

“Kotchubaea” montana Steyerm., Mem. New York Bot. 
Gard. 10(5): 217. 1963. G. Klug 3452; Dec. 1933; 
Peru, Dept. San Martín, Zepelacio, near Moyobamba, 
1,200-1,600 m; NY (ho es 

* Kotchubaea" morilloi Ste is a Bot . 9: 
287. 1974. J. Hoyas & G. Mori A pos 7 Mie 1973; 
Venezuela, T.F. Amazonas, Cerro Asisa (La Momia), 
e Parú; i, een m; MO (2250699), VEN (holo). 

“Kotchubaea” neblinensis Steyerm., Mem. New York 
Bot, Gard. mein 216. 1963. B. Maguire et al. 42084; 

7 Nov. ; Venezuela, T.F. Amazonas, Cerro de 
la Neblina, as 'N of Camp 3; 650-700 m; NY (holo). 

Ladenbergia buntingii Steyerm., Acta . Venez. 6: 
99. 1971. G. S. Bunting 1509; 31 ju 1971; Ven- 
e Cerro La Chapa; MO (holo— 
3399535), MY, VEN. 

Ladenbergia d Aa ru .. Mem. New York Bot. 

(1): 270. 1972. B. a. et al. 42493; 
25 Dec. 1957; Venere. T.F. azonas, Cerro de 
la Neblina; 1 .050- 1,100 m; NY > lo). 

Ladenbergia venamoensis Steyerm., Mem. New York 
Bot. Gard. 23(1): 271. 1972. J. A. Steyermark & S. 
Nilsson 426; 21 Apr. 1960; Venezuela, Bolivar, Cerro 
Venamo; 1,140 m; VEN (holo). 

uod ee d Mem. New York Bot. Gard. 
23(1): 2 1972, gen. nov 

Maguireoc rae neblinae on Mem. New York 
Bot. Gard. 23(1): 230. 1972. N. T. Silva & U. Brazao 


60898; 26 Jan. 1966; Venezuela, T.F. Amazonas, 
Sierra Pirapucu; 1,300 m; NY (holo), VEN. 

M ener qe Steyerm., Mem. New York Bot. Gard. 
10(5): 2 1963, gen. nov. 

MEA jauaensis (Steyerm.) Steyerm. var. 
breweri Steyerm., Fl. Venez. 9(1): 311. 1974. J. A. 
Steyermark et al. 109765; 4 Mar. 1974; Venezuela, 
Bolivar, Cerro Jaua, Meseta del Jaua; 1,750 m; VEN 


(holo). 

Maguireothamnus speciosus (N.E. Br.) Steyerm. subsp. 

airo: ba de Mem. New York Bot. Gard. 23: 

2. J. A. Steyermark 97837; 22-27 Mar. 
1967; pisse Bolivar, Cerro Jaua; VEN (holo) = 
M. jauaensis (Steyerm.) Steyerm., 1974 

Malanea auyantepuiensis Vi aa Mem. New York 
Bot. Gard. 12(3): 256. 1965. J. 4. Steyermark 94103; 
18 May 1964; ld Asie valle de Kamarata, 
Auyan-tepui; 1,530-1,8 ; NY, US, VEN (holo). 

Malanea chimantensis iem ., Mem. New York E 
Gard. 12(3): 259. 1965. J. A. Steyermark 75166 
Apr. 1953. Venezuela, Bolivar, Chimantá Massif, c 
capa-tepui; 1,400 m; NY (holo). 

Malanea ciliolata Steyerm., Mem. New York Bot. Gard. 
12(3): 258. 1965. S. S. Tillett et al. 45081; 10-12 
Aug. 1960; Guyana, upper Mazaruni River basin, Mt. 
Ayanganna, above Thompson Camp; 1,418-1,525 m; 
NY (holo). 

Malanea cruzii Steyerm., Mem. New York Bot. Gard. 
12(3): 253. 1965. J. > de la Cruz 2164; 22 Sep.- 
6 Oct. 1922: Guyana, upper agen River; NY (holo). 

Malanea egleri Steyerm., Bot. Gard. 
12(3): 260. 1965. 4. Egler E J. Murca Pires et al. 
47729; 22 Aug. 1960; Brazil, Terr. Amapá, 0.5 km 
E of confluence with Rio Oiapoque; NY (holo). 


[| 


Malanea a Steyerm., Mem. 
Bot. Gard. 12(3): 253. 1965. F. Cardona 1117; 10 
July 1944; Mores Bolivar, Alto Rio Paraqua, Cerro 
Guaiquinima; 1,200 m; F, US (holo), VEN. 

Malanea hypoleuca Steyerm., 
Gard. 12(3): 251. 1965. f 
Aristeguieta 5; 6-11 Mar. 1962; Venezuela, Bolivar, 
vicinity of km 110, S of El Dorado; 800-1,200 m; 
Vin (holo). 

Malanea jauaensis Steyer 
i 885. 1972. J. 

67; Venezuela, 
m; VEN (holo). 

Malanea macrophylla Bartl. f. cune pe cu Mem. 
New York Bot. Gard. 12(3 ; 250 E C. Smith 
2618; 21-26 Nov. 1937; Guyana, ca 0 mi. from 
mouth, basin of Kuyuwini River cie tributary); 
NY (holo). 

Malanea die de Standl. & Steyerm., des Bot. 
28(3): 5 3. J. A. Steyermark 60875; 7 Dec. 

1944; Visi Bolivar, Carrao-tepui; 2, 130- 2,430 
m; F (holo). 

Malanea pariensis Steyerm., Mem. New York Bot. Gard. 
12(3): 248. 1965. J. A. Steyermark & G. Agostini 
91126; 19 July 1962; Mua Sucre, Peninsula de 
Paria, NE de Guiria, N de Puerto de Hierro, Cerro 
Patao; 800-825 m; VEN ( holo). 

Malanea ptariensis Steyerm., Mem. New York Bot. Gard. 
vn ): 255. 1965. J. A. Steyermark 59548; 30 Oct. 

944; Venezuela, Bolivar, Ptari-tepui; 2,285- 2,405 
m; F (holo), NY, US, Mon 

Malanea sanluisensis Steyerm., Acta Bot. Venez. 6: 133. 

1971. J. 4. on 99363; 2] July 1967; Ven- 


rm., Mem. New York Bot. Gard. 
A. Steyermark 97857; 22-27 Mar. 
Bolivar, Cerro Jaua; 1,922-2,100 


712 


Annals of the 
Missouri Botanical Garden 


ezuela, Falcón, Sierra de San Luis; 1,300 m; VEN 

holo). 
Malanea sarmentosa Aublet E sra Steyerm., 
5: 4. C. 


Mem. New Bot. Gard. Pe es 


Smith 3066; 15 17 Feb. 1938 ea Parabaru 
Savanna, between Rupununi & Kuyuwini rivers; NY 
(holo 

Malanec za lio Aublet f. Mises a Mem. 
New York Bot. Gard. 12(3): 258 . B. Maguire 
24874; 30 Sep. 1914; Suriname, pel River 


headwaters, between camps 3 and 2; NY (holo). 

Malan^a UM Steyerm., Mem. New York Bot. 
Gard. 12(3): 2 1984, nom. nov. 

Malanea ee erm., Mem. New York Bot. Gard. 
12(3): 260 J. A. Steyermark 58558; 24 Sep. 
1944; Venezuela, Bolivar, stream tributary to Rio Ku- 

enan at base of Mt. Roraima; 1,185-1,280 m; F, 
NY (holo), VEN. 

Malanea sipapoensis Steyerm., Mem. New York Bot. 
Ga 12(3): 252. 1965. B. Maguire & L. Politi 
28743; 2 Feb. 1949; Venezuela, T.F. Amazonas, Cerro 
Sipapo (Paraque), near Intermediate Camp; 600 m; 
NY (holo). 

Malanea subtruncata Steyerm., Mem. New hr Bot. 
Gard. 12(3): 258. 1965. 4. Ducke 1943; 29 Apr. 
1946; Brazil, Pará, Belem Bosque Municipal; io (holo), 


US 
Malanen RA e rgen nsis pire Se m., Mem. New York Bot. 
rd. 12(3): 251. . B. Marnie 24530; 28 Aug. 
1944; Surname, Tafelberg. Savanna No. 8; 776 m; 
NY (holo). 
eis a ee nsis Steyerm., Mem. New York ae Gar 
2(3): 255. 1965. J. A. Steyermark & | P ipie 
Pe 20- 22 Apr. 1960; Venezuela, doe between 
ie a Cerro Venamo, vicinity of road campamento 
inity of Cerro Uei; 1, 100 m; VEN (holo). 
a Mutis ex L. sect. ica Steyerm., 
tonia 30: 36. 1978, 
Manettia badilloi Steyerm Men. New wy Bot. 
972. Badillo 3126; 4 July 19 
Venezuela, Aragua NS de Choroni; MAY (holo), VEN. 
Manettia bernardii Steyerm., Acta Bot. Venez. 6: 104. 
971, also Mem. New York Bot. Gard. 23(1): 2 
us L. Bernardi 611; 14 June 1953; Veneiela, 
, Sierra Nevada de Mérida; 2,200 m; NY (holo). 


ç 


Brit- 


Manettia iir Steyerm., Acta Bot. Venez. 6: 103. 
Mem. New York Bot. Gard. 23(1): 244. 
T 3 b Breteler 4089; 11 Aug. 1964; Venezuela, 


Trujillo, a pd. road to Biscucuy; 1,850 m; MO 
(2417037), VEN (holo). 

Manettia cocc had loides Wernham var. glabrior Stey- 
erm., Acta Bot. Venez. 6: 107. 1971, also Mem. New 

k Bot. Gard. 23(1): 248. 1972. J. A. Steyermark 
& 7 Rabe 97071; 31 Aug. 1966; Venezuela, Mérida, 
E of Jaji & SE of La Carbonera; 1,900-2,100 m; VEN 
(holo). 

cour honigii Steyerm., p 31(6): 483. 1975. 

. Tillett & K. W. : 


Venezuela, tr. Rivas Davila km above 
Las Tapias S of Bailladores, Sierra de Tovar, Cordillera 
Mé m 2, 275 m; VEN (holo). 
Vanettia irwinii Steyerm., Brittonia 30 a 
S. Irwin et al. 18688; 17 Jan. 1968; Braal, Minas 
Gerais, 15 N of Corumba de roa 


m 
AO 1,150 m; MO (281760 2 NY. UB (holo), 
VEN. 


Manettia lindenii Sprague f. appendiculata Steyerm., 


Acta Bot. Venez. 6: 102. 1971, also Mem. New York 

Bot. Gard. 23(1): 243. 1972. J. A. Steyermark & M. 

Rabe 97239; 3-4 Sep. 1966; Venezuela, Trujillo, 30 

km from Boconó; 2,300-2,500 m; MO (2270190), 
VEN (holo). 

Manettia paramorum Steyerm., Acta Bot. Venez. 6: 104. 

s , also Mem. New York Bot. Gard. 23(1): 245. 

. L. Bernardi 1248; 16 May 1954; Venezuela, 

THEN Páramo de los E d 3,100 m; NY (holo). 

Manettia tachirensis Steyerm., 


E Mise A Táchira, E af Zumbador; 1,700 m; 
VEN (a lo). 
Manettia tamaensis Steyerm., Acta Bot. Venez. 6: 106. 
D also Mem. New York Bot. Gard. 23(1): 247. 
A. Ste hene et al. 98298; 15-16 May 
; Venezuela, Táchira, entre Bramon y Las Deli- 
jd. 750 m; VEN (holo). 
Manettia tillettii Steyerm., Phytologia t 484. 1975. 
. Tillett & K. W. Honig 746-501; 24-26 June 
197 4; Venezuela, Zulia, Distr. Perijá, d de Perijá, 
Serranía de los Motilones, headwaters of Rio Tocuco, 
Mu ds ine Frontier I; 1,900-2,000 m; VEN (holo). 


Meruma Steyerm., Mem. New York Bot. Gard. 23(1): 
232. 1972. gen. nov. 

Meruma PERS O Mem. New York 
e Gard. 23(1): 233. 1972. B. Maguire & L. Politi 


243; 8 Jan. 1949; dd Amazonas, Cerro 
pian (Paraque); 1,600 m; NY (holo 

Meruma Pea Steyerm., Mem. New York Bot. Gard. 

. 1972. S S. Tillett et al. 43944; 1 es 
1960; Guyana, Merume Mountains, Merume Mt., 
per Mazaruni River basin, Partang River; 1, 140 1 Ud 
NY (holo N. 

Miracarpus frigidus (Willd.) Schumann var. andinus 
Steyerm., Mem. New York Bot. Gard. 23: 779. 1972 
O. Kuntze 1643; 21 May 1874; Venezuela, Distr. 
Federal, Silla de Caracas; NY (holo). 

Mitracarpus, frigidus (Willd.) Schumann var. orinocensis 
Steyerm., Mem. New York Bot. Gard. 23: 780. 1972. 

J. J. Wurdack & L. Adde rley 42698; 30 May 1959; 

et 2 F. Amazonas, Río Orinoco, Sabanita Mo- 


roc -150 m; NY (holo), 
ade arpus. frigidus (Willd. ) Se humano \ bed HE ruvianus 
;'erm., Mem. New York Bot. Gard. 23: 780. 197 E 


L Evinger 513; 21 Aug. 1943; Per u, Dept. Am 


zonas, 10 km E of Balzas; 2,300 m; NY (holo), US 
VEN 


Mitracarpus pum Steyerm., pres 30: 36. 1978. 
I Irwin et al. 27120; 1 M 1970; Brazil Minas 
Gerais, Serra do € hall ca. 8 in W of Joaquim Felicio; 
1,200 m; NY, UB (holo). 

Morinda longipedunculata Steyerm., Ann. Misao Bot. 

. 1988. G. Davidse & J. S. 

7; 9-14 July 1984; Venezuela, T.F 
Dept. Rio Negro, Cerro de la Neblina, Canon Grande; 
350-400 m; MO (holo), VEN 

M Ra eue Steyerm., Mem. New York Bot. Gard. 
10(5): 2 1963, gen. nov 

Td in aracamunianus Steyerm., Ann. Mis- 
souri Bot. Gard.: in press 9. R. L. Liesner & F. 
Delascio 22153; 19 Oct. 1987; roce T.F. Ama 
zonas, Dept. Rio Negro, Cerro i summit, 
Popa Camp; 1,550 m; MO t 

Veblinathamnus argyreus Steyer vd New York 
Bot. Gard. 10(5): 229. 1963. B Wenn 42120; 


LI 


Volume 76, Number 3 
1989 


Taylor 713 
Plants Described by 
Julian A. Steyermark 


6 Nov. 1957; Venezuela, T.F. Amazonas, Cerro de 
E Neblina: NY (holo). 

Neblinathamnus brasiliensis a p Mem. New York 
Bot. Gard. 23(1): 309. 1972. N. T. Silva & U. Brazao 
60907; 26 Jan. 1966; Brazil, Amazonas, Serra Pira- 

â h 


Neblinathamnus glabratı 
Bot. e ae ber 231. 1963. 
30 53; Venezuela, T. F Amazonas, Cerro de 

la tds p (holo). 
Pagamea acrensis Steyerm 


V. A. Ros 


, Brittonia 33: 385. 1981. 
7 706; 10 Feb. 1976. Brazil, Acre, Cruziero 
o RADAM-Ponto 10-SB-18-2D; IAN, 


Pagamea anisophylla Standl. & Steyerm., rada Bot. 
28(3): 584. 1953. J. A. Steyermark 5 ; 8 Sep. 
1944; Venazicls, T.F. Amazonas, bie "100 m; 
F (holo). 

Pagamea auyantepuiensis Steyerm., Fieldiana Bot. 28: 
584. 1953. G. H. H. Es Pas Dec. 1937-Jan. 
1938; Venezuela, Boliv n-tepui; 2,200 m; 
F (holo) = Coryphothamnus cle (Stey- 
erm.) Steyer 1967. 

Pagamea brev ipeduneulata Steyerm., Mem. New York 
Bot. Gard. 12(3): 283. 1965. B. ad 33070; 10- 
12 Jan. 1952; ei Bolívar, Cerro Guaiquinima, 
North Valley; 1,600-1,700 m; NY (holo). 

Pagamea capitata Benth. subsp. d S cus ne 
erm. revi s a . k Bot. € ard. 

| » 281. 1965. B. Maguire et al. peers 20 Nov. 
ee d T.F. Amazonas, Cerro Duida, Rio 
me. above Culebra; 1,400 m; NY (holo 

e var. acuta Steyerm., Mem. 
New York Bot. Gard. 12(3): 273. 1965. R. E. Schultes 
& I. Cabrera 19367; 14 May 1953; Colombia, Vaupés, 
Río Vaupés, between Mitu and Jauarete, Yutica; US 
(holo). 

Pagamea coriacea Spruce var. pubescens Steyerm., pons 
New York Bot. Gard. 12(3): 273. 1965. 4 Ducke 6 
22 Mar. 1941; Brazil, Amazonas, Rio Negro en 

d 


everm 


m., Mem. New York y Gard. 
S. jos & J. urdack 
31241; 4 Feb. 1941: Vales uela, T.F. d. Ser- 


, Fieldiana Bot. 


( G. H. H. Tate 867; l Jan. 1929; 
Venezuela, a F. Amazonas, Mt. Duida, Central Camp; 
1,600 m; F (holo), NY 


Pagamea nn Steyerm d ie 31(2): 226. 1975. 
T. idley 1354 1969; Perú, Dept. 
Huanu SW slope a Río Llulla Pichas watershed, on 
ascent a Cerros del Sira; 1,850 m; NA, VEN (holo). 

Pagamea guianensis Aublet var. macrocarpa Steyerm., 
Mem. New York Bot. Gard. 12(3): 278. 1965. E. L. 
& R. R. Little, Jr. 9533; 4 Mar. 1945; Colombia, 
Comisario de Caqueta, Tres Esquinas, 
Solano, on Río Caqueta below mouth of Rio Orteguaza; 
200 m; NY (holo), US 

Pagamea harleyi Steyerm., Kew Bull. 38(2): 317. 1983. 
R. Harley et al. 17069; Jan. -Apr. 1974; Brazil, Bahia; 
CEPEC (holo), MO (2397731). 

Pagamea jauaensis Steyerm., Mem. New York Bot. Gard. 
23: 883. 1972. J. 4. Steyermark 97928; o di Mar. 
1967; Venezuela, Bolivar, Cerro Jaua; 1,922-2,100 
m; VEN (hol 

Pagamea Ru A Steyerm., 


Mem. New York Bot. 


Gard. 12(3): 285. 1965. B. Maguire 33427; 15 May 
1952; Venezuela, Bolivar, Ilu-tepui, between N Peak 
and Central Plateau; 2,450-2,550 m; NY (holo). 

de cae Wap ER Standl. & Steyerm., o Bot. 
28(3): 5 53. J. A. Steyermark 60888; 7 Dec. 
1944; dude Bolivar, Carrao-tepui; >, ran 2,500 
m; F (holo 

Pagamea plicata Spruce var. multinervia Steyerm., Mer 
New York Bot. Gard. 12(3): 274. 1965. L. hine 

. 1954; Venezuela, Bolivar, region de 
Uriman; 340-380 m; VEN (holo). 

d Pp due a Mem. New York Bot. 
e 12(3): 274 . Ll. Williams 14190; 9 Feb. 

; Venezuela, TT. Amazonas, Alto Rio Negro, 
ou selva de Maroa; 127 m; F (holo). 

Pagamea puberula Steyerm., Mem. New York Bot. Gard. 
12(3): 276. 1965. A. Ducke 8462 (RB22951), 21 
Aug. 1907; Brazil, Pará, Campos do Chicodaca; US 
(holo). 

Pagamea reducta Steyerm., Mem. New York Bot. Gard. 
LAS 283. 1965. K. D. Phelps & C. B. Hitchcock 
366; 1-15 Feb. 1948; Venezuela; Bolivar, Uaipan- 
dur 1,900 m; NY (holo), VEN. 

Pagamea standleyana Steyerm., Fieldiana Bot. 28(3): 
589. 1953. J. A. Steyermark 5963: 4; 1 Nov. 1944; 
Venezuela, Bolivar, Ptari-tepui; 1,600 m; F (holo). 

Pagamea uniflora Steyerm., Mem. New York Bot. Gard. 
12(3): 284. 1965. J. A. Stey dish 74877; 13 Apr. 
1953; Venezuela, Bolívar, Chim e NW part 
of Abacapa-tepui; heparin not | designated 

Pagamea eo ew York Bot. Gard. 

B. Maguire & L. Politi 28536; 
2] Jan. 1949; diit TU Cerro Sipapo 
(Paraque), near Camp CRM 1,500 m; NY (holo). 

du e Steyerm., Mem. New York Bot. Gu 12(3): 


267. 1965, gen nov 
Pai is maguirei i Steyerm, ., Mem. New York Bot. 
Gard. 12(3): 267. 1965. 4. Steyermark & J. J. 


Wurdack 1020; 21 Feb. 19 

Chimantá Massif, os tepui, Cao Mojado; 2,0 30- 

2,150 m; NY (holo), V 
Pagameopsis maguirei Steyern m. 


. New Yor 


subsp. bia A 


k Bot. Gard. 12(3): 


Camp, summit; 
Pagameopsis maguirei Steyerm. subsp. neblinansis 
Steyerm. var. angustifolius Steyerm., Mem. New York 
Bot. Gard. 12(3): 270. 1965. B. Maguire et al. 36948; 
1953; Venezuela, T.F. Amazonas, Cerro de 


maguirei > 
Steyerm. = eer Steyerm., Ann. na 
Bot. Gard. 08. 1987. N. T. Silva & U. Brazao 
60880; 20 e 1966; Brazil, Serra Pirapucú; 1,300 
m; NY (holo). Note: This — is also holotype for 
Psychotria canescens Steyerm., however the dates and 
elevation differ. 

agameopsis maguirei Steyerm. subsp. pusillus Stey- 
erm., Ànn. Missouri a Gard. 74: 106. 1987. R. L. 
ree 16013 8 Feb. 1984; Venezuela, T.F. 
Amazonas, Pon, Ri gu Cerro de la Neblina, Camp 
II, NW Plateau; 1,750-1,850 m; MO (holo— 
3327370), VEN. 

Pagameopsis maguirei Steyerm. subsp. pusillus d 
erm. var. iim Steyerm., Ann. Missouri Bot. G 


74: 108. . E. Cardona 3096; 5 Dec. 1972; 


714 


Annals of the 
Missouri Botanical Garden 


Venezuela, T.F. Amazonas, summit of Cerro Avispa, 
Rio Siapa; 1,510 m; NY (holo), VEN. 
Palicourea aculeifera Steyerm., Acta Bot. Venez. 2: 331. 
isa 60377; 20-21 Nov. 1944; 
Venezuela, Bolív , Auyan-tepui; 1,065-1,220 m; F, 
=P. Desain subsp. chrysorrhachis 
(Bremek. ) Steyerm., 1972. 

Palic ourea amapae id s Steye rm., Mem. New York Bot. 
; 6 Murca din et al. 50366; 
20 ji 1961; ‘Brazil, Terr. Amapá, near Rio Araguari 
between Cachoieras Travessão a Santa Maria; NY 
holo). 

Palicourea angustifolia H.B.K. var. glabricaulis Stey- 

Venez. 6: 174. 1971, also Mem. New 

. 23: 771. 1972. L. Ruíz Terán & L. 
E. Rodriquez Poveda 3746; 10 Mar. 1967; Vene- 
zuela, Lara, Distr. Iribarren, Mun. Juarez, La Cuchilla; 
1,500 m; VEN (holo 

Palicourea ay haves hya Steyerm., Acta Bot. Venez. 
6: Funck & Schlim zd Nov. 1846; 
Venez a de San nd P (ho 

Palicourea buntingii Steyerm., Acta Bot m : 148. 
1971. J. A. Steyermark et at 101900; 5 (rie 1968; 
Venezuela, Apure, ferro La Cristalina; 300 m; NY, 

lo). 


VEN (holo 
Palicourea buntingii Steyerm. var. glabrior Steyerm., 
:ta Bot. Venez. 6: 151. 1971 Bernardi 7437; 
22 Mar. 1959; Venezuela, Zulia, Rio Negro; 1,000 
m; MER (holo), MO ii raids 
Palicourea canaguensis Steyerm., Acta . Venez. 6 


57. 1971. Type not aia in de iind de- 
scription. Note: Fl. Venez. cites the type a 
Steyermark 56412; n.d.; Venezuela, Mérida, Quebra. 
da de Los Salichales, afluente del Rio Canagua, al SO 
ae agua; depository not g e 
Palicourea coheed Steyerm., Mem. New York Bot. 
Gard. 2¢ 72. F. ree 1383: 16-24 Feb. 
1946; Bor E nid 0, pon te de Rio Padauiri 
que cae al Rio - US (holo). 
Palicourea chimo Standl. Steyerm., Fieldiana Bot. 


uh 
rea demissa Standl. var. chink (Standl. & Steyerm.) 


Steyerm., 1971. 
E crocea (Sw.) Roemer & Schultes var. riparia 
. N 


(Benth.) Griseb. f. O ens Mem. New 
York Bot. Gard. 23: 739. 2. J. Lanjouw & LG 
Lindeman 802; 12 Oct. ras Suriname, km 12.2 


Moengo; NY (holo), U. 
Palicourea ctenocalyx Steyerm., Acta Bot. Vene 
16 971. J. A. Steyermark et al. 100763; 12 $^ 
1968; Venezuela, Táchira, cabeceras del Río Quini- 
mari; 2,500-2,800 m; VEN (holo 
Palicourea demissa Standl. var. chimo (Standl. & Stey- 
erm.) Steye rm. f. juu Steyer 
1971. J. A. Ste siiis j M. Rabe 96810; 
28 Aug. 1966; s Táchira, entre Michelena 
y Boca gk Monte; 2,000-2,400 m; olo) 
Pati ourea dunsterv illeorum Steyerm., Acta 
: 175. 1971. J. A. Mobi ioi i al. 
A May 1907; Venez Ns Táchi 
2,300- / 450 m; VEN (holo). 
bc a ac Steyerm., Acta Bot. Venez 
. 1971. A. Steyermark 57192; 13 July un 


Bot. Venez. 
98810; 22- 
ira, Páramo de Tamá; 


Táchira, Montana of Rio Táchira at base 

2,430 m; VEN (holo). 

Palicourea flexiramea Secon, Acta Bot. Venez. 6: 
1 1971. R. Velasco & M. Ramia 627; 2 Apr 
1950; Venezuela, Mérida, entre La Azulita y Ejido; 
MY (holo). 

Palicourea D _Steyerm., Mem. New York Bot. 

777. 2. E. Foldats 3670; 4 Sep. 1960; 

Venezuela, T. $ Amos Santa Cruz, Rio Atabapo; 
NY, VEN (ho 

Palicourea us etu Acta Bot. Venez. 6: 167. 
971. F. Garcia 55; 12 May 1951; Venezuela, Ara- 
gua, Fila del MM Rancho Grande; 1,375 m; VEN 
holo 


Venezuela, 
of Páramo de Tama: 


1olo 

Palicourea glabriflora Steyerm., Mem. New York Bot. 
Gard. 23: 735. 1972 Maguire et al. 37490; 4 
Feb. 1954; Venezuela, T.F. Amazon as, vun ici rios 
Pacimoni- Yatua, Piedra Tururumeri; 110-220 m; NY 
(holo). 

Palicourea guianensis Aublet subsp. occidentalis Stey- 

k Tot Gard. 23: 729. 1972. De 

2; Guyana, between the Dem- 

olo). 


erare anc Jerbice rivers; NY 

Palicourea AEN NR e Te occidentalis =u 
erm. f. glabra Ste , Mem. 
23: 730. 1972. F. "n. pese 21211; 10 Oct. 1943; 


Colombia: Sed Narino, along Rio TRES 1-2 km S 

of Barbacoas; 40-50 m; NY (holo), 

Palicourea pa Aublet subsp. i mus eee 
erm. var. glabrescens Steyerm., Mem. : 
Gard. 23: 730. 1972 itiv j 
July 1964; Ecuador, Prov. Esmeraldas, San Lorenzo; 
UCLA. VEN (holo). 

Palicourea huberi PA dove 9: 8. 1981. C. 
Davidse et al. 17 May 1979; Venezuela, T.F. 

be Caño Y agua at Cucurital de 

N (holo). 
Ec ae insignis Steyerm., Acta Bot. i. a 176. 
2 Wis New York Bot. Gard. 2: . 1972. 
A la ; 22 Aug. 1944; Venezuela, “Trujillo, 
bs a a de Escuque; VEN (holo). 

Palicourea irwinii EM. Mem. New ps Bot. Gard. 
23: 758. 1972. H. S. Irwin et al. 47354; 5 Aug. 
1960; Brazil, Amapá, 5 km SE of Clovelandia NY 
(holo). 

Palice ourea lasseri Steyerm., Acta Bot. Venez 

. T. Lasser 1156; Aug. 1944; Venezuela, Trujillo, 
entre Boconó y Trujillo; VEN (holo). 

Palicourea meridensis Steyerm., Acta Bot 

5 971. J. A. Steyermark & M. Rabe 97090; 1 
Sep. 1966; Venezuela, Mérida, Río Capaz, arriba de 
La Azulita; 2,100- nid m; VEN (holo). 

Palicourea nana Steyer Acta Bot. Venez. 6: 170. 

971. J. A. ds et al. 101055; 16 Jan. 1968; 
Venezuela, Táchira, entre Las Copas y Alto de Fila de 
Tierra Negra; 2,879-2,880 m; VEN (holo). 

Palicourea nubicola Steyerm., Acta we Venez. 6: 168. 

971. J. A. Steyermark & | 97243; 3-4 Sep. 

1966; Venezuela, NM 30 km from Boconó; 2,300- 
2,500 m; VEN (ho 


Palicourea aran cil Wernham var. brevipes 
o He 


Steyer em. ard. 23: 774. 197 
T io 1042; 29 May 1 'enezuela, Miranda, 
Guatire, Hacienda El beds 750-1,200 m 


holo) 


Palicourea perquadrangularis Wernham var. guaya- 


Volume 76, Number 3 
1989 


Taylor 715 
Plants Described by 
Julian A. Steyermark 


nensis Steyerm., Mem. New York Bot. Gard. 23: 774. 
1972. J. A. Steyermark & J “urdack 1172; 27 
Feb. 1955; Ai Bolivar, Torono-tepui, Chimanta 
Massif; 1,880- 0m; depository not designated. 


J. A. Steyermark & L. Nevling 95934; 24 June 1966; 
Venezuela a, Distr. Federal, between Portachuelo and 
Penita (Petaquire) and Carayaca; 1,300- 1,500 m; VEN 
(holo). 


Palicourea p Standl. & Steyerm., Field Mus. 
Nat. Hist PEE Ser. 23(5): 252. 1947. J. A. Steyer- 
mark 41952 26-27 Dec. 1941; Guatemala, Dept. 
Izabal, Cerro San Gil; 1,200 m; F (holo). 


Palicourea puberulenta Steyerm., i Bot. Venez. 6: 
971. J. A. Steyermark 105063; 3 Mar 1971; 
Venezuela, Táchira, 3-8 e A 2, 650- 


2,700 m; NY, VEN (holo). 
a La Steyerm., Acta Bot. Venez. 6: 156. 
o Mem. Ne bi Bot. Gard. 23: 736. 1972. 
y Y el 62225; 19 Apr. 1945; Venezuela, 
Monagas, Quebrada de Pajarral; 600-900 m; F, VEN 
holo 


Palicourea rigida H.B.K. var. iden uid Mem. 

York Bot. Gard. 23: Ll. Williams 

uen 3; 15 May 1939; bah s es Bajo Rio 
ura, Maripa; 80 m; VEN (holo). 

Fu men rigida H.B.K. f. mic rophyll si Steyerm., Mem 

ork Bot. Gard. 23: 732 B. Maguire et et 
al. 447 ^13: 20 Dec. 1959; Brazil; Aue Gerais, Serr 
do a 1,212 m; NY (holo). 

Palicourea tachirensis Standl. & Steyerm., ee 
Bot. 28) 593. 1953. J. A. Steyermark 5744 ; 18 
July 1944; Venezuela, Táchira, Alto de Lirio; 1, i 
2,285 m; F (holo) = Psychotria schlimii Standl., 1972, 

1974 


Palicourea tamaensis (Standl. & Steyerm.) Steyerm. va 
psilophylla Steyerm., Acta Bot. Venez. 6: 153. 1971. 
J. A. Steyermark et at 100940; 15 Jan. 1968; Ven- 

ezuela, cabaceras del Río Quinimari, debajo del Cerro 

lo). 


Las Copas; 2,500-2,700 m; VEN (ho 


Palicourea tepuicola Steyerm., York Bot. 
Ga 972 A. on yermark 74630; 25 
Mar. 1953; Venezuela. Bal , Apacara-tepui, Chi- 


mantá Massif; 400 m; MO (2011490), NY (holo), VEN. 

Palicourea venezuelensis Steyerm., Acta Bot. Venez. 6: 
164. 1971. J. A. Steyermark 99343; 21 E 19685 
Venezuela, Falcón, Sierra de San Luis; 1,300 r 
(holo). 

Palicourea pe boat Steyerm., Mem. New York Bot. Gard. 
23: 734. . S. Vogel 312; 17 Mar. 1956; Colom- 
bia, sde uM 400 m; M (holo). 

Pentagonia grantii d Bol. Soc. Venez. Ci. Nat. 
25(107): 251. 19 M. L. Grant 10567; 2 Nov. 
1944; Colombia, Dent Cundinamarca, Cordillera del 
Monte Cristo, 3 km NW of Mambita; 1,350 m; US 
(holo) 

Pentagonia parvifolia aes Bol. Soc. Venez. Ci. 

25(107): 232. 1964. J. J. Wurdack 2523; 2 
Nov. 1962; Peru, Dept. Loreto, Prov. de Alto Ama- 
zonas, Cerro Campanquiz, Rio Maranon; US, VEN 
(holo). 


Pentagonia wurdackii Steyerm., Bol. Soc. Venez. Ci. 
Nat. 25(107): 233. 1964. J. J. Wur Pss 2435; 26- 
28 Oct. 1962; Peru, Dept. Loreto, Prov. Alto Ama- 


zonas, Pongo de Manseriche, Rio Maranon, 250 m; 
(holo). 
Perama sect. Diperma Kirkbr. & Steyerm., Brittonia 29: 


Perama dichotoma Poepp. & Endl. var. hirsutula Stey- 
erm., Mem. New York Bot. Gard. 10(5): 236. 1963. 
J. A. Steyermark & S. Nilsson 561; 24-25 Apr. 1960; 
Venezuela, Bolivar, vicinity of Camp 150 at km 150 
in valley of Rio Uarama, NE of Luepa; 1,220 m; VEN 


Perama ca nea Poepp. & Endl. var Ted as ees 
Steyerm., . Missouri Bot. Gard. 658. 1987. 
F. d 123: 58; 12-14 Oct. 1984; a T.F. 
Amazonas, Dept. Atabapo, Cerro Huachamacari; VEN 
(holo), MO (3327364). 

Perama galioides (H.B.K.) Poir. var. densipilia Stey- 
erm., Mem. New York Bot. Gard. 10(5): 244. 1963. 
F. Cardona 1630; 18 Sep. 1946; Venezuela, Bolivar, 
Sabanas de Uriman; 400 m; NY, VEN (holo). 

Perama cen s B.K.) dodi var. hispida (H.B.K.) 
Steyerm. f. cinerea Steyerm., Mem. New York Bot. 
Card. 10(5): 243. 1963. B. Maguire et al. 30886; 
14 Jan. 1951; Venezuela, T.F. as, Cerro Mo- 
riche, Rio Ventuari; 800 m; NY oris 

Perama galioides (H.B.K.) Poir. var. intermedia Stey- 
erm., Mem. New York Bot. Gard. 10(5): 243. 1963. 
J. A. Steyermark 59658; | Nov. 1944; Venezuela, 
Bolivar, Ptari-tepui; 1,600 m; F (holo), VEN 


Perama galioides (H.B.K.) Poir. var. galioides f. mac- 


Perama Mti (H.B.K.) Poir. var. galioides f. qua- 
ternata Steyerm., Mem. New York Bot. rn 10(5): 
239. pa 963. LL. William; 15067; 27 Apr. 1942; Ven- 

ipee San Antonio, dis d ced 
121 n m; F (helo), 

Perama galioides UE B K. Poir. var. longiflora Stey- 
erm., Mem. New York Bot. Gard. 10(5): 241. 1963. 

J. J. Wurdack & L. Adderley 43692; 31 July 1959; 
Venezuela, T.F. Amazonas, Sabana de Mayo, Rio Ori- 
noco; 125 m; NY (holo). 

Perama harleyi Kirkbr. & Steyerm., Brittonia 29: 192. 

1977. R. M. Harley et al. 10058: 6 Feb. 1974; Brazil, 
Bahia, Serro do Sincora, 4 mi. S of Mucuge, on road 
from Cascaval by the Rio E 900 m; CEPEC 
(holo), K, NY. 

Perama ica Kirkbr. & bU pola nia 29: 
195. . W. R. Anderson et al. 12 Feb. 
i nu Minas Gerais, Serra 5 Eso nhaco, E 
slopes of Pico do Itambe; 1,550 m; NY, UB (holo), 
VEN 


Perama schultesii Steyerm., Mem. New York Bot. Gard. 
23: 832. 1972. R. E. Schultes & l. Cabrera 20061; 
Apr. 1953; Colombia, Vaupés, near headwaters of Rio 
Kuduyari; 272-300 m; US (holo). 

Perama sparsiflora Standl., Steyerm. & Kirkbr., Brit- 
tonia 29: 196. 1977. Barreto 1134; Brade 4739; 15 

. 1935; Brazil, Minas Gerais, Serra do Cipo; F 
(holo). 

a" duende oo 


: 235. i 


Bol. Soc. Venez. Ci. Nat. 
. J. J. Wurdack 961; 20 June 
i kee Prov. Bongara, 3 km S 
of Pomacocha i 400 m; US (holo). 


716 


Annals o 
Missouri Botanical Garden 


Pittierothamnus Steyerm., Bol. Soc. Venez. Ci. Nat. 23: 
93. 1962, gen. nov. — i ada Standl., 
Pittierothamnus wc Steyerm., Bol. Soc. Venez. 
i. Nat. 23: 92. 19 A. Seyermar 86323; 19 
ie 1960; Va Aragua, acional Henri 
Pittier; 1,300 m; VEN (holo) = “Amphidasya vene- 
zuelensis (Standl.) Steyerm., 1972. 

se Ai aie vg ipiens Woodson & Steyerm., Amer. J. 
Bot . 1952. El. Willia ams 14587; 2 Mar. 
1943; ha ide T.F. Amazonas, San Carlos pg Rio 
Negro; 100 m; A, F the lo), MO, NY, RB, US, VEN. 

e o rant e kei Steyerm., ay a Bot. 39: 422. 

9 ucke 1005; 25 Aug. 1942; Brazil Ama- 
zonas, n Grande; INPA, K, Fine rere NY, 
R, RB, US (holo). 
Platycarpum ea Steyerm., Mem. New York 
ipm Gard. 10(5): 258. 1963. B. Maguire et al. 40666; 
0 Feb. 1955; Guyana, between n Chinowieng ar id Chi 
Chi landing, Mt. Ayanganna, d aima Mountains 
1,000 m; GH, NY (holo), U, 
Platycarpum maguirei E Mor New York Bot. 
ard. 10(5): 255. 1963. B. Maguire et al. 42597; 5 
Jan. 1958; Venezuela, T.F. Amazonas, Cerro Neblina; 
30-700 m; F, GH, K, MICH, MO (1782963), NY 
(holo), RB, U, US, VEN. 

Platycarpum orinocense Humb. & Bonpl. var. grandi- 
florum Steyerm., Mem. New York Bot. Gard. 10(5) 
254. 1963. R. S. Cowan «€ J. J. Wurdack a 
15 Feb. 1951; j 
Cerro Parú, Serranía Pari: 200 m; NY (holo), US, 

'E 

Platycarpum Rr dE Woodson & Steyerm., 
Amer. J. Bot. 39: 423. 1952. F. Cardona P. 860; 2- 
D Aug. e ente la: Bolivar, summit of Cerro 

r Río Paragua; 700 m; F, MO (holo— 
1309009), NY, US, VEN. 

Platycarpum rugosum Steyerm., 

Gard. 10(5) 257. 


Mem. New York Bot. 


Platycarpum schultesii re ex Schultes, Bot. Mus. 
Leafl. 17(3): 96. 1955 . R. E. Schultes de 1. tea 
1992 29; June 1953; Colombia: Comisara del Vau 

ana ee (tributary of Rio Vaupés); 200 n m; 
F (holo), GH, 

Pogonopus specs da E, subsp. sandwithianus 
Steyerm., Acta Bot. Venez. 6: 110. 1971. H. H. Smith 
100; 20 Aug. 1898- 1899; Colombia, Dept. Magda- 
lena, Santa Marta, near Bonda; 45 m; NY (holo). 

Posoqueria ere Steyerm., Mem. New York Bot. 

17(1): 3 1967. Ll. Williams 14525; 2 Feb. 
1942; Le T.F. Amazonas, San Carlos de Río 
Negro; N 2 — 

h Altsonii Steyerm., 
Bot. eee 23: 9558 2, ser. nov. 


Mem. New York 


Psychotria L. ser. Aneurophyllae Steyerm., Mem. New 
ork Bot. Gard. 23: 545. 1972, ser. nov. 

Psychotria L. ser. i gra Steyerm., Mem. New 
ork Bot. Gard. 23(1): 6 1972, ser. nov. 

Psychotria L. ser. Crococ uos Steyerm., Mem. New 
ork Bot. Gard. 23(1): 593. 1972, ser. nov. 


Psychotria L. sect. Didymocarpos peu Mem. New 
York Bot. Gard. 23: 516. 1972, sect. 
. sect. Duriflia si hag die Men. "Nb York 
Bot. Gard. 23: 705. 1972, sect. nov. 
Psychotria L. subg. Heeropssehuri Steyerm., Mem. 
New York Bot. Gard. 23: 484. 1972, subg. nov. 


Psychotria L. ser. Mollirami Steyerm., Mem. New York 
Bot. Gard. 23: se 1972, ser. nov 
Psychotria L. sect. ii Steyerm., Mem. New 
ork Bot. Gand. "231y 6 972, inci; nov 
nd L A Peale ad. Steyerm., Mem. 
w York Bot. Gard. 23(1): 593. de sect. nov 
Psychotria adderleyi d Mem 
ard. 23: 313. 1972. B. Maguire. et al 00073; 4 
Nov. 1965; Brazil, Amaronas, Rio Cauaburi (Rio Ne- 
gro) near base of Cachoeira Caranguejo; NY (holo), 
'E 


P byehotris Miei Steyerm., Mem. New York Bot. 
Gard. 23(1): 599. 1972. B. Maguire et al. 29851; 5 
Dec. 1950; Venezuela, T.F. rou Rio Cunucu- 
numa, Cerro Huachamacari; 1,300-1,600 m; NY 
(holo). 

Psychotria adpressipilis Steyerm., Mem. New York Bot. 
ede 23(1): 648. 1972. R. Spruce 4008; 1855- 

; Peru, Tarapoto; NY (holo). 

d. agostinii Steyerm., 
Gard. 23: 551 . J. A. Steyermark & G. Agostini 
8; 4 Sep. 1960; Venezuela, Aragua, Parque Nacional 
Henri Pittier, Pico Periquito; 1,250-1,600 m; VEN 
(holo). 

Psychotria a ers is Steyerm., Field Mus. Nat 
Hist., eed 23(1): 23. 1943. P. C. Standley 80900; 
Dec. 0; Guatemala (e Chimaltenango, between 
E and San Martin Jilotepeque; 1,500- 
1,700 m; olo). 

P sychotria aligera Steyerm., Mem. New York Bot. Gard. 


Mem. New York Bot. 


S 
— 


572.1972 
1962; Venezuela, Bolivar, 
yuk; NY (holo), VEN 

Psychotria alloantha Steyerm., Wr es 36: 154-160. 

84; J. J. de Granville 3127; 16 July 1979; French 
Guiana, 4 km au sud du Pic E (30 km ENE de 
Saul); CAY, P, VEN (h. alo). 

Psychotria alticola eph Vbi 9: 
Humbert 26739; 6-8 ( 92: doit. 
Sierra del Norte; 2 en 2 Po m; P (holo). 

P sychotria altorum Standl. & Mr Field 

23(2): 86. 

34302; Jan. 1940; kc es Dept 

Montana Chicharro, SE s 


Alto Río Cuyuni, Rio Uis 


81. H. 
[UST 


Mus. Nat. 
Steyermark 

t- Quezaltenango, 

rcd Santa Mar 


P ahis deer Steyerm, Mem. New York Bot. 
ard. 17(1): 431. 1967. J. A. Steyermark 93230; 2 

May 1964; Venezuela, Bo liy r, Auyan- i 1,690 m; 
VEN (holo) = P. pan ae Standl., 

Psychotria amplinoda Steyerm. var. VA Kn Stey- 
erm., Mem. New York Bot. Gard. 17(1): 4 iln 
. S. Tillett et al. 45105; 12 Aug. 1960; 
Mt. ipo Thompson o 1,418 m; NY (holo) 
— P. tapajozensis Standl., 


Psscharia o aura Ann. Missouri Bot. 


Gard. 7 988. G. Davidse 27651; noice July 
1984; Avenir T.F. Amazonas, Dept. Rio Negro, 
3393677) 


lower part of Rio Baria; 80 m; MO (holo 
VEN. 


P Poir anceps H.B.K. var. robustior Steyerm., Mem. 

k Bot. Gard. 23: 466. 1972. J. A. Steyermark 

& M. Rabe 96039; 7 Aug. 1966; Venezuela, Sucre, 

Peninsula de Paria, Quebrada Nivardo; 700-750 m; 
VEN (holo). 

Psychotria WOCHE een Steyerm., Mem. 

Bot. Gard. 23: 547. 


New E 
12. J. A. Steyermark & G. ! 


Volume 76, Number 3 
1989 


Taylor 717 
Plants Described by 
Julian A. Steyermark 


Bunting 97655; 11 Mar. 1967; Venezuela, Yaracuy, 


Cerro La Chapa; 1,200-1,360 m; VEN (holo). 
Psychotria antenniformis Steyerm., Brittonia 33: 3 
1981. J. J. de Granville C. 159; 12 Mar. 1971; French 


Guiana, Monts Atachi Bacca, Camp no 1 et Alentours, 
foret sur la rive droite du Maroni au reiveau de l'ilet 
Assadam Tabiki; P (holo). 

Psychotria apoda Steyerm., Mem. New York Bot. Gard. 
23: 668. 1972, nom. nov 

Psychotria appuniana Steyerm, Mem. New York Bot. 
Gard. 23(1): 660. 1972, nom. nov. 

Psychotria mamaria E a Mem. New York 
Bot. Gard. 23: 588. 1972. /3 y n es et al. 

t amp 12 


d ire M Steyerm., Mem. New York Bot. 
Gard. 2 Led. [penc 047; 14 Sep. 1946; 
eus Aragua, [ee Nacional; VEN (holo). 

Psychotria arenaria Standl. Pe Bot. 
28(3): 594 ; 6 Nov 
1944; Venezuela, Bolivar, pes 2m 2, a " F bolo] 

eratantha Standl., 

Psyc des argoviensis B. i "Mem. New York Bot. 

23: 500. i nom. nov. 
m aud 
1971. Fa 


Psychotria  aroensis nn Mem. N 
Gard. 23(1): 4 12. L. Aristeguieta & F. Pannier 
1952; 6-7 July Ub Venezuela, Yaracuy, Sierra de 
Aroa; VEN (holo). 

Psyc t aubletiana Mi dip 
Gard. 23: 694. 1972, nom. 

Psyc ir aubletiana e var. andina Steyerm., 

lem. New York Bot. Gard. 23: 695. 1972. J. A. 
Steyermark et al. 08683; 22-24 May 1967; Vene- 
zuela, Paramo de Tamá, near Quebrada Buena Vista; 
2,300-2,450 m; VEN (holo 

Psychotria aubletiana Seya: var. andina Steyerm. 
f. p e uae Mem. New York Bot. Gard. 

97 238; 19 June 1953; 


Mem. New York Bot. 


2 200 m; VEN 

Psychotria d ina: var. cacuminis (Stey- 
erm.) Steyerm. f. furca d a Mem. New 
York Bot. Gard. 697. 1972. J. . 4. Steyermark 
75830; 20 June TUN bium Bolivar, Apacara- 
epui; 2,000-2,150 m; F, NY, VEN (holo). 

Psychotria Moria Steyerm. var. centro-americana 
Steyerm., Mem. New York Bot. Gard. 23: 694. 1972. 
L. O. Wi ilic ms & A. M 11237; 30 Dec. 1962; 
Honduras, Dept. Morazán, La Tigra; 1,800 m; F, NY 
(holo). 

Psychotria aubletiana Steyerm. var. WO Me Steyerm., 
lem. New York Bot. Gard. 698. 1972. ljjasz 
425; 15 May 1964; Viena Noc entre la India 
y El Moro; 2,430 m; MAR (holo) 

Psychotria aubletiana Steyerm. var. o i Steyerm., 

Ann. Missouri Bot. Gard. 71: 1176. 1984. R. L. Lies- 

ner & A. Mind eg 10107; 5 s 900; Venezuela, 
Yaracuy, Sierra de Aroa, 15-20 km NW of Cocorote 
on road to ront MO (2777271), VEN (holo). 

Psychotria avia Standl. & Steyerm., Fieldiana Bot. 28(3): 
596. 1953. J. A. Steyermark 58056; 26 Aug. 1944; 
Venezuela, T.F. Amazonas, Cerro Duida; 1,095-1,520 
m; F (holo) = Rudgea avia (Standl. & Steyerm.) Stey- 
rm., 1972. 


= 


Psychotria avilensis Steyerm., Mem. New York Bot. 

Gard. 23(1): 450. 1972. B. Trujillo 1374; 25 June 

i. Venezuela, Distr. Federal, entre Galipan y Los 
Venados; MAR (holo) 

Psychotria ayangannesis Steyerm., Mem. New York 
Bot. Gard. 23(1): 594. 1972. S. 5. Tillett et al. 45059; 

1960; Guyana, Thompson Camp, Mt. 
Aina 1,370-1,525 m; NY (holo), VEN. 

Psychotria a Steyerm. . New York Bot. 
Gard. id 542 . B. Maguire et al. 37483; 4 
Feb. Ped zonas, Piedra ur 
rumeri; EN S not puros 

E egre bernardii pae Aem. New York Bot 
Gard. 23(1): 450. 1 cp. | 1304; 25 Juge 
1954; Venezuela, M Carrizal; 1,400 m; NY (holo). 

Psychotria berteriana DC. subsp. venezuelica Steyerm., 
Mem. New York Bot. Gard. 23: 535. 1972. H. Pittier 
14010; 28 May 1937; Venezuela, Aragua, Parque 
Nacional Henry Pittier, Rancho Grande; 1,200 m; VEN 
(holo). 

Psychotria pa cur & Steyerm., Fieldiana Bot. 
28(3): 596 Holt & E. R. Blake 743; 
2 1931; E Gr Amazonas, Cerro Yapa- 

na; 100 m; F (holo). 

en ieee e Standl. & Steyerm. e pu- 
b y em. New York Bot. Gard. 693. 
1972. L. Ruiz Terán 2980; 6 Apr. 1966; alió 
Mérida, Distr. Arzobispo Chacón, Mun. Libertad, Pe- 
ramo de Canagua; 2,187 m; VEN (holo). 

Psychotria boyanii Steyerm., Mem. New York Bot. Gard. 
23(1): 659. 1972. S. S. & C. L. Tillett 43954; 1 July 
1960; Guyana, Merume Mountains, Portang River; 
1,140 m; NY (holo 

Psychotria brazoi Steyerm., Mem. New York Bot. Gard. 
23(1): 610. 1972. N. T. Silva & U. Brazao 60633; 
15 Dec. 1965; Brazil, Vénus above Igarape Anta, 
Pico de Neblina; 1,400-1,800 m; NY (holo). 

Psychotria n Steyerm., O New York 
Bot. Gard. 23(1): 643. 1972, nom. 

Psychotria breweri aed Acta Bot. y m 14(3): 
251. 1984. J. A. Steyermark et al. 126381; 10 Feb. 
1982; Venezuela, T.F. Amazonas, Cerro Duida; 1,230 
m; NY, VEN (holo). 

e reri buntingii Steyerm., Ann. peces EE Gard. 
o 1987. G. S. Bunting et al. 10876; 25 

3 Mar. RC Venezuela, Zulia, Dept. Perijá a, alrede- 
dotés de la Estación rica a Aricuaisá-Pie de Me nte; 
100-250 m; JBM, MO (holo— 3191936), V 

Psychotria saleiphula qd 
Gard. 23: 471. 1972. J. A. S 
July 1967; Venezuela, Falcón, Sierra de San bus 800- 
900 m; depository not designate 

Psychotria calviflora Steyerm., Mem. w York Bot. 

yard. 23: 686. 1972. Ll. Williams 1 2992; 18 May 
1940; Venezuela, T.F. Amazonas, S of Puerto Aya- 
cucho; 100 m; VEN (holo). 


sychotria canaguensis Standl. & Steyerm., 


Fieldiana 


a f. peuples (Standl. 8 Steyerm.) Stey- 
erm., i 

Psychotria eM Steyerm., Mem. New York Bot. 
Gard. 23(1): 6 1972. N. T. Silva & U. Brazao 
60880; 24 E 1966; Brazil, Comissao de 2 
Marker BR-1, Serra Pirapucú, Rio Negro, Rio Ca 
aburi; 1,250 m; NY (holo). Note: This specimen is ven 


718 


Annals of the 
Missouri Botanical Garden 


holotype for Pagameopsis maguirei Steyerm. subsp. 
neblinensis Steyerm. var. pirapucuensis Steyerm. 
however, the dates AH. elevation differ. 

Psychotria capitata Ruiz & Pavon subsp. amplifolia 
(Raeschel) Steye be f. tomentosa Steyerm., Mem. Nev 
York Bot. Gard. 23(1): 623. 1972. A. de La Rue 13; 

1948; French Guiana, Maripa, 


15 Dec. vallée de 
l'Oyapock; P (holo 

Psychotria capitata "Bf & Pavon n oe 
(Benth. ) Steyerm. var. Bedale gee Stey n. 
New York Bot. Gard. 23(1): 6 E 


G. Britton 2104; 4 Mar. e Denidad. Valencia; 
NY (holo). 

Psychotria capitata Ruiz & Pavon f. d Stey- 
erm., Mem. New York Bot. Gard. 23(1): 617. 1972. 
S. Vogel 290; 12 Mar. ; Colombia, Río Zanza, 
Sierra Macarena; M (holc a 

Psyc e caquetensis Steyerm., Mem. New York Bot. 
Gard. 23(1): 653. 1972. S. Vogel 81; 12 May 1956; 
O Intendencia del ea near Leguizamo 

caya); 150 m; M (holo), N 

ib carrenoi Steyerm., Fl. Juss. 9(3): 1658. 
974. J. A. Ste, yermark et al 109428; 24 Feb. 1974; 
Venezuela, Bolivar, Cerro Jaua, Meseta del Jaua; 1,800 
m; VEN (holo). 

Psychotria pr Steyerm., Mem. New York Bot. Gard. 

B. Maguire et al. 60511; ec 
2,425 m; 


23: 7 

1965; ber poenas Serra de Neblina; 

NY (holo). 

Psychotria cerronis Steyerm., Mem. New York Bot. Gard. 
23: 886. 1972. J. A. Ste yermark 97852; 22-27 Mar. 
1967; Venezuela, Bolivar: Meseta de Jaua, Cerro Jaua; 
1,922-2,100 m; VEN (holo). 

— chapensis Steyerm., Mem. New York Bot 

J. A. Steye rmark etal. 100217; I 
7, Cerro La Cha- 


Gard. 23: 552. 1972. 
9 P <a 1967; Venezuela, Yar 
pa; 1,200-1,400 m; VEN (holo 
Psyc ex clivorum Standl. & Sterea Field Mus. Nat. 
Hist., Bot. Ser. 23(2): 87. 1944. 4. F. Skutch 2073; 
7 Jan. 1935; ba des. Dept. Suchitepequez, Finca 
Moca; 1,020 m; F (ho 
Psychotria a cip H. B. K. 


2 


bsp. pd Stey- 


erm., Mem w York Bot. Gard. 23: 1972. Re. 
Sc homburgk 282; 1841-1842; b on Pirara; P 
(holo). 


Psychotria costanensis Steyerm., bing New York Bot. 
Gard. 23 3(1): 658. 1972, nom 

Psychotria costanensis Steyer: pore pri Si 
erm., Mem. New York Bot. Gard. 23(1): 6 1972 
L. Bernardi 1296; 23 June 1954; bud edis 
El Carrizal, del Rincón; 1,450 m; NY (holo). 

Psychotria pedis Steyerm. var. Imc Stey- 
erm., ven, ew York Bot. Gard. 23(1): 6 1972. 
TA a yermark 95314; 29 Mar. ee Ven- 
ezuela, Carabobo, S of Borburata, entre Quebrada no. 
2 y Quebrada de los Verros; 750-900 m; MO 
(2270193), VEN (holo). 

Psychotria coussareoides Standl. var. ciliata Steyerm 
Mem. New York Bot. Gard. 23: p 1972. J. A. 
Ste yermark el a 92929; 10 Jan. 1964; Venecia. 

namo; 950-1,000 m; NY, VEN (holo) 

ar. coussareoides, 1974 

Psychotria crassa Benth. f. diis Steyerm., Ann. Missouri 
Bot. Gard. 71: 331. 1984. J. 4. Steye 

1983; Vene 

o de Chimantá; 2,000 m; 

Pob à crassa Benth. f. angustior Steyerm., Ann. 


Missouri Bot. Gard. 71: 331. 1984. J. A. Steyermark 
al. 128992; 1-9 Feb. 1983; Venezuela, Bolivar, 
Distr. Piar, Macizo de Chimantá; 2,450 m; VEN (holo). 
Psychotria dps Steyerm., ida m. New York Bot. 
Gard. 23: 690. 1972, nom. 

Psychotria abdis Standl. & CN Fieldiana Bot. 
28(3): 598. 1953. J. A. Steyermark 60380; 20-21 
Nov. 1944; TAA Bolíva ar, Quebrada O-pau-ma; 
1,065-1,220 m; F (holo) = P. ea Ruiz & Pavon 


= 


subsp. inundata ipt ob var. roraimensts 
(Wernham) Steye 197 

Psychotria KA Ss Steyerm. Brittonia 36: 155. 
1984. C. Sastre 6463; 27 Dec ; French Guiana, 
fleuve Maroni: Pompidou- Papaichton; CAY, P, VEN 
(holo). 


" 


P pepe “davidsae” Steyerm., Ann. Missouri Bot. 
: 1177. 1984. J. A. Steyermark et al. 97; 
10 May 1980: Nemmel, T.F. Amazonas, Distr. Atures, 
along Rio Cataniapo, N side of river at dam site; ` MO 
(2899787), VEN (holo). Note: P. ia is on the 
label, which is spelling that conforms with 1988 ICBN. 
egi Er: Steyerm., Mem. New York Bot 
Gard. 2 . 1972. J. 4. Steyermark & G. W Al 
Boer 0 13 Nov. 1967; Venezuela, Yaracu uy, 
Sierra de Aroa, Cerro Negro; 1,100-1,250 m; VEN 
(holo). 
Psyc TOME deflexa DC. es cubensis A us Mem. 
rk Bot. Gard. 505. 1972. J. . Hi d 
8805; 6-18 Feb. 1911: ca Oriente, La P 
holo). 
Psychotria dudleyi Steyerm., Phytologia 31(2): 
5. TR. Dudley 13089; 22 July 1969; Peru, Dept. 
ados: SW slope of Rio Llulla Pichas watershed, on 
the ascent of Cerros del Sira; 1,400 m; NA (holo) 
Psychotria dunstervilleorum Steyerm., Mem. New York 
Bot. Gard. 23: 702. 1972. J. A. Steyermark et al. 
100762; 12 Jan. 1968; Venezuela, Táchira, Río Quin- 
imari, SE of Santa Ana; 2,500-2,800 m; VEN (holo). 
Psychotria duricoria Standl. & Steyerm., Fieldiana Bot. 
28(3): 599. 1953. J. 4. Steyermark 58048; 25-26 
Aug. 1944; Venezuela, T.F. Amazonas, Cerro Duida; 
1,665 m; F (holo 
Psychotria duricoria 1 Steyerm. x i epe Steyerm., 
Mem. New York Bot. Gard. 2 . & C. 
Maguire 35092; 8 Feb. ps Venema, T.F. Ama- 
zonas, Serrania Yutaje, Rio Manapiare, near Camp 
Yutaje; NY (holo). 
Ecce essa Steyerm., Ann. par Bot. 
Gard. 7 da . Liesner 19595; 5 Nov. 


997 


adl. 


1985; Piae la, Boliva 0-3 km W Ree E Polo (8 
km W of El Pauji); 650- 800 m; MO (holo — 3393682), 
VEN. 

Psychotria embirensis oo rm., Mem. New Yor 


Gard. 23(1): 656. 1972. B. A. K rukoff 4940; ds i 
1933; Brazil, Amazonas, basin of Rio Jurua, tributary 
of Rio Tarauaca, near mouth of Rio Embira; NY (holo). 

Psychotria everardii Wernham subsp. Ware Stey- 
erm., Mem. New us Bot. Gard. 23 Es 72. B. 
Maguire et al. 42266; 1-2 Dec. 1957; reli 
T.F 


Amazonas, i de la Neblina; NY (holo). 


k Bot. Gard. un: 540 72. B. 
Maguire & L. Politi 27864; 23 Dec. 1948; iati 
T.F. Amazonas, Cerro Sipapo (Parana): 1,400 m; NY 
(holo). 

E ye dien s ii Steyerm., Mem. New York Bot. 

ard. 23: 554. 1972. B. Trujillo & Fernandez 766; 


Volume 76, Number 3 
1989 


Taylor 719 
Plants Described by 
Julian A. Steyermark 


Jan. 1950; Venezuela, Yaracuy, Aroa, Pitiguao; 1,600 
holo 


;M 
Psychotria fimbriflora Steyerm., Brittonia 33: 386. 1981. 
B. S. Pena s.n.; 14 Nov. 1975; Brazil, Amazonas, 
Operacao four Ponto 86-SC-18-XD; VEN (holo). 
Psychotria ca Steyerm., Brittonia 33: 387 
1.0. C. Nascimento et al. 20; 2 Apr. 1975; Brazil, 


usc Vaupés, Rodovia Peut Norte; IAN, 
VEN (holo). 
Psychotria fosbergii Steyerm., Mem. New York Bot 


Gard. 23(1): 630. 1972. F. R. Fosberg & N. C. Pasce 
21810; 17-18 Apr. 1944; Colombia, Dept. Santander, 
Cordillera Oriental, Jordan; 1,100 m; US (holo). 
ica galbaoensis Steyerm., Brittonia 33: 387. 
1 . J. de Granville 2383; 11 Mar. 1975; French 
De Pente nord-est des Monts Galbao, a 10 km au 
SE de Saul; 500-600 m; MO (2798770), VEN (holo). 
Psychotria glabristipula Steyerm., Mem. New York Bot. 
. J. A. Steyermark 98939; 16 
July 1967; Venezuela, Falcón, Sierra de San Luis; 
1,450 m; VEN (holo). 
Psychotria glandulicalyx Steyerm., 


Mem. New York 


rono-tepui, Chimantá Massif; 2,165-2,180 m; F, MO 
(2012057), NY, VEN (holo). 

Psychotria glandulicalyx Ps f. be in Stey- 
erm., Fl. Venez. 2(3): 1364. 1984 A. Steyermark 
106037; : E oa 45 km al N de 
las cabeceras del Rio Orinoco; 1,250 m; VEN (holo). 

Psychotria glandulicalyx Steyerm. subsp. y nsis 

New York Bot. Gard. 23: 538. 1972. 

Mag . 1951; Poenis 
T.F. Amazonas, Cerro Camani; 1.500 m; NY (holo). 

di e paca Steyerm. a e 

ork Bot. Ga Pd 538. 1972. 
J. A Steyermark 08057: . 1967; s 
zuela, die Cerro Jaua; 1,922- 2.100 m; depository 
not designat 

Psychotria pee llei Steyerm., Brittonia 33: 389. 1981. 
J J. de Gr 1975; French Guiana, 

MO 


air 2353; 3 Mar 
aul, sommet des Monts la Fumée: 410 m; CAY, 

T 98769), NY, P, VEN (holo). 
Psychotria guadalupensis (DC.) Howard subsp. vene- 
y New York Bot. Gard. 23: 570. 
ag cia 
Venezuela, Aragua, Parque Nacional Henry Piter. 

Rancho Grande; VEN (holo). 

Psychotria aa ii Steyerm., Ann. Missouri Bot. Gard. 
74: 109. 1987. F. Guanchez & E. Melgueiro 3644; 
15 Nov. 1984; Venezuela, T.F. Amazonas, Rio Sipapo 


junto a la communidad indigena de la etnia Piaroa; MO 
(3327369), TFAV, VEN (holo). 
Psychotria heterocarpa Standl. & Steyerm., RP 


Bot. 28(3): 600. 1953. J. - P. 60191; 15 
Nov. 1944; Venezuela, Bolíva Eh ii; to 
1,980 m; F (holo) = P. Rien Rui on subsp. 
E a (Benth.) Steyerm. var. roraimensis (Wern- 
ham) Steyerm., 1972. 

Psychotria [bet Steyerm., Mem. New York Bot 
Gard. 23: 714. 1972. N. T. Silva & U. Brazáo 60607: 
14 Dec. 1965; o Amazonas, Rio Maturaca, Rio 

up to Pico de Neblina; 1,350- 


Sororopan 


Psychotria Vade dn agi (Willd. ex Roemer & 
Schultes) Muell. Arg. var. celsa Steyerm., Mem. Ne 
ork Bot. Gard. 23(1): 608. 1972. H. S. Irwin et al 


14771; 13 Apr. 1966; Brazil, Bahia, Planalto do Brasil, 
Rio Corrente drainage, Rio Piau; depository not des- 
ignated. 
Psychotria VE e oa (Willd. ex Roemer & 
Schultes) Muell. Arg. var. f. pubescens Steyerm., Mem. 
972. R 


1962; G 

;orge, Kaieteur Plateau, near Kaieteur Falls; 425 m; 
US, VEN (holo) 

Psychotria horizontalis Sw. var. Dabo de Steyerm., 
Mem. New York Bot. Gard. 23: 472. I. H. 
Smith 393; May 1898-1899; a Dent Mag- 
dalena, Santa Marta, near Masinga Vieja; 300-450 
m; NY (holo). 

Psychotria ae Steyerm., 
Gard. 23 T2.. J As 
s ion Venaouélá. Sucre, Peninsula de 

o de Humo; 800-1,000 m; VEN (holo). 

Bon hein hyalina Steyerm., as New York Bot. Gard. 


Mem. New York Bot. 
Steyermark 95075; 5 
Paria, Ce- 


~ 


23: p 1972. J Steyermark 95246; 29 
Mar. 1966; Ven ezuela, esce Rio San Gian, S of 
eei 0-900 m; MO (2223248), VEN (holo). 

Psychotria edilicia Muell. Arg. subsp. V eg 
Steyerm., Mem. New York Bot. Gard. 23(1): 645. 
1972. J. Murca Pires 122; 30 Nov. 1946; Brand. 
Amazonas, Maues; NY (holo). 


Psychotria irwinii Steyerm., Mem. New York Bot. Gard. 
23: 453. 1972. S. Irwin et al. 55706; 13 Sep. 
1963; Suriname, Wilhelmina Gebergte; 350 m; NY 
(holo), VEN. 

Psychotria peers ^W ee Mem. 
Gard. 23: 889. A. Steyermark 98098; 

7 Mar. 1967; dos iN Bolivar, Cerro Jaua; 1,922- 
an m; VEN (holo 

Psychotria fe TEE Steyerm. 28(3): 

5: Steyermark : . 1944; 
Venezuela, B a, Gran rl “1,065 m; "E (holo) 
— P. boliviana Standl., 1972. 

Psychotria lateralis Steyerm., Brittonia 33: 391. 1981. 

. J. de Granville 2360; 9 Mar. 1975; French Guiana. 

sommet des Monts Galbao, a 10 km au SE de Saul; 
650 m; VEN (holo 

Psychotria leiantha Steyerm., 


New York Bot. 


22- 


~ 


pr As 


Mem. New York Bot. 
9 
18-19 Dec. 1957; , T.F. 
de la Neblina; 1,900-2,000 m; MO (3194441 
(holo), V 
Psychotria lilacina Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(5): 252. 1947. J. A. Steyermark 
46564; 14 July 1942; Guatemala, Dept. Huehuetenan- 
go, Cerro Huitz; 1,500-2,600 m; F (holo). 
Psychotria liogieri Steyerm., Mem. New York Bot. Gard. 
3: 700. 1972, nom. nov. 
Psyc hotria e Steyerm., 
,ard. 23: 586 . B. Maguire et « 
. Am 


azonas, 


NY 


— 
E 


Mem. New York Bot. 
ıl. 29786; 29 
Cerro Hau- 

chamacari; 600 m; NY (holo). 
Psychotria lupulina Benth. in Hook. subsp. rhodoleuca 
. Ar . var. maypurensis (Humb. & 


A. Steyermark 86861; 5 Aug. 1960; Ven- 
ndun Bolív ar, Cerro Cotorra (El Vigia); 350 m; VEN 
(holo). 

Psychotria macrophylla Ruiz & Pavon subsp. anomo- 

Schumann & Donn. Sm.) Steyerm. f. js 


Mem. New York Bot. Gard. 23 


thyr rsa 
tella Steyerm., 


720 


Annals of the 
Missouri Botanical Garden 


1972. C. Játiva & C. Epling 735; 
Ecuador, Prov. Esmeraldas, San € Pd (holo). 
Psychotria maguireorum Steyerm., Mer 
Bot. Gard. 23(1): 625. 1972, nom. nov. e Ma R. 
Spruce 3440; 1853-1854; Venezuela, Amazo- 
nas, Vasiva et Pacimoni; depositor a not designated. 

Psychotria manaraeana Steyerm., ot. Venez. 6: 

.197 0: 

zuela, Lara, Distr. Palavecino, La Tomi la Loma 
Redondo: 1,100-1,200 m; VEN (holo). 

Psychotria rae nsis Steyerm., Mem. New York Bot. 
Gard. 23: 516. 2. E. Ule 5386; Jan. 1901; Brazil, 
Amazonas, Val “HBG holo). 

P peus ipd diee Ste yerm., Mem. New York Bot. 
na 3:578.19 . T. Silva & U. Brazáo died 

pum 1966; in i Áiiisends. Rio ca, be- 
tween eds Salesiana and Serra Pirapucu; 50- -400 
m; NY (ho 

POL megacephala P cua Mem. New York 

. B. A. Krukoff 0858; 12 

.19 i ae pea Rio Livramento, 

Mun. ae NY (holo). 

Psychotria meridensis Steyerm., 
Gard. 23: 531. 1972. 

1944; Venezuela, 


2.745 
- Í 


— 


d. Z3: 


Mem. New York gr 
Steyermark 56014; 
Mérida, above La Isla, a 
m; F, NY (holo), VEN 
Psyc aga merumensis wa ., Mem. New York Bot. 

Gard. 574. 1972 Tillett 43996; 4 July 

1960; Guyane, Partang a Merume Mt.; 1,140 m; 

NY (holo). 

Psychotria microbracteata Steyerm., Brittonia 33: 391. 
1981. J. J. de Hie ua ee 9 Mar. 1975; French 
Gi e pe des Monts Galbao, a 10 km au SE de 
Saul; , MO (2 198772), P, VEN (holo). 

ED nicrudon (DC.) Urban var. mananahi 

, Mem. New York Bot. Gard. H. Pittier 7911 

- 1916; Koarer Carabobo, Hacienda de Cura, 

San Jo ; VEN (holo). 

Psychotria minari apr E Steyerm., Field Mus 
Hist., Bot. Ser. 23(5): 2 1947. J. A. Stey juu 
43106; 21 Jan. 1942; Conia Dept El Progreso, 
Sierra de las Minas ra and summit of 
Volcán Siglo; 2 .000- 3, 000 1 m; F (hok »). 

Psychotria moroidea Steyerm., i New York Bot. 
Gard. 23: 485. 1972 


e 


near 


nom. no 

Psyc are multiramosa eS "s rm., "Mem. New York Bot. 
Gard. 23: 568. Tillett et al. = 9; 10 
Aug. qunm Guyana, Tm Audet 1,418-1,525 m; 
NY (holo). 


Psychotria muscosa (Jacq.) 


ue 


Stey erm. up brev ie 
23 ‘ 19 


Stey verm., Mem. New York Bot. € : 672. 
E Broadway 9823; 10-26 Jan. 1929: iet 
heihts of Aripo; NY (hole): 


Psychotria oe Steyerm., se New York 

Bot. Gard. 23: 512 2. J. Ewel 11 1; 18 Apr. 1964; 
, Amazonas, cerca ade la Neblina; 400 m; MAR, 
lo). 


a. 


Psychotria nora Steyerm., Mem. New York Bot. Gard 
23: 703. 1 . Steyermark et al. 101030; 16 
Jan. Vn Mis Táchira, SE de Santa Ana, entre 
el Páramo de Pata de Judio y el Páramo de Cobre; 
2 600- 2 2.800 m; VEN (holo). 
Psychotria occide do Steyerm., 
Gard. 23: 545 
May ; V Sad 


896 
Orinoco; NY (holo). 


9 
Ú < 


Mem. New York Bot 
. H. Rusby & R. Sq 


Terr. 


uires s.n.; 
Delta Pe uro, lower 


12 July 1964; 


Psychotria officinalis ecd Sandw. subsp. wilhelmi- 
nensis Steyerm., Mem. N ot. Gard. 23: 615. 
. B. Maguire et d. 5444 2 Aug. 1963; Sur- 
iname, Juliana Top, Wilhelmina ee 1,200 m; 
NY (holo). 

Psychotria oiapoquensis Steyerm., Mem. New e» Bot. 
Gard. 23: 647. 1972. S. Irwin et al. 47393; 7 

Aug. 1960; Brazil, Terr. Amapá, Rio DUM NY 
(holo). 

Psychotria pachecoana Standl. & PVT Field Mus 
Nat. Hist., Bot. Ser. 22(3): 205. 1940. P. C. dla 
69922; 30 Mar. 1939; Guatemala, Dept. Baja Vera- 

, N of Santa Ros 1,650 m; F ere 

Boo D. ia ie 

i Gard. 


. New York 
. B. Mus et al. 40573; 
» Feb. Tus Co Pakaraima aaa Mt. 
bisce 1,000-1,500 m; NY (ho i 
Psychotria pallidinervia Steyerm., Mem. 


ew York Bot. 


Gard. B: e oni et p 41990; 1 
Nov. 1957; Neues. T.F. azonas, Cerro de la 
Neblina; 500-700 m; NY nes 


Psychotria pariensis Steyerm., Mem. New York Bot. 
Gard. 23: 513. 1972. J. 4. Steyermark 95019; 4 
Mar. 1966; Venezuela, Sucre, Cerro de Humo; 1,250- 

3 m; VEN (holo). 

Psychotria parimensis Steyerm., Mem. New York Bot. 
Gar 691. 1972. F. Cardona P. 133 33; 11-23 
Mar. 1946; Venezuela, T.F. Amazonas, Sierra Parima; 
550 m; US, VEN (holo). 

Psyc hotria puro ibractea Steyerm., Mem. New York Bot. 
es : 581. Steyermark & S. Nilsson 

; 14 Apr. 1960; enema la, Bolívar, Cerro Vena- 
mo; EN 100- Wo m; NY (holo), VEN 

b ena? d Standl. & Steyerm., Fieldiana Bot. 
que ) 601. 195: A. Steyermark 56411; 9 May 

944; ae Mérida; Que ^brada de Los Salichales; 

1 950-2,135 m; F (holo 
Psychotria patria Standl. & Ste eyerm. var. tov arensts 
'yerm New York Bot. Cara: 23: id 1972 

, Ara- 
VEN. 

pap 

19; 23 


12. 


ho lo) 
. 28(3): 6 


Psychotria pectinata Steyerm., 


Ann. De Bot. c: 

i5: e 1988. 2 E Liesner 15 5; 17 July 198: 
Venez azonas; a à la acid 150 
m; MO Thao — 3303684 ), VEN. 

Psychotria perferruginea Steyerm., A 39: 
1981. . de Granville B.4640; 20 De 
French Guiana, Montagne la Fumea, PK: 
(holo). 


Psychotria perijaensis Steyerm 
Die 


93. 
72; 


, P 


19 
CAY 


, Acta Bot. Venez. 8: 
L3. J. A. Steyermark e al. 105749; 3l Ma ar. 
1972; Venezuela, Zulia, Sierra E Perija; 1,140-1,400 


; VEN (holo). 
aude PU n ra Steyerm., Mem. New York 
Bot. Gard. 23: . 1972, nom. nov. ME Crue- 
ger s.n. o Bot. Gard. Herb. n. 3127. 


veleta pin SITES Mem. 


New York Bot. 


Gard. 23: . 1972, nom. nov. 
Psyc hotria lo EA Standl. & Steyerm., a 
B A. Steyermark 58117; 2 


Oct. 1944; Venezuela, Bolivar Gran Sabata; 1,005 
) 


1,065 m; F (holo 


Volume 76, Number 3 
1989 


Taylor 721 
Plants Described by 
Julian A. Steyermark 


Psychotria Acad Standl. € Steyerm. var. an- 
TA or Stey ., Mem. New es Bot. li 23:591. 
l Cardon na P. 2967; ; Venezuela, 
Bolivar. Ríó Pauo afluente del io! C aura; M m; VEN 
(holo). 

Psychotria mae Steyerm., Mem. New York Bot 

2. J. A. Steyermark 98070; 22- 

ay 1967; SPEM Bolivar, Cerro Jaua; 1,922- 

2.100 m; VEN (holo). 

Peyaholria B Steyerm., Mem. New York Bot. Gard. 

624. 1972. J. Murca Pires & P. B. Cavalcante 
Ve 23 July 1962; Brazil, Terr. Amapá, Agua Azul; 
NY (holo). 

P lana plowmanii i ko Ann. redi Bar Gard. 
71: 1178 4. 4. oo et al. 122132; 9 
May 1980; Venezuela, T.F. Amazonas, Dept. Atures, 
along Rio Cataniapo, 44-45 Ds SE of Puerto Aya- 
eod MO, VEN (holo). 

Psychotria prancei Steyerm., Mem. New Eius ey Gard. 

56. 1972. G. T. Prance et al. 2 Dec. 
1968; Brazil, Amazonas, Reserva P inus e 
Manaus-Itacoatiara road; NY (holo). 


Psychotria psittacina Steyerm., Mem. New York Bot; 
,ard. 23: 665. 1972. S. S. Tillett et al. 45007; 


Aug. 1960; Guyana, Mt. Ayanganna, upper aude 
sidad basin; 800-900 


m; NY (holo 
Psyc abr pr Standl. & Serem Fieldiana Bot. 
Y 1953. J. A. Steyermark 59440; 27 Oct. 


1944; dud Bolivar, Ptari-tepui; 1,250-1,520 
m; F (holo) = P. imthurniana Oliv. f. ptariensis (Standl. 
& Steyerm.) Steyerm., 197 

Psychotria Jr bri Svar rm., Mem. New Yor 

2 E Maguire & D. B. Pa ae 

326 18: 16 Nov T Guyana, upper Mazaruni River, 
Pakaraima Mausia 550 m; depository not desig- 
nated. 

Psychotria pungens Steyerm., Mem. New York Bot 
Gar 677. 1972. Leprieur 118; French Guiana; 


P (holo). 
Psychotria quadribracteata uv ., Mem. New York 
, 23: 614. 1972 T. Silva 81; 2 Feb. 
1972; Brazil, Belem, Para, along road to Barrucào de 
Borracha, Inst. Agron. do Norte; NY (holo). 

Psychotria ronaldo Steyerm., Ann. Missouri Bot. Gard. 
74: 111. 1987. R. L. Liésnér rens 4 Mar. 1985; 

. Amazonas, Dept. 0, Cerro Hua- 
750 m; MO are 332736 ) VEN. 

Psychotria rosacea Stee erm., Mem. New York Dol. Gard. 

972. J. A. Steyermark 105045; 3 Mar. 
971; aede Táchira, entre bado y Queni- 
quea; 2,650-2,700 m; V (holo). 

Psychotria rosea (Benth. ) Muell. Arg. r oo scens Ste 
erm., Mem. New York Bot Hiro . 1972. . d. 

] 4 De ec. E prin ezuela, 
Amazonas, Rio Yama prese. af Rio Yacibo; ae 
140 m; NY (hol 

a rosea Bent ) Muell Arg. var. AR 
Steyer . Mem. New York Bot. Gard. 2 8 
O. iim 2190: 30 res T E Dept. S 
de Santander, Med of Dorado Creek, vicinity of Paro 
Berrio; 250 m; NY (holo) 

Psychotria ruiz- di ranii Steyerm., Mem. New York Bot. 
Gard. 23: 702. 1972. L. Ruíz Turin 3199; 2] June 
1966; Venezuela, xd Distr. Rangel, Las Escaleras; 
2,800 m; VEN (holo). 

P sychotria bou nsis Steyerm., Mem. New York Bot. 

;ard. 23: 4 1972. J. A. Steyermark 98951; 16 


July 1967; oe Falcón, Sierra de San Luis; 
1,450 m; VEN (holo). 

Psychotria sanmartensis Rusby var. a Stey- 
erm., Mem. New York Bot Cad 23: 72. y 
A. Sema 91631; nus Venezuela, ri be- 
tween Petare and Guarenas, 16 km of Caucaquito; 
1,200-1,500 m; VEN (holo). 

Psychotria sastrei Steyerm., Brittonia 33: 394. 1981. 

7. Sastre 2392; 6 Oct. 1973; Colombia, Amazonas, 
Rio Igara-Paraná, La Chorrera, Cerro Adofiki, 35 km 
en aval de La Chorrera; P (holo). 

Psychotria saulensis Steyerm., Brittonia 33: 395. 1981. 

. J. de Granville 2361; 9 Mar. 1975; French Guiana, 
sommet Monts Galbao, a 10 km au SE de Saul; 650 
m; CAY, MO (2798713), VEN (holo 

Psychotria sc ice Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(1): 24. 1943. C. L. Lundell 6193; 
June-July 1936; Belize, El Cayo Distr., Valentin; F 
(holo 

Psychotria sciaphila S. bd subsp. a X aur 
erm., Mem. New York Bot. Gard. 23: 641. M. 
S. Irwin et al. 10161; 10 Nov. 1965; Sune ra 
Distr. Federal, Planalto do Brasil, Parque Mun. do 
Gama; 1,150 m; NY (holo). 

Psychotria silv s Steyerm., Mem. New York Bot. Gard. 
23: 581. 1972. N. T. Silva & U. Brazáo 60872; 24 
on 1966; ery Amazonas, Serra Pirapuci; 1,250 

1 NY (holo 

Bou sipapoe ES Steyerm., Mem. New York Bot. 
Gard. 23: 708. 1972 . Maguire & L. Politi 28778; 
2 Feb. 1949; i T.F. Amazonas, Cerro Sipapo 
(Parque); 600 m; NY (holo). 

m M ace. nsis Standl. & Steyerm., 
Bot. 1 Steyermark 60175; 15 
Nov. pe svi | Mn Bolivar -tepul; 2130- 
2,250 m; F mea = P. sessilis var. angustifolia ( (Muell. 
Arg.) Steyer 1972 

Pescara spectabilis Seyer Mem. New York Bot. 
Gard. 23: 664. 1972, nom. nov. 

Psyehoiria speluncae eue & e Pares Bot. 
up ): 6 Steyermark 59521; 29 Oct 

944; a B Ptari- -tepui; 1, 810 m; F (holo). 

oo speluncae oe & Steyerm. subsp. brevi- 
calyx ar Mer w York Bot. Gard. 17(1): 
436. 1967. B. uo et all 37148; 10 Jan. 1954; 
Venez ss TF. Amazonas, Cerro de la Neblina; 1,700 
m; NY (holo). 

Psychotria speluncae Standl. & Steyerm. subsp. exserta 

em. New York Bot. Gard. pes 436. 

n. Maguire et al. 36921; 27 Dec. 1953; Ven- 

uela, T.F. Amazonas, Cerro de la Neblina, T Yatua; 
1,700 m; NY (holo). 

Psychotria iro ra Steyerm., Brittonia 36: 156. 
1984. J. J. Granville B.5288: 12 June 1975; 
French a sur la Riviere Comte, en foret sur la 
Montagne Soufflet; CAY, P, VEN (holo). 

Psychotria standleyana Steyerm., Fieldiana Bot. 28(3): 

07. 1953. J. A. Steyermark 56681; 19 May 1944; 
Venezuela, Mérida, Montaña de San Jacinto; 1,675- 
2,195 m; F (holo). 

Psychotria steinii Steyerm., Ann. Missouri Bot. Gard. 
75: 345. 1988. B. 4. Stein & A. Gentry 1553; 

1984; Venezuela. T.F. Amazonas, Cerrc 
Neblina, Camp 2; 2,100 m; MO (holo — 
3393676 

Psychotria ihiihi Steyerm., Ann. Missouri Bot. 


Gard. 74: 400. 1987. J. A. Steyermark & B. Manara 


~ 


Fie nete 


ororopan 


1967. 


summit, 


Annals of the 
Missouri Botanical Garden 


125483; 31 Oct. 1981; Venezuela, Miranda, Parque 
Nacional Guatapo, a lo largo del Río Santa Cruz de Río 
Grande, al Sureste de Los Alpes; 400 m; VEN (holo). 
Psychotria subpedic sre Eas beng Mem. New York 
P Gard. 23: Ta 972. G. Klug 2019; Mar.-Apr. 
931; Peru, Dep eto, Rio Ares at mouth of 
Re Zubineta, Florida; 2 m; NY (holo 
Psychotria sucre nsis Steyerm., Mem. New York Bot 
Gard. 23: 1972. Steyermark 94956; 3 
1966; Venezuela Siere, Cerro de Humo; 1,060- 
1,97 3 m; VEN (holo). 
Psyc hotria tamae nsis Standl. & Steyerm.,  Fieldiana Bot. 


` Man m. New York 
E Cardona P. 1496; 
jet orillas del Rio 


US, VEN (holo). 


1946; 
Castanho 100 140 m; 


Psychotria rider d moe Mem. New York 
Bot. Gard. 23: 565. . B. Trujillo 1955; ie an. 
1954; Venezuela, ded Terpanas 1,450 m; MAR 
(holo). 

Psychotria e Steyerm., Mem. New York Bot. 
Gard. 23(1): 654. 1972. Forest Dept. Rec. 7832 (field 


no. RB 8), 5 July 1957; Guyana, Chinoweg; NY (holo). 
Psychotria thesceloantha Steyerm., Ann. Missouri Bot. 
yard. 75: 347. Liesner 16963; 4 Dec. 
ibus Venezuela, T. F. A nazonas, u Caio Baria, 
amp area between Rio Mawarinuma and headwaters 
ul Bio Baria; 130 m; MO (holo— 3393681), 
Psychotria tillettii no Mem. New York Bot. Cani: 
23: 496. 1972. S. S. & C. L. Tillett 43946; 12 Sep. 
1960; Guyana, upper Mazaruni River basin, near Suru- 
a-gu-puh; 470 m; NY (holo). 
Psychotria trichotoma Mart. & Gal. var. T 
Steyerm., Mem. New York Bot. Gard. 23: 475. 1972. 
L. Bernardi 1720; 10 Dec. 1954; Musa. Barinas, 
Mun. Pedraza, Rio Curbatico; 250-350 m; NY bee 
P iyeka trujilloi Steyerm., Mem. New York Ek 
: 475 Trujillo 1921; 24 Jan. 
a, Terepaima; 1,450 m; MAR (holo). 


acuyensts Stey- 


J. rs 


C. Espinoza 123742; 6 Dec. 1980; Venezuela, Yara- 
cuy, Dept. San Felipe, le Gamelotal; 1,300 m; 
VEN (holo). 

Psychotria ube rta Standl. 


& Steyerm., | Fieldiana Bot. 


Y) 
e 
a 
o 
c 
= 
= 
E 
= 
a 
= 
zm 
I 
y 
Sl 
l 
E 


ATY de Palmar, foot 
of Páramo de Tamá; m; F (holo). 

Psychotria ulviformis E Mem. New York Bot. 
Gard. 23: 638 1972, nom. nov. 

Psyc ld banana Steyerm., Mem. New York Bot. 
Gard. 23: 671. 1972, nom. nov. 

o urceolata Steyerm., Brittonia 33: 395. 1981. 

. J. de Granville 2393; 11 Mar . 1975; French Wwe. 
Pente NE des Monts Galbao a 10 km au SE de 
500-600 m; CAY, P, VEN (holo). 

Psychotria a gy Ste uw . New York Bot 
Gard. 23: 556. 1972. Tillett et al. 45008; 
Aug. 1960; pem pe ME River basin, m 
Ayanganna; 800— m; NY (holo). 

Psychotria varesc Re cil Mem. New York Bot. 
Gard. 23: 706. 1972. V. Vareschi 7790; 2 Nov. 1962; 


E 


, 


Venezuela, T.F. Amazonas, Rio Casiquiare, Cano Mo- 
nomi; VEN (holo). 

Psychotria variegata Steyerm., Mem. 
Gard. 23: 638. 1972, nom. nov 

Psychotria Ea Steyerm., Mem. New York 
Bot. Gard. 23: . 1972. B. Maguire et al. 42031; 
17 Nov. 1 a PEREA T.F. Amazonas, Cerro de 
la Neblina; 1,600-1,700 m; depository not designated. 

Psychotria ventuariana Standl. & Steyerm., Fieldiana 
Bot. 28(3): 609. 1953. Ll. Williams 14983; 20 Apr. 

42; Venezuela, T.F. Amazonas, Delta del Ventuari; 
124 m; F (holo). 

Psychotria ra ee ., Brittonia 36: 158. 

J. de Granville 2793; 1 Mar 11; French 
Guiana, Saul, sommet des Monts Calbso: 670 m; CAY, 
P, VEN (holo). 

Psychotria wesse d boeri Steyerm., Mem. New York Bot 
Gard. 23: 467. 1972. J. G. Wessels-Boer 962; 18 
Mar. 1963; Ende Oelemari; U (holo). 

Psychotria wilhelminensis pa Mem. New York 
Bot. Gard. 23: 487. 1 . H. S. Irwin et al. 55048; 
à Aug. 1963; Suriname, Wilhelmina Gebergte, 12 

N of Lucie River; NY (holo), VEN. 
Poxchotria w ee Lr io, Mem. New York Bot. 


New York Bot. 


No) 


Gard. . B. eed et bs 42475; 24 
Dec. o ‘Vite 20 T.F. Amazonas, Rio Yatua, 


Cerro de la Neblina; 1,100 m; aa not desig- 
nated. 

Psychotria yapac aen PISTES ., Pittieria 9: 10. s 
O. Huber 1791 lay 1978; Venezuela, Dept. A 
bapo, Cano € eta y el Cerro Yapacana; 100 m; V EN 
(holo). 

Psychotria yaracuyensis Steyerm., Mem. New 107% ies 
Gard. 23: 447. 1972. Sievermark 
Bunting 97688; 11 Mar. 1967; Venezuela, D 

of Nirgua, Cerro La Chapa; 1,200-1,360 m; VEN 
holo). 

ebria yavitensis > yerm. , Mem. New e E 

d. : 456. 1972 i & 
Bunting 102933: 22 "Apr. Eros ato the TE 
Amazonas, 7-9 km from Yavita; 128 m; VEN (holo). 

Psychotria yutajensis Steyerm., Ann. Missouri Bot. Gard. 
15: 348. 1988. R. L. Liesner & B. K. Holst 21649; 
4 Mar. 1987; Vanini, T.F. Amazonas, Dept. Atures, 
summit E of unnamed peak, 8 km NW of Yutajé, W 

ia Yutajé 1,500-1,700 m; MO (holo — 

N. 


Randia diode L. var. Hu E Steyerm., Mem. 
New York Bot. Gard. 23: 34 A. Steyermark 
P June 1960; jd Bolivar NE of Tu 
between Hato Corumo and Las Chicharras; 
holo). 
Randia ar ata L. f. minor Steyerm., Mem. New York 
Gard. 341. 1972 Britton & W. 
Hess 2702; 5 Apr. 1913; Pieris Rico, Mayaguen, 
Monte Mesa; NY (holo). 
Randia amazonasensis Ss) erm., Mem. New York Bot. 
Gard. e 336. 1 B. Maguire et al. 29469; 7 
Nov. ; Ven se T.F. Amazonas, Rio Orinoco, 
Rio Gin 'unuma, Playa Alta; 100 m; (holo) 
Randia aristeguietae Steyerm., Bol. Soc. Venez. Ci. Nat. 
97 .. Aristeguieta 4193; May 1960; Ven- 
ezuela, Dip Carretera El Rastro, Calabozo; VEN 
(holo), NY. 
d brevipes Steyerm., Acta Bot. 
1971. Pistillate: L. aa 5343; 


6624 1; 


inp Me 


29: 22. 1 


Venez. 6: 126. 
Apr. 1964; 


Volume 76, Number 3 
1989 


Taylor 723 
Plants Described by 
Julian A. Steyermark 


Venezuela, Bolivar, 
minate: N. L. Britton et al. 
chacare; NY (holo). 

Randia Vb dp m Row & pun Field Mus. 


Puerto Ordaz; VEN (holo). € 
2684; Trinidad, Chaca- 


Nat. Hist., Bot. S 1): 25. 1943. P. C. Standley 
76321; Git- Nov. vA Guatemala, Dept. Jutiapa, 
vicinity of Jutiapa; 850 m; ). 

Randia pubiflora Steyerm., Brittonia 33: 3 81 


Cavalcante 2511; 25 Feb. 1970; Brazil, Pará, 
Parque Indigena do Tumucumaque, Rio Parú de Oeste 


Missão Triyo; NEN we lo). 


ol. Soc. Venez. Ci. 
197 "i J. I. Wirdack & J. V. Wonachino 


7 June 1956; Venezuela, Bolivar, betw 


Chiriquí, Quebrada Velo, vicinity of Finca Lerida l, 631 
m; F. (holo), MO. 

Ravnia panamensis Steyerm., Ceiba 3: 22. 1952. Col- 
lector and collection number not indicated; 29 June 
1946; Panamá, Prov. Coclé, North Hills, El Valle de 

Antón; F (holo), MO 


cus A hs Steyerm., Mem. New York Bot. Gard. 
23: 197: Maguire. et al. e 16 Sep. 
1957. k E T. F. Amazonas, NW base of Cerro 


Yapacana; 125 m; NY (holo), VEN. 

Re ee gs sej Steyerm., Ann. Missouri Bot. Gard. 71: 
332.1 4. P. E. Berry & L. Chesney 2116; 26 Feb.- 
2 Mar. rper Venezuela, Amazonas, Estación 
Experimental de Santa Barbara del Orinoco, a 1-2 km 
5 de Trapichote; 130 m; VEN (holo) 

Remijia delascioi Steyerm., Ann. Missouri Bot. Gard. 
71: 332. 1984. J. A. Steyermark et al. 130339; 1 
Mar. 1984; Venezuela: T.F. s Cerro Vinilla; 
440 m; MO (holo — 3223165), VE 

"E did in Benth. f. ctrl bM Mem. 

Gard. 23(1): 265. . B. Maguire 
& B. Maguire, Jr. 29079; 22- d ji 1949; Ven- 
ezuela, T.F. Amazonas, Cerro Duida; 1,600 m; NY 
(holo). 

Remijia densiflora Benth. var. bids Enn a jo 
New York Bot. Gard. 23(1): 265. 1972. J. . 
ermark 90269. 27 Dec. 1961; E IP 
Sierra Ichun; 625-725 m; VEN (holo 

Remijia i iid wo ., Mem. New You Bot. Gard. 
23(1): 2 . B. Maguire et al. 42469; 24 Dec. 
1957; dni a TF. Amazonas, Cerro de la Neblina; 
1,100 m; NY (holo 

Remijia marahuacensis Steyerm., Ann. 
Gard. 74 J. A. Steyermark & 


Missouri Bot. 
B. K. Holst 
130440; Venezuela, T.F. Amazonas, Dept. Atabapo, 
Cerro Marahuaca, Sima Camp; 1,140 m; MO (holo — 
3327367), VEN. 
Remijia morilloi Steyerm., Fl. Venez. 9(1): 115. 1974. 
C. Morillo et al. 4192; 29 Apr. 1974; Venezuela, 
T.F. Amazonas, 12-15 km NE de San ur de Rio 
Negro, carretera San Carlos-Solano; VEN (holo). 
Remijia pilosinervula SCR Mem. New York Bot. 
Gard. 17(1): 232 Steyermark & J. J. 
Wurdack 713; 17 Feb. 1955; ety Bolivar, Chi- 
manta Massif; 2,150 m; VEN (holo). 
, Ann. Missouri Bot. Gard.: in 
> G. Carnevali 22407; 
25 Oct: 1987; Vexsanela, T.F. Amazonas, Dept. Rio 


gro, Cerro ko aie summit, Proa Camp; 1,400 
m; MO (holo), VE 

Remijia roraimae run Schumann var. adpressa 
5 rm., Mem. New York Bot. Gard. 23(1): 262. 

A. Steyermark 90429; 29 Dec. 1961; Ven- 
enel: Bolívár, Sierra Ichun; 500-625 m; VEN (holo). 

Remijia roraimae (Benth.) Schumann var. auyante- 
puiensis Steyerm., Mem. New York Bot. Gard. 23(1): 
263. 1972. J. A. Steyermark 93361; 4 May 1964; 
Venezuela, Bolivar, Auyan- tepui; 1,760 m; VEN (holo). 

Remijia roraimae (Benth i 
Steyerm., Mem. New York Bot 
1972. B. Maguire et al. 46152-A; 9 Sep. 
Guyana, Pakaraima Mountains, E Savanna; 
1,210 m; depository not designa 

Remijia sessilis Steyerm., Ann. Missouri Bot. Gard. 75: 

1988. M. Nee 30967; 20 Feb. 1985; Venezuela, 
.F. Amaronas, SW side of Cerro de E Neblina; 400- 
500 m; MO (holo), NY, VEN. 

Remijia sipapoensis Steyerm., Mem. New York B 

Gard. 23(1): 255. 1972. B. Maguire & L. Po lit 
: é . Amazonas, 
Cerro Sipapo bios uid NY (holo). 

Remijia stenopetala $ : pe Bot. 
28(3): 610. 1953. T A. Steyermark 60 359; 20-21 
e 1944; Venezuela, Bolivar, Quebr ade 0- -paru-ma; 

65-1,220 m; F (holo) = R. densiflora sabap. steno- 
Eus (Standl. & Steyerm.) Steyerm., 1972. 

Remijia vaupesiana Steyerm., Mem. New York Bot. 
Gard. 23(1): 253. 1972 R. de Lemos Fróes 21225; 
20 Oct. 1945; Brazil, Amazonas, Rio Negro, Vaupés, 
Jauareté; depository not designate 

Remijia p Steyerm., Mem. New York Bot. Gard. 

3(1) 259. 1972. J. J. Wurdack & L. Adderley 
42824; 5 o 1959; Venezuela, T.F. Amazonas, Caño 
Cupaven; 125-150 m; NY (holo) 

Retiniphyllum glabrum Steyerm., Mem. New York Bot. 
Gard. 12(3): 243. R. E. Schultes & J. Murca 
Pires 9100; 14-15 Nov. 1947; Brazil, Amazonas, Rio 
Vaupés, between Ipanore and confluence with Rio Ne- 
gro; US (holo). 

Retiniphyllum is Steyerm., Mem. New York 
Dot, Gard. 12(3): 242. 1965. S. 5. die ott et al. 44824; 

5 July 1960; Guyana, upper Mazaruni River basin, 

Partang River, SE side of Merume Mt: 1,140 m; NY 

holo). 


( 

Retiniphyllum laxiflorum (Benth.) N. E. Brown var. 
brasiliense Steyerm n. New York Bot. Gard. 12(3): 
234. 1965. B. A. "Krukoff 3; 9 Oct. 1942; Brazil, 


Bahia, foris Sincora, Gorge of Zesuserra; 1,0 


Y (holo). 
Retiniphyllum laxiflorum (Benth.) N. E. Br. var. lon- 
pola Steyerm., Mem. New York Bot. Gard. 12(3): 
5. B. Maguire & L. Politi 27869; 23 Dec. 1948; 
a T.F. Amazonas, Cerro Sipapo (Paraque); 
1,400 m; NY (holo). 
c o ur longiflorum Steyerm., oo 33: 396 
oP. Vaupés, 1 712; Mar. 9; Brazil, rie 
zonas, Rio , Taracua; VEN T 
Retiniphyllum m e Standl. var. retic uloin Stey- 
erm., Mem. New York Bot. Gard. 12(3): 239. 1965. 
B. Maguire 24361; 13 Aug. 1944; Suriname, Ta- 
felberg, Savanna #1; 565 m; NY, VEN (holo). 
Retiniphyllum parvifolium Steyerm., Brittonia 33: 397. 
1981. J. Murca Pires et al. 6063; 12 Dec. 1956; 


724 


Annals of the 
Missouri Botanical Garden 


Brazil, Pará, Serra do Cachimbo; 425 m; NY (holo), 
VEN. 


Retiniphyllum scabrum Benth. var. 
erm., Mem. New York Bot. Gard. 12(3): 

1965. S. S. Tillett et al. 45194; 8 Aug. 1960; wee 
Mt. Ayanganna, above Thompson Camp; 1,370-1,5 
m; NY (holo). 

Retiniphyllum schomburgkii (Benth.) Muell. Arg. jd 
des on le Steyerm., Mem. New York Bot. 

1 2(3): 240. 1965. B. Maguire & J. J. Wurdack . 34531. 

T.F. Amazonas; NW b 
of Cerro ) Yap #3; 150 m; 

Pira ue sc honhurghi (Benth.) Muell. Arg. subsp. 
occidentale Steyerm. hirticalyx Steyerm., Mem. 
New York Bot. Gard. 123 is 241. 1965. B. Maguire 

; 12 Oct. ; Colombia, 0.5-1.5 km 
N of Puerto Colombia pane Maroa); 130 m; NY 
(holo). 

o tepuiense Steyerm., Mem. New York Bot. 

ard. 12(3): 237. 1965. B. Maguire & L. Politi 

En 20 Jan. 1949; Venezuela, ' Amazonas, 
upper E Basin, Cerro Sipapo (Paraque); l, 600- 1,800 
m; NY (holo). 

is cit truncatum Muell. Arg. var. i he bh 

Steyern 1. New York Bot. Gard. 12(3): 242. 

Bre «€ R. Bl es 46-357; 8 Nov. 

1946; Colombia, "fic da Cotuhe; US (holo). 

s New York Bot. 

puram & C. Mon- 

1954; Venezuela, Yaracuy, Aroa; 


AENEA 
: 246. 1967. L. 
; Feb. 


Rondeletia brachistantha Standl. & Steyerm., Field Mus 
Nat. Hist., Bot. Ser. 23(1): 25. 1943. J. 4. a 

1940; Guatemala, Dept. San Marcos, Río 

Vega, near San Rafael and Guatemala - Mexican bound- 

Tacaná; 2,500-3,000 m; F (holo). 

ae letia O nsis Lie & Steyerm., Field Mus. 

at. Hist., BOE Ser. 23(5): 254. 1947. J. A. Steyer- 
mark 4563 2 A r. 1942 Guatemala, Dept. Alta 
Verapaz, Cane Chinaja; 150-700 m; F (holo). 
p oid ju ae Standl. & oa. Field Mus. 
, Bot. Ser. 23(1): 26. 1943. J. A. Si ab ed 
29906; [ a 39. Guatemala, Dept. Zacapa, Que 
brada Alejandría, summit of Sierra de las Minas, vicinity 
of Finca Alejandria; 2,500 m; F (holo). 

ee iz ees nsis Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 22(4): 286. 1940. P. C. Standley 
72891; 3 May 1939, Guatemala, Dept. Izabal, Escoba; 
sea level; F (holo). 

Rondeletia larensis Steyerm., Mem. New York Bot. Gard. 
1 7(1): 245. 1967. J. Saer 727; Jan. 1931; Venezuela, 
Lara, Distr. Torres, Río Sicariqua; VEN (holo). 

Rondeletia macroc aly x Pie a Steyerm., Field ied 
Nat. Hist., aer Ser. BMS): 2 1947. J. A. Stey 
mark 418 41; DLE Dept. laia, 
Cerro San Cn -300- 900 m; F (holo). 

Rondeletia myriantha Standl. & Steyerm., Field Mu 

. Hist., Bot. Ser. 22(4): 288. 1940. 4. F. Skutc , 
1569; 31 Oct 1934; Guatemala, Dept. Suchitepéquez, 
Finca Moca; 960 m; F (holo). 

Rondeletia He Wis Steyerm., Mem. New York ie 
Gard. 17 1967. LI. Williams 13329; 15 Jur 
1940; de Bolivar, Cerro El Tigre, Rio C ch 
vero; 90 m; VEN (holo). 

Rondeletia purdiei Hook. 

k Bot. 


5 


f. var. glabrior Steyerm., Mem. 
Gard. 17(1): 243. 1967. H. H. Smith 


ayangannense 
236 


320; 17 July 1898; Colombia, ORAS near Ma- 
singa, Santa Marta; 78 m; (holo 
Rondeletia skutchii Sens & Steyerm, Field Mus. Nat. 
Hist., Bot. Ser. 22(4): 2 4. F. Skutc h 2110; 
10 Jan. 1935; M. es Suchitepéquez, Finca 
Moca; 1,260 m; F (holo). 
Rondeletia venezuelensis d 


Mem. New York 


Bot. Gard es 243. . H. Pittier 11995; 25 
ec. 1925; Venezuela, nu El Tinaco; NY (holo). 
Rondeletia vulcanicola $ andl, ES Steye "i. Mus. 
Nat. His st, € Ser. s A. ee 


mark 33220; 31 Dec br eL Dept. 
paltenangn; Finca Pireneos, Volcán Santa María; l, 300- 
1,500 m; F (holo). 
Rondeletía: zolleriana Standl. & Steyerm., Field 
Hist., Bot. Ser. 22(4): 292. 1940. J. . CMS 

1940; Guatemala, Dept. San 

d tid Finca El Porv and Loma Corona, 
Volcán Taimules; 1,300-2 000 1 m; F is olo). 

Rudgea TA et Steyerm., Mem. New York Bot. 
Gard. 17(1): 410. 1967. S. Ls Tillett et al. 45182; 7 
Aug. 1960; ae Mt. Ayanganna, upper Mazaruni 
River; 900-1,100 m; NY (holo). 

Rudgea sto Steyerm. & Dwyer, ape E 12, 1981, 
P. E. Berry & C. Uhl 1558; 24 Sep 5; Venezuela, 
T.F. ret 4 km from San e Ph Rio Negro, 
on road to Solano; 120 m; VEN (holo), MO. 

Rudgea bolivarensis e Mem. New York Pon 
Gard. 17(1): 415 7. J. A. Steyermark 59118; 2 

. 1944; Vsnsuels, Bolivar, Gran Sabana; 1,005 
l, 065 m; VEN (holo). 

Rudgea bremekampiana Steyerm., Mem. New York Bot. 
Gard. 17(1): 407. . R. S. Dues 38429: 19 Nov. 

1954; inm Amapa, Rio Araguary, Serra do Navio; 
10-300 m; NY (holo). 
etin buntingi Steyerm., Acta Bot. Venez. 6: 17 
. J. A. Steyermark et al. 100252; 9-10 Nov. 
e d Venet uela, Yaracuy, Cerro La Chapa; 1,200- 
1,400 m; VEN (holo), NY 

Rudgea cardonae O Mem. New York Bot. Gard. 
17(1): 403. 1967. F. Cardona P. 1503; 16-24 Feb. 
1946; ala T. F. Amazonas, Rio Castanho; 100 
140 m; VEN (holo). 

Rudgea PW Standl. & Steyerm., Fieldiana Bot. 28(3): 

d. Williams 14674; 3 July 1942; Ven- 
ezuela, ir. Amazonas, San Carlos de Rio Negro; 100 
m; F (holo). 

Rudgea corocoroensis Steyerm., Ann. Missouri Bot. Gar 

75: 349. 1988. R. L. Liesner & B. K. Holst 21827: 
10 Mar. 1987; Venezuela, T.F. Amazonas, 5-8 km 
NW of Yutajé settlement, 3 km W of Rio Coro Coro; 
700-1,000 m; MO (holo — e VEN 

Rudgea costanensis Steye Mem. New York Bot. 
Gard. 17(1): 420. 1967. H Pieler 13498; May 1934; 
Venezuela, Distr. Federal, near El Limón, Valle de 
Puerto Cruz, El Camaticaral; 950-1,000 m; US (holo). 

Rudgea enervia Steyerm., 
17(1) 418. 1967. S. S. Tillett et al. 45117; 12 

1960; Guyana, Mt. Ayanganna, Thompson Camp, up- 
per Mazaruni River; 1,418 m; NY (holo). 

iu d Standl. & Steyerm., ON Bot. aes 

953. J. A. Steyermark 55509; 12 Feb. 1944; 

VS Lara between Buenos Hin to ud of 

El Callado, above Humocaro Alto; 2,285-2,740 m; F 
(holo). 

Rudgea lucentifolia Standl. & Steyerm., 


w 
= 


Fieldiana Bot. 


Volume 76, Number 3 
1989 


Taylor 725 
Plants Described by 


Julian A. Steyermark 


28 


eS 


3): 616. 1953. J. A. ue 56502; 14 May 
i el Trapiche and El 


u eda marcano- hor Steyerm., Pittieria 9: 12. 1981. 

L. Marcano Berti & I Pena 532-979; Venezuela, 

Táchira, Mun. Lo Pa era, Parque iix di Palo Grande- 

Minas de Carbon de Lobatera; 1,700 m; MER, VEN 

(holo). 

Rudgea merumensis Steyerm., Mem. New York Bot. 
Gard. 17(1): 402. 1967. S. S. Road et al. 44825; 
5-6 July 1960; Gu uyana, Merum 


Rio Ventuari; 800 m; NY in 


, Brittonia 
/; 31 Aug. 1966; French 
Guiana, Grand Matoury, Ilede oe P (holo). 
Rudgea phaneroneura Steyerm., Mem 
l 


= 


P á Massif, Central Section; 1, 925 
m; MO (20 011378, 72011681), VEN (holo 

Rudgea prancei Steyerm., Brittonia 33: 398. 1981. G. 
T. Prar nce et al. 14739; 14 Sep. 1971; Brazil, Ama- 

y Manaus, Reserva Ducke, vicinity of Iga- 
rapé Acara; NY, VEN (holo). 

Rudgea ruiz-teranii Steyerm., Pittieria 9: 13. 1981. L. 
Ruíz Terán 4703; 12 Oct. 1967; Venezuela, Mérida, 
Distr. Sucre, Azulita y Lagunillas; MER, VEN (holo). 

Rudgea o Le res Mem. New York Bot. Gard. 
17(1): 420. 1967. L. Mei ca 3938; Aug. 1959; 
Venezuela, v ara, oen 1,50 ; VEN (holo). 

Rudgea simiarum Standl. & Fon Field Mus. Nat. 
Hist., Bot. Ser. 22(5): 389. 1940. J. A. Steyermark 
38839; 5 Apr. 1940; e Dept. Izabal, Mon- 
taña del Mico; 50-500 m 

Rudgea sipapoensis Gieren: : 
Gard. 17(1): 411. 1967. B. Maguire & L. Politi 
28483; 21 Jan. 1949; Venezuela, T.F. Amazonas, 
Cerro Sipapo (Paraque); 125-600 m; NY (holo). 

, Mem. New York Bot. 


acd New York Bot. 


man 39036; 31 Dec. 1954; Suriname, Nassau Moun- 
tains, Marowijne River, Plateau A; 430-520 m; NY 


(holo). 
Rudgea P Steyerm., Mem. New York Bot. Gard. 
967. S. S. Tillett et al. 43934; 28 June 
1960; abb Porkknocker un 2 on Partarg Tt pss 
Merume Mountains; 625 m; NY (holo) = P. s 
sareoides Standl 
Rudgea trujilloi Ste a Bot. Venez. 6: 178. 
1971. B. Trujillo 5555; ny Jus 1964; Venezuela, 


bor y Sanare; 1,000 m; MY (holo us 
Acta Bot. Venez. 6: 


Lara, entre Qui 

Rudgea ne Steyerm., 
1971. V. Vareschi 1386; 30 May 1952; e 
Mérida, Selva de Mucuy; 3,120 m; MER (holo). 

Rudgea wurdackii Steyerm., Mem. New York Bot. Gard. 
17(1) 413. J. J. Wurdack & L. Adderley 
42727; 1 June 1959; Venezuela, T.F. P Rio 
Orinoco, above mouth of Rio Atabapo; 125 m 


(holo). 


Rustia d ced SUE & Steyerm., ra Bot. 

28(3): 6 Steyermark 56333; 6 May 

1944; V d Mérida, below El Bao. Pue Mu- 
cuchachi and Canagua; 1,065-1,820 m; F (holo). 

Sabicea aristeguietae Steyerm., Mem. New York Bot. 
Gard. 17(1): 309. 1967. L. Aristeguieta 1049; Feb. 
1953. Venezuela, Barinas, Ciudad Bolivia (Pedraza); 
VEN (holo). 

Sabicea bariensis Steyerm., Ann. Missouri Bot. Gar 
75: 350. 1988. 4. Gentry & B. Stein 47314; 9 May 
1984; Venezuela, T.F. Amazonas, upper Rio Baria; 
140 m; MO (holo —3393683), VEN. 

Sabicea brachycalyx Steyerm., Mem. New York Bot. 
Gard. 17(1): 313. 1967. J. J. Wurdack & L. Adderley 
see 6 June 1959; Venezuela, T.F. Amazonas, San 

o de Atabapo; 125 m; NY (holo). 
moe grandifolia Steyerm., Fl. Venez, 9(1): 514. 1974. 
orillo et al. 4070; 27 Apr. 1974; Venezuela, 
Es Amazonas, alrededores de San Simón de Cucuy, 


2 km N 


md & R. L. ics 118876; Venezuela, Tá- 


Sabicea morillorum Steyerm., Fl. Venez. 9(1): 511. 1974. 
G. Morillo et al. 3903; 24 Apr. 1974; Venezuela, 
T 


San Carlos de Rio 


Sastre 
tequia, 12 km en amont de Villavicencio, Quebrada de 
la Mendoza; P (holo). 

Sabicea tillettii pur pigs s mE 484. 1975. 
S. S. Tillett et al. p 


Venezuela, eo nas, p cai Fr ca. 1 km 
ds from mouth of Cano Chamuchina; 130 m; VEN 


Sabicea velutina Benth. subsp. ew rar E 
ew York Bot. Gard. ES . J. A. 

JJ. ut pr 10 M 1955; 
Venezuela, Bolivar. Amuri-tepui (Chimantá Massif); 
1,365 m; MO (2011341, 2012047), VEN (holo). 

Sabicea Mie qe Benth. subsp. pr RT 

k Bot. Gard. 17(1): n e [me 
& B. Mss Jr. 29137; 22 uh A 949. Ven- 
ezuela, T.F. Amazonas, Culebra Peak, fae Duida; 
1,600 m; NY (holo 

Sabicea venez zuelensis 2 Mem. New York Bot. 
Gard. 17(1): 309. 1967. F. Cardona P. 707; 21 July 

1943; Venezuela, Bolívar. Raudal Aguacarita, Rio Pa- 
ragua; VEN (holo). 

Schradera andina Steyerm., Acta Bot. Venez. 6: 122. 
1971. J. A. Steyermark et al. 103752; 30 Sep. 1970; 
Venezuela, D above Escuque, El Paramito; 1,650 
m; MO (2217413), NY, VEN (holo). 

Schradera eurer Po m., Mem. New York 
Bot. Gard. 10(5): 2 dH. Mcd. et al. 40814; 
20 Aug. 1962; Venezuela, Bolívar, Rio Uiri-yuk, alto 
Río Cuyuni; 500-600 m; NY (holo), VEN. 

Schradera se Stey , Mem. New York Bot. Gard. 

26 p. pon & L. Politi 27895; 

e E nd T.F. Amazonas, Cerro Si- 

m; NY d 

w Yo rk Bot. Gard. 

1963. E. L. Ekman 0368: 7 July 1918; 

Cuba, ` Prov. Oriente, Sierra Maestra, prope Nanacal, 

prope Alto de Comején; 1,100 m; NY (holo). 


726 


Annals of the 
Missouri Botanical Garden 


Schradera glabriflora Steyerm., Mem. New York Bot. 
Gard. 10(5): 267. 1963. J. A. Ste; 
tini 17; 4 Sep. 1960; Venezuela, Aragua, Parque N 
cional Henri Pittier, trail above Pittier National Mon- 
ument on Pico Paraiso to summit; 1,250-1,600 m; 

(holo). 

Schradera hilliifolia Steyerm., Mem. New York Bot. 
Gard. 10(5): 269. 1963. J. J. Wurdack & L. S. Ad- 
derley 43316; 4 July 1959; Venezuela, T.F. Amazo- 
nas, Maroa, Río Guainía; 130 m; NY (holo). 

Schradera maeut i Steyerm., Mem. New York Bot. 
Gard. 10(5): 271. 1963. B. Maguire 32974; 4 Jan. 
1952. lak. T.F. Amazonas, Cerro read 
North Valley; 1,600-1,700 m; NY (holo), VEN. 

Se hradera marahuacensis Steyerm., Ann. Missouri Bot. 

A. Steyermark & B. K. Holst 
130721; 21-22 Feb. 1985; Venezuela, T.F. Amazo- 

. Atabapo, Cerro Marahuaca, Sima Camp; 
1,140 m; MO (holo— 3327359), VEN 

Schradera nilssonii Steyerm., Bol. Soc. aes. Ci. Nat. 

: pem J. A. Steyermark & S. Nilsson 196; 

Bolivar, Fila de La Danta; 


Da 


“ 


~ 
5 
z 


15 Apr. 1 960; Venezuela, 
1,200 m; NY, V Yolo 
p diio puberula Sie yerm., Mem. New bs Bot. 
0(5): 273. 1963. J. es 15182; 19- 
28 Aug. 1943; ía Dept. El Valle, Cordillera 
Occidental, bosques Hoya del Río Digua, Piedra de 
Moler; 900-1,180 m; F (holo). 
Schradera pulverulenta Steyerm., das ^ut Venez. 4(1): 
113. 19 Cuatrecasas 21086; 18-22 May 1946; 
Valle, Rio Calima (d del Chocó); 
VEN. 


Colombia, Dept. del V 


30-50 m; US (holo), 
Schradera ou Steyerm., Mem. New York Bot. 
Pod nado 275. 1963. R. A. Howard 12302; 2 
SA dias Asa Re ublic, Prov. Barahona, 


a Nueva to Loma Alta; 1,500 m; NY (holo). 

Sc E. ra ternata Steye . New York Bot. Gard. 
10(5): 275. 1963. S Tillett et al. 44992; 2 Aug. 
1960; Guyana, NE side of Mt. Ayanganna; 884-915 
m; NY (holo). 

Schradera vahlii Steyerm., Mem. New York Bot. Gard. 

0 1963, nom. nov 

Schradera vahlii Steyerm. var. acutifolia us ta 
New York Bot. Gard. 10(5): 277. 1963. 4. C. Smith 
10289; 26 Mar. 1956; Dominica, vicinity of inh 
Water Lake, near Laudat; 450-600 m; NY (holo), US. 

ee hea inde Steyerm., Mem. New York Bot. 
Gard. 10(5): 276. 1963. B. & C. K. Maguire 35356; 
21 Feb. 1953; Venezuela, T.F. Amazonas, Cano 
taje, Cerro Yutaje; 1,500 m; NY (holo). 

tekina aristeguictae Steyerm., Bol. Soc. Venez. Ci. 

. 1954. L. ms 2 F. Pannier 

4; ] July 1953; Venezuela, Yara 
Aroa; VEN (holo) = Simira a. Giao. ) 
Steyerm., 7 

Simira erythroxylon (Willd.) oru var. meridensis 
Steyerm., Acta 971. E. E. Lütle 

| PUO San Juan 


: 277. 


O 


st, 000 m; NY 


bore 
~ 
= 
= 
cS 


erm., Acta Bot t. Venez. PR 112 
& J. V. Monachino 39858; 13 n. 
Bolivar, Orinoco River, Isla Sta. im 80-90 m; NY 
(holo), VEN. 
Simira ignicola Steyerm., Ann. Missouri Bot. Gard. 75 
1086. 1988. J. 4. Steyermark et al. 131659; 9 Sep. 
1985; Venezuela, Bolivar, Distr. Cedeno, 1 km S of 


Quebrada La Flore, AS of Rio Ore, peris of Rio 

Paraguaza; 85 m; 2e 3295642), VEN. 
ae e eye Acta Bot. 

Torres a zamae 203; 15 Apr. 1972; 

zuela, Barinas, Distr. Obispos, Mun. Cruz Pele 
360 m; VEN (holo). 

Simira 1 e kii Steyerm., Mem. New 
23(1): 304. 1972. J. J. W e 2461; 
1962; se Dept. Loreto, Prov. Alto Amaz 
Marañon, Pongo de Meda 250 m; US 

EN. 


Simira lezam 
9 


York Bot. s 
26-28 Oc 
onas, Rio 


(holo), 


s a Aublet subsect. d Steyerm., Mem. 
w York Bot. Gard. 17(1): 264. 1967, Erbe nov. 
Sipanea Aublet subsect. rhea Steyerm., Mem. 
ew York Bot. Gard. 17(1): 270. 1967, subsect. nov. 
Sipanea Aublet subsect. Ma Steyerm., Mem. New 
Bot. Gard. 17(1): 268. 1967, subsect. no 
M dk Aublet subsect. ad "rotricha Steyerm., Mem, 
w York Bot. Gard. 17(1): 266. 1967, lack: nov. 
CAE a ayangannensis 2 Mem. New York Bot. 
,ard. 17(1): 269. 1967. S. S. Tillett et al. 45114; 
12 Aug. 1960; Guyana, upper Mazaruni River, Mt. 
Ayanganna; 1,418 m; NY (holo 
s carrenoi Ste Pyerm., Ann. Missouri Bot. Gard 
98 A. Steyermark et al. 115598; 22 
1,130 


eb. erudi: Bolívar, Gran cdd 
m; MO (3223163). VEN (holo). 

Sipanea cowanii E Mem. New York Bot. Gar 

. S. Cowan & T. R. $ Saderstrom 
1863; 18 Feb. ie. Guyana, Kaieteur Plateau, 
Johnson’s View, Kaieteur Falls, Potaro Gorge; 425 3 m; 
US (holo). 

Sipanea gleasonii Steyerm., Mem. New York Bot. Gard. 
17(1): 265. 1967. H. 4. Gleason 431; 18 June-8 
July 1921; Guyana, Tumatumari NY (holo). 

Sipanea glomerata H.B. . paucinervia pede 

lem. New York Bot. Card. 17(1) 282. B. 
Maguire et al. 30458; 28 Dec. 1950; adieu T. F. 
Amazonas, 30 km above Playa Alta; 200 m; NY (holo). 

Sipanea ovalifolia wo var. villosissima Steyerm. 

dU 36: 159. 1 . G. Cremers 6460; 26 Apr. 
80; French Qu. a ka SE de Saul; 600-700 m; 
ue (holo), VEN. 

Sipanea prancei Steyerm., Brittonia Es: 400. 1981. G. 

. Prance et al. 10528; 14 Feb. 1971; Brazil, Roraima, 
Indian trail from Surucucu, near Maita Indian village; 
NY (holo), VEN. 

Sipanea pratensis Aublet var. dichotoma An B.K.) Stey- 


~ 


erm rachycarpa Ste 1 m. ' York Bot. 
Gard. 17(1) 276. i M e L. Politi 
27324; 19 Nov. 1948; Venezuel: Mr d 


; i . 
Danta (Tapir) Falls, Rio Cuao, Rio CERA 125 1 
VEN (holo). 

Sipanea pratensis Aublet var. = Ape n B.K.) Stey- 
erm. f. breviflora Steyerm., 1. New k Bot. Gard. 
17(1): 277. 1967. A. EE 89276; 1-11 
Feb. 1961; Venezuela, Bolivar: cerro between Las Nieves 
and base of slopes SE of uis Picacho Altiplanicie de 
Nuria; 100-300 m; VEN (ho 

Sipanea pratensis Aublet var. dic atom (H.B.K.) Stey- 

erm. f. Mie a m., Mem. New York Bot 

Gard. Vx 216. 1967. J. J. Wurdack & L. Adderley 

42916; n.d.; MR T.F. Amazonas, Yavita—Pi- 

michin trail near Yavita, Río Atabapo; 125-140 m; 
(holo). 


"e a pratensis Aublet var. dichotoma (H.B.K.) Stey- 
rm. f. glabrior Steyerm., Mem. New York Bot. Gard. 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


ae 2777. 1967. J. J. Wurdack & J. V. Monachino 
39823; 12 Dec. 1955; Venezuela, Bolivar, Cerro San 


Sipanea pubinoda Steyerm., Mem. New York Bot. Gard. 
17(1): 278. 1967. L. Williams 15656: 29 May 1942; 
Venezuela, T.F. Amazonas, Capihuara, Alto Casiquiare; 

120 m; VEN (holo). 

orae PM Steyerm., Ann. Missouri Bot. Gard. 74: 
114. O. Hub m al. 8246; 3l Aug. c 
ae Bolivar. Distr. Piar, Cerro El Venado, 
km E of Canaima; 1,300 m; NY (holo 

E Steyerm., Mem. New York Bot. Gard. 17(1): 
84. 1967, gen. nov. 

Sipaneopsis foldatsii Steyerm., Mem. New York Bot. 
Gar : 288. 1967. E. Foldats 3678; 4 Sep. 
1960; Venezuela, T.F. Amazonas, Santa Cruz, Rio Ata- 
bapo, Rio Atacavi; VEN (holo). 

ogg huberi Steyerm., Ernstia 23: 37. 1984. O. 
Huber & E. Medina 5786; 6 Feb. 1981; Venezuela, 

.F. Amazonas, Dept. Río Negro; 350 m; MO 
0 VEN (holo). 
paneopsis maguirei Steyerm., Mem. New York Bot. 
ice 17(1): 287. 1967. B. Maguire et al. 41438; 
13 Sep. 1957; Colombia, Cacagual Savanna, Rio Ata- 
o, between San Fernando de Atabapo and Cano 
Temi; 135 m; NY (holo). 

Sipaneopsis morichensis oe Mem. New York Bot. 
Gard. 17(1): 286. 1967. B. Maguire et al. 30878; 
14 Jan. 1951; Veriernelà: azonas, Cerro Mo- 
riche, Rio Ventuari; 800 m; NY (holo) 

Sipaneopsis pacimoniensis Steyerm w York 
Bot. Gard. 17(1): 289. 1967. B. Maguire et al. 37570; 
1 Feb. 1954; Venezuela, T.F. Amazonas, Rio Pacimoni; 
100-140 m; NY (holo). 

E odes w aa Steyerm., Mem. New York Bot. 
Gard. 17(1): 288. 1967. B. Maguire et al. 37639; 9 
Feb. 1954; d. T.F. Amazonas, Cano Hechi- 
moni; 100-130 m; NY nid 

Rr perijaensis Steyerm., Acta Bot. Venez. 8: 
252. 1973. J. A. Sana be et ur 1057 47; 31 Mar. 

1972; Venaxielix. Zulia, Sierra de Perija; 1,140 m; 
VEN (holo). 

Sphinctanthus insignis Steyerm., Brittonia 33: 400. 
1981. R. P. Belem & R. S. Pinheiro 2738; 8 Oct 
1966; Brazil Bahia, Guaratinga; VEN (holo), NY. 

Stachyarrhena acutiloba Steyerm., New York 
Bot. Gard. 12(3): 220. 1965. 4. Ducke 1844: 24 Nov. 


D s, river mar 
gin between San Fernando de Atabapo and Cacagual; 
130 m; NY (holo). 

Tocoyena brevifolia Steyerm., Mem. New York Bot. Gard. 
12(3): 196. 1965. Ll. Williams 16039; 7 May 1943; 
Venezuela, T.F. Amazonas, baja Sanariapo, laja Ca- 
restia; 120 m; VEN (holo), F. 

Tocoyena costanensis Steyerm., Bol. Soc. Venez. Ci. Nat. 
25(106): 73. 1963. F. Delgado 160; 22 d 1938; 
Venezuela, Distr. Federal, Avila; VEN bs lo). 


; LR n 1433; 

n.d.; Venezuela, Mérida, E. al VEN el. 
Tarena cuatrecasasii Steyerm - Bot. Venez. 4(1): 
113. 1964. J. Cuatrecasas 919 
lombia, Comisaria del Caqueta, ae Oriental, Su- 
cre, Quebrada de la Calana; 1,000-1,100 m; US (holo). 


6 Apr. 1940; Co- 


Tocoyena guianensis K. Schum. var. lesa Stey- 
erm., York Bot. Gard. 12(3): 1 1965. 
H.-S. buin et al. 47620; 18 Aug. 1960; Brad Terr. 
Amapa, Rio Oiapoque opposite Pedro Alice; NY (holo). 

Tocoyena guianensis K. Schum. var. pans ula Stey- 
erm., Mem. New York Bot. Gard. 12(3): 1 1965. 
B. Maguire & J. J. Wurdack 34923; Venecie T.F. 
Amazonas, Piedra Cucuy, Rio Negro; 100-200 m; 
depository not designated. 

Tocoyena orinocensis Standl. & Steyerm., Fieldiana Bot. 
28(3): 617. 1953. J. A. Steyermark 58442; 8 Sep. 
1944; Venezuela, T.F. Amazonas, below mouth of Rio 

'anariapo, along Orinoco River; 100 = F (holo). 
M en thyrsiflora Steyerm., Bol. . Venez. Ci. 
25(107): 248. 1964. 4. L. dar 56294; 23 
Nox 1956; Venezuela, Miranda, selvas pluviales del 
o 400-600 m; NY (holo 

Yutajea dn Ann. Missouri Bot. Gard. 74: 676. 
1987, . nov. 

Yutajea liesneri Steyerm., Ann. Missouri Bot. Gard. 74: 
676. 1987. R. L. Liesner & B. K. Holst 21826; 10 
n 1987; Venezuela, T.F. Am oe Pr 

8 km NW of Yutaje settlement, 1 W 
Coro Coro, W of Serrania de Yutaje; MO (holo), E EN. 


RUTACEAE 


Amyris ignea Steyerm., Fieldiana Bot. 28(2): 272. 1952, 
nom. nov. 

Casimiroa emarginata DE & ng Field Mus. 
Nat. Hist ke Ser. 23(4): 165. 1944. J. A. Steyer- 
mark 369. ; 20 Feb. 1940; Guatemala: Dept. San. 
Marcos, Tajurnuleo, Volcán Tajumulco; 2,300-2,800 
m; F (holo 

Diomma frutic osa Steyerm., Fieldiana Bot. 28(2): ue 
1952. J. A. Steyermark 60820; 20 Nov. 1944; Ver 
a a Rio Karuai, between La Laja and Sania 
Teresita de Kavanayen; 1,220 m; F (holo). 

Esenbeckia echinoidea Standl. & Steyerm., Field Mus. 

Nat. Hist., Bot. Ser. 23(4): 164. 1944. P. C. Standley 
74456; 15 Oct. 1940; Guatemala, Dept. Chiquimula, 
between Ramírez and Cumbre de Chiquimula; 400- 
600 m; F (holo 

Galipea guatemalensis Standl. & UM. Field Mus. 

, Bot. Ser. 23(4): 165. 1944 A. Steyer- 

mark 39448, 14 Apr. 1940; Cuatemala, Dept. Izabal, 

Rio Dulce; F (holo). 

Lubaria szczerbanii o pom 32: 17. 1980. 

Steyermark et al. ; 23-24 May 1978; 

Venezuela, Bolivar, T eee a iu largo del 
afluente de Río Carapo, río arriba del S 

730-750 m; US, VEN (h 

Rauia subtruncata Steyerm., 
75: 315. 1988. R. L. Liesner : 
1-3 Aug. 1978; d Bolivar, 20-25 kr 
of Manteco, on roa ro de las Dos Bocas; 
200 m; MO EE 3890481), VEN. 

Ravenia ruellioides p. var. ptariana Steyerm 
Fieldiana Sn 2802) 2 Shevermark 
59511; 944; a Bolivar, Ptari-tepui; 
1,810 m; F (holo) 

E aracaensis Kallunki & Steyerm., pisa 
39: 409. T. Prance et al. 29016; 12 Feb. 
1984; B ds Serra Aracá, e of N 
Massif, S extreme of N part of Aracá; F, INPA (holo), 


olo). 


Ann. juo Bot. Gard. 
zale 


Raveniopsis breweri Steyerm., Brittonia 32: 48. 1980. 


728 


Annals of the 
Missouri Botanical Garden 


A. Steyermark et al. 116166; 28 Feb. ; Ven- 

ezuela, Bolívar, Auyan-tepui, N de la Misión ER Ca- 

; VEN (holo). 

Raveniopsis Die qe Hey & Luteyn, Ann. Mis- 

1984. J. Luteyn 9413; 7 

984; ea Río Negro, Cerro de la Neb- 
lina: l, e m; VEN (holo). 

Raveniopsis qaeri Steyerm., Soc. Venez. Ci. 
Nat. 32(132/133): 338. 1976. . Steyermark et 

974; Vene abla: Bolívar, Meseta 


Bol. 


oL 410 m; VEN (holo). 
Rav eniopsis liesneri Steyerm., Ann. e Bot. Gard. 
75: 317. 1988 L; Liesner 16605 N 


1984; Venezuela. T.F. Amazonas, De Rio Negro, 
Cerro de la s Camp 4; 780 m; MO (holo — 


3405796), V 
Hutanebliua oA & Luteyn, Ann. Missouri Bot. 
Gard. 71: 1984, gen. nov. 


Rutaneblina pusilla Steyerm. & Luteyn, Ann. Missouri 
Bot. Gard. 71: 314. 1984. J. A. Steyermark 129798; 
8 Feb. 1984; Venezuela, . Amazonas, Cerro de 
Neblina; 1,880 m; MO (32231 49), VEN (holo). 

Zanthoxylum aguilar Standl. & Steyerm., Field Mus 
Nat. Hi 22(3): 146. 1940. P. C. Standley 
63209: 21 Jan. 1939: Guatemala, Dept. Sacatepéquez, 
above Dueñas; 1,600-1,800 m; F (holo). 

Zanthoxylum nion 29 > "wu pu 
Nat. Hist., Le 22(4): 2 1940 


ist., Bot. Ser 


Field Mus. 
Steyer- 


mark 314 Ls Nov. 1939; gere Dept. Chi- 
quimula, Voled de ocre eps ue, e mi. NE of 
uezaltepeque; d. 500-2,000 m; F (holo). 


Peur: Bot. 28(2): 
ormark 5 ; 18 May 
2.745 m; F 


Za iat y n go Se erm., 
213 A ME 


deris TREE kid Tabay, ese 


(holo). 
Zanthoxylum tachirense Steyerm., Fieldiana Bot. 28(2): 
274. 1952. J. A. Steyermark : 57111; 7 July 1944; 


Venezuela, Táchira, above La Grita, below Páramo de 
la Negra; 2,430-2,510 m; F (holo). 


SABIACEAE 
Meliosma in Standl. & ds Field Mus. Nat 
Ser. 23(2): 60. 1944 A. Sanat 
46773; May 1 1942; Guatemala, Dept. Er gn 
Volcán de Santa Clara, above Chicacao; 1,250 r 
(holo). 

Meliosma maxima Standl. & Steyerm., ve Mus. Nat. 
ist., Bot. $ 23(2): 61. 4. Steyermark 
38170; Mar. 1940; Guatemala, Dept. aun betw 
Bananera and La Presa, Montana del Mico; 300 m; E 
holo 

Me ema nubicola Steyerm. & Lasser, Bol. Soc. Venez. 
4. Nat. 26: 463. Steyermark & M. 
Farinas 90967; 21 Oct. 1962; “Venezuela, Distr. Fed- 


Aa 


eral, Altos de No León; 2,400 m; VEN (holo). 
uet V du Steyerm., er Bot. 28(2): 
54. 1952 . Steyermark 61524; 18 Mar. 1945; 


Vnd Jinsodiesui. Quebrada Hk 1,000- 1,100 
m; F (holo). 

cede tepuiensis Steyerm. & Maguire, Mem. New 
n A Gard. 17(1): 448. 1967. J. 4. Steyormark 
7570 19 June 1953; Venezuela, Bolivar, Apacara- 
tepul, cae Massif; 1,900-1,950 m; “NY (holo). 


SALICACEAE 


ed €— Muehlenb. f. mollis erum & Steyerm., 
. Missouri Bot. Gard. 25: 77 8. J. A. Stey- 


ermark 12491; 4 Aug. 1936; U.S.A., Missouri, Dent 
Co.; MO (holo— 1174271). 

Salix puero Muehlenb. f. aiu i Palmer & Stey- 

Ann. Oe Bot. Gard. 25: 770. 1938. J. 4. 

J 10; 28 Sis 1936; U.S.A., Missouri, 
Phelps Co., e of lake at Yaney Mills Spring; MO 
(holo) = S. rigida Muehlenb. f. subintegra (Palmer & 
Steyerm.) Steyerm., 1960. 

Salix sericea Marsh f. a Palmer & Steyerm., 
Missouri Bot. Gard. 2 
12548; 4 Aug. 1936; ts 
(holo 1130660). Note: The original description er- 
roneously cites two different collections with the number 
12918 as both the holotype and paratype. 


Ann. 
A. Steyermark 
a Missouri: Dent Co.; MO 


SANTALACEAE 
Thesium tepuiense Steyerm., Peldisna Bot. 28(1): 225 
1951. J.. Steyermark 00237; 15-17 Nov 
Venezuela, Bolivar 


, between Pin -tepui and Sororopan- 
tepui; 1,615 66). 


m; F (holo), MO (162126 


SAPINDACEAE 


md parimensis Steyerm., Ann. osea Bot. Gard. 
333. 1988. J. A. Steyermark 107€ 33; 18 Apr. 
23 May 1973; Venezuela, T.F. iE Sierra Pa 
rima, vecinidades de Simarawochi, Rio Matacuni; 795 
830 m; MO (holo — ipie VEN. 
Cupania andina Steyerm., Bol. Soc. Venez. Ci. MET eit 
463. 1966. Steyermark 56301; 4 May 1944; 
V enezuela, Mérida, between Mucuty and e 


1,065-2,430 m; VEN (holo 


Cupania kavanayena Steyerm., Fieldiana Bot. 28(2): 
343. 1952. J. 4. S mE 60912 ; 11 Dec. 1944; 


V enezuela, Bolivar, Quebrada « Hie 1,200- 
1,220 m; F (holo) = Matayba heh 'anayena (Steyerm.) 
Steyerm., 

C cies E a Steyerm., ldiana Bot. 28(2): 344. 
1952. J. Steyermark : 9182; 2 Oct. 1944; Ven- 


ezue ich. Bolivar, Gran "die 1,065 m; F (holo 
C i roraimae Steyerm., T Bot. 28(2): 346. 
52. J. A. Steyermark < )/; 1 Oct. 1944; Ven- 


i Bolivar, Gran cai 1,065 1,220 m; F (holo). 
finan tepulensis Steyerm. & 
York Bot. Gard. 17(1): 
& J. J. Wurdack 1133; 
Bolivar Sey ps Chane Massif; 1,880- 
1,95 5 m; MO (3¢ 63), NY (holo). 
Dipterodendron venezue bin Steyerm., Fieldiana Bot. 
28(2): 346. 1952. J. A. Steyermark 56569; 15 May 
1944; Venezuela, Mérida, between San Isidro Alto 2 


1955; Venezuela, 


Santa Cruz de Mora; 760-1,800 m; F (holo) = Dilo- 
dendron elegans (Radlk.) Gentry & Steyerm., 1987. 
Llagunoa venezuelana Steyerm., Fieldiana Bot. 28(2): 
347. 1952. J. A. Steyermark 56507; 14 May 1944; 


Venezuela, Mérida, above El Molino; 2.010- 2,135 m; 
F (holo). 
Matayba affinis Pa es Ann. Missouri i Gard. 75: 
067. 1988. Davidse et al. 17011; 30 Apr.-1 
May 1979; E T.F. Amazonas, Dept. Atabapo, 
al de Caname, S bank of s middle part of 
Caño Caname; 100 m; MO (holo —2744469), VEN 


Matayba chimantensis Steyerm., Mes. New York Bot. 
Gard. 17(1): 447. 1967. J. A. Steyermark 74656; 25 
ar. 3; Venezuela, Bolivar, Río Apacara, Apacara 


tepui, Chimantá Massif; 400 m; MO (2011418), VEN 
(holo). 


Volume 76, Number 3 
1989 


Taylor 729 
Plants Described by 


Julian A. Steyermark 


Maytayba jauaensis Steyerm., Bol. Soc. Venez. Ci. Nat. 
32(132/133): 347. 1976. J. A. Steyermark et al. 
09673; 28 Feb.-1-2, 5 Mar. 1974; Venezuela, Bo- 
livar, Meseta del Jaua, Cerro Jau ua; l, pod 880 m; 
VEN (holo) = M. oligandra Sandw 
hr longipes Radlk. subsp. bn “Ste 


nn. Missouri Bot. Gard. 75: 1069. 1988. R. S. n 
. J. Wurdack 31450; 13 Feb. ie Venezuela, 
T.F. Amazonas, Serranía Parü, Caño Asisi, Top Camp 


to Cano Camp: 1,400 m; MO Tune UN NY. 
Matayba oligandra Sandw. var. occidentalis Steyerm. 
Ann. Missouri Bot. Gard. 75: 1069. 1988. B. Maguire 
& L. Politi 28399; 17 July 1949; Venezuela, T.F. 
RE Cano Grande, Rio ^ Río Orinoco; 125 


O (holo— 2934861), N 
"ns ba iie Dor. Fein Bot. 28(2): 350. 
1952 mark 60203; 15 Nov. 1944; Ven- 


ua. Bolivar. Salto de cee a -meru; 1,015 m; F 

(holo). 

Matayba ptariana di e subsp. guaiquinimae Stey- 

erm., Ann. Missouri Bot. Gard. 635. 19 
Huber 10388; 27 Mar. 

. Heres, Meseta de T 1,400 r 
(holo— 3382924), NY, VEN. 

Matayba reducta Steyerm., Muere Bot. 28(2): 351. 
1952. J. A. Steyermark 60365; 20-21 Nov. 1944; 
Venezuela, Bolívar, Quebrada O» paru ma; 1,065-1,220 
m; F (holo). 

Matayba a Steyerm., Ann. Missouri Bot. Gard. 
75: 1070. 1988. - C. K. di 35143; 11 Feb. 
1953; Venezuela, .F. Amazonas, Serrania Yutaje, 
NW Ridge; 1,400 m; MO olo 2084874) NY, VEN. 

Mata yba sororopaniana Steyert a Bot. 28(2): 
352. 1952. J. Steyermark 602 ; 15 Nov. 1944; 
s Bolivar, Sororopan- a i ,656-1,980 m; 
F (holo). 


1985. la; Bolívar 
; MO 


ar Bot. 
d Nov 


Matayba venezuelana Steyerm. 28(2): 
353. 1952. J. A. Steyermark 606 1944; 
Venezuela, Bolivar, Ptari-tepui; ooa F (hola). 

Paullinia manarae Steyerm., Acta Bot. Ve nez. 3: 202. 

Manara s.n.; 4 Oct 1965; Venezuela, 

Distr. Federal, Naiguatà, Lon de Las Delicias, Que- 

ee de Basenilla y Quebrada Guayoyo; 1,390-1,500 
; VEN (holo). 

Paullinia mexiae Steyerm., Field Mus. Nat. Hist., Bot. 
Ser. 22(3): 155. 1940. Y. Mexia 8855; 19 Nov. 1937; 
Sn Guerrero, Distr. Adama, Temisco, Barranc 

Sierra Madre del Sur, N of Rio Balsas; 
F (holo), MO (1181481). 
Paullinia aco Steyerm., Fieldiana Bot. 28(2): 353. 
Steyermark 60579; 25 Nov. 1944; Ven- 
ezuela, Ren Quebrada O- -paru- ma; 915-1,065 m; 
holo 


Paullinia naigualensis Steyerm., Bol. Soc. Venez. Ci. 
f A. Stey "venari 91874; 2 Nov. 
1963; Venezuela, Distr. Federal, Cerro Na VEN 
(holo 
Paullinia nuriensis Steyerm., Acta Bot. Venez. * 179. 
1968. Steyermark 89005; 3 Feb. 1961; Ven- 
ezuela, Bolívar, Cerro El Picacho, Lu de Nuria; 
620 m; NY (holo). 
Paullinia toxicodendroides Steyerm., 
26: 460. 1966. J. 


Bol. Soc. Venez. 


.«& EC Field M 
ia 171. 1944. J. A. reps 
43926; 19 Feb. 1942; Guatemala, Dept. Alta Verapaz, 


along road between San Cristóbal and Chixoy; 1,200- 
1,300 m; F (holo). 

Serjania e Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(1): 14. 1943. P. C. Standley 73715; 
Oct ne Cas. Dept. Chiquimula, near th 
divide on road from Zacapa to Chiquimula; 660 m; F 
(holo). 

Serjania macrocarpa Standl. & Steyerm., Field Mus. 

t. Ser. 23(1): 15. 1943. W. A. Kellerman 


7532:-23 Fe b 1908; Guatemala, Dept. Izabal, Los 


Amates; F (holo). 

Serjania ds Hin Standl. & P Field Mus. 
Nat. Hist., Bot. 23(4): 171. 1944 A. Steyer- 
mark 50908; Ti gom 1942; Duatema " ept. Hue 


huetenango, along Rio pud ali Gullo and San 
Juan; 1,200-1,300 m; F (ho 

Talisia caudata Steyerm., Dn Missouri Bot. Gar 
1072. 1988. B. Maguire & L. Politi 28615; 25 dor 
1949; Venezuela, T.F. Amazonas, Cerro Sipapo, trail 
from Base Camp; 125 m; MO (holo— 3304133, 
3304132), NY. 

Talisia eM es a 2d Ann. Missouri Bot. Gard. 

l Mori et al. 15027; 1 Oct. 1982; 
French dp E Monte La Fumée; 200-400 m; 
MO (holo —3427739), NY. 

Talisia iri Steyerm., Ann. Missouri Bot. Gard. 
75: 1073. 8. R. L. Liesner & A. González 11151; 
n.d.; E Boliv: ^, Repressa Guri islands, 6-18 
km S of dam; 220-240 m; MO (holo— 2926530), 
VEN 


Talisia maruayana Steyerm., Ann. Missouri Bot. Gard 
75: 1070. 1988. R. L. Liesner & B. K. Holst nn 
26 Apr. 1986; Venezuela, Bolivar, Amaruay-tepui; 550- 
810 m; MO (holo —3446405, 3446406, 3446407, 
3446408), M da e — T. amaruayensis). 

Talisia morilloi S ot. Venez. 10: 237. 

975 Morillo. jr a EUN in 2 1973; Vene 

), Quebrada Río 


rande; 90 
Talisia pentantha Steyerm., Mustela Bot. Gard. 
075. 1988. F. Ehrendorfer 74104-23; 4 Oct. 
1974; Venezuela, Bolivar, Canaima, W of Avensa Camp; 
500 m; VEN (holo 

Talisia uo Steyerm., Ann. Missouri Bot. Gard. 
795: 2 de 988. L. Marcano Berti & P. Alcedo 119- 
979; 11- T Mar. 1970; Venezuela, s Amazonas, 
E San Carlos and Solano; MER (holo). 

Talisia tiricensis Steyerm. & ba wk Mem. New 

448. 1967. J. A. Steyermark « J. 

055; Venezuela, Bolivar, 
Rio Tirica (Rio Aparuren), above Techine-mero; 470 
m; NY (holo). 

Toulicia anomala Steyerm., Ann. Missouri Bot. Gard 
75: 1076. 1988. J. J. Wurdack & J. V. Monachino 
41253; l7 Jan. 1956; Venezuela, Bolivar, Rio Sua- 
pure, middle Orinoco; 110-120 m; MO (holo— 
3310903), NY. 


SAPOTACEAE 


Eoo ficoides Steyerm., Acta Bot. Venez. 2: 
279. Steyermark 934 431; 5 May 1964; 
cala. D Auyan-tepui; VEN (hol 
Chrysophyllum monachinoanum Steyerm., Bo oL S oc. Ve- 
nez. Ci. Nat. 26: 432 A. si aad et al. 
92333; 29-30 Dec. 1963: Venezuela, Bolivar, Cerro 
Venamo; Tu l, 150 m; VEN (ho 
Pouteria auyan is Steyerm., rom Bot. Venez. 2: 


730 Annals of the 
Missouri Botanical Garden 
Is 1967. J. 4. Steyermark 93917; Venezuela, Bo- idend 95418; 1 Apr. 1966; Venezuela, Carabobo, 


var, rune -tepui, 1,850 m; pane not designated. 
Hxc spectabilis Steyerm., Bol. Soc. Venez. Ci. Nat. 
A. Steyermark n S. Nilsson 768; 
24-25 Apr. 1960; Venezuela, Bolivar, near headwaters 
of Rio T NE of Luepa; 1,200 m; VEN (holo). 
Pouteria 1 , Bol. Soc. Venez. Ci. Nat. 
26: 434. 1966. ra 4. Stbyeraiak et al. 92414; 29- 
30 Dec. 1963; Venezuela, Bolivar, Cerro Venamo; 
950-1,150 m; VEN (holo) 


SARRACENIACEAE 


Heliamphora heterodoxa Steyerm., Fieldiana Bot. 28(1): 
239. 1951. A. Steyermark 59766; 2 Nov. 1944; 
Venezuela, Bolivar, Ptari- -tepui; 2,000-2,200 m; F 
(holo). 
Heliamphora heterodoxa Steyerm. var. exappendicu- 
aguire & Steyerm., Mem. Ne 
29: 54. 1978. J. J. W 
TR Bolívar, Chimantá- EE NW slope oft ihi 
ri-tepui; 2,050 m; NY (holo 
eras ie heterodoxa Bio m. var. ui re 
mn ta apice Steyerm., Ann. Missouri Bot. 
LL. . J. A. Steyermark 93712; 11 vie 
1904: oe Bolivar, Auyan-tepui; 1,600- 1,800 
; VEN (holo 
He dor minor Gleason n ferg Steyerm., Ann 
Missouri Bot. Gard. 71: 311. 1984. F. Cardona 2661: 
Jan. 1949; Venezuela, Ao Cerro Auyan; 2,100 
m; VEN (holo). 


SAXIFRAGACEAE 


Heuchera n rula Mack. & Bush f. glabrata n erm., 


Rhodora A. Steyermark 6 5: 9 
Oct. Due U.S SA., Missoni: Shannon = len Up 
Bluff along Jack's Fork of Current River, 4 mi. N of 
Teresita, 6 mi. NW of Monteer; F (holo). 
SCROPHULARIACEAE 
Acta Bot. "e 10: 


es eolaria gaa Steyerm., 
1975. J. A. Ste yermark et al. dde 22-23, 

> Mar 197 2; Vene zuela, Zulia, rra de Perija; 
1,440 l, 460 m; US, V EN (holo) = C ds xicana Benth. 
subsp. prostrata (Kranzlin) Molau, 

C ‘astilleja ms a 


ied 


e Steyerm., Mus. Nat. 
Hist., Bot. Ser. 23(2): 85. 1944. Steyermark 
49908; Aug. 1942; cabe Dept. Huehuetenango, 


Cerro Chemalito, Sierra de los Cuchumatanes; 3,100- 
3,150 m; F (holo). 

Pedicularis canadensis L. f. albescens Steyerm., Rho- 
dora 54: 258. 1952. J. 4. Steyermark 67499; | May 
1949; U.S.A., Missouri, Wright Co., along route 5, 
2.5 mi. S of Mansfield; F (holo). 

in digitalis (Sweet) Nutt. f. baueri Steyerm., 
Rhodora 43: 663. A. Steyermark Un 1 
July 1939; U.S.A., Miss Osage Co., 3 mi. NE of 
Westphalia, along Maries River: E (holo »). 

Russelia Ted onm P 
Ser. 22(5): € 940. J. . | Steyarmark 37 982: 16 
Mar. 1940; a ine San Marcos, above Finca 
El Porvenir, along Rio Cabüs near Cueva de las Pal- 
omas, Volcán Tajumulco; 1,300-1,500 m; F (holo). 


SELAGINELLACEAE 


Selaginella vicine f & A. R. Smith, es 
lissouri Bot. Gard. 73: 209. Das 4. A. & ( ey- 


entre. Los Tanques y La Toma, Rio San Jian; 750 m; 
Us (holo), VEN. 


SIMAROUBACEAE 


Picramnia caracasana oy a A w Fs Muriel 
Fie iana Bot. 2 n 274. J. eye mark 
019; 25 June 1944; Vene s e | dis 
of El Junquito; l, a 2 130. m; F agi 
Picramnia nuriensis Steyerm., Acta Bot. Venez. si 7 
Sone 88591; 19-23 Jan. 1961; 
Venequelá, Bolivar, Cerro El Picacho; ia: de 


Nuria; 450-620 m; NY (ho E 
Picramnia pou Steyer | Fieldiana Bot. 28(2): 
275. 1952. Ll. Williams 13918; 2 23 Jan. 1942; Ven- 


ezuela, re Amazonas, camino de Yavita; 128 m; F 


(holo). 


SMILACACEAE 


SHADE auraimensis Steyerm., Bol. Soc. Venez. Ci. Nat. 
2 12. 1966. ata 90823; 16 Jan. 
1962: Venezuela, Bolivar. Sierra Auraima; 400 m; 
VEN (holo). 

dines chimantensis Steyerm. & Maguire, Mem. New 
Yo t. Ga 1 7(1): 440. xdi J. A. cag da ei 

25 Mar. 1953; Vene , Bolívar, Río Apa- 

Chimantá prie 400 1 m; NY 


y 74652. 
cara, Apacara-tepul, 
(holo 

Smilax duidae a Marre e 28(1): 154. 1951. 

4. S rmark 5 ; 26 Aug. 1944; Venezuela, 
T F. focii in Duds; Caño Negro; 260-610 
m; F (holo). 
Smilax jauaensis Steyerm. & Maguire, Mem. New York 
. 1972. J. A. Steyermark 98073; 
22-27 Mar. 1967; Venezuela, Bolivar, Cerro Jaua; 
1,922-2,100 m; VEN (holo). 

Smilax lasseriana Steyerm., Fieldiana Bot. 
1951. J. Steyermark 60121; Venezuela, 
Sororopan- tepui; 2,255 m; F (holo). 

Smilax pud meyer ae Bot. 28(1): 155. 

ER 6025 -17 Nov. 1944; 
V ee Bolivar, on mesa ede Ptari-tepui and 
Sororopan-tepui; 1,615 m; F (holo). 
Smilax staminea Griseb. f. obtusata Steyerm., Fieldiana 
J. 


-~ 


28(1): 154. 


Bolívar, 


Bot. 28(1): 156. 1951. Steyermark 59695; 1 
Nov. 1944; Venezuela, Bolivar, Ptari- -tepui; 1l, 700- 
1,800 m; F (holo). 
SOLANACEAE 
Athenaea loc me s pee o n Field Mus 


Be Bot. Ser. 22(5): : 1940. gode sine 

7953; 16 Mar. 1940; e Dept. San d iue 
ed Finca El — along Rio Cabus, near Cue 

de las Palomas, Volcán Tajumulco; 1,300- 1,5 


F (holo) — Witheringia stramonifolia H.B.K., 
Athenaea macrocardia Standl. Sí Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 22(5): 375. 1940. J. A. Steyer- 


mark 30004; 14 Oct. 1939; Guatemala, Dept. Zacapa, 
Río Lima, Sierra de Las Minas, below Finca Alejandría; 
2,000 m; F (holo) = 4. viscosa (Schrader) Fern., 1974. 

Cestrum aristeguietae Steyerm., Acta Bot. Venez. $ 
1971. J. A. & C. Ste armani 95463; 2 Apr. 1 
Venezuela, Carabobo, Rio San Gean, S of a 
350 m; US VEN peta 

Cestrum pariense Steyerm., Acta Bot. Venez. 1(2): 62 
1966. J. A. Stey ad & G. Agostini 91026; 18 


Volume 76, Number 3 


Taylor 
Plants Described by 
Julian A. Steyermark 


July 1962; Venezuela, Sucre, Cerro Patao; 850 m; 
VEN (holo 
C yphomandra bolivarensis Steyerm., Bol. . Venez 
Ci. x 26: 441. 1966. J. 4. Steyr. pon 13 
'. 1944; Venezuela, Bolivar, Sororopan-tepui; 2,225 
2.255 m; VEN (holo). 

Cyphomandra depa ps in pd Field Mus. 
Nat. Hist., 22(5) 376. 1940 Steyer- 
mark 337 43: 8 p^ 1940; e Dept. Quezal- 
tenango, Volcán Santa María; 1,300-1,500 m; F n 

Cyphomandra villosa Steyerm., ud 9(6): 3 
1964. M. Acosta Solis 7743; 12 Apr. 1944 M ne 
Prov. Loja, region central, Las Chinchas; 2.9 50m;F 
(holo). 

Lycianthes jage Standl. & eyer Field Mus. 
Nat. Hist., 23(5): 228 194 . J. A. Steyer 
mark 46653; D Máy 1942; used Dept. Such- 
itepéquez, Volcán de Santa Clara; 1,250-2,650 m; F 
(holo). 


Lycianthes chiapensis (Brandegee) Standl. var. sparst- 


stellata Standl. & a Field Mus. Nat. Hist., B 
Ser. 22(4): 274. A. Steyermark 31555; 10 
Nov. 1939; — Mur Chiquimula, Cerro Tixixi; 


500-1,500 m; F (holo) = L. chiapensis (Brandegee) 
ipe var. "iapens 1976 


Lycianthes andl. & Steyerm, Field Mus. Nat. 
Hist. e Ser. a > 229. Steyermark 
52104; Sep. 1942; ees Dept. ¿himalte- 
nango, iw slope Volcán Fuego; 1,200-1,600 m; F 
(holo). 


Markea costanensis Steyerm., Acta Bot. Venez. 3: 
1968. J. . Steyermark 94382; 12 Oct. 1965; d 
ezuela, Distr. Federal, between Colonia Tovar- Junquito 
road and Hacienda El Limón; 1,300-1,500 m; VEN 
(holo). 

Markea grandiflora Steyerm., Phytologia 9(6): 349 

6 Acosta Solis 7009; 18 Jan. 1944; Fonade 
Prov. Pichincha, baja e Loche a Condor Mackay, 
Cordillera oci 2,700- 3, 100 m; F (holo). 


olivar, Abacapa-tepui, Chi- 

manta Massif; 1,200-1,600 m; VEN (holo). 

Nierembergia e dre Steyerm., pec 9(6): 349. 
96: “spinosa 1076; 1946; Ecuador, 


rov. Loja, Namanda; 1,500 m; F (h holo). 
Phy an amica Standl. & ra Field V Nat. 
Hist., Bot. Ser. 23(5): 231. 194 E 


30175; 20 Feb. 1940; ela. Dept. San Marcos, 
between La Vega ridge and NE slopes Volcan Tacana, 


along Rio Vega; 2,500-3,000 m; F (holo). 

Phy salis enr nn Standl. & Steyerm., Field Mus. Nat. 

st. 23(5) 231. 1947. P. C. Standley 
07488; 4 Mar. 1939; Guatemala, Quezalte- 
nango, Fuentes Georginas, W Volcán Zunil; 2,850 m; 
F (holo). 

Phy salis campanula Standl. & Steyerm., Field Mus. Nat. 
Hist., Bo 23(1): 18. 1943. Pai ln 
36067; Feb. 1940: Guatemala, De t. San Marcos, 
Quebrada Canjula, between Sibinal and oe Volcán 
Tacaná; 2,200-2,500 m; F P n n 

Physalis carnosa Standl. 5 , Field Mus. Nat. 
Hist., Bot. Ser. 23(1): e A MEN 
37766; Mar. 1940; Guatemala, UM San Marcos 


Ocos; F (holo). 
Physalis chimalteca Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 23(25): 232. 1947. P. Standley 


61803: 3 Tan. 1939; Guatemala, Dept. Chimaltenango, 


Volcán de Acatenango, above Las Calderas; 2,400- 
2,700 m; F (holo) = P. calidaria Standl. & Steyerm., 
974, 
Phy. igo lassa Standl. & Steyerm., Field Mus. Nat. Hist., 

. Ser d 19. 1943. J. . ed 32868; 


temala, Dept. Jala a, betw 


and Montaña Ue ae 1,500-2,000 m; F (holo). 
Solanum alpicum Standl. & Steyerm., Field Mus. Nat. 
23(5): 232. 19 A. Steyermark 


50303; 8 Aug. 1942; Guatemala, De pt. Huehuetenan- 
go, Cerro Chemal, Ris de los Cuchumatanes; 3,700- 


3,800 m; F (holo) = S. demissum Lindl., 1974 
Solanum bi urtatum 2 VAR , Field Mus 
Nat. , Bot. Ser. 23(1): 20. 1943 Lo: Dal 


2012; Mar. t Guatemala, Petén, Lao de Yaxha; 
F (holo) = S. blodgetii Chapman, 197 
Solanum finum Standl. & Steyerm. 


UE Mus. Nat. 


Hist., Bot. Ser. 23(5): 234. 1947. P. C. Standley 
67473; 4 Mar. 1939; Guatemala, Dept. Quezaltenan- 


go, Fuentes Georginas, W Volcán Zunil; 2,850 m; F 
(holo). 

Solanum huehuetecum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 234. 1947 . Steyer- 
mark 51039; 23 Aug. 1942; Guatemala: Dept. 
Huehuetenango, above Democracia on trail to Jutal; 
1,000 m; F (holo). 

Solanum — Standl. & Steyerm., eae Mus. Nat. 
Hist., Bot. Ser. 22(4): 275. 1940. Muenscher 
ne 0; ie ii 1937; Guatemala, ug  Sololá, near 

; 3,150 m; F (holo). 

ma nitens Steyerm., Bol. Soc. Venez. Ci. Nat. 
26: 444. 1966. J. 4. Steyermark & S. Nilsson 185; 
ue 17 Apr. 1960; Vne Bolívar, Fila de La Dan- 

; 1,200 m; VEN (hol 


Solanun rufistellatum ida Bol. Soc. Venez. Ci. 
. Os NE Say En. & S. Nilsson 
17 8; 15-17 Apr. 1960; V enezuela, Bolivar, Fila de 


La Danta; 1,200 m; VEN (holo 
Solanum d ense Standl. a Users Field Mus. 
22(4): 2 1940. P. C. Sandy 
69756; 30 m 939: D ile Dept Baj a Vera- 
paz, N of Santa a F (holo) = 5. torvum je 
1974 


STAPHYLEACEAE 
cce api c b bre, Por Bot, 28(2): 341. 
1952. J. . eyermark 56455; 10 May 1944; Ven- 
dedos to El Molino, Cuesta de Rincón 


1,925-2,175 m; F (holo) 


ezuela, PR 
and Cuesta del Barro; 


STERCULIACEAE 
> Ac ta Bot. Venez. 
A. Steyermark 87782; 26 Nov. 
1960; Venezuela, Terr. Delta ion betwee 
Margarita and Puerto Miranda, Rio Acure; 80-100 


“Buettneria” venezuelensis Steyerm., 
3: 3. 1968. J. 


m; NY (holo). Note: Byttneria is the correct spelling. 
Melochia humboldtiana Steyerm., Fieldiana e . 28(2): 
952. J. 4. Steyermark 62436; 1 I 1945; 


Venezuela, Sucre, between Cocollar and b umi: 1,200 


m; F (holo). 


STYRACACEAE 

ara 4: 12. vu 
Veillo uela, Mérida, es o 
a Carbonera; 2.600 : m; n VEN (holo). 

Styrax costanus Steyerm., Fieldiana Bot. 


Styrax andinus Steyerm., 
197 


28(3): 490. 


732 


Annals of the 
Missouri Botanical Garden 


1953. J. 4. Steyermark 61575; 20 Mar. 1945; Ver 
ezuela, Anzoátegui, Bergantin, Cerro Peonia (Cerro Los 
Pajaritos); 1,600-1,800 m; F (holo). 

Styrax duidae Steyerm., Fieldiana Bot. oe : 491. 1953. 

Steyermark 58242; 2 Sep. 1 ; Venezuela, 
T.F. Amazonas, Cerro Duida; 1,025- Xo m; Mea 

Styrax as Steyerm., Bol. Soc. Venez. Ci. e 
390. 1976. J. A. Steyermark et al. 109574; I 
1974; Venezuela, Bolivar, Meseta del Jaua, Seamer 

; VEN (holo). 

Styrax longipedicellarus r us Acta Bot. died 10: 

V. Be 3; Apr.-June 1950; Vene- 
uela, T. F. Aman, so del Duida y Marahmaca, 
VEN (holo). 

Styrax teputense eii pot rm. & P Mn Mem. New York 
Bot. Gard. 17(1): 456. 19 Steyermark & J. 
J. Wurdack 1028; + m 1955: a Bolivar, 
Torono-tepui, Chimanta Massif; 2,030-2,150 m; NY 
(holo 

Styrax tepuiensis Steyerm. & Maguire subsp. guaiquin- 
imae Maguire & Steyerm., Mem. New York Bot. Gard. 
29(1): 218. 1978. B. Maguire 32758; 25 Dec. 1951; 
Venezuela, Bolivar, Rio Paragua, Cerro Guaiquinima; 
NY (holo). 

Styrax tepuiensis Steyerm. & Maguire subsp. huacha- 
macarii Maguire & Ste: yerm., Mem. New York Bot 
Gard. 29(1): 218. 1978. B. Maguire et al. 30001; 6 
Dec. 1950; Venezuela, Terr. Cerro Hua- 
chamacari; NY (holo) 

Styrax vulcanicola Sandi: Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 22(4): 2 1940. J. . pei 
36801; 27 Feb. 1940; le Dept. San dune 


ari 197% . urnes 


Amazonas, 


between town of Tajumue and Tecutla, NW 
of Volcán Tajumulco; 1,800-2,500 m; F (ho Pu 
Styrax wurdackiorum Save, Acta Bot. Venez. 10: 
241. 75. . Steyer mark et al. oe 6573; 7-10 
197 ic Minis sla, Bolivar, carretera El Dorado 


1,200- e m; VEN 


ene Sania Elena de Uairen; 


(holo). 


SYMPLOCACEAE 


Symplocos abietorum Standl. & Steyerm., Field Mus. 
Nat. Hist., Bot. Ser. 23(5): 221. 1947 M yer- 
ma a 48482; 9 July 1942; Guatémala. Dep Hue 
om jupes de Teja, Sierra de los Cae 

; F (holo). 

—€— ac Poe nsis Steyerm., Ann. Missouri Bot. 
Gard. 74: 1987. J. A. Steyermark et al. 131868; 
173.20 M s Venezuela, Bolivar, vicinity of 
Guadequen, Rio Acanán, affluent of Rio Carrao, W of 
Cerro Los Hue 470 m; MO (3514025), VEN 
(holc 

Symplocos chimantensis Steyerm. & Maguire, jie 

w York Bot. Gard. 17 7(1): 459. 1967. J. 4. 
E & J. J. Wurdack 1100; 23 Feb. 1955; Ven 
ezuela, Bolivar, Torono-tepui, Chimanta Massif; 1,95 

2,090 m; NY (holo). 

Sy plo os culminicola Ol. & B Cw 
Nat. E a Bot. Ser. 23(5): 222 4. Steyer- 
mark . 69; 8 Nov. 1939; TP Dept. Chi- 
nde ud de Que meer a 2,000 m; F RA 

Symplocos jauaensis Steyerm. & n guire, Mem. New 

« Bot. Gard. 23: 876. 1972 Steyermark 


tanes; 3,33 


Field Mus. 


98072 n 27 Mar. 1967; MGR Bolivar Cerro 
Jaua; 1,922 22.100 m; VEN (holo), NY. 


Symplocos lasseri Steyerm., Fieldiana Bot. 28(3): 492. 


1953. T. Lasser & J. A. Steyermark 55103; 30 Dec 
1943; Venezuela, Distr. Federal, Corera a enl 
above Caracas; 1,585-1,820 m; F (ho 
Sy dd os neblinae Maguire oa. Mem. New 
Yo Gard. 29(1): 229. 1978. B. Maguire et al. 
423 78; 13 Dec. 1957; Venezuela, Terr. Amazonas, 
Cerro de la Neblina; 2,200 m; NY (holo). 
Symplocos pittieriana Steyerm., Fieldiana Bot. 28(3): 
92. 1953. J. A. Steyermark 57335; 15 July 1944; 
Venezuela, Pickin, base of Paramo de Tama; 22.500. 
o 


Symplocos roraimensis Steyerm., Fieldiana Bot. 28(3): 
494. 1953. J. A. des i 58761; 27 Ld 1944; 
Venezuela, oliva . Mt. Roraima; 2,255-2,620 m; F 


<= 


(holo) = S. Savon oan Sleumer, 1978. 

Symplocos tamana Steyerm., ws Bot. 28(3): 495. 
1953. Steyermark 573 5 July 1944; Ven- 
ezuela, quce base of i dei. de Tamá; 2,500- 
2,895 m; F (holo). 

Symplocos vatteri Standl. & Steyerm., Field Mus. Nat. 

ist., Bot. Ser. 23(5): 1947. J. A. Steyermark 
48463; 9 July 1942; Guatemala, Dept. Huehuetenan- 
go, Rancho de Teja, Sierra de los Cuchuamatanes; 
3,330 m; F (holo). 

Symplocos yapacanensis Steyerm., Acta Bot. Venez. 10: 
de 1975. J. A. Steyermark & G. S. Bunting 1031853 

5-7 Mar 1970; Venezuela, T.F. Amazonas, Cerro 
pacana; 1,000-1,200 m; VEN (holo 


TAXODIACEAE 


Taxodium distichum Rich. f. confusum Palmer & Stey- 
Missouri Bot. Gard. 25: 169. 1938. J. 4. 
USA: Missoni. 
Wayne C , Markham Springs, 3 mi. W of Williams- 
ville; MO th olo). 


IEFFUIAINIBHALLAF, 


E ec Maguire & Steyerm., Mem. New York 
Bot. Gard. 29(2): 8. 1981, fam. nov 

er W gs & d Mem. New York Bot. 

ard. 29(2): 9. 1981, gen. : 

ui. aracensis Bur & Maguire, Acta Ama- 
zonica 16/17: 219. 1986/1987. J. Pipoly et al. 6838; 
18 Mar. 1984; Brazil, Amazonas, Serra Acaca, summit 
of re massif; 1,200 m; INPA (holo), MO 
(3327363), NY. 

alas e nsis Erg & Steyerm., Mem. 
New York Bot. Gard. 29(2): 1981. J. A. Steyer- 


livar, Auyan-tepui; 1,940 m; NY, VEN Mid 
Tepuianthus abe aie Maguire & Steyer 
ork Bot. Gard. 32: 16. LR. 
d d Cabrera 15056; 22-25 Jan. 1952; Colombia, 
aupés, near summit of Cerro Varii GH (holo). 
Merida sarisariname nsis "s & Steyerm., 


Mem. New York Bot. Gard. 32 "1981. J. A. Stey- 
ermark et al. 109237: 19-20 a 1974; Venezuela, 
Meseta del Jaua, Cerro Sarisarinama; 1,350 m; NY, 
"EN (holo). 

Tepuianthus : sari is Maguire & oo subsp. 
duidensis Maguire & Stey erm., Mem. New York Bot. 
Gard. 32: 18. 1981. B. Maguire et »i 29543; 19 


'. 1950; Venezuela, T.F. Amazonas, Cerro Duida; 
) 


Tepuianthus savannensis Maguire & Steyerm., Mem 


New York Bot. Gard. 29(2): 14. 1981. O. Huber et 


Volume 76, Number 3 
1989 


Taylor 733 
Plants Described by 


Julian A. Steyermark 


79; Venezuela, T.F. Amazonas, 


holo); NY. 


ul. 3759; 7 May 19 
Dept. Atabapo, Cano Yagua; 120 m; VEN 


~ 


Tepuianthus yapacanensis Maguire & Steyerm., Mem. 
New York Bot. Gard. 29(2): 12. 1981. B. Maguire 


e 30683; 3 Jan. 1951; Venezuela, T.F. Amazonas, 
Cerro Yapacana; 120 m; NY (holo) 


THEACEAE 
4copanea Steyerm., Ann. Missouri Bot. Gard. 71: 323. 
1984, gen. nov 
de an ahogadoi Steyerm., Ann. Missouri Bot. Gard. 
po J. A. Steyermark et al. 129924; 14-16 
84; Venezuela, Bolivar, Piar, Macizo del Chi- 
7 920 m; MO (3234986), NY, VEN (holo). 
Bonnetia a Steyerm., Ann. — En Gard. 
74: . O. Huber 9818; 19 Nov. 1984; Ven- 
ezuela, E Ptari-tepui; 2,400 m; MO, ae (holo). 
Bonnetia chimantensis Steyerm., Ann. Missouri Bot. Gard. 
71: 326. Steyermark & J. J. Wurdack 
080; 9 Feb. 1955; Verwiuela: lo Chimantá Mas- 
ono-tepui; 2,090 m; F, NY, N (holo). 
la EV Kobuski & Steyerm., 6 Arnold Arbor. 
29: 41 8. J. 4. Steyermark 58186; 1 Sep. 1944; 
tati E Amazonas, Cerro Duida, summit, Broc- 
chinia Hills; 1,700-1,980 m; F (holo), NY = Neo- 
gleasonia duidae (Kobuski & Steyerm.) Maguire, 1972. 
omita euryanthera Steyerm., Ann. Missouri Bot. Gard. 
74 47. 1987. J. . Steyermark et al. 124311; 14 
. 1981; Venemela, Bolívar, Meseta de Jaua; 2,020 
m; MO, NY (holo), VEN 
Bonnetia guaiquinimae Steyerm., Ann. Missouri Bot. 
Gard. be 102. 1987. J. A. Steyermark 117421; 26 
May 1978; Venezuela, Bolív ar, Cerro Guaiquinima; 
1.2 25 ^ m; MO, VEN (holo). 
Bonnetia huberiana Steyerm., Ann. Mise Bot. Gard. 
tle 326 e Steyermark "urdack 
955; Venal Bolívar, Chimantá Mas- 
US 


, VEN 


ds 


ma ns 


633; 9 Feb. 
sif, oo fe 2,165-2,180 m; F, NY, 


(ho 

nata liesneri Steyerm., Ann. Missouri Bot. Gard.: in 
press, . R. L. Liesner & F. Delascio 22105; 18 
Oct. 1 - Veneadeld. T.F. Amazonas, Dept. Rio Ne- 
gro, ie Aracamuni, Popa Camp; 1,550 m; MO 

(holo), VEN. 

Bonnetia maguireorum bi gs 

d. 71: 330. 1984, nom. 

Bonnetia mis Steyerm. 
71: 330. 1984, nom. nov. 

Bonnetia mo Steyerm. . Missouri Bot. Gard. 
74: 102 


Ann. Missouri Bot. 


pri Missouri Bot. Gard. 


87. J. A. Md et al. 115645; 23 
Feb. 197 78; Venezuela, Bolívar, Ptari-tepui; MO, “VEN 


(holo). 

a tepuiensis Kobuski & Steyerm., J. Arnold Ar- 
bor. 29: 399. 1948 A. Steyermark 60871; 5-6 
944; Venezuela, Bolivar, pam tepui (Ptari- 

epui); 1,675-1,980 m; F (holo), N 

oua tepuiensis Kobuski & uu. subsp. minor 

'erm., Ann. Missouri Bot. Gard. 71: 326. 1984 

4. Stey 'ermark & J. J. Wurdack 990; 20 Feb. 

, Chimantá Massif; 1,975 m; F, NY, 


Dec. l 


US, VEN (holo). 

Bonnetia toronoensis Steyerm., Ann. Missouri Dot. Gard. 
71: 327. 1984 Steyermark & J. J. Wurdack 
1047; 21 Feb. 95 js: Venezuela, Bolivar, Chimantá 
Massif, Torono-tepui; 2,030-2,150 m; F, NY, US, 
VEN (holo). 


s tristyla Gleason subsp. nervosa Steyerm., Ann. 
Missouri Bot. Gard. 74: 103. 1987. J. A. Steyermark 
LG. Ss. Bunting 103153; 5 May 1970; Venezuela, 
T.F. Amazonas, Cerro Yapacana, below summit; 825 
m; VEN w^ Note: Superfluous acu ed in: Ann. 
Missouri Bot. Gard. 74: 649. 1987. B. Maguire et al. 
30632; 2 Jan. 1951: Venezuela, T. F. Arona Cerro 
pacana; NY, VEN (holo) 
as fruticosa (Schrader) Kobuski var. chimantae 
nn. Missouri Bot. Gard. 74: 650. 1987 
A Steyermark & J. J. Wurdack 879; 17 Feb. 
1955; Venezuela, Bolivar, Chimantá Massif, central 


upper falls; 2,090 m; F, NY, VEN (holo). 

Laplacea pubescens Planchon & Linden ex Triana & 
Planchon var. minor Steyerm., Ann. Missouri Bot. 
Gard. 74: 652. 1987. B. Maguire et al. 30285; 17 

1959; Venezuela, T.F. Amazonas, Cerro Hu- 
achamacari, between Summit Camp amd SW escarp- 
ment; 1,850 m; NY (holo). 

ibm acrodantha Kobuski & Steyerm., Fieldiana 
Bot. 28(2): 381. 1952. J. 4. Steyermark 56582; 18 
May 1944; Venezuela, Mérida, above Tabay; 2,285- 
2,745 m; F (holo), GH 

Ternstroemia camelliifolia Linden & Planch. var. minor 
Kobuski & Steyerm., Field. Bot. 28(2): 383. 1952. J. 


2,130 em F (holo). 

Ternstroemia uniloc Pr Kobuski & Steyerm., Fieldi- 
ana Bot. 28(2): 383 52. J: A. = yermark 61626; 
20 Mar. 1945; Vene UR Anzoátegui, Cerro Paeonia 
(Cerro Los Pajaritos); 1,800-2,000 m; F (holo), GH. 


THYMELAEACEAE 


P dircoides Steye 1 Bot. 28(2): 
52. J. d: pulum 601 ; 14 Nov. 1944; 


y EE Bolivar, Sororopan- E 2255 m; F (holo). 
Daphnopsis ficina Standl. & Steyerm., Field Mus. Nat. 
Hist., Bot. Ser. 22(4): 254. 1940. J 
1442; Jan. 1932; Guatemala, Dept. Guatemala, La 
Cienaguilla, San José Pinula; 1,600 m; F (holo 

Daphnopsis guaiquinimae Steyerm., Ann. vos Bot. 

Gard. 74: 652. 1987. J. A. Steyermark et al. 117977; 
pr. 1979; Venezuela, Boliv ar, Cerro He di 
1,650 m; MO, VEN (holo). 

Daphnopsis d Standl. & an Field 
Mus. Nat Bot. Ser. 23(2): 68. 1944. J. 4. 
Steyermark pU 18 July 1942; Pun Dept. 
Huehuetenango, Cerro Cananá, Sierra de los ea 
matanes; 2,500-2,800 m; F (holo). 

Daphnopsis nevlingiana Steyerm., Ann. Missouri Bot. 
Gard. 74: 653. 1987. J. 7 Steyermark et al. 109199; 


ed 18 Feb. os e Venezuela, Bolívar, Cerro Sarisa- 

ama, Mes os summit; 5 NIU, VEN (holo) 
Daphnopsis retife ra s idl. n., Field Mus. Nat 
Bot. Ser. ipee y 254 d P rmark 


939; Cintemála. 
tween railroad station of Mita and ea Mita: 6 650 
m; F (holo) = D. monocephala Donn. Sm., 2 

c es amaz a Steyerm., Ann. Missouri Bot. 
Gard. € r1 . G. Wessels-Boer 1953; 5 Au 

ds s mazonas, 2.5 km S of Perte 


Ayacucho; m (PER MY. 


t 
7a S 
> 
c 
& 


734 


Annals of the 
Missouri Botanical Garden 


TILIACEAE 


E d zu & rU ., Field Mus. Nat. 
ist., Bot. Ser. 22(2): 157. 1940, gen. nov 
Mortoniodendron nemas Standl. & Steyerm., Field 
Mu ot. Ser. 22(3): 157. W.R 
Hatc h & C. L. Wilson 54; 21 4 y 1936; Guatemala, 
Dept. Isabal, Río Dulce: F (hok 
Mecanica Flur vid en Stodi. & Steyerm., 
i lus. Nat. Hist., Bot. Ser. 22(3): 158. 1940. 1 
G. Yuncker et al. 8519; 19 July 1938; Honduras, 
Dept. Atlántida, above Puente Alto stop on S.F. Co 
RR, E of La Ceiba; 240 m; F (holo). 


TRIGONIACEAE 


s costane nsis hs erm. & Badillo, Acta Bot. Ven- 
ci Madriz 37; 8 Oct. 1967; Vene- 
ds Ya cuy, Vah de Nirgua; 1,200 m; MY (holo). 
Trigonia bes im & Steyerm., Field Mus. Nat. Hist., 
Bot. Ser. 23(2): 59. 1944. P. C. Standley 78584; 28 
Nov. 1940; Guatemala, Dept. Santa Rosa, Rio Panal, 
Volcán Tecuamburro; F (holo) = T. eriosperma (La- 
marck) UR : Sons subsp. membranacea (A. C. 
Smith) Lleras, 1978. 


TURNERACEAE 


ies ipei standle yl Steyerm. ., Field Mus. Nat. Hist., Bot. 

| G. Yuncker et al. 8323; 

d 1938; Honduras, Dept. Atlantida, Salado Riva, 

above Salado, vicinity of La Ceiba; F (holo) = E. odo- 
rata Seeman, 1979. 

Erblichia xylocarpa var. ign Standl. & Steyerm., 
Field Mus. Nat. Hist., Bot. Ser. 22(5): 355. 1940. J. 
A. Steyermark 33448: D 1940; Guatemala, pi 

ada San Gerónimo, Finca 
ía; 1,300-2,000 m; F (holo) 

rata Seeman var. mollis (Standl. & Steyerm.) 


L. O. Williams, 1979 


URTICACEAE 
Boehme ria ramiflora i var. vinacea Steyerm., Acta 
3: 197 ven . J. A. & C. Ste yermark 
95187; 21-28 Mar. 1966; Venezuela, Carabobo, Río 
San Gian, Borlinrsta. "m Electrica, Los Teques, and 
La Tomia; 790 m; V EN (h olo), 


415. 1952. J. A. Steyermark 447 573 Nn. d: 
Alta Verapaz, alon and 
Cubilquitz to Quebrada n 250- 350 m; F (holo). 
Pilea mimema $ y , Fieldiana Bot. 24(3): 
417. 1952. YA R. Batek & Ç. ya Wilson 162; n.d.; 
Guía nala, Alta Verapaz, Sonenn, Cerro Sillab; F (holo). 
Urera lira St andl. & S ., Fieldiana Bot. 24(3): 
42 A. F. Sk ; Guatemala, Que- 
uc. Volcan Zunil; F (holo). 


VALERIANACEAE 


Vale riana delicata Standl. & Steyerm., Field Mus. Nat. 
Bot. Ser. 23(5) 255. 1947. J. . 
1942; Guatemala, Dept. 
nango, Pume Papal; 1,400-3,000 m; F (holo) = I. 
robertianifolia Briq., 1976 
Valeriana texana Steyerm., 
19: 393. 1932. J. A. 
3528; 21 July 1931; 


= 
es 
= 


Ann. Missouri Bot. 
Moore & J. 4. ol 
U.S.A., Texas, € ulberson Co., 


dalupe Mountains, upper McKittrick Canyon; 2,000 
m; - GH (holo), MO (1011707). 


VELLOZIACEAE 


Vellozia Fara Steyerm., m Bot. 28(1): 157. 
95]. dO ded 562 2 Sep. 1944; Ven- 
1azonas; Cerro s 1,025-1,200 m; 
tubiflora (A. Rich.) H.B.K., 1969. 


ezuela 


F eh, T = 


VERBENACEAE 

tegiphila dea Standl. & Steyerm., Field Mus. 
. Hist., Bot. $ 23(5): 227. 1947. J. A. Steyer- 
mark 48843; = 16 July 1942; Guatemala, Dept. 
Huehuetenango, near Maxbal, Sierra de los Cuchu 
mantes; 1,500 m; F (holo) = ET Lue nicotanifolia 
(Mart. & Gal.) L. Williams, 1975 in the Rubiaceae not 

Verbenaceae. 

Clerodendron mimicum Standl. & Steyerm., 
Nat. Hist., Ser. 23(5): 227. 1947. 
mark 48631; 14 July 1942; Guatemala.. 
huetenango, hin Huitz, Sierra de los Cuc E. 

O m; F (holo) = Gibsoniothamnus mimicus 
(Standl. 8 Steyerm.) L. Williams, 1974 in the Scroph- 
ulariaceae not Verbenaceae. 

Clerodendron pithecobium Standl. & Steyerm., Field 
Mus. Nat. Hist., Bot. Ser. 22(5): 373. 1940. P. C. 
Standley 67887; 8 Mar. 1939; Dustemála, Dept. Que- 
zaltenango, La Shuya, SW of San Martín C hile Verde; 
1,620 m; F (holo). 

Lantana ruiz-teranii Lopez-Palacios & Steyerm., Ernstia 
36: 10. 1986. L. E. Ruiz d &S López Palacios 
10907; Venezuela, Bolivar, Distr. Roscio, Gran Sa- 
bana, Santa Elena de Uairén: 900 m; MER (holo); 
VEN. 


Field Mus. 


Ta ea 


VIOLACEAE 


Hybanthus sylvicola Standl. & Ste Vas ., Field Mus. Nat 
Hi er. 23(4): 176 Wilson 329; 
1939; Guatemala, Dept. Ms Verapaz, Pila- 
pec, Finca Los i d 2 m; F (holo). 
Orthion Standl. & Stey , Field Mus. 
Ser. 22(4): 249. 1940, a nov. 
Rinorea oraria Steyerm. & Fernandez, Brittonia 30: 43. 
9 Morillo et al. 3355; 17 July 1973; Vene- 
zuela, Distr. Federal, Cordillera Costanera, S of Camuri 
Grande; 700-800 m; COL (holo), VEN. 
Viola triloba Schwein f. albida P bg B reo 54: 
A. Steyermark 67440; 29 Apr. 1949; 
U.S.A., bou Stone Co., along b 13, 0.5 mi. 


ist., Bot. S 


20 Mar 


Nat. Hist., Bot. 


78. G. 


NE of Lampe; F (holo). 
Viola venezuelensis Steyerm., Siga: Bot. 28(2): 403. 
Steyermark 57077; 7 July 1944; Vene- 


ichs “Mérida, Páramo de la Negra; 2,745-3,045 m; 
F (holo 
Viola viarum Pollard f. pilifera Palmer & Steyerm., 
dus Y 116. 1958. . Steyermark dens 
Apr. 1955; U.S.A., YEN Gentry Co. alor 
ird ribuar to ravines on N and E side of Grand 
River, 1.5-2 mi. S of McFall; F (holo). 


VISCACEAE 


Prode ndron aguilarii Standl. 7 PeP ., Field Mus 
. Hist., Bot. EM er. 23(2): 40. 4. J. A. min 
31889; 18 Nov. 1939; econ Dept. Jutiapa, Vol- 


Volume 76, Number 3 Taylor 735 
1989 Plants Described by 
Julian A. Steyermark 


cán Suchitán, NW of Asunción Mita; 1,000- 2,050 m; Marcano Berti var. angustior (Steyerm.) Marcano Ber- 
holo ti, 1969. 


, 


Pharadendran aphyllum oi ium Bol. Soc. Venez. Ci Vochysia pauciflora Pu des Bol. Soc. Venez. Ci. Nat. 
lat. 2 : Steyermark & S. Nilsson 26: 424. 1966. J. A . Steyermark et al 92902; 9 Jan. 
473; 20-22 Apr. 1960: ene Bolivar, Corto Uei 1954; Venezuela, Bolívar, Cerro Veraimo: 900 m; VEN 
1,100 m; VEN (holo). (holo). 
Phoradendron huehuetecum Standl. & Steyerm., n 
Mus. Nat. Hist., Bot. Ser. 23(2): 41. 1944. WINTERACEAE 
Standley 82597; Jan. 1941; Guatemala, Dept. Hue- 
Hueteñanko, near Tachique, E of Bodl celeb 1.900 Drimys tebe ies Miers var. ptaritepute nsis Steyerm 
Ft a n a Bot. 28(1): 235. Steyermark 
ni eu S 37; E Oct. 1944; Venezuela, Bolivar, Ptari- -tepui; 
Prades insigne Steyerm., Bol. Soc. Venez. Ci. > ud 285 m: F (holo); 
Nat. 26: 415. 19 J. 4. Md & L. Ariste- 2 22 
guieta 59; 9 Mar. 1962; Venezuela, Bolivar, Rio Uara- u oe 
ma valley below Uarama-tepui; 1,200 m; VEN (holo) een 
= P. semivenosum Rizz., 1982. us gleasoniana Steyerm., Fieldiana Bot. 28(4): 
Phoradendron karuaianum Steyerm., aie Bot. . 1957, nom. nov. 
28(1): 222. 1951. J. A. Steyermark 606 27 Nov. Fors macrostachya Spruce var. robustior Stey- 
1944; Venezuela, Bolívar, Ptari- -tepui; M F (holo). erm., Fieldiana Bot. 28(1): 104. 1951. J. 4. Steyer- 
Phoradendron ptarianum Steyerm., ER Bot. 28(1): mark 59349; 26 Oct. 1944; Venezuela, Bolivar, NW 
222. 1951. J. A. Steyermark 59483; 29 Oct. 1944; 


- 


o Rio Karuai, Gran Sahana: 1,220 m; F (holo 
Venezuela, Bolívar, Ptari- -tepul; 1, B m; F (holo). 


). 
rw ptaritepuiana Steyerm., D md Bot. 28(1): 
Phoradendron SO Steyerm. & Maguire, Mem. 1951. J. A. Steyermark 59760; 2 Nov. 1944; 
New Bot. Gard. 17(1): 442. 1967. J. A. Stey- Venda, Bolivar, Ptari- ul 2 000 2 a m; F (holo) 
ermark 75426; Venezuela, Bolivar, Chimanta-tepui = 


— 


(Toronto-tepui), "Chimantá om VEN (hol 


Maguire, 1958. 
0). 
Phoradendron tepuianum Steyerm., Fieldiana Bio. 28(1): 


anthe dabis ue d 
rry, Ann. Missouri Bot. Gard. 


trnitisopea S Ste 


297. 1981, Aen no 
sias 1951. J. 4. Steyermark 39688; 1 Nov. 1944; ws albescens Steyerm. ` Fieldiana Bot. 28(1): 105. 
zuela, Bolívar, Ptari- tepui; 1, P m; F (holo). 1951. J. A. Steyermark S 9734; 2 Nov. ; Ven- 
Plaani treleaseanum Stand ds Steyerm., Field ezuela, Bolivar, Ptari- i UA m; F (holo) 
Mus. Nat. Hist., Bot. Ser. ve 1944. W. A. Xyris capillaris Steyerm., Fieldiana d 28(1): 
Kellerman 7630; Jan. 1908; Cuna Dept. Baja 1951. J. 4. Steyermark 5 Min 22 Aug el ^ud A en- 
Verapaz, Sierra à las Minas, opposite El Rancho (El 


c 
N 
e 
HGB 
HM: 
N 
© 
=| 
D 
k 
E 
& 
J 
tr 
Uu 
= 
Z 
i 
m 
£5 
m» 
E 
w 
in E 
& 
^ 
£e 
= 
= 
z] 
p 


Progreso); 700 m; F (holo). E 
Phoradendron uie anum Steyerm., Fieldiana Bot. 28(1): Xyris culmenicola Steyerm., eres Bot. 28(1): 108. 

223. 1951, nom. nov. = P. undulatum (Pohl) Eichl., 1951. J. A. Steyermark 58233; 2 Sep. 

ezuela, T.F. Amazonas, Cerro Duida, Brokehinis Hills; 

1,700-1,980 m; F (holo). 


1982. 


VITACEAE Xyris esmeraldae Steyerm., Fieldiana Bot. 28(1): 109. 

SS 1951. J. A. Steyermark 57821; 22 Aug. 1944; Ven- 

Cissus venezuelensis Steyerm., Bol. Soc. Venez. Ci. Nat ezuela, T. F. Amazonas, between Esmeralda Savanna 

J. A. Ste yermark et al. 92987; 11 and SE base of Cerr da; 200 m; F (ho 

Feb. 1964; Venezuéla, Bolivar, km 119 and 123 S of Yyris jauana L. B. Smith & Steyerm., Bol. Venez. 

El Dorado; 1,030-1,100 m; depository not designated. Ci. Nat. 32: 277. 1976. J. A. Steyermark et al. 109390; 

22-28 Feb. 1974; Venerei, Bolívar, Meseta del Jaua, 

"TD Cerro Jaua; 1,750-1,800 m; depository not designated. 

IO Xyris lanulobractea Steyerm., ' Fieldiana Bot. 28 28(1): 109. 

Euphronia acuminatissima Steyerm., Ann. Missouri Bot. 51. J. A. Stey ermark 59336; 26 Oct. 1944; Ven- 
sard. 74: 91. 1987. J. J. Wurdack & L. S. Adderley ezuela, Bolívar, Gran Sabana; 1,220 m; F (holo). 

43762; 5 Aug. 1959; Venezuela, T.F. Amazonas, Rio Xy e ale d Fieldiana Bot. 28(1): 110. 1951. 

Orinoco, Sabana Cumare on right bank of Cano Cu 


mark 59919; 7 Nov. 1944; Venezuela, 
Bolivar. Prari. -tepul; carts 2,450 m; F (holo) 
Xyris scabridula Steyerm., Fieldiana Bot. 28(1): 111. 
ue S arpa Steyerm., Bol. Soc. Venez. Ci. Nat. . J. A. Steyermark 58168; 1 Sep í 
966. A. 


mare, Río Atabapo; 125-150 m; MO (holo — 2427924), 
NY. 


p. 1944; Ven 
Steyermark 90467; 30 Dec. ezuela, T. F. Amazonas, Cerro Duida, Brocchinia Hills; 
m n ud Boliv ar, Rio Pn debajo del Salto 1,700-1,980 m; F (holo). 
ría Espuma; 500 m; VEN (holo) — Ruizterania Xyris schneeana Smith & Steyerm., Acta Bot. Venez. 
jp arpa (Steyerm.) Marcano Berti, 1969. 2 


C 2: 149. 1967. L. Schnee 1577; 19 Apr. 1956; Ven- 
marg ferruginea Steyerm., Fieldiana Bot. 28(2): 295. ezuela, Bolivar, Auyan-tepui; MY (holo) 
952. J. A. Steyermark 60914; 11 Dec. 1944; Ven- 


Xyris setigera Oliver var. elongata Steyerm., Fieldiana 
ezuela, Bolivar, Quebrada de Kavanayen; 1,200- 1,220 f 


see Bot. 28(1): 112. 1951. J. 4. Steyermark 59660; 1 
m; F (h o) = Ruizterania ferruginea (Steyerm.) Mar- Nov. 1944; Venezuela, Bolivar, Ptari-tepui; 1,600 m; 
cano Ber 969. 


F (holo). 
Yyris stenostachya Steyerm., » Fieldiana Bot. 28(1): 113. 
ies J. A. Die rmark 58437; 8 S ep. 1944; 
uela, T.F. Amazonas, below mouth of Río ere 
ue Úsinaca poem 100 m; F (holo). 


Qualea urbe Stafleu var. m stior Steyerm., Acta 
Bot. Venez. 2 967. J. . ine ano d 
18 May A Venezuela, Bolivar, Auyan-tepui; 1,00 
m; VEN (holo) = Ruizterania perci SPUR 


736 


Annals 
Msn Bond Garden 


ZINGIBERACEAE 


Costus zamoranus Steyerm., Phytologia s 339. 1964. 
J. eyermark 54664; 14 Oct. 1943; Ecuador, 
Santi tiago-Zamora, along Rio Vali nee Tambo 
Valladolid; 2,000 m; F (holo), NY, 

Renealmia puberula Stey sr pe 9(6): 

964. J. 4 Steyermark < ; 6 Nov 
uador, Prov. Napo- Bona pon Rio Pastaza s 
Finca Black; 1,212 m; F (holo), MO, NY. 

Renealmia stellulata Steyerm., Phytologia 2 

1964. J. 4. Steyermark 53769; 11 Au 943; Ec- 
dador, Prov. El Oro, along tributary to Rio Palma, near 
Pampa do los Cedros S of Cerro Chivo-turco; 2,135 


2,285 m; F (holo), NY. 


340. 


BIBLIOGRAPHY 


ANONYMOUS. 1945. Botanical adventures in Venezuela. 
ull. vta Nat. Hist. Mus. 16(7-8): 7 
947. Important Ecuador ed received 
in erum, Bial, red Nat. Hist. Mus. 
— useum botanist to explore ud s 
‘Lost Cane Bull. Chicago Nat. Hist. Mus. 24(3): 3 
———. 1954. Plant from Guatemala makes U.S. de 
liit: pet Chicag o Nat. Hist. Mus. 25(4): 7. 
———. Bugs to explore d ^u orld’ again. 
Bull. c hai Nat. Hist. Mus. 26( 
——. 19 A boost from Missouri i bot tany's 
Niscounendk 1(2): 3. [Excerpt from a letter written 


Suggestions for beu a survey: 
ideas of Dr. Steyermark. Missouriensis 3(2): 10-12. 
ATWOOD, E L., JR. & J. A. STEYERMARK. 1937. Second 


record for rare freshwater jellyfish for Missouri. Amer. 
Naturalist 71: 280. 

BabiLLo, V. M. & J. A. STEYERMARK. 1973. Una nueva 
especie de Gunnera. Acta Bot. Venez. 8: 311-3 

Berry, P. E. & J. STEYERMARK. 1985. Florula de 
los bosques deciduos de Caracas. Mem. Soc. Ci. Nat 
La Salle 43(120): 1 : 

Brewer-Carias, C. 1965. The Lost World and the 
tropical forest in bir e Caracas. [Steyermark 
authored two cha te 

¿DA 


1976. dem eg pi 


snuff on of the Yanomamo Caburiew-Teri in the 
Cauabu i River, razil. Econ. 30: 57- 
BUNTING, G S . & J. A. STEYERMARK. 196 A new 


American species of Schismatoglottis (Araceae). 
Brittonia 21: 187-19 0. 

CARNEVALI, G. & J. A. STEYERMARK. 1984. Additions 
in the areon fiora of Cerro Marahuaca. Phoil 

55(5): 289- 

Dix G. id "Me ember highlights: MU re- 
turns to his native Missouri. Missouriensis 5(1): 
39 


= 


Drew, W. B. Book review — Spring Ps of 
Missouri by Jolian A. Steyermark. Rhodora 42: 215- 
21 


GENTRY, A. H. & J. A. STEYERMARK. 1987. A revision 
of Dilodendron (Sapindaceae). Ann. Missouri Bot. 
Gard. 74: 533-538. 

GONDELLES A., R., J. R. Garcia A. & J. A. STEYERMARK. 
1977. Los parques nacionales de Venezuela. Insti- 
tuto de la Caza Fotográfica y Ciencias de la Natu- 


raleza, Centro Iberoamericano de Cooperación, Ma- 
drid, Spain. 

HENDERSON, A. & J. A. STEYERMARK. 1986. New palms 
from pera dd 38: 309-: Me 

Huser, O., J. A RMARK, G. T. PRANCE & C. ALÈS. 
1984. TM Acn of the Sierra Parima, Venezuela 
Brazil: some results of recent exploration. Brittonia 
36: 104-139. 

Jackson, R. E. & J. A. STEYERMARK. 1978. Lentinus 
velutinus en Venezuela. Acta Bot. Venez. 13: 117 
124, 

KALLUNKI, J. A. & J. A. STEYERMARK. 1987. A new 
ni of Raveniopsis (Rutaceae), a genus new to 

azil. Brittonia 39: 409-411. 
Kobuskt, C. Studies in the Theaceae, XVII. 
review of the genus Bonnetia. J. Arnold Arbor. 
29: ES 

LASSER, N & J. A. STEYERMARK. 1974. 
Catalogo de las vee s que crecen en el Jardin Bo- 
tanico del Ministerio de Agricultura y Cria. Acta Bot. 
Venez. € m 

GUIRE. 1950. A report on the PA 
of the Phelps Cerro Yavi Expedition of 1947. Brit 
tonia 7: 75-90. 

LOPEZ-PALACIOs, S. Una =, ee de Lan- 
tana de Venezuela. rasta 36: 

. A. STEYERMARK. E m nueva es- 
cie de. Lantana de Venezuela. Acta Bot. Venez 
l 


MacuinE, B. 1957. Domi of the Chimantá Massif, I. 
Gran P Venezuela. Mem. New York Bot. Gard. 
9(3): 393-440. 

——— qo 179 eus to the Rapataceae. Acta 
Amazonica iu 269. 

——— & . Wurpack. 1957. Botany of the Phelps 
Guayana Expeditions II. Uaipan-tepui, Estado Bo- 
livar. Mem w York Bot. Gard. 9(3): 477-484. 

COLLABORATORS. 1948. Plant explorations 
in Pisis in 1944, chiefly to the Tafelberg and the 
on Plateau, VI. Bull. Torrey Bot. Club 75(6): 
633 67 
& COLLABORATORS. 1957-1989. The xe 
of the Su ee Highlands: parts 2-13. Mer 
e Bot. Gard. 
Tes 8(2): 87-159. (no Steyermark taxa) 


^ 1957. 9(3): 235-392. (no Steyermark taxa) 
3. 1958. (1): 1-156. (no Steyermark taxa) 
4(1) 1960. 10(2): 1-37. (no Steyermark taxa) 
4(2). 1961. 10(4): 1-87. (Steyermark: 17, 19) 
5. 1964. 10(5) 1-278. (Steyermark: 186- 
6 1965. 12(3): 1-285. (Steyermark: 178- 
285) 
7 1966. 17(1): 1-439. (Steyermark: 229- 
436) 
8 1967. 18(2): 1 E (no Steyermark taxa) 
9 1972. 23: 1-832. (Steyermark: 139-203) 
10 1978. 29: 1-288. (Steyermark: 139-203) 
. 32:1 


391. (Steyermark: 4-20, 306 


8 
12. 1984. 38: 1-84. (Steyermark: 46-82) 
L3. 1989. 50: in press. 
MOHLENBROCK, R. H. 


1979. Thren months with Dr. 


Volume 76, Number 3 
1989 


Taylor 737 
Plants Described by 


Julian A. Steyermark 


, J. A. STEYERMARK & M. SIVADASAN. 
M pip cie (Marantaceae), a 
om Venezuela and Brazil. 


NicOLSON, D. 
: 


w genus er aS 
Beenie 33: 22=2 
OLiva-EsTEVa, F. s J STEYERMARK. 1987. 
meliaceas n Vanala, native and cultivated. E. Ar- 


Bro- 


Fern and 


105- 


J. A. STEYERMARK. 1932. 
fern allies of Missouri. Amer. Fern J. 22(4): 
122 


————,. 19: An annotated catalogue of 
une flowering plants ul Missouri. Ann. Missouri Bot. 
ard. 375-15 
1938. Additions, corrections, and 
revisions to the “Annotated catalogue of the flowering 
plants of Missouri.’ Ann. Meus Bot. Gard. 25: 
94. 


775-7 

—————, 1938. New varieties and forms 
from Missouri. Aon: Missouri Bot. Gard. 25: 769- 
774. 


Notes on Geoc d mini- 


——— & ———. 1950. 
mum Mackenzie. Bull. Torrey Bot. Gard. 268- 
273. 


& ——. 1952. New al te records 
from MOREL iier: Fern J. 42(2): 6 5. 
1955. Plants new to Missouri. 


310- 319. 

o 1958. Plants new to Missouri. 
Brittonia 10: 109-120. 

REESE, G. A. 1986. Conversations with Julian Steyer- 

mark: the flora of Missouri years. Missouriensis 7(1): 


Rhodora 57: 
& 


R. E. y aa Austro-Americanae, 
pr ue ee 15(2): 29-7 
— 55. Plantae Colombianae, XIL Bot. Mus. 
Lea a 65-100. 
zu p B. 1967. Dos especies Bromeliaceae bei 
a la Ciencia. Acta Bot. Venez. 2(5-8): 380-382. 
96 ia on Bromeliaceae, XXVII. Phy- 


SCHULTES, 
VII 


teles 16: 62 
new species of Connellia from Ven- 
ezuela. J. Bromel. Soc. 35(2): 52-53 
1986. Revision of the Guayana Highland Bro- 
melts, Ann. Missouri Bot. Gar 
STANDLEY, P. C. & J. STEYERMARK. 1940. Studies 
of d —€— plants, I-II. Field Mus. Nat. 
Hist., Bot : d 


es) 
Oo 


-721. 


——— p: — 43. Studies of Camal American 
ien HI. Field e Nat. Hist., Bot. Ser. 23(1): 


ie —— ———,. 1944. Studies of Central American 
plants, IV- VI. "Field Mus. Nat. Hist., Bot. Ser. 23 
(2/4) 29-192. 

& 1945. The vegetation of Guate- 
mala, a brief review. /n: Plants and Plant Sciences 
in Latin America. Chronica Botanica, Waltham, Mas- 
sachusetts. 


47. Studies of Central American 


& —— —. 4 
eh m a d Nat. Hist., 


Bot. Ser. 23(5) 
: e Flora of Guatemala. Fieldi- 
ana Bot. 24(4): 1 
& ; ue Flora of Guatemala. Fieldi- 
ana Bot. 24(5): 1-502. 
a Flora of Guatemala. Fieldi- 


24(6): 1- 


ana Bor. 


————— & —————. 1952. Flora of Guatemala. Fieldi- 
ana Bot. 24(3): 1-432 
& . 1957. A new Lisianthus from 


Flora of Guatemala. Fieldi- 


Guatemala. Bull. Torrey Bot. Club 84(1): 46 
& 195 


ana Bot. 24(1): 1-478. 
& O. WinLiams. 1952. 
Americanae, IV. Ceiba 3: 100- 

STEYERMARK, J. A. 1931. A study of m distribution 
in relation to the acidity ie various soils in Missouri. 
Ann. rca Bot. Gard. 1-55. 

31 


S PAS Centrali- 


on Mi ssouri plants found on 
certain prae formations. Bull. Missouri Bot. Gard. 
-56. 


19 Some new and interesting a from 
Missam. Bull. Missouri Bot. Gard. 19: 56-57 
1932. A revisio n of the genus Manda 
re Missouri Bot. Gard. 19: 87-1 
2. Some new w spermatophytes from Texas. 
hives Missouri Bot. Gar 
933. Notes on Missouri plants. Rhodi 35: 


D» »^ 


283- "f 
1933. A monograph of the North American 
species of the genus. ee lia. ee Dissertation, 


Washington Univ., , Missour 
Studies in Grinde elia, "n New ecies 
nietos qa oa of Grindelia. Ped Mis- 
. 21: 227-230. 


souri Bot. 
1934. $ Studies in Grinde lia, II. A monograph 

genus Grin- 

33-608. 


Recent additions to the flora of St. 


Loue County, Missouri. Rhodora 36: 
934. ri virginiana in Missouri. Rho- 


3(o-21 


ds so 352-3 
——— ——. 1934. A grass new to Missouri. Rhodora 36: 


. Rm virginiana in Md 
Rhodora e 97-10 
934 Le extension in Missouri. for 
TR Mob vulgatum. Rhodora 36: 22-23. 
1937. eco) in Grindelia, Hl. Ann. Missouri 
-262 


Bot. Gard. 24 


1938. ae of the American flora, I. Field 
Mis. irs os Bot. Ser. 17(5): 411-443 

19 combinations and undescribed 
er Piel Mimi. “Rhodora 40: 177-179. 
we tomentose form of Cladrastis lutea. 


dara 40: 
1938. a gracillima in Missouri. Rhodora 
40: e 


—————,. 1938. Pi ue ribed plants from Arkansas. 
a 40: - 


a onia nuttallii in Missouri. 
> 


Rho- 


38. ro = po ae nlite 
& Steyermark' s ed catalogue of t 
ering plants of Me 'Rhodora 40: 131 iis 
1938. Plants new to Missouri. Rhodora 40: 


249- 255. 
1939. Some newly described forms from Mis- 
souri. Rhodora 


41: 
udies at the vegetation of Missouri, 
I. Natural plant associations and succession in the 


738 


Annals of th 
Missouri Ae Garden 


from rr Rho 
— D 


Ozarks o zum Field Mus. Nat. Hist., 
9(5): 35 


Bot. Ser. 


lon Determination of Amphicarpa, 
Strophostyles, racer and Apios by vegetative 

== Rhodor d 213-215. 
Viola cucullata in Missouri. Rhodora 


42: E 
1940. 


Ludwigia microcarpa in Missouri. 
NEL SA 42: 144. 
19 


40. Plants new E or a unreported 
: 97-105 


Mr s a in de urls of south- 
n. Lidl Rhodora 42: 32-33 
19 The first miedo occurrence of Di: 

tichlis spicata in the central interior of the United 
States. Rhodora 42: 22-24. 

41. Studies in the vegetation of Missouri, 
II. Phanerogamic ie of ds water springs in Aa 
Ozarks of Missouri. Field Mus. Nat. Hist., Bot. Ser. 
9(6): 479-6 618. 
1941. Notes on Missouri plants. Rhodora 43: 
6: 58 663. 

Aquatic varieties of Poa annua. Rho- 


——, 194]. 
dove 43: 630-632. 


1941. A study of Arenaria patula. Rhodora 
43: 3 25-333. 
1942. T 


. wo albino Wes of Echinacea from 
en Rhodora 44: -151. 
: ediscovery y du^ ier hellum at 
its nortan limit. Rhodora 

)44. Exploración rod a e regiones 
es de Venezuela. Bol. Soc. Venez. Ci. Nat. 
10(67): 259-276. 
A native Oxalis as a foliage ns for 
Missouri zx dens, Bull. Missouri Bot. Gard. 35(9): 
)1 - 208. 


“Lost World” botanizing in the Gran 


Sabana, Bull oe Nat. Hist. Mus. 17(3-4): 5 


7. Speciation ir E the Venezuelan Guayana 

abstract Amer, J. Bot. 34(suppl.): 6 

A rare oa plant is SN by mu- 

seum eaa Bull. Chicago Nat. Hist. Mus. 18(3):7 
947. ‘What’s ina Lt ?' in the di adit 

Bull. Chicago Nat. Hist. Mus. 18(5): 4- 

1947. A unique pee a. vegetable 

L5. 


ivory. Bull. Chicago Nat. Hist. Mus. 18(7 


varieties. Lay dia 
— | 


a Bl Chicago Nat 


947. Spring and early summer lower rec- 
m 1947. Bull. Chicago Nat. Hist. Mus. 18(8): € 
l 947. Notes on drying plants. Rhodora 49. 


220-227. 
1947. Herman C. Benke. Rhodora 49: 142- 
143 


1948. eto chimboracensis and its 
14-20. 


The cobra a what is it? Bull. Chi- 
cago o Nat Hist. Mus. A 
Stewartia she s America’ of 
the dri Bull Chicago Nat. n Mus. 19(10): 5 
ho llo azaleas — plant aris- 
. Hist. Mus. 19(12): 6-7. 
. „Lentibulariaceae. Bull. Torrey Bot. Club 
í tae a a 
c Bl luets as summer flowers. Bull. Mis- 
souri Bot. Cani. 36: 93-94. 
194 Hens PE roosters of the plant world. 
36: 140. 


8. 
Bull. Missouri Bot. Gard. 


1949. Effects x Auer. Ozark springs. Bull. 

Missouri Bot. Gard. 3 -48. 
The ie A Missouri's forests. Bull. 

Missouri Bot. Gar : 126-127 
949, The persimmon enm Bull. Missouri 
Bot. n 37: 192-195. 
ea Missouri records (1946-1948). 
115-119 
U dere er orchid. Bull. Chicago Nat. 
Hist. we 20(5): 8 
The quest for plants. Bull. Chicago 
Nat. A Mus. 20(8): 6 

19 P lovers in late fall. Bull. Chicago 
Nat. Er Mus. 2 DO, 

940 


os melissaefolia. Rhodora 51: 


m 5l: 


153- o 
Plants new to Illinois or to the Chicago 
area in Illinois. Rhodora 51: 147-149. 

1950. Flora of Guatemala. Ecology 31: 308 
372 


50. Blue mushrooms. Bull. Chicago Nat. 


inox. ON 
Hist. Mus. 21(2): 3, 7. 


1950. Tree damage by ice costs millions. Bull. 
Chicago Nat. Hist. 21(4): 4-5. 
1950. Hawthorns ARA but a headache to 
taxonomists. Bull. Chicago Nat. Hist. Mus. 21(7): 3. 
1950. Evergreen aC hristmas: some legends of 


holiday plants. Bull. Chicago Nat. Hist. Mus. 21(12): 
5-7 


1950. The genus Oreobolus in South Amer- 
ica. Bol. Soc. Venez. Ci. Nat. 11(74): 306-311. 
1950. A new Utricularia from Honduras. 

26 


. 1950. Plant Pour found in Missouri towns. 
Bull. Wei. Bot. Gard. 38: 55-60. 
Wild orc eed of Missouri. Bull. Missouri 
Bot. e 38: 61-64 

1950. The yellow in the ae land- 
scape. Bull. Missouri Bot. Gard. 38: 

1 Scarlet m in Missouri. cal [OM 
Bot. er 38: 143- 

951. When e Pai hazel bloom —it’s 

spring. Bul Chicago Nat. Hist. Mus. 22(4 8. 


Venezuelan ues pith used in native 


e) 
en 


= 


wn 


di Bail ag Nat. Hist. Mus. 22(12): 5. 
—— New 


y Rubiaceae from Panama. Ceiba 
3(1): Ds x 
. 1951. The ipe S ii (Gentiana- 
ceae). verae 14(1): 5 
951. vc i survey of Missouri. Bull. Mis- 
souri Bor Gard. 
951. “The snowy MT Bull. Missouri Bot. 
Card. es 5 
51. p nical apes in is eri Ozarks. 
Bull. ede Bot. Gard. 
5l. labrous vus b Span tere- 
Binden. Rhodora 53: 133- 
1952. The genus um (Rubiaceae). 
Ames, Y Bot, 418-423. 
9! dt pan records (1949- 
1951), pom 54: 250- 

952. List of native ee ri wild flowers and 
FUN that should be protected and should not be 
used in flower shows. [Manuscript. ] 

52. A rare grass Api in ED area. 
23(3): 5 


Bull. Chicago Nat. Hist. Mus. 


Bot. 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


739 


—————. 1952. 
54: Kc 34 
New ETT species of Utricularia. 


952. 
Bull. Tore Bot. Club 79: 310-311. 
Ra 


orci pm Missouri Bot. 


are S nnd plants IL The Ozark 
ard. 40: 


ded EE Missouri Bot 77-80. 
————. 1952. A 


new Ceres wm Guatemala and 


Honduras Caha 3: 23-24. 

952. New Rubiaceae from Panama. Ceiba 

3: 18-22. 

1952. An example of how dams destroy valu- 

able uera records. Sci. Monthly 74(4): 231-233. 
9 are Missouri plants, HI. The Ozark 

‘illum: i Missouri Bot. Gard. 40: 80-82 

Pera P in 1952. Bull. Missouri 

Bot. Card 40: 169- 

————. 195 Breaking a through ‘Lost World’ 

of m Bull. Chicago Nat. Hist. Mus. 24(9): 

3-4, 

1953. The discovery and destruction of Cal- 

fura americana in Missouri. Rhodora 55: 238- 

241. 

1953. Dodecatheon amethystinum and for- 
ma margaritaceum in the Missouri Ozarks. Rhodora 
226—227. 

1953. 

55: 156. 

1953. Color form of Helianthus mollis. Rho- 
108. 


Elymus riparius in Illinois. Rhodora 


dora po 

953. Another € m relict in the Mis- 
souri ien region. Rhodora 55: 15-17. 

1953. A second speci of Schismocarpus. 

Bull. Torrey Bot. Club 8 

95 otes on some roses in the Gray’s 
D range. Rhodora 56: 70- 19. 
Una nueva especie de Sic MC > 


mad E Soc. Venez. Ci. "Nat. 15(81): 3 


nderwater hid. Aquarium 20 

26-2 T Lenripied from Bull. Chicago Nat. Hist. Mus.] 

"n 4. Una nueva especie de Ilex en Vene- 

T3 Bol. Soc. Venez. Ci. Nat. 15(82): 176-180. 

. 1955. Attack on Chimantá. Nat. Hist. 64(9): 
482-4 


: r past, presens and 
future. Bull. Missouri Bot. Gard. 43(1 =13, 
1955 i nais helleborine in Illinois. Rho- 


durs 51:13 


The effects of the three year wo 
on Missouri plant life. Bull. Missouri Bot. Ga rd 
54- 


1955. Wildflower contrasts between northern 
- southern Missouri. Bull. Missouri Bot. Gard. 43: 
——. 1955. ad of spring in Missouri. Bull. 

Misses. Bot. Gard. 43: 95-96. 
——— ——. 1955. What EA Bull. Missouri Bot. Gard. 
43: ee 

55. Discoveries in 1955. Bull. Missouri 
A 43: 150-151. 

955. New Pe from El Salvador and Mex- 

ico. “Ceiba 4: 300-303. 


Color forms of the may-apple. Rhodora 


are valen plans E bap) fringed 
ard. 40: 39-41 


Soc. Venez. Ci. Nat. 
————. 196 


—. 1956. Book review — Vascular plants of Il- 
linois, by ha am & G. D. Fuller. Amer. Midl. 
Nat 


p ssouri's crop m "a annuals and 
p Missouri Bot. Gar : 41- 
. Review piod ra of W uds County, 
Rhodora 58: 213-214. 
— : is e E hazel. Bull. Missouri Bot. 
Gud en S d 


t 
95 


Z. 


as 
19 


en Missouri plants, IV. Whorle 
Poonia « or M five-leaved orchid. Bull. stone 
Car i 
re Missouri ri plants, V. Umbrella plant 
A riogonum i ails ). Bull. Missouri Bot. Gard. 
44 


Pe dade d flowers in Missouri. Bull. 
wen Bot. Gard. 44: 116-118. 
956 


Rice, Mcd ho! Bull. Missouri Bot. 


Card: m 136-137 
57. White- Dabei da of Liatris and 


ed Rhodora 59: 
1958. Bs Hh E olo x Wauconda 
bans pu Gard. Club Amer. 46(1): 57-58. 
. An albino form of psa us Rum estris. 
Rhodora 60: 174-175. 
958. An unusual botanical area in Missouri. 
PN 60: 206-208. 
19 


Floerkea ai a in Mis- 
souri. Brittonia 10: 150-15 
958 


X Carex deamii in Missouri. Rhodora 


60: Sd 
: 1958. The Machris Brazilian Expedition. Bot- 

any: Phanerogamae, Euphorbiaceae, Lentibulari- 
aceae, and B biaceae. Los Angeles Co. Mus. Contr. 
Sci. = 1-23. 

958. p side a Missouri ia ^ press swamp. 
Bull. Missouri Bot. Gard. 46(1): : 

958. Another station B e qi mint- 
mum. E Torey Bot. Club 85: 124-127. 

195 Ophioglossum vulgatum in Mni 
Aiar en J. 48(2): 7 
959. New M of lo recurvatum. 
S 61: 124. 

cp uei AE of the Ozark for- 
Missouri Stud. 31: 1-138 

e taxonomic status e Saxifraga 
palmeri. Brittonia 11: 71-77. 

1960. Two new mes 's of S ae is (Rubi- 
ac "m from South America. Bol. 
Nat. 21(96): 240-242. 

An unusual hybrid Helenium. Rhodora 


est. Univ. 
1959 


Venez. Ci. 


62: td 


0. 
Me flora. Rhodor 


B ur CEN of the Guayana. 
-41. 


New popuii and forms in the 
131. 


Bull. Brome Soc. 11(3): 
— Bo 


tanical Bn and botanical insti- 

tution os e Venezuela. Bull. Missouri Bot. 
Gard. 49(1): 1-4. 

1961. Mere the Mire: paradise. Bol. 
22(100): 291-297 
Informe sobre los aioe del oriente 
de la Guayana Venezolana, Bosques. Reconocimiento 
Agropecuario-Forestal del Oriente de la Guayana Ve- 
nezolana. Consejo Bienestar Rural y Ministerio de 
Agricultura y Cria 5: 139-184. 


Annals of the 


740 
Missouri Botanical Garden 
———. 1961. A neglected Camassia. Brittonia 13: 1967. Flora of Ayuan-tepui. Acta Bot. Venez. 
206-211. 2:1 se 
1962. Bootheel forests. Missouri Log 15: 23-  ———— 67. Rubiaceae. /n: B. Maguire & collab- 
28. orators, n botany of the kou. Sir ca VII. 


Bol. Soc. Venez. Ci. Nat. 
—.. 19 


1962. Pittierothamnus, a new genus of Ru- 
ico, Bol. Soc. Venez. Ci. 23(101): 92 Ed 
963. Standley and the rra ae. Pp. 90- 
94 1 in L. O, Williams (editor), Homage to B. 
Papers in "Honor of Paul C. Standley. Chicago Nat. 
Hist. , Chicago, Illinois. 
sl 36: n New species of Rubiaceae from Parque 
Nacional * e Pittier.” Bol. Soc. Venez. Ci. Nat. 
25(106 JH 76 
Ba. New species from = xenezuelan 
Guayana Bol. Soc. Venez. Ci. Nat. 25(106): 
86 


1963.  Dipic = ndron, new genus of Rubi- 
aceae. we Soc. Venez. Ci. Nat. 25(106): 23-28. 
963. Flora of Misssuri. lowa State Univ. 
Press; ros Iowa. 
1963. Rubiaceae. /n: B. Maguire, J. Wurdack 
& collaborators, The botany of p "eror pd 
land, part M m. New York Bot. Gard. 10: 
209, 250-259. 
^a A new Tresanthera noe Venezuela. 
25(107): 248-249 
A new Pe OR from ia EE Bol. 
Mite Ci. Nat. 25(107 50- 
. 1964. Spring Flora of Mun a Broth- 
ers, Columbia, Missouri. 
19 New species of Rubiaceae from Peru 


collected by John o ck. Bol. Soc. Venez. Ci. 


25(107): 231-2 
1964. Nov eis e Ae er Rubiaceae e C d 
a de Cuatrecases. a Bot. Venez. 4(1): 


113 
Status of the genus Fuosmia Humb. 


& Bonpl. Bol. Soc. Venez. Ci. Nat. 25(107): 217- 
224. 


New species of cow tree Lo mella) 
from Distrito Federal, Ver Bol. Soc. Venez. 
Ci. Nat. 25(107): 240-2 

5. Una nueva Ln de Rytidostylis 
para onem Bol. Soc. Venez. Ci. Nat. 26(109): 
148- 
io Venezuelan bry T — by J. 
A. Steyermark. Acta Bot. Venez ) 73-84. 
l Genero Hippotis. Acta E Venez. 


t 
O 


n 85-102. 
966. C d :iónes a la flora de Venezuela, 

parte 5. Acta Bot. 1(3-4): 1-256 
1966. ees di ‘novel in ibe. region of 
Starea de Lima, Estado Bolivar, Venezuela, II. Bol. 
Soc. Venez. Ci. Nat. 26(110 ): 411-452. 
. 1966. Nuestra vegetación. El Farol. Ano 
27(216): 6-15. 
1966. The genus Bathysa in Venezuela. Bol. 

83 


ae. Venez. Ci. Nat. 26(110): 480-4 
196 


The genus Borojoa in Venezuela. Bol. 
fos. Venez. Ci. Nat. 26(110): 474-479. 

1966. Botanical novelties from upper Rio 
Paragun, Estado Bolivar, Venezuela, III. Bol. Soc. 
Venez. Ci. Nat. 26(110): 472-473. 

1966. Novedades de la Cordillera NETS e 
y los Andes. Bol. Soc. Venez. Ci. Nat. 26(110): 45 


Mem. New York Bot. Gard. 17 


Contribuciones a a Ta a " Sierra 
de [mataca, Altiplanicie de Nuria y region adyacente 
del Territorio Federal Delta Amacuro al Sur del Río 
Orinoco. Acta Bot. Venez. 3: 49-175 

196 Novedades de la Cordillera Costanera, 
IL PRA Bot. Venez. 3: 197-212. 
1968. El genero Lennoa. Acta Bot. Venez. 
38. 


1968. Otra especie de Borojoa en Venezuela. 
ota Bot. Venez. 13: 193-195. 
19 Nuevas especies de la Sierra de Imataca 
y Altiplnicie d de Nuria del Estado Bolivar y E Terri- 
toria EA curo. Acta Bot. Venez. 3: 177-192. 
6 pos on the use of formaldehyde for 


7 


Ai reparation of herbarium specimens. Taxon 17: 


A new species of Be sleria (Gesneri- 

aceae) finm Venezuela. Brittonia 22: 303 

1971. Notes on the genus Sarcorhac his Trel. 
(Piperaceae). Pittieria 3: 29-38. 

1971. Rubiaceas nuevas de los Andes y Cor- 
diera e E Costa de Venezuela. Acta Bot. Venez. 

99 

s e Una nueva a de la sección 

Foliosa e. Acta Bot. Venez. 6: 89-92. 


Lennoaceae. Fl. Venez. 3(1) 187- 


— 


194. 
1971. Tres especies nuevas de Rubiaceae para 


55 la ciencia. Bol. Soc. Venez. Ci. Nat. 29(119-120): 
26. 


Novedades Vid cnp en los generos 

Peperomia y Cestrum. Acta . Venez. 6: 81-88. 
197 na nueva e e de Trigonia de la 

Cordillera Coutanera Venezolana. Acta Bot. Venez. 
-80 


1971. 


1971. Ponencia sobre ge qd de areas 
naturales de Venezuela. il t Primer Con- 
greso etagi de Botánica, pd 

972. Descripción y deralada de doi especies 
raras de Venezuela, Pittie : 27-32 
A e ¡edades "ie E Andes Vssesilunos: 
P "e 9- 

1972. n on Rubiaceae. Bradea 1(16): 
145-150. 
1973. Preservemos las cumbres de la Pe- 
A de Paria. Defensa de la Naturaleza 2(6): 33- 
35 


. 1973. ney tae oe de Venezuela. 
Acta je Venez. 8: 253. 
bes Vo Acta Bot. Ve- 
nez. e an 9. 
974. Rubiaceae. Fl. Venez. 9(1): 1- 2070. 
1974. ción actual de las exploraciones 
TAN en Venezuela. Acta Bot. Venez. L- 
974. Relación floristica entre la Cordillera 
de " Co osta y la zona de Guara ana y Amazonas. Ácta 
Bot. TE 9: 245-252. 
974. Una nueva especie de Kotchubaea 
xoa od. de e pps Venezolana. Acta Bot. 
enez. 9: 287 


Volume 76, Number 3 Taylor 741 
1989 Plants Described by 
Julian A. Steyermark 


974. The summit vegetation of Cerro Au- Diversification in the Humid Tropics. Columbia Univ. 
tana. Biotrópica 6: 7-13. Press, New 
1975. Notes on leaf variation 5 Lantana . 1l New p of Rubiaceae from Ven- 
aristeguietae. Phytologia 30(3): 209-2 ezuela. Pittieria 9: 5- 
1975. New species of "nin E Ven- 198 Erroneous: citations of Venezuelan lo- 
ezuela. m 31(6): 483-485. calities. Taxon 30: 8 l 
== region del Tama debe ser conser- —————. 1981. New species er Rubiaceae from French 
pis. Natura 57. 5-8. Guiana, Brazil and Columbia. Brittonia 33: 385-400. 
975. New taxa in Peruvian Rubiaceae. Phy- ————. 1982. yellow fat Vriesea capituli- 
lois 31(2): 226-228. gera. J. Broil: Soc. 32(3): 1 113. 
——. 1975. Revision a P genus Joosia (Rubi- . 1982. Warning: a T to jd wise is sufh- 
m Brittonia 27: 251-26 cient. Missouriensis 4(1): 7 


975. Anew Paulini from Venezuela. Phy-  ————. 83. Valores un Natura 74: 34-38. 


en a ): 486- 1983. Dr. Leon Croizat-Chaley. Natura 74: 


5. Novedad Venezolanas. Acta Bot. A 
Venez p 235-246. w Pagam em from 
1975. Dun. sobre la flora del Cerro Autana. Babia, fr E Bull 38(2): 3 -3 
Aia ES Venez. 10: 219-234. —— ———. 1984. Piperaceae. Fl. s 22): 1-619. 
Flore de la Sierra de San Luis (Estado ————. i Flora of the cons re I. 
m Venezuela) y sus afinidades fitogeográficas. Ann. C Bot. Gard. 71: 297-34( 
Acta Bot. Venez. 10: 131-218 ————. New ae eee Venezia Ann. 
976. Tropical rainforests of Venezuela need- Missouri p Gard. 71: 1175-1 
ing protection. In: L. S. Hamilton (editor), Tropical |—————. 1984. Taxones nuevos de e Ernstia 
Rainforest Use and Preservation: A Study o G4 23: va 39. 
lems and Pru. in ru Sierra Club Spe . 1984. New Rubiaceae from French Guiana. 
Publ., Intl. Ser. 4: endix III: 1-15. Brittonia 36: 154-160. 
. 1977. A NA bo: of the Rubiaceae from 1984. A new species of ue a ELA 
jr Geaphila croatit. Phytologia 35(6): 401— Poin Bahia, Brazil. Kew Bull. 731 
1986. A new species P. E eperonia Pipers 
. 1977. New combination in Witheringia. Phy- eae) from Venezuela. Brittonia 38: 220-221. 
tologia 37(5): 482-484. —————. 1986. Holstianthus, a new genu bol Rubi- 
1978. A new Dess af Dulacia from Ven- aceae from the uiae Highland. dna Missouri 
ela: ue 38(3) 217-218. Bot. Gard. 73: 495-497. 
Novedades ia del genera ————. 1986. Speciation and endemism in the flora 
Sum. Pittieria 7 To 13-106. se Veneen tepuis. Pp. 317-373 in F. Vuilleumier 
l . New Rubiaceae from Brazilian Planalto. & M. Monasterio (editors), High Altitude Tropical 
Brtiónia ae 34-38. Biogeography. Oxford Univ. Press, New York. 
—. 1978. Notes on px. of Avila and Naiguata 1986. Comments on Family sy nopsis for the 
mountains. Pittieria 7: 24. Missouri flora, 1.' Missouriensis 7(1): 21-22 
1978. The s. palustris complex in 86/1987. Notes on the meee of ns 
Von. — 11-25-30. Aracá. pex Amazonica 16/17: 219-2 
978. w taxa from Cerro Marahuaca, Ven- . 1987. sie rdi new pude d f Rubi- 
ezuela. Brittonia 30: 50-54. aceae from the Guayana Highland. Ann. a. 
. 197 Plant E and dispersal centers in Bot. Gard. 74: 676- 678. 
Venezuela: their relict and endemic elements. Pp. ] 87. New i of Rubiaceae from Venezuela. 
185-221 in K. Larsen & 4 ae Nielson (editors), Ann. Missouri Bot. Gard. 74 8-400. 
m Botany. Academic Press, New York. l sem review of Prat ts of the E oun 
979. A new species of Roupala D ee Flor a by Marc C. c an Roosmaler. Natl. 
fron TARE Phytologia 44: 321-32 Res. E ): 140-14 
1 Flora of the Guayana mm id; e —————. 1987 n nueva Hillia (Rubiaceae) para la 
dou of the generic flora of a oe of i ciencia. Po 


'enezuelan tepuis. Taxon 28: 45 


"7 


Cn in floras of islands and atolls 
of ee Opera Bot. 92: 235-252 
. 19 Flora of the 2 d E IL 
Ann. ME Bot. Gard. 74: 85-116. 
987. Flora of the V Mio. Guayana, Ill. 
inn VR Bot. Gard. 74: 609-65 
1988. Flora of the Venezuelan Guayana, IV. 
bv. 


A new species of prat (Rubiaceae) 
from Venezuela. Ann. Missouri Bot. Gard. 66: 902- 
904. 


80. New species from the summit ad Cerro 
Guaiquinima, Venezuela. Brittonia 32: 17 
1980 


s ). Una nueva T: del genero p iu Missouri Bot. Gard. 75: 311 
melia (Rubiaceae). Mem. Soc. Ci. Nat. La Salle 988. Flora of the Venezuelan Guayana, V. 
40(113): 83-85. ra Missouri Bot. Gard. 1058-1080. 
. new species of Raveniopsis (Ruta- A new species of Peperomia (Pipera- 
ceae) from Venezuela. Brittonia 32: 47-50. deae) on the Venezuelan CS highland. Brit- 
Relationships of some aa for- tonia us 294-295. 


1980. 
est refuges with lowland t boi fores 988. Flora of the Venezuelan M VI. 


s. Pp. 182- 
220 in G. T. Prance (editor), The Model a Biological dun ed Bot. Gard. 75: 1572- 


742 


Annals of the 
Missouri Botanical Garden 


aiu Bot. Gard. 


AGOSTINI. Notas sobre algunas 
especies E de Trichilia. Bol. Soc. Ven. Ci. 
Nat. 26(109): 143-147. 

& 1966. Exploración botánica del 

rro Patao y zonas adyacentes a Puerto Hierro en 

la Peninsula de Paria, Edo. Sucre. Acta Bot. Venez. 
1(2): 7-80 

. ARISTEGUIETA & J. J. Wukrback. 1965. 

pos nueva Mr p dg de d s i enezuela. Bol. 

s 'enez. Ci. Nat. 26(109): 1 

. F. Austin. 1970 ecies of 
Dic ranosty les (Conv p ae uid from V Ec Ann. 

155- 


iia ARIAS. a La vegetación 
de la € cima Ee Macizo de Jaua. Bol. Soc. Venez. Ci 
Nat. m 133): 179-406. 

S BUNTING 1975. Revision of the genus 
Froesia (Quinace ae). Brittonia 27: 172- 

ELASCIO CHITTY. eG ontribuciones 
a la Hors de la Cordillera de Perijá, Estado 
Venezuela. Bol. Soc. Venez. Ci. Nat. 40(143): 153- 
294. 


G. DUNSTERVILLE. 1980. The lowland floral 
element on the summit of Cerro Guaiquinima and 
other cerros of the ss Highland of Venezuela. 

03. 


J. Biogeogr. 7: 285 
—— & 


grasses. Rhodora 63: 24- 
——— & 


Horsr. 1986. Una nueva especie de 
Navia ilivomelisteno) de Venezuela. Ernstia 38: 44- 
46. 
. HUBER. 1978. Flora del Avila. Flora y 
a de las montanas del Avila, de la Silla y 
del Naig uatá. Soc. Venez. Ci. Nat., Caracas. 


. H. KIRKBRIDE, JR. 1977. Review of the 


genus Pe rama (Rubiaceae). iden: 29: 191-198. 
——, 197: 


e genus Wittmackian- 
4-509. 


thus. Ann. Missouri Bot. Gard. 62: 504- 
& ( 


New combinations in 


. LASSER. 1981 ri new species of Eugenia 
(Myrtaceae) from Ve o Brittonia 33: 25-27. 
& ——. 198 yellow-flowered i of 
Erythrina Büeppisiana, Fito 48(4): 2 
LIESNER. 1983. Revision of the genus 
Sterigmapetalum A Ann. Missouri 
Bot. Gard. 70: 179-193. 


D Notes on reed in South 
Pen Ann. Missouri Bot. Gard. 
& . LUTEYN. 1980. Rec of the genus 
Oc hthoc osmus (Linaceae). Brittonia 32: 128-143 
5 di smithiana, a 
new Connellia from the Guayana Highland. J. Bro- 
mel. Soc. 35(4): 152-153. 


B. MacuiRE. 1967. Botany of the "PR 


Vin II. Mem. New York Bot. Gard. 17(1): 4 


d L. B. Marcano Berti. 1966. Una especie 
nueva be Sloanea. Bol. Soc. Venez. Ci. Nat. 26(110): 
67 M 

^ . MEYER. 1944. Informe de la Misión 
m Bol. Soc. Venez. Ci. Nat. 
ae 190. 

. Moore. 1933. Report of a botanical 
expen into the mountains of western Texas. Ann. 
Missouri Bot. Gar 91-8006. 

)RTEGA. 1980. 


2 ris torrestana 
in suela, Amer. Fern J. 70: 80 


—b—— & 
from ala Rbudaia 83: 141- 
———— & 


—— & 
North Pug Rhodora 62: 223-2 
—— & F. 


— & k TRUJILLO. 


leyi. Bull. Bromel. Es 6(1): 6- 
& 958 


New records of tree ferns 
43. 


A. SCHMOLL. Panicum tucker- 
mani, à variety of Panicum philadelphicum. Rho- 
dora 41: 86-90. 

SMITH. 1986. A remarkable new 
ao = Venezuela. Ann. Missouri Bot. Gard. 
9 


& ums 1967. A new variety of Vriesea 
Des ns from Venezuela. Bull. Bromel. Soc. 17(3): 
53 


Lindmania holstii, a new 
species from the Venezuelan Guayana. J. Bromel 
Soc. 51-53 

974. A new Drosera from Ven- 
ezuela. Rhodora 76: 491-493 
& STEVENS. 1988. Notes on Rhodo- 
gnaphalopsis and Bombacopsis — ae Wer in 
the Guayanas. Ann. Missouri Bot. Gard. 
S. STEYERMARK. 1960. e in 
SWINK. 1952. o new Ane Illinois 
and to Pthe pee region. Rhodora 54: 208-213 
1955. Plants new to Tid and 
to thg Chicago region. Rhodora 57: 265-268 
—— Calycera Pune 
in the United States. BaS T: 
& ——— ida drin var. 
kansana i in New York state. n hodora 58: -198 
. Plants e to lia and 
to the Chicago region. Rhodora 61: 24-2 
. W. THIERET. prs a nts new 
to Ilinois and Indiana and the Chicago region. Rho- 
l- 


dora 5 


1964. A new Sic ydium from 
the conata t ange ai Bol. Soc. Venez. Ci. 
Nat. 25(107 k 245 

&. ATTER, de 


1956. _Tillandsia stand- 


Pitcairnia heterophylla. 
Bull. Bromel, Soc. pes 22. 
rd GHT & R. H. MourENBROCK. 1959. 
Prateni biological status of Geoc pt nium 
Mackenzie. Bull. Torrey Bot. Club 86: 23 
, B. MAGUIRE Y COLABORADORES. a Flora 
de la Guayana Venezolana. Acta Bot. Venez. 14(3): 
1-117 


a Guayana Venezolana: 5-52. 
e de la cumbre 


uevos taxa de la 

II. Pn preliminar sobre la flor 

erro Marahuaca: 53-90. 

III. dee JC sobre la flore del Cerro Ru- 
tani: l 

, 5. NILSSON k COLLABORATORS. 1962. Botan- 

TA novelties in the region of Sierra de Lema, Edo. 

Bolivar. Bol. Soc. Venez. Ci. Nat. 23(101): 59-83. 

& COLLABORATORS. 1963. Botanical 

novelties in the region of Sierra de Lema, Estado 


Bolivar, II. Bol. Soc. Venez. Ci. "Nat. 25(106): 42- 
49. 
& COLLABORATORS. 1951. Contributions to the 
a of Venezuela. Botanical exploration in Vene- 


. 28(1): 1-242 

52. Contributions to the 
ad of j sse Botanical exploration in Vene 
piii th hrough Uinbelliteras, 


zue Fro 
Fieldiana Bot. 28(2): 24 


Volume 76, Number 3 
198 


Taylor 


y 743 
Plants Described by 


Julian A. Steyermark 


& COLLABORATORS. 1953. sn eran to the 
flora of Venezuela. Botanical explora in Vene- 
zuela, . From Ericaceae rough poste 
Fieldiana Bot. 28(3): 449-67 

———— & COLLABORATORS. 1957. 

ra ak dec Botanic 


Contributions to the 
in Vene 


OLLABORATORS. 1902. Botanical noveles 

he upper Rio Pm "e Bolivar, Venezuela, I. 
. Soc. Venez. Ci. 23(101): 8 . 

& COLLABORATORS. 1962 : Notes: on the flora 

l. Soc. Venez. 


of Parque Nacional de Guatopo, I. Bol. Soc. 
Ci. Nat. 23(101): 84-88. 

& COLLABORATORS. 1963. Botanical novelties 
from the upper Rio Paragua, Edo. Bolivar, Vene- 
zuela, II. Bol. Soc. Venez. Ci. Nat. 25(106): 50-55. 

—————— & COLLABORATORS. 1964. Notes on Ecuador 
plants. a ogia 9(6): 331-350. 

———— & COLLABORATORS. 1972. Flora of the n 

de Cerro Jaua. Mem. New York Bot. Gard. 23: 8 


8 COLLABORATORS. 1978. New taxa from the 


Avila and a mountains, Venezuela. Brittonia 
30: 39-4 
& COLLABORATORS. 1981. 
Venezuelan Guayana. Brittonia 33: 
& COLLABORATORS. 1989. Flora of the Vene- 
zuelan Guayana, VIII. Contributions to the flora of 
ES Fo Aracamuni, Venezuela. Ann. Missouri Bot. 


New taxa from the 
28-36. 


en 
TILLETT, De S. J. A. STEYERMARK. 1982. Contribu- 
ciones a la flora del Cerro Marahuaca, Territorio 
Federal Amazonas, Venezuela. Ernstia 9: 1-9. 
Webber, W. R. 1981. Flora of Missouri project: com- 
ments by Steyermark and Weber. Missouriensis 23) 
5- 


wo. C. GILLESPIE & J. A. STEYERMARK. 
a Systematic of Croizatia (Euphorbiaceae). 
. Bot. 1-8. 
uu T: x & A. STEYERMARK. 193! 


tological apace of Grinde lia species. Bull. “Torrey 
Bot. Club 62: 69- 

Wurpack, J. J. & J. EN 1972. Notes on 
the genus Pe viia uuu. Bradea 1: 151-152. 


744 Annals of the 
Missouri Botanical Garden 


APPENDIX I 
Eponymy. Plants named in honor of Julian Steyermark. These are all the names found while examining the 
literature cited in the bibliography. It is not a complete list. Additions or corrections would be most welcome 


TRIBES 
Steyermarkochloeae Davidse & Ellis Poaceae 
GENERA 
Aristeyera H. Moore = Asterogyne H. Wendle Arecaceae 
Jasarum Bunt ing Araceae 
Steyerbromelia L B. Smith Bromeliaceae 
Steyermarkia S Standl. ubiaceae 
Steyermarkiella H. Robinson Musci 
Steyermarkina R. M. King & H. Robinson Asteraceae 
Stey ermarkochloa Davidse & Ellis Poaceae 
SPECIES TYPE 
Abuta steyermarkii (Standl.) Standl. Menispermaceae 
Achnopogon steyermarkii Aristeg. Steyermark 93512 Asteraceae 
Aegiphila steyermarkii Mold. Steyermark 59960 Verbenaceae 
Amanoa steyermarkii Jabl Steyermark 93238 Euphorbiaceae 
Anomospermum steyermarkii Krukoff & Barneby Fróes 21466 Menispermaceae 
Anthurium julianii Bunting Steyermark 97325 Araceae 
Anthurium steyermarkii Bunting Steyermark 74850 Araceae 
Aphanocarpus steyermarkii (Standl.) Steyerm. Rubiaceae 
Aristolochia steyermarkii Standl. = Aristolochia Steyermark 33455 Aristolochiaceae 
arborea J. Linden 16 
Arthrostylidium acia McClure Steyermark 59926 Poaceae 
Asketanthera steyermarkii Markgraf Steyermark 99348 Apocynaceae 
Aspidosperma steyermarkii Woodson Steyermark 59858 Apocynaceae 
Axonopus steyermarkii Swallen Steyermark 58001 Poaceae 
Azorella julianii 2 & Const. Steyermark 55894 Apiaceae 
Befaria steyermarkii A. C. Smith Steyermark 62711 Ericaceae 
Begonia ep L. B. Smith & B. G. Schubert Steyermark 75502 Begoniaceae 
Beloperone steyermarkii Leonard Steyermark 56869 Acanthaceae 
Besleria steyermarkiorum Wiehler Steyermark 111182 Gesneriaceae 
Bidens steyermarkii Sherff Steyermark 50998 Asteraceae 
Blepharodon julianii Morillo Steyermark 111304 Asclepiadaceae 
Blepharodon steyermarkii R. Holm Steyermark 60813 Asclepiadaceae 
Bonnetia steyermarkii Kobuski Steyermark 59570 Theaceae 
Bouvardia steyermarkii Standl. Steyermark 29671 Rubi 
Brachionidium julianii Carnevali pi Ramirez Steyermark 129512 Orchidaceae 
Brachionidium steyermarkii Foldat Steyermark 98592 Orchidaceae 
Brocchinia steyermarkii L. B. Smith Steyermark 60347 Bromeliaceae 
Bulbophyllum steyermarkii Foldats Steyermark 100818 Orchidaceae 
Bursera steyermarkii Standl. Steyermark 30068 Burseraceae 
Byrsonima steyermarkii Anderson Steyermark 97979 oo 
Caladium steyermarkii Bunting Steyermark 102035 Ara 
Calceolaria steyermarkii Pennell = Calceolaria fusca Steyermark 54475 ig m 
Pennell, 1988 
Campylocentrum steyermarkii Foldats Steyermark 96715 Orchidaceae 
Capparis steyermarkii Standl. Ste) memari 39387 Capparidaceae 
Carex steyermarkii Stanc eyermark 48542 Cyperacea 
Castil s rkii Pennell Steyermark 57137 Scrophulariaceae 
Cecropia steyermarkii Cuatr. ) rmark 109842 Cecropiaceae 
E in arp us lacey ile Gilly Steyermark 56940 Cyperaceae 
steyermarkii Standl. Steyermark 36237 Rubiaceae 
C diia i stey le Monachino Steyermark 57717 a 
Citharexylum stey ermarkii Mo Steyermark 31433 ne 


Cladium steyermarkii T. Koyama = Rhynchoclad- Steyermark & Wurdack 1225 sii rac 
ium steyermarkii (T. Koyama) T. Koyama, 1972 


Clidemia steyermarkii Wurd. s ermark 96653 b MEE 
Clitoria steyermarkii Fantz ned. abace 

Clusia steyermarkii Maguire cm ermark 75977 Clusiaceae 
Cobaea steyermarkii Standl. Steyermark 37759 Polemoniaceae 
Coccoloba steyermarkii Standl. Steyermark 39533 Polygonaceae 


Columnea steyermarkii Morton Steyermark & Nilsson 41 Gesneriaceae 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


745 


Connarus steyermarkii Prance 
Conomorpha stey UH iie d Cybianthus 
it.) Agost., 1988 
udi 


G S 5 pt 
Cottendorfia steyermarkii s. is Smith) L. B. Smith 


andl. 
Cuchumatanea | stey a Seidensch. & Beaman 
Cyathea steyermarkii Tryon 
Cybianthus steyermarkianus (Agost.) Agost. 
Cyclanthera steyermarkii Standl. 
o 


Daphnopsis steyermarkii Nevling 

Davilla steyermarkii Kubitzki 

Dendrophthora steyermarkii Rizz. = 
squamigera (Benth.) O. Kuntze, 1 

Diacidium steyermarkii (Maguire) em 

Dicella julianii (Macbr.) Anderson 

el steyermarkii Prance 

Dicran steyermarkii Bartr. 

T i a R. H. Maxwell 

Distictis steyermarkii Gentry 

Duranta i Re: dii 

Echeveria steyermarkii Standl. 

Elaphoglossum te Mickel 

Elleanthus steyermarkii Barringer 

Elvasia steyermarkii Ma e 

Epidendrum steyermarkii A. D. Hawkes 

Epiphyllum steyermarkii patel 

Eriocaulon stey 'ermarkii Mold. 

Eriosorus paucifolius var. steyermarkii A. F. Tryon 

Erythroxylum steyermarkii Plowman 

Espeletia steyermarkii Cuatr. = E. brassicoidea var. 
contracta Cuatr., 1964 


Pe drophthora 


Greigia steyermarkii L. 
Guettarda steyermarkii Étandl. 
Guzmania steyermarkii L. B. Smith 
Helianthostylis e AM ada C. C. Berg 


"wa l 97 (Es 


ydrangea e sey Marie Sand. 


Ow 
Ae 2 
ef 


andl.) Standl., 1 
Hypericum steyermarkii Standl. 
Hex julianii Edwin 

llex steyermarkii Edwin 


Steyermark 91995 
Cardona Puig 2723 


Steyermark 31341 


Steyermark 60025 
Steyermark 60002a 
Steyermark 30448 
Beaman 3962 
Steyermark 105194 


Steyermark 34047 


Steyermark & Wurdac 781 


Steyermark 11 
Steyermark E. : A 


Steyermark 89875 
a ÓN 74980b 


oe mark 105274 
Steyermark 62102 
Steyermark 43145 
Stey Mik 128500 
i out 54873 
Pose 96342 
Moritz 

ER 108741 


Steyermark 57217 


Steyermark 36210 


Steyermark 48814 
Steyermark 36426 


Steyermark 5 59942 


i ermark 39454 


Steyermark 34760 
Steyermark 91538 
Steyermark 75501 


Connaraceae 
Myrsinaceae 


Asteraceae 


scl 
Bud 


Loranthaceae 


Malpighiaceae 

Malpighiaceae 

Dichapetalaceae 
usci 

Fabaceae 

Bignoniaceae 

V eae 


Erythroxylaceae 
Asteraceae 


Myrtaceae 


raceae 
Heliconiaceae 
Melastomataceae 
Araceae 


Malpighiaceae 


us 
(A 

pia 
o 


Menispermaceae 


Clusiaceae 
Aquifoliaceae 
Aquifoliaceae 


746 


Annals of the 
Missouri Botanical Garden 


Jasarum steyermarkii Bunting 
Juglans steyermarkii Mannin 
Justicia steyermarkii Standl. & Steyerm. 
Lagenocarpus steyermarkii Gilly 
Leandra steyermarkii Wurd. 
,edothamnus steyermarkii A. C. Smith 
Leothrix steyermarkii Mold. 
Lepanthes steyermarkii Foldats 
Lepanthopsis steyermarkii Foldats 
Lepidopilum steyermarkii m 
Licania steyermarkii Magu 
Lindmania stey erick L . Smith 
Lyc opodium steyermarkii Ollgaard, ined. 
Lysimachia steyermarkii Standl. 
Macrocentrum steyermarkii Wurd. 
Macrolobium steyermarkii Cowan 
alaxis steyermarkii Correll 
Mendes illa Mo Woodson 
Mapania steyerma . Koyama 
Marcgravia stey nont de Roon 
Marsdenia steyermarkii Woodson 
Matelea steyermarkii Woodson 
Maxillaria steyermarkii Foldats 
Meriania steyermarkii Gleason 
Miconia steyermarkii Gleason 


Microlicia steyermarkii Gleason = Microlicia ben- 


thamiana Triana ex Cogn., 1 
Micropholis steyermarkii Monachino 
Monstera ste yermarkii Bunting 
Mouriri ste yermarkii S Stan 


My rcia ES Mc Vaugh; dedicated to Dr. Juli- 
k 


an Stey 
My rioc ladis Torovi Swallen 
lavia steyermarkii L. B. Smi 


Nothoc hlaena steyermarkii Vareschi 
Werff 


Oreopanax steyermarkii A. C. Sikh 

Ormosia steyermarkii Rudd 

Ouratea steyermarkii S Sastre 

Oxythece steyermarkiana Monachino 

Oyedaea steyermarkii Blake 

Pac hyphyllum steyermarkii Foldat 

Paepalanthus perplexans var. B Mold. 

Paepalanthus steyermarkii Mold. 

Pagamea steyermarkii Standl. 
ermarkit (Standl.) Steyerm., 1965, 1 


Peperomia steyermarkii Yuncker 


Perissocarpa steyermarkii (Maguire) Steyerm. & Ma- 


2 
steyermarkii C. K. Allen 

Phainantha stey inb a 

ilacra steyermarkii Magu 
Philodendron steyermarkii Bunting 
Pho ebe steyermarkiana 

anii va er 
Phoradendron steyermarkii Rizz. 
Phthirusa steyermarkiana Rizz. 
Piper na ermarkii Yuncker 
Piptocarpha jedes Aristeg. 


= Aphanocarpus stey- 
967 


K: Allen = Ocotea juli- 


Steyermark 105500 


ark 516 7 


Steyermark 59331 


Stey ermark 60650 
)5 


9446 
d 60879 


Steyermark E LIUM 819 
Steyermark 5 
td 75 


Stey ark & Wanda 921 


Steyermark 115640 
Steyermark 59369 
Steyermark 59744 


Steyermark F n iai 954 
Ste yermark 9 9. 
Steyermark a 


Steyermark 37061 
Steyermark 108984 
Steyermark 93844 
Steyermark 102603 
Steyermark & Wurdack 921 


Steyermark 115564 
Steyermark 112735 
Steyermark 59996 

Steyermark 91450 


Araceae 
Juglandaceae 
aia 
Cypera 
Melastomataceae 
Ericac 
enr 
Orchidaceae 
Orchidaceae 

usci 
Chr di sai 
Bromeliaceae 


Melastomataceae 
Fabaceae 


rchidace 
Apocynaceae 


nan 
Melastomatacea 


pee 


Melastomataceae 


Myrtacea 


Poaceae 


romeliaceae 


Menispermaceae 
e 
Orchidac 


Fup horiaca 


Poaceae 
Bignoniaceae 
Piperaceae 
Ochnaceae 


uraceae 
Melastomataceae 
Ochnaceae 
Araceae 
Lauraceae 


Vis 

Lorant om 
Piperaceae 
Asteraceae 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


Pitcairnia steyermarkii L. B. Smith 
Pleurothallis steyermarkii C. Schwein. 
Podocarpus steyermarkii Buchh. & Gray 
Psittacanthus julianthus Rizz. 

Psychotria steyermarkii Standl. 
Pourouma steyermarkii Standl. & Cuatr. 
Pterozonium steyermarkii Vareschi 
Quararibea steyermarkii Cuatr. 

Rapatea steyermarkii Maguire 

Rauw d steyermarkii Woodson 

Remijia stey ermarkii Standl. 

CH lh d steyermarkii Bunting 
Rhychocladium steyermarkii Ko oni Koyama 
Rhy nchospora stey ermarkii M 
Roupala steyermarkii Sleume 

Russelia steyermarkii € ae 1 
Schistocarpha steyermarkii H. Robinson 
Selaginella steyermarkii Alston 
Selenipedium steyermarkii Foldats 
Sematophyllum steyermarkii Bartr. 
Senecio stey ermarkiana Robinson 
Senecio steyermarkii Greenman 

Senecio steyermarkii Cuatr. = Senecio meridianus 


zuat 

Sideroxylon A Standl. 
Simsia steyermarkii H. Robinson & Brettell 
Sipapoa stey aA Maguire — Diacidium steyer- 

markii (Maguire) pas l 
Sloanea steyermarkii C. E. Smith 
Stegolepis stey mei mum 
Stelis steyermarkii 
Stenopadus podi Ari 
Stenospermation steyermarkii Buming 
Stomatoc haeta ste iter ~ ris 


Taralea stey markii F ‘hery = T. reticulata (Benth.) 
Ducke 
schia steyermarkii Math. & Const. 

Telipogon steyermarkii Foldats 

Ternstroemia steyermarkii Kobuski 

Tetrapterys julianii Macbr. = Dicella julianii 
Macbr.) Anderson, 1981 

Thalictrum steyermarkii S 

Tibouchina steyermarkii Wu 

Tillandsia steyermarkii b. B Smith 

Topobea steyermarkii Wur 

Trichipteris steyermarkii Tryon = Cyathea venezue- 
lensis A. R. Smith, 1981 

Turnera steyermarkii Arbo 


Pu. steyermarki Cuatr. = W. velutina f. 
ii (Cuatr.) Berna rdi 
W einmannia ae f. steyermarkii (Cuatr.) Bernardi 
obium steyermarkii Foldats 


Steyermark 126510 


Steyermark 96161 


Steyermark 87853 


ibi 50. 50 1 


Steyermark 50208 
Steyermark 33858 


Steyermark 42931 
Steyermark 98016 
Steyermark 9357 1 
Steyermark & Aristegueita 17 
Steyermark 9866 1 
Steyermark 93808 
Steyermark 87511 
Steyermark 97917 
Steyermark 89491 
Klug 2010 
Steyermark 60822 
Steyermark 50280 
Steyermark 98568 
Steyermark 61665 
Klug 347 


ey ermark 3057 25 


Ste 
Stey ermark 94923 


ined. 
Steyermark 93837 


Stey ermark 59759 


Steyermark 62165a 


Note: this collection number is incorrectly cited in the literature. 


Xyris steyermarkii Maguire & L. B. Smith 


NAMES NOT VERIFIED 
Dolio steyermarkii 
Ecclinusa steyermarkii Aubr., ined.? 


Steyermark & Wurdack 1096 


IS 


jn aaa 
Aster 

Selaginellac eae 
Ore metas 

Mus 


Asteraceae 
Sapotaceae 
Malpighiaceae 
nsu sigs 


Rapateace 


Orc HARI 
e 


aceae 
Dichapetalaceae 
Fabaceae 


Apia 
Ore hida eae 
Thea 
Maaien 
Ranunculaceae 
Melastomataceae 
omeliaceae 
Melastomataceae 


jvatheaceae 


Turneraceae 
Lentibulariaceae 
Erica 


Cunoniaceae 


Cunoniaceae 
)rchidaceae 


Xyridaceae 


Sapotaceae 


748 


Annals of 


| the 
Missouri Botanical Garden 


Lindsaea steyermarkiana 

Lueheopsis julianii Macbr 
Phragmipedium steyermarkii 
Plagiochila steyermarkii H. Robinson 
Serjania steyermarkii 


APPENDIX II 

Collectors Index. This cross-reference is arranged al- 
phabetically by the collectors’ last names; these are fol- 
lowed by initials ans full name in parentheses. Then appear 
sequential listing of collection numbers with taxa and 
author. Taxa listed in Appendix I are included. Collector 
eponymy is shown with italics. A few duplicate numbers, 
resulting from manuscript names or posibles orania, are 

av 


T 


also italicized. Most co-collectors 


I 
when they collected under vene aba series. 


COLLECTORS NOT GIVEN 


s.n. Orthrosanthus chimboracensis var. in- 
termedius Steyerm. 
s.n. Ravnia panamensis Steyerm. 


ABRAMS, L. R. (LeRoy) 
3957 Grindelia hallii Steyerm. 


ACOSTA SOLIS, M. 


5054 Clethra rugosa Steyerm. 

700 Siparuna rugosa Steyerm 
1009 arkea grandiflora Steyerm 
7743 Cyphomandra villosa Steyerm. 
8233 Siparuna verrucosa Steyerm 


ADAMS, W. (Wayne) 


s.n. Trillium recurvatum Beck f. foliosum 
Steyerm. 
s.n. Trillium recurvatum Beck f. petal- 


oideum Steyer 
AGOSTINI, G. (Getulio) 


diras ere subsp. vene- 


zuelica Steye 
84 i. ae Steyerm. 
1120 Piper agostiniorum Steyerm. 


AGUILAR, J. I. (José Ignacio) 


10 Calyptranthes aguilarii Standl. & Stey- 
erm. 
1757 Eugenia rheedioides Standl. & Steyerm. 
ALLART, A. 
225 piene cole x 'occa subsp. venezuele- 
s Steyer 
ALLEN, P. H. (Paul Hamilton) 
2153 icis e nii SN 
2240 Voyria allenii Steyerm. 
3560 Chimarrhis decurrens Steyerm. 
4576 Coussarea darienensis Steyerm. 
4744 Ravnia longifilamentosa Steyerm. 


ALSTON, A. H. G. (Arthur Hugh Garfit) 
6167 Piper arieianum var. yaracuyense f. nu- 
bicolum Steye 
6533 Galium canescens n alstonii Steyerm. 


Pennstaedtiaceae 
Tiliaceae 
Orchidaceae 

usci 


Sapindaceae 


ANDERSON, W. R. (William) 


ARISTEGUIETA, I 


5264. 


Perama irwiniana Kirkbr. & Steyerm. 
. (Léandro) 
eo An suia Uem 

trionalis var. pubens Steyer 
Sabicea aristeguietas Sievert rm. 
Sickingia aristeguietae m 

eperomia aroensis Steyern 
Peperomia poe Steyerm. 
Psychotria aroensis Steyer 
Flaegia indere E 
Rondeletia ar EE Steyerm. 
rn 


Borreria capitata f. glabra Steyarin. 

sa lane acuminata var. obtusiuscula 
Steye 

Randia ae Steyerm. em 

Borojoa universitatis ye 

Psychotria patria var tovarensis Stey- 
erm 


AYMARD, G. (Gerardo) 


4631 
BADILLO, Y 

312 

4806 


Zinowiewia aymardii Steyerm. 


M. (Victor) 


Manettia badilloi Steyerm 
Gunnera pittieriana Badillo & Steyerm. 


BALDWIN, J. T., J 


4077 
BARNES, V. 
BARRETO 

1134 


BARTEL, K. E. 


S.n. 


l 


BARTLETT, 
8300 


BAUER, B. 
s.n. 

BAUTISTA, J. 
5037 


A. 


Utricularia baldwinii Steyerm. 
Styrax longipedicellatus Steyerm. 


Perama sparsiflora Standl., Steyerm. & 
Ki 


rkbr. 


Liatris cylindracea f. bartelli Steyerm. 
Dipsacus sylvestris Huds. f. albidus 
Steyerm. 


1. (Harley Harris) 
Diodia hy d eal var. linearis f. gla- 
briuscula Steyerr 


Ruellia pedunculata f. baueri Steyerm. 
(John Louis) 


Peperomia glabella var. obtusa Stey- 
erm 


Volume 76, Number 3 


Taylor 749 


1989 Plants Described by 
Julian A. Steyermark 
BEAMAN BRADE 
3962 Cuchumatanea steyermarkii Seidensch. 4739 Perama sparsiflora Standl., Steyerm. & 
& Beaman Kirkbr 
BELEM, R. P. BRANDEGEE, K. (Katherine, or Mary Katherine Curran 
2738 Sphinctanthus insignis Steyerm. nee Layne) 


BERLANDIER, J. L. (John Louis) 
766 (2186) Grindelia tenella Steyerm. 


BERNARDI, A. L. 


3180 Hoffmannia bernardii Steyerm. 
5829a Tresanthera thyrsiflora Steyerm. 


BERNARDI, L. (Luciano) 


611 Manettia bernardii Steyerr 
985 Arachnothrix reflexa var. meridensis 
Steyerm. 
1248 Manettia paramorum Steyerm 
1296 vchotria costanensis subsp. ania 
Steyer rm. 
1304 Ps ola bernardii Steyerm. 
1690 Pagamea plicata var. multinervia Stey- 
erm. 
1720 Psychotria trichotoma var. trichophylla 
Steyerm. 
2011 Guettarda bernardii Steyerm. 
2325 Coussarea bernardii Steyerm. 
0746 Neea bernardii Steyerm. 
7437 Palicourea buntingii var. glabrior Stey- 
erm. 
BERRY, P. E. (Paul) 
094 Piper otto-huberi var. eciliatum Stey- 
erm. 
1558 Rudgea berryi Steyerm. & Dwyer 
2096 Faramea berryi Steyerm. 
2116 Remija berryi rm 
2152 Ouratea macurucoensis Maguire & 
Steyerm. 
3670 Eugenia mevaughii Steyerm. & Lasser 
3825 Piper berryi Steyerm. 
3923 Pereromia berryi Steyerm. 
BLANCO, C. (Carlos) 
1158 Guapira sipapoana Steyerm. 
BLANKINSHIP, J. W. (Joseph William) 
s.n. Grindelia nana var. altissima f. puberula 
'erm. 
BLYDENSTEIN, J. 
250 Diodia teres subsp. prostrata f. latifolia 
Steyerm 


BOLANDER, H. N. (Harry Nicholas) 


389 Grindelia hirsutula f. cacumena Stey- 
erm. 
BONO, J. (José) 
3797 Hillia bonoi Steyerm. 
BOOM, B. (Brian) 
5274 Ferdinandusa boomii Steyerm. 
5278 Faramea boomii Steyerm. 
6417 Peperomia boomii Steyerm. 


BORDENER, V. (Valerie) 
s.n. Podophyllum peltatum f. biltmoreanum 
Steyerm 


s.n. Grindelia rubricaulis var. latifolia f. pu- 
escens Steyerm 

L. (Lucy) 

s.n. Silphium Mr nea E var. lucy- 
brauniae Steye 


BRAUN, E. 


BRETELER, F. 


3463 Hoffmannia pauciflora subsp. venezue- 
lensis Steyerm 

4082 Piper veraguense var. venezuelense 
Steyerm. 

4089 Manettia breteleri Steyerm. 

4095 Diodia radula subsp. venezuelensis Stey- 
erm. 

4204 Diodia teres subsp. prostrata f. leiocar- 

a Steyerm. 
4832 


P 
Ouratea inconformis Maguire & Stey- 
erm 


BREWER, C., JR. 


248 Navia breweri L. B. Smith & Steyerm. 
BRINKLEY 
250 Bi is Te var. arkansana 
Steye 
268 Clematis v versicolor f. pubescens Stey- 


BRITTON, N. L. (Nathaniel Lord) 


2104 Psychotria capitata subsp. inundata var. 
septentrionalis Steyerm 

2684 Randia brevipes Steyerm. oe 

2702 Randia aculeata f. minor Steyer1 


BROADWAY, W. E. 


9823 Psychotria muscosa subsp. breviloba 
Steyerm. 
BUCHTIEN, O. (Otto) 
1719 dera M nc var. schwackei f. 
dentata Steyern 


BUNTING, G. S. "is 


2332 Piper deliciasanum Steyerm. 
4461 Peperomia choroniana var. heterodoxa 


Steyerm. 


4509 Ladenbergia buntingu Steyerm. 
4805 Monstera steyermarkii Bunting 
10876 Pyschotria buntingii Steyerm. 


BUSH, B. F. (Benjamin) 


7041 Diospyros virginiana f. pumila Palmer 
& Steyerm. 
10332 poyas americ ana f. indehiscens Palm- 


CÁRDENAS DE GUAVARA, L. (Lourdes) 


805 Arachnothris rugulosa var. tachirensis 
Steyerm 
CARDONA P., F. (or F. Cardona Puig) (Felix) 


K maguirei op pusillus 
V glabrus Steyer1 


750 


Annals of the 
Missouri Botanical Garden 


707 Sabicea venezuelensis Steyerm. 

819 Duroia bolivarensis Steyerm. 

860 Platycarpum rhododactylum Woodson 
& Steyern 

1051 Sloanea merevariensis Pittier ex Stey- 
erm. 

1115 Faramea cardonae Standl. & SENE 

1117 alanea guaiquinimensis Steyer 

1333 Psychotria parimensis Steyerm. 

1383 Palicourea cardonae St teyerm. 

1480 Ilex cardonae Steyerm. 

1496 Psychotria tapirapecoana Dn 

1503 udgea cardonae Steyern 

1610 Gleasonia duidana var. el 
Steyerm. 

1630 ir glalioides var. densipila Stey- 

2190 Stenopadus taloumifolius var. magnifo- 
lius Steyerm. 

2204 Ouratea ramosissima Maguire & Stey- 
erm. 

2288 Quelchia cardonae Steyerm. 

2661 Heliamphora minor f. laevis Steyerm. 

2123 Conomorpha steyermarkiana Agost. 

2943 llex magnifructa var. minor Steyerm. 

2967 Psychotria phaneroloma var. angustior 
Steyerm 

CARLSON, M. ( 
1744 Ardisia carlsonae Steyerm. 


CARNEVALI, G. (Germán) 

040 Peperomia carnevalii Steyerm. 
CASTELLANOS, E. 

46 Borreria aristeguietaeana Steyerm. 


CAVALCANTE, P. 


712 Retiniphyllum oe Steyerm. 
2511 Randia pubiflora Stey 
CHARDON PALACIOS, €. E. (Carlos Moser) 
179 Arachnothrix venezuelensis Steyerm. 
CLARK, H. L. (Howard) 
6742 Catostemma pubistylum Steyerm. 
6980 de clarkii Steyerm. 
7794 suapira sanc arlosiana Steyerm. 
8111 E hoepfia d Steverm. 
8117 tostemma sancarlosiana Steyerm. 
8126 Catostemma a clarkii Steyerm. 


CONGDON, J. W. 
s.n. Grindelia aggregata Steyerm. 
CONICIT, E. 
2663 Piper bredemeyeri f. escabridum Stey- 
erm. 


COOPER, G. P. 


586 Bertiera aequaliramosa Steyerm. 
CORE 
1367 Joosia umbellifera var. caucana Stey- 
erm. 


COWAN, R. S. (Richard Sumner) 
1863 Sipanea cow 
1897 Psy 


i Steyerm. 
'chotria onsen f. pubes- 
cens Steyer 


31061 


8 
31405 


31447 
31450 


31455 
31462 
31473 


38429 
39036 


CREMERS, ( 


6460 


CROAT, 1 


59478 
CROIZAT, 
624 
CRUEGER 


s.n. 


Orthaea paruensis Maguire, Steyerm. & 
Luteyn 

Ouratea paruensis Maguire & Steyerm. 

Chondrococcus laevis Steyerm 


Rhamnus longipes Steyerm. 

Schefflera m. nayensis subsp. ies 
agui ire, Steyerm. & Frodi 

Duroia acu Steyerm. 

Matayba longipes subsp. tepuiensis 
Steyerm. 

ioa galioides f. macrocephala Stey- 


reine brevicalyx Maguire & Stey- 
erm. 

Platycarpum orinocense var. grandiflo- 
rum Steyerm. 

Rudgea bremekampiana Steyerm. 

Rudgea standleyana Steverm. 


Sipanea ovalifolia var. villosissima Stey- 
erm 


B. (Thomas) 


Geophila croatii Steyerm. 
Coccocypselum croatit DE 
llex gransabanensis Steyern 
Ilex abscondita ci 


L. (Léon Camille Marius) 


Ouratea croizatii Maguire & Steyerm. 


Psychotria perpapillifera Steyerm. 


Cl UD d 


13694 


14827 


21086 


CURRAN, H. 


S.n. 


CUTLER, H. 


DAVIDSE, 


17359 


Toc a cuatrecasasit i i 
Elaeagia multinervia Stey 
Cuatrec nae pi tE 


Steye 

Joosia umbellifera var. occidentalis 
Steyerm. 

Schradera puberula i deb" 

ipid odendron cuatrecasasit Steyerm. 
iatrecasasiodendron spec ie Stey- 


Se ides pulverulenta Steyerm. 
Elaeagia cuatrecasasti Steyerm. 
Elaeagia alterniramosa Steyern 
Arachnothrix reflexa var. slve Stey- 


Faramea occidentalis var. brachycalyx 
Steyerm 


C. (Hugh Carson) 


Utricularia cutleri Steyerm. 
Utricularia cearana Steyerm. 


G. (Gerrit) 


Alibertia davidsei Steyer1 
Coccocypselum A Steye rm. 
Guapira davidsei Stey 

Neea davidsei Steyerm 

Matayba affinis Steyerr 

Palicourea huberi Don 


Volume 76, Number 3 
1989 


Taylor 751 
Plants Described by 


Julian A. Steyermark 


17427 Botryarrhena venezuelensis Steyerm. 
22679 Ilex polita Steyerm. 
22719 Ilex davidsei Ste 


yer 
27327 Morinda o e Steyerm. 


27651 Psychotria anartiothrix Steyerm. 
27719 llex davidsei Steyerm. 
27189 Sloanea davidsei Steyerm. 


DAVIDSON, A. (Anstruther) 
736 (96) Grindelia arizonica var. microphylla 
Steyerm 

DAVIDSON, M. E. 

Croton standleyi Steyerm. 


DAWSON, E. Y. (Yale) 


14163a Declieuxia dasyphylla f. ciliata Steyerm. 
14594 Euphorbia kenn i a 
14672 Diodia angustata Steyer 
14685 Croton wur ca. 
14770 anaes ia dawsonii Steyerm. 
14776 Phyllanthus dawsonii Steyerm. 
14918 Phyllanthus websterianus ipii 
14973 dne dum. Steyerm 
DE GRANVILLE, J. J. (Jacques) 
Cals Psychotria antenniformis Steyerm. 
2353 Psyc :hotria granvillei Steyerm 
2360 Psychotria lateralis Steyerm 
2361 Psychotria saulensis Stey 
2362 sychotria microbracteata Steyerm 
2383 Psychotria galbaoensis Steyerm 
2393 sychotria urceolata Steyerm 
2793 Ee hotria viridibractea Sun erm. 
3127 Psychotria alloantha Steyer 
B.4640 rd perferruginea Steye erm. 
B.5288 Psyc :hotria squamelligera Steyerm. 
DE LA CRUZ, ]. 5. 
1638 Palicourea guianensis subsp. occidental- 
is Steyerm. 
2164 Malanea cruzti Steyerm. 
DE LA RUE, A. 
13 Psychotria a subsp. amplifolia f. 


tomentosa Stey 


DE LEMOS FRÓES, R. (Ricardo) see Fróes, R. L. 


DEAM, C. C. (Charles Clemon) 
40 Mikania guatemalensis Standl. & Stey- 
erm. 


DELASCIO CHITTY, F. (Francisco) 


8471 Chomelia delascioi Steyert 

12358 erama dichotoma var. monocephala 
Steyer m. 

13161 Peperomia delascioi Steyerm. 


DELGADO, E. 


160 Tocoyena costanensis Steyerm. 
233 Linociera avilensis Steyerm. 


DUCKE, A. (Adolpho) 


81 Henriquezia verticillata var. apiculata 
` teyerm. 
222 Burdachia duckei Steyerm 
683 Pagamea coriacea var. pubesc ens Stey- 


erm. 
799 Duroia nitida Steyerm. 


Platycarpum duckei Steyerm. 
1141 Borojoa duckei Steyern 


1167 Ferdinandusa duckei Stey 

1332 Duroia eriopila var. bre DS Stey- 
erm. 

1557 Kotchubaea duckei Steyerm. 

1844 Stachyarrhena acutiloba Steyerm. 

1943 Malanea subtruncata Steyer 

8402 


Pagamea puberula e (RB 
22951) 

DUDLEY, T. R. 
13089 
13543 


Psychotria dudleyi Steyerm. 

Pagamea dudleyi Steyerm. 

K. (Geoffrey) 

pe steyermarkii Garay & Dun- 


rville 


DUNSTERVILLE, G. C. 


DUQUE, J. M. 
1559 Elaeagia barbata Steyerm. 


EASTWOOD, A. (Alice) 


s.n. Grindelia hirsutula var. brevisquama f. 
edunculoides Steyerm. 
794 Grindelia rubricaulis var. latifolia f. mi- 


r Steyerm. 


EGLER, W. A. 
46683 Hillia irwinii Steyerm. 
47729 Malanea egleri Steyerm. 
EHRENDORFER, F. (Friedrich) 
04-23 Talisia pentantha Steyerm. 
EKMAN, E. L. (Erik Leonard) 
9568 Schradera cubensis Steyerm. 


ESPINOSA, R. 
1076 


EVINGER, E. L. 


Nierembergia espinosae Steyerm. 


513 Mitracarpus frigidus var. peruvianus 
Steyerm 
EWEL, J 
114 Psychotria nematostachya Steyerm. 
135 Pochota ewelii Steyerm. 
209 Brunellia neblinensis CU & Cuatr. 


FAIRBROTHERS 

2. Psychotria aristeguietae Steyerm. 
FANSHAWE, D. B. 

877 apis brevipes Steyerm. 

ra potaroensis Steyerm. 

TRES pues ei Steyerm. 
FARINAS, M. 

433 Phyllanthus tepuicola Steyerm. 
FERRIS, R. (Roxanne) 

3391 Grindelia squarrosa var. nuda f. angus- 

tior Steyerm. 
FLORSCHUTZ, J. 

Faramea costata Steyerm. 

FOLDATS, E. (Ernesto) 

2784 Ouratea subamplexicaulis Maguire « 

Steyerm 


752 Annals of the 
Missouri Botanical Garden 
a 0 Palicourea foldatsii Steyerm. GENTRY, H. S. (Howard Scott) 
3678 Sipaneopsis foldatsii Steyerm 2525 Tinantia erecta f. puberula Standl. & 
3758 Kol ubaea micrantha Steye Steyerm 
7100 Chimantaea cinerea f. glabra Steyerm. 
7192 Hillia foldatsi Steyerm. GHIESBREGHT 
AN 871 Arenaria chiapensis Standl. & Steyerm. 
CAP 41 Appunia sparsiflora Steyerm. GLEASON, H. A. (Henry Allan) l 
WB 388 Isertia parviflora var. hirta Steyer 431 Sipanea gleasonii Steyerm. 
WB 500 Appunia tenuiflora var. leiophylla Stay: GONZÁLEZ, T. (Tomás) 
erm. à : a 
G 569 Piram Step s.n. Colubrina venezuelensis Steyerm. 
F 877 Faramea brevipes Steyerm. GRANT, J. M. 
FOREST DEPT. (Record No.) s.n. Grindelia stricta var. aestuarina Stey- 
7227 Ouratea fasciculata Maguire & Stey- SEN 
erm. GRANT, M. L. 
7832 10567 Pentagonia grantii Steyerm. 


Psychotria tetramera Steyerm. (Field 
No. R.B. 8) 


FOSBERC, F. R. (Francis Raymond) 


21211 


21418 


21810 


23141 
FRÓES, R. 
21225 


21466 


22885 


Palicourea guianensis subsp. occidental- 
is f. glabra Steyerm. 

Arachnothrix reflexa var. breviloba 
Steyerm. 

Psychotria fosbergii Steyerm. 

Amphidasya intermedia Steyerm. 

Arachnothrix fosbergii Steyerm. 

Arachnothrix lojensis Steyerm. 


L. (Ricardo) 


emijia vaupesiana Steyern 
a venal Krukoff 
Barneb 
€ V itida var. subcuneata 
Steyer 


FUNCK & SCHLIM 


1500 
GARCIA, F. 

55 
GARCIA, J. 

80 

128 
GARCIA, T 

147 


Palicourea bostrychostachya Steyerm. 
Palicourea garciae Steyerm. 


Faramea garciae Steyer 
Borojoa venezuelensis da 


Chimarrhis microcarpa var. speciosa 
Steverm 


GARCIA SALAS, J. 


1442 
GATTINGER 


s.n. 


GENTLE, P. E 


116 
871 
2264 
2619 
GENTRY, 
46694 
6887 


47314 


Daphnopsis ficina Standl. & Steyerm. 


Arenaria patula f. media Steyerm. 


1. (Percy) 


Psittacanthus mayanus Standl. & Stey- 
erm. 
Lina heterodoxa Standl. & Stey- 
m. 
Piper vaccinum Standl. & Steyerm. 
Phyllant thus longipes Steyerm 


A. (Alwyn) 


Schefflera simplex Steyerm. & Holst 
Piper gentryi Steyerm. 
Sabicea bariensis Steyerm. 


L. (Edward Lee) 


s.n. Grindelia humilis f. reflexa Steyerm. 


GREENE, E. 


GREGG, J. 
44 Menodora iS var. engel- 
mannii Steyerr 
GUANCHEZ, F. (Francisco) 


3644 Psychotria guanchezii Steyerm. 


HALL, F. (Elihu) 
ies en var. virgata f. vil- 
losa Steye 
HALLE, F. 
1018 Coussarea hallei Steyerm. 
HARLEY, R. M. 
15006 


Geophila harleyi Steyerm. 
16038 Perama harleyi i Kirkbr. & Steyerm. 


17069 Pagamea harleyi Steyerm. 
HATCH, W. R. 
54 Mortoniodendron guatemalense Standl. 
& Steyerm. 
162 Pilea mimema Standl. & Steyerm. 


HAUGHT, O. 


2190 Psychotria rosea var. trichophora Stey- 
erm. 
3068 Faramea occidentalis subsp. lonchocalyx 
Steyerm. 
5400 Hippotis grandiflora Steyerm. 
HELLER, A. A. (Amos Arthur) 


12579 Grindelia nana var. turbinella Steyerm. 


HENDERSON, A. (Andrew) 
501 Prestoea humilis Henderson & Stey- 
erm 
HENDERSON, L. F. 
2791 Grindelia howellii Steyerm. 
. (Lionel) 


Ochthocosmus micranthus Steyerm. 


HERNÁNDEZ, I 
348 
HEYDE & LUX 


6392 Tinantia longipedunculata Standl. & 
teyerm 


Volume 76, Number 3 
989 


Taylor 753 
Plants Described by 
Julian A. Steyermark 


HOFFMANN 


s.n. Grindelia camporum var. abbreviata 
Steyerm 
HOLST, B. (Bruce Kendall) 
3426 Schefflera yutajensis Steyerm. S Holst 
3523 Paepalanthus Aolstii Steyerm. 
3539 Lindmania holstii Steyerm. & L. 
Smit 
3617 Ilex summa Steyerm. 
HOLT, E. ( 
743 Psychotria blakei Standl. & Steyerm. 


HOWARD, R. A 


12302 Schradera subsessilis Steyerm. 
HOWELL, J. T. (John Thomas) 
4352 Grindelia rubricaulis var. permixta Stey- 
erm. 
5201 


Grindelia camporum var. interioris f. 
foliacea Steyerm. 


5221 Grindelia hirsutula var. brevisquama f. 
labrata Steyerm. 
5338 Grindelia hirsutula var. brevisquama 
Steyerm. 
5473 Grindelia arenicola Steyer 
5496 Grindelia camporum var. Sardis 
yerm. 
6574 Grindelia savages var. elata Steyerm. 
7545 Grindelia stricta var. aestuarina f. elon- 
gata Steyerm. 
8106 Grindelia arenicola var. pachyphylla 
eyerm. 
10805 dn delia humilis f. pubescens Steyerm. 
11414 md hirsutula var. subintegra Stey- 
11658 Grindelia maritima f. anomala Steyerm. 
HOYOS, J. (Jesüs) 
46 Kotchubaea morilloi Steyer rm. 
90 Ouratea asisae Maguire & Steyerm. 


HUBER, O. (Otto) 


41 Maytenus huberi Steyerm. 

43 Croton huberi Steyerm. 

1229 Maytenus insculpta Steyerm. 

1447 Ouratea ayacuchae Mo & Stey- 

erm. 

1456 Neea ignicola Steyerm. 

1693 Neea robusta Steyerm. 

1791 Psychotria yapacanensis Steyerm. 
1848 Piper otto-huberi Steyerm. 

1904. Ouratea huberi Maguire & Steyerm. 
2504 Ouratea deminuta Magui re & Steyerm. 
3759 Tepuian anthus savannensis Maguire & 

yerm. 
4437 Ilex paruensis Steyer1 
5058 Navia huberiana L. B. Smith, Steyerm. 
| H. inson 
5481 — a i un re & Steyerm. 
5536 i mila . heterodoxa Ma. 
gui one 

5763 Ouratea eee Maguire & Steyerm. 
5186 Sipaneopsis huberi Stey 
6136 Panopsis parimensis Saeed 


6184 Navia culcitaria L. B. Smith, Steyerm. 
& Robinson 
6945 Aphanocarpus steyermarkii f. elongatus 
8246 Bas "e Steyer 
9123 Coccoc ypselum huberi poc 
9467 Stegolepis huberi Steyerm 
9818 d bolivarensis Steyerm 
10388 ayba ptariana subsp. guaiquinimae 
St eyerm. 
11849 Sy smbolanthus yaviensis Steyerm. 
UMBERT, H. (Henri) 
26739 Psychotria alticola Steyerm. 
HUTCHINSON, P. C. 
6799 Hillia wurdackii Steyerm. 
IJJASZ 
425 Psychotria aubletiana var. producta 
teye 


IRWIN, H. S. (Howard) 
10161 Psychotria sciaphila subsp. longicalyx 
Steyerm. 


14771 Psychotria hoffmannseggiana var. celsa 
Steyerm. 

18688 Manettia irwinii Steyerm. 

23537 Hindsia irwinii Ste 1 

27120 Mitracarpus pusillus Steyerm 

47354 Palicourea irwinii Steyerm 

47393 Psychotria oiapoquensis Steyern 

47620 Toc oyena guianensis var. communis 

teyerm. 

54949 Mrs wilhelminensis Steyer 

55048 d ies wilhelminensis Ste yerm. 

55299 ertiera diversiramea Steyerm. 

55706 Psyc pem irwinii Steyer 


JANGOUX, J. (Jacques) 
10157 Borreria jangouxii Steyerm. 


10193 Piper perijaense Steyerm 
JÁTIVA, C. (Carlos) 
682 Palicourea guianensis subsp. occidental- 
is var. glabrescens Steyerm. 
735 Psychotria macrophylla tus anomo- 


thyrsa f. tomentella Steye 


JENSEN; L- P: 


s.n. Cassia fasciculata f. jensenii Palmer & 
Steyerm. 


JOHNSON, H. 
Poikilacanthus setiferus Standl. & Stey- 
erm 


JOHNSTON, J. R. 


1026 Verbesina agricolarum Standl. & Stey- 
erm. 

1255 Clethra johnstonii Standl. & Steyerm. 

1480 Eugenia jutiapensis Standl. & Steyerm 

1525 Berberis johnstonii Standl. & Steyerm 

1643 Thalictrum johnstonii Standl. & Stey 
erm. 

1812 Hypericum arbuscula Standl. & Stey- 
erm. 


754 


Annals of the 
Missouri Botanical Garden 


1887 Struthanthus johnstonii Standl. & Stey- 
erm. 
JOLY, C. 
5630 dis arielanum var. ae a f. 
barbulaense Steyer 
JOOR 
s.n. Grindelia littoralis Steyerm. 


JÓRGENSEN, P. 
2643 
3694 


Menodora pinnatisecta Steyerr 
Genipa americana 
senii Steyer 


KELLERMAN, W. A. (William MT 


5175 paisa pantomalaca Standl. & Stey- 
erm. 

7223 Schistocarpha steyermarkii H. m 

7223 da steyermarkiana H. Robinson 

7532 ee macrocarpa Standl. & Ste ey- 

7630 


ne an ee treleaseanum Standl. & 
Steyerm 


KILLIP, E. P. (Ellsworth Paine) 


6585 Polygala santanderensis Killip & Stey- 
erm. 
26780 Geophila cordifolia var. peruviana Stey- 
erm. 


KIRKWOOD, J. E. (Joseph Edward) 


20 Menodora coulteri var. minima Stey- 
erm. 
KLUG, ( 
347 Tetrapterys po Macbr. 
1969 Mabea klugt 
2010 Tapura a Macbr, 
2019 Psychotria subpedicellata Steyerm. 
2064 Mabea standleyi Steyerm. 
2164 Bertiera guianensis subsp. pubiflora 
Steyerm. 
2362 Coussarea klugii Steyerm 
3084 Hippotis subelongata Steven 
3206 Mabea elata Steyerm. 
3452 Kotchubaea montana Steyerm. 
KOYAMA, T. (Tetsuo) 
1506 Mapania steyermarkii T. Koyama 


KRUKOFF, B. A. (Boris Alexander) 


Retiniphyllum laxiflorum var. brasiliense 


'yerm. 

1983 Mabea pirioides Steyer 

4940 Psychotria embirensis DU 
6210 Drypetes amazonica Steyern 

6297 Omphalea elaeophoroides Steyerm 
6455 Pseudoconnarus krukovii Steyerm 
6858 Ps negacephala Steyerm 
8396 Conceveiba krukoffit Steyerm 
8513 Richeria submembranacea Steyerm. 
8556 Rourea Arukovii Steyerm. 

8616 Conceveiba simulata Steyerm 

869 Conceveiba magnifica Steyerm. 
10650 Elaeagia microcarpa Steyerm. 

KUNTZE, O. (Carl Ernst Otto) 
1643 Mitracarpus frigidus var. andinus Stey- 
erm. 


var. caruto í jorgen- 


KUYLEN, H 
151 Clethra bimatris Standl. & Steyerm. 
LANJOUW, J. (Joseph) 
802 Palicourea crocea var. riparia f. hetero- 
doxa Steyerm. 


LASSER, T. (Tobias) 


1042 Palicourea perquadrangularis var. brev- 
ipes Steyerm. 

1156 Pa oua lasseri Steyerm. 

1221 areik insignis Steyerm. 

1417 Coccoc um gulanense var. patens 
ea 

1682 Coussarea loss i St teyer 

2047 Psychotria araguensis Ma 

LASSER, T. & J. A. STEYERMARK 


55103 Symplocos lasseri Steyerm. (possibly a 
Steyermark number) 


LEPRIEUR, F. R. M. 


118 Psychotria pungens Steyerm. 
LESUER, H. (Harde) 
1016 Grindelia confusa Steyerm. 
LIESNER, R. L. (Ronald) 
409 Neea mapourioides Steyerm. 
5991 Rauia subtruncata Steyerm 
6222 Piper pseudoacreanum Steyerm 
6405 Piper liesneri Steyerm. 
7729 Peperomia falconensis i iia 
8176 Piper cerronianum Steye 
8211 Piper wingfieldii Steyerm 
8959 eea sebastianii Steyerm. 
9699 iper sierra-aroense Steyerm 
9767 Piper cordiforme Steyerm 
10022 Piper linguliforme Steyerm. 
10107 Psychotria aubletiana var. villipila Stey- 
erm. 
10494 Piper arieianum var. yaracuyense f. 
puberulum Steyerm. 
10555 Peperomia fundacionensis Steyern 
10891 Piper longiappendiculatum veni 
11151 Talisia heterodoxa Steyerm. 
11277 Neea bracteosa Steyerm. 
11479 Sloanea bolivarensis Steyerm. 
11647 Piper fundacionense Steyerm 
11873 Piper fete Steyerm. 
15866 Navia liesneri L. B. Smith, Steyerm. & 
: nos 
15955 Psychotria pectinata Steyer 
16013 Pagameopsis maguirei alps pisie 
yerm. 
16025 Navia litera L. B. Smith, Steyerm. & 
obinson 
16026 Navia crassicaulis L. B. Smith, Stey- 
m. & H. Robinson 
16355 Pochot ta mawarinumae Steyerm. 
16502 Neea lies Ste 
16605 Raveniopsis lio ^sneri fi Steyerm. 
16688 llex glabella Stey 
16720 Maytenus Puls a "Ste eyer 
16809 Sauvagesia marahuacensis m nebli- 
nensis o. 
16901 Peperomia gentryi eoi iiia) 
16960 Faramea paludicola Steye 
16963 Psychotria thesceloantha a 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


17493 Coussarea evoluta Steyerm. 
17599 Ilex a et Steyerm 

17677a Sauvagesia mecha Steyerm. 
18214 O ronaldii Steyerm 
18403 Neea huachamacarae Steyerm. 
18428 Neea OS cae Steye 

18452 Pochota fuscolepidota Steyerm. 
18455 Sloanea longiaristata Steyer 
18469 Ilex liesneri Steyerm. 

18595 Ilex acutidenticulata Steyerm. 
18637 ochota liesneri Steyerm 
18796 


19595 Psychotria edaphothrix Steyerm. 

19878 Ilex longipilosa Steyerm. 

20394 Talisia maruayana Steyerm. 

20394 Talisia amaruayiensis Steyerm., ined. 
mss. 

20514 Neea amaruayensis Steyerm. 

20800 Guapira bolivarensis Steyerm. 

21248 Pochota redmondii Steyerm. 

21640 Faramea y 


21826 Yutajea liesneri Steyerm 

21827 Rudgea corocoroensis Steyerm 

21904 es siapensis L. B. Sub & Stey- 

21968 Bon: mosaicum Steyerm 

21998 — terramarae L. B. Smith & Stey- 

22041 Navia | ria L. B. Smith & Steyerm. 

22062 Navia Sarda L. B. Smith & Stey- 
erm. 

22064 Cephalodendron aracamuniensis Stey- 
erm. 

22075 Sauvagesia mp subsp. aracamu- 
niensis Steyer 

22079 Symbolanthus aracamuniensis Steyerm. 

22105 Bonnetia liesneri Steyer 

22153 Neblinathamnus aracamunianus Stey- 
erm. 

22197 atea aracamuniana Steyerm. 

22203 Navia iosothrix L. B. Smith & Steyerm. 

22202 Coussarea liesneri Steyer 

22322 jga carnevalii L. B. Smith & Stey- 

22407 Remijia reducta Steyerm. 

22473 Ilex aracamuniana Steyerm 

22559 Pochota aracamuniana Steyerm. 

22622 


e steyermarkii Maas & 
Maa 


LIGGETT, W. E. (William) 
l Echinacea purpurea f. liggettii Stey- 
erm. 


LINDHEIMER, F. (Ferdinand Jacob) 
9 


Grindelia microcephala var. adenodonta 
Steyerm. 


LITTLE, E. L. (Elbert Luther) 

Pagamea guianensis var. macrocarpa 
Steyerm. 

Psychotria aubletiana var. andina f. pu- 
bescens Steyerm. 

Simira erythroxylon var. meridensis 
Steyerm. 


15238 


15354 


15372 Guettarda steyermarkii f. pilosior Stey- 

erm. 
LUNDELL, C. L. (Cyrus Longworth) 

2012 Solanum decurtatum Standl. & Stey- 
erm. 

2213 Annona primigenia Standl. & Steyerm. 

3193 Lisianthius petenensis Standl. & Stey 

6193 Kocani schippii Standl. & Steyerm. 


LUTEYN, J. L. (James Leonard) 


9413 Raveniopsis cowaniana Steyerm. & Lu- 
teyn 
9610 Connellia smithiana Steyerm. & Luteyn 


MADRIZ, A. 


Trigonia costanensis Steyerm. & Badillo 


MAGUIRE, B. (Bassett) 


2874 Catostemma ebracteolatum Steyerm. 
3558 Dendrosipanea wurdackii yerm 
4612 Borreria cataractarum Steyerm 
23072 Coussarea fanshawei Steyerm 
23204 Utricularia kaieteurensis Steyerm 
23249 tricularia maguiret Steyerm. 
24361 Retiniphyllum pb var. reticula- 
tum Steye 
24393 groom Miei var. tafelbergensis Stey- 
24530 shines tafelbergensis Steyerm. 
24874 Malanea sarmentosa f. tomentosa Stey- 
erm. 
27324 Sipanea pratensis var. dichotoma f. 
brachycarpa Steyerm 
27485 Ouratea ornata Maguire & Steye 
21691 — sipapoensis mh & Stey- 
er 
27674 ee sipapoana Steyerm 
27756 Schefflera a on sipapoen- 
si , Steyerm. & Frodin 
27864 -— gane. subsp. sipapoensis 
Stey 
27869 ati phys laxiflorum var. longilo- 
m Steyerm. 
27895 Schradera brevipes Steyerm 
27971 Chimarrhis brevipes s Steyerm. 
28013 Remijia Won Steyer 
28243 eruma coccocypseloides o — 
28349 ince sina Maguire, Steyerm. 
Lut 
28399 ccs oligandra var. occidentalis 
Ste 
28409 Pa lineis cuaoensis Steyerm 
28454 Schefflera pauciradiata Mascus Stey- 
erm. & Fre din 
28458 Retiniphyllum tepuiense Steyerm. 
28471 Schefflera sipapoensis Maguire, Stey- 
erm. & Frodin 
28483 Rudgea sipapoensis Steyerm 
28516 Piper politii subsp. as Steyerm. 
28536 Pagamea velutina Steyerm 
28615 Talisia caudata Ste 1. 
28626 Chimarrhis bathysoides a 
28656 Rhamnus sipapoensis Steye 
8741 Catostemma ebracteolatum [A 
28743 Malanea sipapoens is Steyerm. 
28718 Psychotria sipapoensis Steyerm. 


756 Annals of the 
Missouri Botanical Garden 
29079 Remijia densiflora f. glabricalyx Stey- 33427 Pagamea magniflora Steyerm 
erm. 33700 — guianense var. cunea- 
29137 Sabicea velutina subsp. duidense Stey- tum Stey 
erm. 34531 Retiniphyllum. schomburgkii subsp. occi- 
29242a Borreria pygmaea var. robusta Stey- dentale Steyerm. 
erm. 34923 Med ngu var. glabriuscula 
29253 Ouratea evoluta Maguire & ee Steyer 
29469 Randia amazonasensis $ 35092 PR duricoria var. longiloba Stey- 
29529 Perissocarpa umbellifera eco & erm. 
Maguire 35143 atayba yutajensis Steyer 
29543 Tepuianthus sarisarinamensis subsp. 35149 Thibaudia dolichandra aie Stey- 
duidensis Maguire & Steyerm. erm. & Luteyn 
29546 Schefflera pedicelligera Maguire, Stey- 35154 Sc o reticulata subsp. yutajensis 
erm. & Frodin Maguire, Steyerm. & Frodin 
29564 ma UMS Maguire, Steyerm. 35287 Hillia oa Steyer 
& Lu 35304 Peperomia yutajensis Bout rm 
29573 Pagamea pum bing conferta f. 35344 Gleasonia duidana var. latifolia Stey- 
breviloba Steye erm. 
29786 diede ELS Steyerm. 35356 Schradera yutajensis Steyerm. 
29851 Psychotria adenophora Steyerm. 35635 Schefflera pimichinensis Maguire, Stey- 
29859 ED huachamacariensis Stey- erm. & Frodin 
erm. 36188 Portulaca pygmaea Steye 
30001 Styrax tepuiensis zr huachamacarii 36259 Stachyarrhena reticulata Steyerm 
Maguire & Ste 36518 Psychotria rosea f. calvescens Steyerm 
30253 Schefflera aca Maguire, Stey- — 36657 Ouratea discophora subsp. pervenulosa 
“rodin uire & Steyerm. 
30285 Laplacea pubescens var. minor Stey- 36800 Ferdinandusa neblinensis Steyerm. 
erm. 36889 Peperomia foveolata um 
30458 oe glomerata var. paucinervia 36891 Peperomia yatuensis Steyerm. 
Steyerm. 36921 ‘a :hotria speluncae aa exserta 
30557 Ochthocosmus multiforus var. angusti- Steyerm. 
folius Steyerm. & Luteyn 36948 ge are maguirei nq neblinen- 
30632 Bonnetia tristyla subsp. nervosa Stey- sis var. angustifolius Steyerm. 
erm. (superfluous, illegitimate) 36975 Geophila orbicularis var. neblinae Stey- 
30683 Tepuianthus yapacanensis Maguire & erm. 
Steyerm 37011 blinathamnus glabratus Steyerm 
30766 Ouratea rotundipetala Maguire & Stey- 37040 Rhamnus neblinensis Maguire & Stey- 
erm. erm. 
30878 Sipaneopsis morichensis Steyerm. 37040 Rhamnus roin Maguire & 
30881 Rudgea morichensis Steyerm. eyerm., ine 
30886 en pue var. hispida f. cinerea 37103 Macrocarpaea dua: Maguire & 
Steyerm. 
30945 TRA Lode Maguire, Stey- 37130 Pagameopsis maguirei subsp. neblinen- 
erm. & Lute sis Steyerm. 
31740 nas pe Maguire, Stey- 37148 Psychotria speluncae subsp. brevicalyx 
rodin Steyerm. 
31823 Psychotria glandulicalyx subsp. caman- 37230 Rhamnus psilocarpa Maguire & Stey- 
iensis Steyerm. erm. 
32219 Calycophyllum spectabile d E 37254 Cavendishia neblinae Maguire, Steyerm. 
32329 Faramea fanshawei Steye Luteyn 
32339 Orthaea pubifolia Maguire. peut. & 37328 Dioscorea neblinensis Maguire & Stey- 
uteyn erm. 
32351 Mapania maguireana T. Koyama & 37354 Schefflera globulifera Maguire, Steyerm. 
Steyerm. Frodin 
32357 aramea maguirei Steyern 37483 chotria barnebyana Steyerm. 
32430 Cavendishia salicifolia Mis Stey- 37490 Palicourea glabriflora Steyerm. 
erm uteyn 37570 Sipaneopsis pacimoniensis Steyerm 
32456 Gonzalgunia dicocea var. guianensis 37592 Dendrosipanea revoluta Steyerm 
Stey 37639 Sipaneopsis wurdackii Steyerm. 
32618 Psy houia puberulenta Steyerm. 40009 Alibertia myrciifolia var. tepuiensis 
32629 Ouratea gillya oil pachypoda Ma- Steyerm. 
guire & S N 40513 Psychotria pakaraimensis Steyerm 
32758 Styrax ue ES guaiquinimae 40637 Chalepophyllum longilobum ek: 
Maguire & Steyer 40666 Platycarpum b up nid Steyerm. 
32974 Schradera maguirei Ste eye 40799 Ixora aluminicola Steyerm. 
33070 Pagamea la a 41438 Sipaneopsis a Steyerm. 


Volume 76, Number 3 Taylor 757 
1989 Plants Described by 
Julian A. Steyermark 


41496 Remijia argentea 60503 Schefflera pallens Maguire, Steyerm. & 
41860 iris dar hombre me occi- Frodin 
entale var. hirticalyx Steyerm 60511 Psychotria celiae Steyerm. 
41909 Ms brevipedic ae ne & 65567 Baccharis schomburkkii subsp. occiden- 
rm. talis Steyerm. & Maguire 
41996 Payo Noia pallidinervia Steyerm 65587 Chionolaena breweri Steyerm. & Ma- 
42025 Amaioua gulanensis var. ans guire 
Steyerm. 65593 Tapeinostemon breweri Steyerm. & 
42031 Psychotria venezuelensis Steyer Maguire 
42062 Dendropanax neblinae Maguire, "Stey- 65616 Mikania marahuacensis Steyerm. & 
erm. & Frodin aguire 
42068 Guapira neblinensis Maguire & Stey- 65641 ME marahuacae L. B. Smith, 
erm. Steyerm. & H. Robinson 
42072 Neea neblinensis Maguire & Steyerm. 65705 Navia pues L. B. Smith, Steyerm. & 
42084 Kotchubaea neblinensis Steyerm. H. Robinson 
42096 Celiantha bella Maguire & Steyerm 
ule f - MANARA, B. (Bruno) 
42120 Neblinathamnus argyreus Steyerm. x . 
42134 MM longiflorum Maguire & "n Paullinia manarae Steyerm. 
Steyerm. s.n. Peltastes manarae Steyerm. 
42226 Weinmannia neblinensis Maguire & sn Peperomia manarae Steyerm 
Steyerm: s Piper pubivaginatum Steyerm 
42266 Psychotria everardii subsp. neblinensis 51 wa manarae Steyerm. 
Steyerm. MARCANO BERTI, L. (Luis) 
42319 Dioscorea abysmophila Maguire & 54-979 a elabriflora Steyerm. 
49350 0 ie en lackii Maguire. S 119-979 Talisia Sancarlosiana Steyerm. 
m picis SURE. Magure, Steyerm, 209 Guapira marcano-bertii Steyerm. 
uteyn . . 
42378 Symplocos neblinae Maguire & Stey- as anis nie i a 
f PIM: Sloanea subpsilocarpa Steyerm. 
42379 Schefflera tremuloidae Maguire, Stey- 519-979 Hillia marc ae berri Steyerm. 
49449 Ps i a el TN 532-979 Rudgea marcano-bertu m 
- is ciantia teyerm, 551 Guapira amacurensis Steyern 
42407 sinit globosum Steyerm. 
42469 Remijia maguirei Steyerm. MATUDA, E. (Eizi) 
42415 Psychotria wurdackii aai 1741 Galactia acuminata Steyerm. 
42493 Ladenbergia puberula Steyer 2260 onki standleyi Steyerm. 
42496 Rhamnus acuminata e. & Stey- 5330 Schismocarpus cie Steyerm. 
erm. = 
42496 Rhamnus nu mista Maguire & MEME . T mE 
Steyerm:. ined: Schefflera rugosifolia Maguire, Steyerm. 
42515 Faramea neblinae Sum & Frodin 
42597 Platycarpum maguirei Steye MEDINA, E. (Ernesto) 
44113 Palicourea rigida f. nen Stey- 34 Ouratea medinae Maguire & Steyerm. 
erm. 
46152a Remijia roraimae var. guianensis Stey- MELLO BARRETO, H. L. 
erm. 226 Mendoncia mello-barretoana Steyerm. 
46814 Schradera ao ulata Steyerm. E 
46864 Cephaelis tepuier didus m. pou LED . o ; 
46872 Psychotria aligera ES e: 744 Grindelia arizonica var. dentata Stey- 
46876 Sterigmapetalum anes ion erm 
eyerm. Jesner MEXIA, Y. (Ynez) 
93562 Cephaélis sandwithiana Steyerm 8783 Salpianthus standleyi Steyerm. 
53608 roy tite cannoides Nic dor Stey- 8855 Paullinia mexiae Steyerm. 
rm. & Sivadasan . 
53676 Duroia strigosa Steyerm MOORE, G. E. (George) 
52731 Maytenus longistipitata cip s.n. Anemone virginiana f. plena Palmer & 
53864 Macrocentrum steyermarkii Wurdack " o. : : T 
54440 P sye horria ofie inalis Mri vilhelm: s.n. Cirsium altissimum f. moorei Steyerm. 
e sis Steyerm. * MOORE, J. A. 
6007 1 a maguirei Steyerm. 3488 Haplopappus gymnocephalus f. albus 
60073 Psychotria adderleyi Steyerm. eyerm. aen 
60181 Scleronema neblinense Steyerm. 3515 Polveals nous sola oct 
60295 Amphidasya neblinae Steyerm 3528 Valeriana texana Steye 
60477 Befaria ee Maguire, Speen. 3547 Laphamia llos pw erm. 


3607 Grindelia havardii Steyern 


758 


Annals of the 
Missouri Botanical Garden 


3686 Grindelia blakei Steyerm. 

3687 Grindelia stricta var. procumbens Stey- 
erm. 

3768 Grindelia squarrosa f. depressa Stey- 
erm. 

3784 Grindelia revoluta Steyerm. 

3795 Cladothrix lanuginosa var. carnosa 
Steyerm. 

MORALES, R. J. 
1106 die E f. pubescens 
ndl. & Steye 


MORI, S. (Scott) 
5027 


Talisia glandulifera Steyerm. 


MORILLO, G. (Gilberto) 
3339 Talisia morilloi sus aie 
3355 inorea oraria Steyerm. & Fernandez 
3903 Sabicea morillorum eus 
4070 abicea grandifolia Steyerm 
4136 Piper papilliferum Steyerm 
4192 emijia morilloi Steyerm 
5510 Piper morilloi Steyerr 
6716 mea morilloi St 
6811 


10102 Cynanchum julianii Morillo 


MORITZ, J. W. K. (Johann Wilhelm Karl) 


1065 Epidendrum steyermarkii A. D. Hawkes 
MUENSCHER, W. C 
12360 


Solanum muenscheri Standl. & Stey- 
erm. 
MUNZ, P. A. (Philip Alexander) 

11157 Grindelia camporum var. australis Stey- 
MURCA PIRES, J. (or J. M. Pires) 

22 


Psychotria iodotricha subsp. multiflora 
Steyerm. 


6063 ia jene Steyerm. 

15010(33) ea piresi . Smith, Stey: 
r ute 

15014(37) Nave y piresu L. B. Smith, Steyerm. & 
H. Robinson 

50285 Coussarea amapaensis Steyerm. 

50386 Palicourea EEA Steyerm. 

50632 Ixora piresii Ste 

51210 Ixora amapaenis oun. 

51371 Psychotria araguariensis Steyerm. 

51601 Duroia amapana Steyerm 

92250 Psychotria piresii Steyerm. 

52451 Genipa spruceana Steyerm. 


NASCIMENTO, O. €. 

2 Psychotria formosissima Steyerm. 
NEE, M. (Mike) 

30967 Remijia sessilis Steyerm. 
NELSON, E. W. (Edward William) 

3 Grindelia sublanuginosa Steyerm. 

6864 Grindelia nelsonii Steyerm. 
NELSON, J. C. 


Grindelia integrifolia f. dentata Steyerm. 


OLDEMAN 
2 Chomelia glabriuscula Steyerm. 
OLDERMAN, R. A. A 
B-627 


Rudgea oldemanii Steyerm. 


OLIVA ESTEVA, F. (Francisco) 


s.n. Brocchinia oliva-estevae Steyerm. & L. 
B. Smith 
ORCUTT, C. R. (Charles Russell) 
120 Grindelia greenei Steyerm. 
OWNBEY 
2739 Joosia macrocalyx Standl. ex Steyerm. 


PALMER, E. (Edward) 


163 Grindelia palmeri Steyern 

316 Grindelia oxylepis f. eligulata Steyerm. 

429 Menodora decemfida var. longifolia 
eyerm. 

409 Grindelia microcephala var. pusilla 
Steyerm. 

471 Grindelia greenmanil Steyern 

520 Grindelia oxylepis var. las Stey- 


PALMER, E. J. (Ernest Jesse) 


16342 Quercus Xinaequalis Palmer & Stey- 
erm. 

18928 Sambucus canadensis f. rubra Palmer 

Steyerm. 
21923 id aestivale var. pubescens Palm- 
r & Steyerm. 

25421 De cus Xvaga Palmer & Steyerm. 

26069 Quercus X mutabilis Palmer & Steyerm. 

31083 Kee scabra var. ramosissima 

rm. 
33590 Gia microcephala var. adenodonta 
gustior Steyerm. 

35387 ei lutea f. tomentosa Steyerm 

39039 me setigera f. inermis Palmer & Stey- 

40317 Amelanchier ANE f. nuda Palmer 
& S 

41450 Nodes alternifolia v var. pubescens 
Palmer & Steyer 

41642 Rosa setigera f. serena a Palmer & Stey- 
erm. 

55411 Phlox divaricata var. laphamii f. candi- 
a Palmer & Steyerm. 

59655 Rubus flagellaris f. roseo-plenus Palmer 

m. 
61259 Vernonia bladwinii f. albiflora Steyerm. 
65070 


Phacelia hirsuta f. albiflora Palmer & 
Steyerm 


PARISH, S. B. (Samuel Bonsall) 
9795 


Menodora spinescens var. mohavensis 
Steyerm. 


PARRY, C. C. (Charles Christopher) 
570pp Menodora helianthemoides var. magni- 
flora Steyerm. 
o7lpp 


Menodora bobecihsmoltiss var. humilis 
erm 


Volume 76, Number 3 
1989 


Taylor 759 
Plants Described by 
Julian A. Steyermark 


PEGLER 
950 


PENNELL, F 


1676 


PHELPS, K. 


366 


PHILIPSON 
2072 


Menodora heterophylla var. australis 
Steyerm 


Psychotria fimbriflora Steyerm. 


w. (Francis Whittier) 


Coccocypselum brevipetiolatum Steyerm. 


D. (Kathleen Deery) 


Pagamea reducta Steyerm. 


Joosia umbellifera subsp. macarensis 
Steyerm. 


PINKUS, A. S. 


PIPOLY, J. 
6838 


PITTIER, 
5178 
7911 


9829 
9967 


10948 


11995 
12192 


13498 
14010 


14103 


PLOWMAN, 
13610 


Hiraea tepuiensis Steyerm. 

Calycolpus roraimensis Steyerm. 

Ouratea pseudotatei Maguire & Stey- 
erm 


Tepuianthus aracensis Steyerm. & Ma- 
uire 


H. (Henri Francois) 


€ guianensis var. leiophylla Stey- 


NN microdon var. meridionalis 

Steyerm 

Borreria laev vis var. edentata Steyerr 

Guettarda crispiflora var. il pio 
S 


eyerm 

m alar var. epedunculata 
Steyerm. 

Models venezuelensis Steyern 


Faramea occidentalis var. 


2 m 


meridionalis 


ostanensis Steyer 
Psychotria berteriana ae: p 
teyerm. 
Hippos lasseri Steyerm. 


r. (Timothy) 


Maytenus neblinae Steyerm. 


PRANCE, G. T. (Ghillean) 


59234 


PURPUS, C. 


1318 


5722 
12091 
16201 


Duroia irsutissima AN 
Geophila prancei Stey 
"dpud pranc el Seul 
Sipanea prancei — 
oa prancei Ste 
Raveniopsis aracaénsis $ Kallunki & 
Steye 


yerm. 
Duroia prancei Steyerm. 


A. (Carl Albert) 
Menodora intricata var. purpusii Stey- 
erm. 
Grindelia robinsonii picis 
Aster bimater Standl. 
Tinantia eme f. hos Standl. & 
Stever 


RAMIA, M. (Mauricio) 


5254 


Chomelia ramiae Steyerm. 


REDMOND, P. (Parker) 
s.n. Erythrina poeppigiana f. redmondii 
Steyerm. & Lasser 


ROADHOUSE, F. F 


Grindelia hirsutula var. calva Steyerm. 


ROHL, E. 

103 Chomelia venezuelensis Steyerm. 
ROSA, N. A. 

106 Pagamea acrensis Steyerm. 


RUÍZ-TERÁN, L. E. 


44] Elaeagia ruiz-teranii Steyerm 
1433 Toc d costanensis subsp. pm 
erm 
2980 hy HUS ble pharophora var. pubens 
Steyerm. 
3199 Psyc Dus ruiz-teranii Steyerm 
3746 Palicourea i. ES var. dbi 
Steye 
4703 Rudgea r ruiz-teranii Steyern 
6461 Peperomia acuminata f. a ora a Steyerm. 
10907 Lantana ruiz-teranii López-Palacios & 
Steyerm 
RUNYON 
875 jeee a chimboracensis var. 
exsertus f. albus Steyerm. 
RUSBY, H. H. (Henry Hurd) 
s.n. Psychotria occidentalis Steyerm. 
SAER, J 
4 Chiococca alba subsp. DN var. 
micrantha f. pilosa Stey 
635 Gonzalagunia ciliata S m 
121 Rondeletia larensis Steyer 
SASTRE, C. (Claude) 
9 Sabicea sastre O 
2392 Pac hotria sastrei Stey . 
6463 Psychotria cypellantha Sam 


SCHIPP, W. A. 


Cynanchum stenomeres Standl. & Stey- 
erm 


SCHLIM, L. 
531 


SCHMITT, W. F. 


Faramea pseudospathacea Steyerm. 


s.n. Grindelia arenicola f. trichophora Stey- 
erm 

SCHNEE, L. (Ludwig) 

577 ion schneeana L. B. Smith & Stey- 
rm. 


SCHOMBURGK, R. (Robert Hermann) 
Psychotria cordifolia subsp. perpusilla 
Steyerm. 
Chomelia schomburgkii Steyerm 
Ferdinandusa goudotiana var. eciliata 
Steyerm 


924 
955 


SCHULTES, R. E. (Richard Evens) 
46-357 dps serae truncatum var. angusti- 
folium Steye 


760 


Annals of the 
Missouri Botanical Garden 


8156 — hirticalyx var. glabrior Stey- 


9100 Retiniphyllum ro sii 
13075 Hillia schulte 


14475 ere tas sc du Ul i Ste eyerm. 

15031 Stenopadus bn Cuatr. & Stey- 
erm. 

15056 Tepuianthus colombianus Maguire & 
Steyerm. 

19367 Pagamea coriacea var. acuta Steyerm. 

19929 Platycarpum schultesii Steyerm. ex 
Schultes 

20061 Perama schultesii Steyerm. 


SHAFER, J. A. (John Adolf) 
Psychotria deflexa subsp. cubensis Stey- 
erm. 


SHANTZ, H. L. (Homer LeRoy) 
"T 


591 Grindelia inornata var. angusta Stey- 
erm. 
SHREVE, F. (Forrest) 
8035 Viguiera shrevei Steyerm. 
SILVA N. T. (Nilo) 

l Tokan quadribracteata Steyerm. 
60606 grandis Maguire & Steyerm. 
60607 Pola eddie SONG 
60633 Psychotri ria brazoi Ste 
60678 Mollinedia ee Mice & Stey- 

erm. 
60700 Faramea silvae Steyer 
60719 iode grandipetala Maguire & Stey- 
60842 uq maturacensis Steyerm. 
60865 Sloanea parvifructa Steyerm. 
60872 P tria silvae Steyeri 
60873 Ilex attenuata Steyerm rm. 
60880 Pagameopsis maguirei subsp. neblinensis 
var. pirapucensis Steyerm. 
60880 Psychotria canescens Ste M 
60898 Maguireocharis neblinae Steyer 
60903 ree pees ea subsp. ne us Ma- 
uire, Steyerm. & Frodin 
60907 Een Pn brasiliensis Steyern 
60941 


— plurispicata Maguire, Stey- 
. & Frodin 
SILVERIO LEVEL, J. 


145 Faramea parvibractea Steyerm. 
147 Duroia kotchubaeoides Steyerm. 
SKUTCH, A. F. (Alexander Frank) 
935 Chamaedorea skutchii Standl. & Stey- 
erm. 
982 Urera killipiana Standl. & Ste 
1367 Phoebe padiformis Standl. & Po du 
1453 Clethra skutchii Standl. & Steyerm 
1569 Rondeletia myriantha Standl. & Stey- 
erm. 
1660 Fraxinus cavekiana Standl. & Steyerm. 
1811 Diolena calc aps ue & Steyerm. 
2008 Mabea excelsa S . & Steye 
2073 Psychotria elivorum ime & e ern 
2110 Rondeletia skutchii Standl. & Steyerm. 
SMART 
213 Menodora scabra var. longituba Stey- 
erm. 


SMITH, A. 
F177 Licania costaricensis Standl. & Stey- 
erm. 
SMITH, A. C. (Albert Charles) 
Appunia longiloba Steyer 
2618 Malanea macrophylla f. cuneata Stey- 
rm. 
2192 Duroia eriopila f. glabra Steyerm. 
3066 Malanea sarmentosa f. sparsipilosa 
Steyerm. 
10289 Schradera vahlii var. acutifolia Stey- 
SMITH, C. P. (Charles Piper) 
1016 i ssi encia var. latifolia f. vil- 
sa Ste 


SMITH, H. H. (Herbert ced 
10 Pogonopus speciosus subsp. sandwithi- 
nus Steyerm. 


320 Rondeletia purdiei var. glabrior Stey- 
erm. 

393 Psychotria horizontalis var. psilophylla 
Steyerm. 

2628 Diodia teres var. angustata f. latior 
Steyerm. 

SMITH, REV. L. C. 

135 Grindelia microcephala var. montana 

Steyerm. 


SMITH, R. F. (Robert) 
V-6649 Coussarea terepaimensis Steyerm. 
SPAULDING, REV. 
s.n. Grindelia nana f. longisquama Steyerm. 
SPRUCE, R. (Richard) 
37 Ixora sandwithiana Steyern 
3440 Psychotria 1 


ertiera guianensis var. glabriflora Stey- 
1. 


STANDLEY, P. C. (Paul Carpenter) 


7884 Ipomoea ophipides Standl. & Steyerm. 
12459 Siphonoglossa hondurensis Standl. & 
erm. 

58025 radesciota guatemalensis f. alba 
Standl. & Steyerm. 

58597 Archibaccharis i Standl. & 
Steyerm. 

8674 Aphanac tis standleyi Steyerm 

58732 Clematis caleoides Standl. & Steyerm. 

59219 des nc megistanthus Standl. $ 
Steyer 

59321 Salmea m :ephala Standl. € Stey- 
erm. 

59358 So aguensis Standl. & Stey- 

59934 Diospyros johnstoniana Standl. & Stey- 
er 

60082 Jacobinia umbrosa f. erythrantha 
Standl. & Steyerm. 

00304 es antiquorum Standl. & Stey- 

61100 


IN plethadenium Standl. & 
Steyerm. 


Volume 76, Number 3 Taylor 761 
1989 Plants Described by 
Julian A. Steyermark 
61425 s inornatum Standl. & 15307 Pachecoa guatemalensis Standl. & 
ote te Steyer m. 
61803 Physalis an Eo & Steyerm. 75590 Zexmenia pinetorum Standl. & Stey- 
62109 IE pacifica S Standl. & Steyerm. erm. 
62513 jns e viscosissima Standl. & Ste 75788 np um jejunum Standl. & Stey- 
erm. rm. 
63104 ias pachecoana Standl. & 76007 Borreria vegeta Standl. & Steyerm. 
Steyerm. 16162 Fraxinus vellerea Standl. & Steyerm. 
63209 Zanthoxylum aguilarii Standl. & Stey- 76321 Randia habrophlebia Standl. & Stey 
rm. erm. 
63743 Eupatorium carmonis Standl. & Stey- 76561 Schultesia guianensis f. lutescens 
erm. Standl. & Steyerm. 
63878 Hyperbaena vulcania Standl. & Stey- 18534 ee guatemalense Standl. & 
erm. Steyerm. 
65053 Erigeron aquarius Standl. & Steyerm. 78584 ‘Tvizonia rasa Standl. & Steyer 
65123 Clethra pachecoana Standl Steyerm 79287 Ipomoea santae-rosae Standl. & Bu 
65221 Mahonia volcania Standl. $ Me erm. 
6547 Buddleja amplexicaulis Standl. & Stey 80408 Fraxinus bicolor Standl. € Steyerm. 
80900 Psychotria aguilarii Standl. & SER 
65798 Eupatorium huehuetecum Standl. & 80907 Euphorbia guatemalensis Standl. & 
Steyerm. Steyerm. 
65862 Eupatorium ovillum Standl. & Steyerm. 81097 Gnaphalium standleyi Steyerm. 
67195 Lobelia laxiflora f. lutea Standl. & Stey- 81113 Gentiana guatemalensis Standl. & Stey- 
erm. 
221 Eupatorium rojasianum Standl. & Stey- 81651 Garrya corvorum Standl. & Steyerm. 
rm. 82039 Agave huehueteca Standl. & Steyerm. 
67473 Tm fontium Standl. & Steyerm. 82330 Cimacorachi guatemala Standl. & 
07488 Physalis calidaria Standl. & Steyer Ste 
67540 Verbesina apleura var. foliolata Standl. 82430 SM M brachybotrys Standl. & 
& Steyerm. Steyerm. 
67887 Clerodendron pithecobium Standl. & 82538 Trixis amphimalaca Standl. & Steyerm. 
Steyerm. 82597 ea huehuetecum Standl. $ 
68152 Conostegia volcanalis Standl. & Stey- Steyern 
erm. 82871 Vernonia mima Standl. & Steyerm. 
68931 B dd quetzalteca Standl. & 82921 Verbesina petzalensis Standl. & Stey 
Steyer erm. 
69568 Eugenia ll Standl. & y un 83332 Cardamine balnearia Standl. & Stey- 
69756 So A num verapazense Standl. & Stey erm. 
83528 Abutilon pachecoanum Standl. & Stey- 
69922 Pons pachecoana Standl. & Stey- erm. 
erm. 84001 ho cupressorum Standl. & 
70009 Phoebe macrophylla Standl. & Stey- teyerm. 
erm. 84404 Eupatorium vetularum Standl. & Stey- 
70317 rici es gleasoniana Standl. & Stey- rm. 
84556 o lewisiae Standl. & dod 
11125 Eupatorium hastiferum Standl. & Stey- 85146 Eupatorium mimicum Standl. & Stey 
rm. erm. 
71421 ea verapazensis Standl. & oM 85228 Tagetes sororia Standl. & Steyer 
72891 RE. izabalensis Standl. & Ste 85603 Bonifazia quezalteca Standl. & Seyerm 
erm 86226 Bouvardia pachecoana Standl. & Ste 
72793 Sburaubés micrantha Standl. & Stey- erm. 
erm. 86270 Rubus hadrocarpus Standl. & Steyerm. 
73714 Aeschynomene tricholoma Standl. & 86890 Chamaedorea aguilariana Standl. 
Steyerm. Steyerm. 
73715 Serjania lobulata Standl. & Steyerm. 87159 Chamaedorea quezalteca Standl. & 
73841 Mirabilis pulchella Standl. & Steyerm. Steyerm. 
74367 ee saxosum Standl. & Stey- 87473 Vernonia medialis Standl. & Steyerm. 
rm. 87781 Pedilanthus camporum Standl. & Stey 
74420 Mulas orientale Standl. & Steyerm. erm. 
74456 Ese is ckia echinoidea Standl. & Stey- 89560 Aphelandra gigantiflora f. lutea Standl. 
& Steyerm. 
14569 Lonchocarpus phlebophyllus Standl. & 90160 Chamaedorea carchensis Standl. & 
Steyer Steyerm. 
74623 Corallocarpus guatemalensis Standl. & 90328 Piper frioense Standl. & Steyerm. 
Steyerm. 90532 ir dl macrantha Standl. & 
74673 Euphorbia steyermarkii Standl. Stey 
75245 Bursera permollis Standl. & Steyerm. 90524 Rhee dia p Standl. & Steyerm. 


762 


Annals of the 
Missouri Botanical Garden 


90762 
91207 


Cecropia sylvicola Standl. & Steyerm. 
Stillingia cruenta Standl. & Steyerm. 
STEIN, B. A. (Bruce) 
Psychotria steinii Steyerm. 
STERGIOS, B. (Basil) 
2804 Chomelia stergiosii Steyerm. 
8611 Neea cedensis Steyerm. 
STEYERMARK, J. A. & G. AGOSTINI 
Psychotria agostinii Steyer 
Le Schradera glabriflora Sean 
STEYERMARK, J. A. & L. ARISTEGUIETA 
Calycophyllum Mage iden Steyerm. 
5 Malanea hypoleuca Ste 
! uu. stey ermarkii Maguire 


c 
3 


25 Platy um rugosum Sin 
59 Pad insigne Steyer 
102 Elaeagia maguirei var. s Siyen: 


STEYERMARK, J. A. & S. NILSSON 


Columnea steyermarkii 1n 


135 Psychotria parvibractea Steyerm. 

Ld Piper vitaceum var. DE Stey- 

178 Solanum rufistellatum Steyerm. 

185 Solanum puberuloba Steyerm 

196 Schradera nilssonii St 

237 Selenipedium steyermarkii Foldats 

382 Duroia retrorsipila St 

385 Malanea ueiensis Steyer 

392 Hillia psammophila Steyerm 

426 Ladenbergia venamoensis Steyerm 

428 Guzmania steyermarkii L. B. Smith 

429 Schefflera quinquestylorum Steyerm. 

449 Ilex wurdackiana Steyerm. 

450 Chorisepalum acuminatum Steyerm. 

473 Phoradendron aphyllum Steyerm. 

483 Schefflera quinquecarinata Steyerm. 

561 dicm dichotoma var. hirsutula Stey- 
er 

732 Lisianthius acutilobus Steyer 

767 mmeorrhiza umbellata Bi septen- 
trionalis Steyerm. 

768 Pouteria end Steyerm. 


STEYERMARK, J. A. & J. J. WURDACK 
Talisia tiricensis Steyerm. & Maguire 


330 Rudgea phaneroneura Steyerm. 
347 Ilex brevipetiolata Steyerm. & Wurd. 
374a c ee chimantensis Steyerm. & 
Mag 
377 Adenamthe eee Maguire, Stey- 
. & Wurd 
387 epus tatei var. witinanténs Steyerm. 
agui 
426 Tetrapterys alos Maguire 
Steyerm. 
458 Notopora smithiana Steyerm. & Ma- 
guire 
$11 rir ou similis Maguire, Steyerm. 
574 CRIAM espeletoidea Maguire, Stey- 
erm. & uu 
633 Bonnetia huber 


640 P. os e” 


652 Stenopadus chimantensis Maguire, Stey- 
erm. & Wurd. 
669 Vaccinium chimantense Maguire, Stey- 
erm. uteyn 
675 Quelchia eriocaulis Maguire, Steyerm. 
& Wurd. 
680 Bonnetia chimantensis Steyer 
681 ee virgatus Mu. Stey- 
. & Wurd. 
683 Quelchia bracteata Maguire, Steyerm. 
& Wurd. 
713 oe Darii Steyerm. 
737 Ledot nisi ns Maguire, 
Steye ^ m 
747 a humilis Marcos; Steyerm. 
Wurd. 
748 Chimantaea rupicola Maguire, Steyerm. 
& Wurd. 
756 lees x grandifolia Maguire, Stey- 
. & Wurd. 
781 i dut steyermarkii Nevling 
819 Myriocladus steyermarkii Swallen 
821 Chimantaea mirabilis Maguire, Steyerm. 
Wurd. 
825 Mycerinus chimantensis Maguire, Stey- 
erm. uteyn 
826 Notopora chimantensis Steyerm. & Ma- 
guire 
827 Lindmania steyermarkii L. B. Smith 
851 Verbesina angusta Maguire, Steyerm. 
& Wurd. 
861 Macrocarpaea rugosa Steyerm. 
879 gp ced fruticosa var. chimantae Stey- 
921 Phoebe steyermarkiana C. K. Allen 
921 Ocotea julianii van der Werff 
953 Ledothamnus luteus Maguire, Steyerm. 
& Luteyn 
954 Panicum steyermarkii Swaller 
990 Bonnetia tepuiensis subsp. minor Stey- 
erm. 
1020 Pagameopsis maguirei Steyerm 
1028 Styrax tepuiense Steyerm. & Maguire 
1036 Panopsis ornatinervia Steyerm. 
1047 Bonnetia toronoensis uii ius 
1059 Pterozonium steyermarkii schi 
1096 Xyris Pa d Meri & rnt 
1100 Symplocos chimantensis Steyerm. & 
aguir 
1103 Euplassa ies Steyerm. 
1104 Neea tepuiensis Steyerm. 
1105 Roupala chimantensis Steyerm. 
1133 Cupania tepuiensis Steyerm. € Maguire 
1136 Gongylolepis fruticosa Maguire, Stey- 
erm. & Wurd. 
1172 Palicourea perquadrangularis var. gua- 
yanensis Steyerm. 
1197 Piper politii subsp. toronoense Steyerm. 
1225 Cladium steyermarkii T. Koyama 
1227 Tapeinostemon rugosum Maguire & 
Steyerm. 
1340 Sabicea velutina subsp. chimantensis 
Steyerm. 
1346 Piper tepuiense Ste 
1370 


Passiflora bomareifolia 'Steyerm. & Ma- 
uire 


Volume 76, Number 3 
1989 


Taylor 


y 
Plants Described by 


763 


Julian A. Steyermark 


STEYERMARK, J. A. (Julian Alfred) 


11345 


11390 
11461 


12318 


12414 


1249] 


12548 


18628 
20910 


22499 


25927 


29313 


29490 
29505 


29640 


29906 


30004. 


30066 


30068 


30448 


Houstonia pusilla f. rosea Steyerm. 

Hypericum pseudomaeulatun f. flavi- 
dum Steyerm. 

Psoralea tenuiflora f. alba Steyerm. 

Ruellia caroliniensis f. alba Steyerm. 

Rhododendron roseum f. albidum Stey- 
erm. 

Taxodium distichum f. confusum Palm- 
er & Steyerm. 

Rudbeckia hirta f. homochroma Stey- 
erm. 

Apios americana f. pilosa Steyerm. 

Carya = Mu var. i iae f. glabra 
Palm 


Quercus facta E Tae Palmer & 
C oce bulbosa f. fontinalis Palmer 
mE cordata f. mollis Palmer & Stey- 


Salix s sericea f. glabra Palmer & Stey- 
erm. 

Trillium ozarkanum Palmer & Steyerm. 

Salix cordata f. subintegra Palmer 


Steyerm. 
= scilloides f. petersenii Stey- 


Lrapatiens pallida f. dichroma Steyerm. 
a Steyerm. 


Camassia scilloides f. variegata Stey 


Ec mS pallida f. albida Steyerm. 

Mimosa zacapana Standl. & Steyerm. 

— erecta var. RSEN UMS f. 
cana Standl. & Ste 

Commelina erecta f. PAS Standl. & 

Steyerm. 

donne eriocarpa Standl. & 
Steyerm. 

Diospyros zacapana Standl. & Steyerm. 

Verbesina guatemalensis var. glabrata 
Standl. & Steyerm. 

Galactia minarum Standl. & Steyerm. 

a n. 


adl. ye 
Hieracium minarum Standl. & Steyerm. 
Perymenium inamoenum Standl. & 
yerm. 
Mice p disgrega f. glandulosa 
andl. & Steyerm. 
Ti eee cacuminis Standl. & Stey- 
erm. 
Rondeletia cordovana Standl. & Stey- 
erm. 
Athenaea macrocardia Standl. & Stey- 
erm. 
dee guatemalensis Standl. & 
teyer 


E stey ermarkit Standl. 
Ipomoea saxorum Standl. & Steyerm. 
Pectis flava Standl. & Steyerm 
Hydrocleis standleyi Steyerm. 
Crumenaria stey nas! Standl. 


Machaerium fruticetorum Standl. & 


Steyerm. 
Verbesina dd Standl. 
Stevia chortiana Standl. & Steyerm. 


oc cacuminum Standl. & Stey- 


Bo amaranthina Standl. & Stey- 
erm. 

Clusia lusoria Standl. & Steyerm. 

Saurauia veneficorum Standl. & Stey- 
erm. 

Ageratum chortianum Standl. & Stey- 
erm. 

Eugenia sasoana Standl. & Steyerm. 

Cosmos steyermarkii Sher 

Chiococca semipilosa Standl. & Stey- 
erm. 

Zanthoxylum nubium Standl. & Stey- 
erm. 
Citharexylum steyermarkii Mold. 

Eugenia chiquimulana Standl. & Stey- 


Sapo culminicola Standl. & Stey- 
erm. 

Eupatorium macram Standl. & Stey- 
erm. 


Lycianthes a im var. sparsistellata 
Standl. & Steyerm. 

Miconia tixixensis Standl. & Steyerm. 

Viburnum mortonianum a e & Stey 
erm. 

Tradescantia standleyi Steyerm. 

Daphnopsis retifera Standl. & Steyerm. 

Eupatorium salinum Standl. & Steyerm. 
sclepias woodsoniana Standl. & Stey- 


erm. 
Phoradendron aguilarii Standl. & Stey- 


erm. 

Bouvardia nubigena Standl. & Steyerm. 

Orthrosanthus UM var. cen- 
tro-americanus Steye 

Schaetzellia standleyi EU 

ene guatemalensis Standl. & Stey- 


Pare jalapanum Standl. & Stey- 
erm. 

Hieracium jalapense Standl. & Stey- 
erm. 

Echeveria macrantha Standl. & Stey- 
erm. 

Cacalia guatemalensis Standl. & Stey- 
erm. 

Physalis lassa Standl. & Steyerm. 

Rhynchosia jalapensis Standl. & Stey- 
erm. 

Arenaria altorum Standl. & Steyerm. 

Clethra licanioides Standl. & Steyerm. 

Canavalia munda Standl. & Steyerm. 

Rondeletia vulcanicola Standl. & Stey- 


erm. 

Philodendron rojasianum Standl. & 
Steyerm. 

Roupala steyermarkii Sleumer 

Erblin 08 os arpa var. mollis Standl. 


"id hia : ste ever Standl. 


764 Annals of the 
Missouri Botanical Garden 
33479 Chamaedorea rojasiana Standl. & Stey- 36690 Hieracium ostreophyllum Standl. & 
erm. Steyerm. 
33501 Gymnanthes guatemalensis Standl. & 36787 Vernonia corae Standl. & Steyerm. 
eyerm. 36801 Styrax vulcanicola Standl. & Steyerm. 
33533 Pasudocalomme dre SIM 36931 San emarginata Standl. & Stey- 
33005 anda standle 
33015 arsdenia cd “Standl. & Stey- 37004 C alea dics Steyerm. 
erm. 37061 Persea steyermarkii C. K. Allen 
33067 Hillia macrocarpa Standl. & Steyerm 37187 Campelia standleyi St m 
33743 Cyphomandra rojasiana Standl. & Stey- 37276 Buddleja astralis a & Steyerm. 
erm. 31381 hamaedorea stricta Standl. & Steyerm. 
33858 Sideroxylon steyermarkii Standl. 31440 Fitiecciobiun bue pn Standl. & 
34047 :lanthera steyermarkii Standl. Steyerm. 
34273 Sia ydium glabrum Standl. & Steyerm 37583 Ficus cabusana Standl. & Steyerm. 
34302 sychotria altorum Stan Steyerm. 37661 Gonolobus steyermarkii Woodson 
34304 accen guatemalensis Standl. & 37735 Chamaedorea vulgata Standl. & Stey- 
Steye erm. 
34590 Mu POR Standl. & Stey- 37738 Rondeletia zolleriana Standl. & Stey- 
erm. erm. 
34609 Centaurium rosans Standl. & jdn 37759 Cobaea steyermarkii Standl. 
34744 Eupatorium zunilanum Standl. & Stey- 37766 Physalis carnosa Standl. & Steyerm 
erm. 37945 Calathea coccinea Standl Steyerm 
34757 Stevia vulcanicola Standl. & Steyerm. 37953 Athenaea locuples Standl. & Steyerm 
34760 Hypericum steyermarkii Stanc 37963 Anthurium titanium Standl. & Steyerm 
34772 Lysimachia steyermarkii Star idi; 37974 Deppea inaequalis Standl. & Steyerm. 
34815 Hieracium culmenicola Standl. & Stey- 37982 Russelia laciniata ig ik & Steyerm. 
erm. 38123 Chamaedorea fusca Standl. & Steyerm. 
34860 Hieracium guatemalense Standl. & 38170 Vm maxima i Stand & Steyerm. 
, Steyerm. 3827 Eugenia simiarum Standl. & Steyerm. 
34933 acalia pinetorum Standl. & Steyerm. 38526 Lisianthius elatus Standl. & Steyerm. 
35001 mec stenoptera Standl. & Steyerm. 38533 Jacquemontia pinetorum Standl. & 
35020 Odontonema steyermarkii Leonard steyerm. 
35489 Gentiana pumilio Standl. & ia rm. 38658 Calea savannarum Standl. & Steyerm. 
35810 Matelea steyermarkii Woodso 38839 Rudgea simiarum Standl. & Steyerm. 
35835 VOR superba Standl. & Ph 39165 Eugenia musarum Standl. & Steyerm. 
36019 rsdenia steyermarkii Woodson 39185 Chamaedorea brachypoda Standl. & 
36044 i dpi steyermarkii Standl. Steyerm. 
3606; vysalis campanula Standl. € pa 39288 Pancratium guatemalense Standl. & 
36075 nem saxorum Standl. & Ste Steyerm. 
erm. 39238 Lonchocarpus modestus Standl. & Stey- 
36110 Pernettia saxicola Standl. & Steyerm. erm. 
36128 Hieracium tacanense Standl. & Stey- 39375 Chione guatemalensis Standl. & Stey- 
erm. erm. 
36137 Juniperus standleyi Steyerm. 39382 Duroia genipifolia Standl. & Steyerm. 
3617 Physalis amica Standl. & Steyerm. 39387 Capparis steyermarkii Standl. 
36181 Polymnia standleyi Steyerm. 39446 Mouriria steyermarkii Standl. 
36207 Persea vesticula Standl. & Steyerm. 39448 —— guatemalensis Standl. & Stey- 
36210 Eugenia steyermarkii Standl. rm. 
36237 Chiococca stey ermarkii veal 39452 ace ied mollis Standl. & Steyerm. 
36258 Thalictrum standleyi Ste 39454 Hyperbaena ste dotium Stanc 
36268 iin PASE Stand. & Stey- 39456 Mascagnia excelsa .& Bien 
39516 dencia oai Standl. & 
36286 Rondeletia brachistantha Standl. & Steyer 
Steyerm. 39533 dos "seeds Scan 
3629] Hélico cereus heterodoxus Standl. & 39611 Crudia lacus Standl. & Steyerm. 
Steyerm. 39874 Dichapetalum llei Standl. & Stey- 
30341 Fasdtoriu mendax Standl. & Stey- erm. 
erm. 39923 Sem atophyllum steyermarkii pud, 
36381 llex anodonta Standl. & Steyerm 41543 esmopsis izabalensis Standl. & S ey- 
36386 ies psum medialis Standl. & Stey- erm. 
41748 Saurauia persifolia Standl. & Steyerm. 
36426 Hoffmannia steyermarkii Steyerm. 41819 Diolena rosiflora Standl. & Steyerm. 
36543 Liabum tajumulcense Standl. & Stey 41864 Rondeletia macrocalyx Standl. & Stey- 
erm. erm. 
36575 Thalictrum steyermarkii Standl. 41893 Chamaedorea stenocarpa Standl. & 
36576 Fissidens steyermarkii Bartram Steyerm 


Volume 76, Number 3 


Ta 


ylor 
Plants Described by 


765 


Julian A. Steyermark 


41910 


41923 
41928 


41952 


42212 


42600 


42649 


42980 
42985 
43106 


45686 
45710 


45712 


45738 


45827 


Hoffmannia racemifera Standl. & Stey- 
erm. 


Ardisia apoda Standl. & Ste 
Piper philodendroides Standl. e Dar 


erm. 


Palicourea phanerandra Standl. & Stey- 


erm. 


m coibionii Standl. & Steyerm. 
Per 


ea sessilis Standl. & Steyerm. 
Verbesina minarum Standl. a oi 
a steyermarkii 
Hoffnannia quadrifolia ‘Standl. P Stey- 
erm. 
Oreopanax steyermarkii A. C. Smith 
Piper minarum Standl. & Steverm. 
Simsia steyermarkii Robinson & Bret- 
tell 
Russelia steyermarkii Carlson 
Calliandra carcerea Standl. & Steyerm. 
Psychotria minarum Standl. & Stey- 
erm. 
Echeveria steyermarkii Standl. 
Microtropis ilicina Standl. & Steyerm. 
Mikania concinna Standl. & Pur ade 
Vaccinium minarum Standl. & Stey 


erm. 
Phyllanthus minarum Standl. € Stey- 


erm. 
Abutilon tridens Standl. & Steyerm. 
Eupatorium montigenum Standl. & 
teyerm. 
Chamaedorea nubicum Standl. & Stey- 
erm. 
Piper punctulatum Standl. & Steyerm. 


Buddleja euryphylla Standl. & om 
lay" 


Peperomia minarum Standl. & S 
erm. 

Mimosa canahuensis Standl. & Stey- 
erm. 

Hoffmannia tuerckheimii var. glabra 
Stanc p 4 e 


Galactia rsiflora Standl. & m 


Pilea klipa Standl. 

Anthurium radicosum Standl. 8 Stéy. 
erm. 

Piper ixocubvainense Standl. & Stey- 
erin. 


Eugenia ne eds Standl. & Steyerm. 


Sloanea petenensis & Steyerm. 


e dl. $ 
iro zrandifolia $ Sod & Stey 
ern 


5 


Cam calc er^ Standl. " Steyerm. 
ea crassifolia Standl. & Steyerm. 
Rondeletia oe Standl. & Stey- 

erm 


Oyedaea steyermarkii S. F. Blake 

Eugenia savannarum Standl. & Stey- 
erm. 

Phoebe savannarum Standl. & Stey- 
erm. 

Peperomia clavigera Standl. & Stey- 
erm. 

Trichilia privigna Standl. & Steyerm. 


46355 
46398 
40024 
46653 


46696 


48643 


486062 


49055 
49104 


49189 
49270 


49410 


19810 


49855 


Guarea chirriactensis Standl. & Stey- 
erm. 

Peperomia treleasei Standl. & Steyerm. 

Piper santae-clarae Standl. & Steyerm. 

Lycianthes barbatula Standl. & Stey- 


Piper lane iferum Standl. & o 

Meliosma divers Standl. & Steyerm. 

Souroubea tó Standl. & Steyerm, 
ikania petrina Standl. & Steyert 

m ue vulcanicola Standl. & € 


Cacala pudica Standl. € Steyerm. 
x tunimanensis Standl. & Steyerm. 
P juniperorum Standl. & Stey 
erm. 
Viburnum euryphyllum Standl. & Stey- 
erm. 


Symplocos vatteri Standl. & pu d 

Symplocos abietorum Standl. & Ste 
erm. 

Monochaetum guatemalense Standl. & 
Steyerm 

Carex ste yerinarhi Standl. 

Carex venosivaginata Standl. & Stey- 
erm. 

Ardisia vatteri Standl. & Steyerm. 

Psychotria lilacina Standl. & Steyerm. 

cun haematinum Standl. & Ste 


Pao huitzensis Standl. & Stey- 
erm. 
pem retiferum Standl. & Stey- 


Clerodendron mimicum Standl. & Stey- 


erm. 
Solenophora abietorum Standl. & Stey- 
erm. 
Mikania huitzensis Standl. & om 
Spigelia carnosa Standl. & Stey 
Diolena stenophylla Standl. & Steyerm. 
Coussarea mediocris Standl. & Ste 


Pelami simiarum Standl. & Stey- 
erm. 

Hidalgoa steyermarkii Sherft 

Miconia echinoidea Standl. & Steyerm. 

Aegiphila hoffmannioides Standl. & 
Steyerm. 

Justicia steyermarkii Standl. & Stey- 
erm. 

Carex huehueteca Standl. & Steyerm. 

Daphnopsis malacophylla Standl. & 
Steyerm. 

Ocotea laetevirens Standl. & Steyerm. 

Mendoncia guatemalensis Standl. & 
Steyerm: 

Peperomia dorstenioides Standl. & 
Steyerm. 

Ruyschia longistylis Standl. & Steyerm. 

Senecio steyermarkii Greenman 

Piper pinetorum Standl. & o 

Saurauia cuchumatanensis Standl. 
Steyer 

Vibucnum a Standl. & Stey- 
rm. 


766 


Annals of the 


Missouri Botanical Garden 


49859 
49908 
50061 
50065 
50114 
50132 
50143 
50144 
50150 
50157 
50160 
50243 


50258 


50501 
50569 
50672 


50739 
50818 


50876 
50908 


50934 


50936 


51340 


cn 


51398 
51498 


51646 


51666 


51672 
51683 


51697 


52104 


53576 


Rubus hadrocarpus f. adenophorus 
Standl. & Steyerm 

Castilleja altorum Standl. & Steyerm. 

Pinguicula clivorum Standl. & Steyerm. 

Hieracium clivorum Standl. & Steyerm. 

Siegesbeckia nudicaulis Standl. & Stey- 
erm. 

Verbesina calciphila Standl. & adie 

Cardamine eremita Standl. & S i dun 

Romanschulzia alpina Standl. & Stey 
erm. 
arex i arr Suns & Steyerm. 

Cirsturü nigriceps Standl. & Steyerm. 

Hypericum "dn Standl. & Stey- 
erm. 

Drymaria minuscula Standl. & Stey- 
erm. 

Oxalis calcicola Standl. & Steyer 

sl NE alpestris Standl. & Sic rain 
auschia steyermarkii Math. & Const. 

Solanum ña Standl. & Steyerm. 

Mor calcicola Standl. & Steyerm. 

Dahlia « coccinea var. steyermarkii 


ier 
Selaginella stey d Alsto 

Cas damine jejuna Standl. & Sis erm. 

St cubano pep Standl. € Ste 


Euphorbia senilis Standl. & Steyerm. 

Eucnide guatemalensis Standl. & Stey- 
erm. 

Xylosma trinervium Standl. & Steyerm. 

Serjania phaseoloides Standl. & Stey- 
erm. 

Echeveria huehueteca Standl. & Stey- 


erm. 

Valeriana delicata Standl. & Steyerm. 

Eugenia papalensis Standl. & Steyerm. 

Bidens steyermarkii Sherff 

Smilacina crassipes Standl. & Steyerm. 
iuehueteca Standl. & Steyerm. 

Solanum Tuus eu Standl. & Stey- 

erm. 

Juglans steyermarkii Mannin 

Aristolochia mollis Standl. & Steyerm. 

SUAE heterodoxa Standl. & Stey- 


Gphellantha steyermarkii Stand 


ee crocodilium Stand! & Stey- 


Strutt tenuifolius Standl. & Stey- 


Euphorbia cuchumatanensis Standl. & 
Steyerm 
Claigaedubuá aequalis Standl. & Stey- 
s 
is steyermarkii Correll 
eu icd digitata Standl. & Stey- 
erm. 
Hoffmannia huehueteca Standl. & Stey- 
erm. 
Lycianthes orogenes Standl. & Stey- 
erm. 
dida ec er i 2: erm. 
ias Steye 
e Steyerm. 
Brunellia z zamorensis Steyerm. 


53709 


55339a 


n 
ene 
oO 
— 


55980 
61 


56242 


56251 


56333 


56344 


Renealmia stellulata Steyerm. 


pere aequatoria Steyerm- 
Steyerm. 
Tapeinostemon zamoranum Steyerm. 


Costus zamoranus Steyerm. 
Roupala obovata var. ecuadorensis 
Ste) 


Stellaria ovata f. par cado E 


Cephaélis tinctoria Standl. & Steyerm. 
Tofieldia sessiliflora var. robustior Stey- 
erm. 


Conomorpha Teram midides e 
isis va 


Hypericum pseudocaracasanum Stey- 
erm. 
Palicourea chimo Standl. & Steyerm. 
Sabazia urticifolia var. venezuelensis 
eyerm. 
TA adpressipilis Steyerm. 
Cono a larensis Steyerm. 
nima idis venezuelensis Standl. & 


anara ane Steye 
entregan i ied Steyerm, 
zarex larensis Stey 
a cti » var. lanuginosa 


dece nd ae & Steyerm. 
Cajaphora larensis Steyerm. 
ernardia venezuelana Steyerm. 
r 


Euphorbia latazi var. glabra Steyerm. 

Guettarda steyermarkii f. latior Stan 
& Steyerm. 

Conomorpha jajiensis Steyerm. 
ypericum pseudomaculatum f. flavi- 
dum Steyerm. 

Heliconia sehneeana Steyerm. 

Croton croizatii Steyerm. 

Lepidopilum steyermarkii Bartram 

Fuchsia meridensis shee hate 

Croton chamanus Steye 

Oreopanax capitatus var. minor Stey- 
erm. 


Eugenia _ Steyerm. 


Senecio steyermarkii Cuatr. 
VLA um meridense Steyerm. 
Verbesina blakeana Steyerm 


C Mine pen ina Steyerm. 

Ardisia meridens erm. 

Rustia cel Standl: & Stey- 
erm. 

Saurauia meridensis Steyerm. 


dl. 


Volume 76, Number 3 Taylor 767 
Plants Described by 
Julian A. Steyermark 


56411 Psychotria patria Standl. i UR 57276 Casearia tachirensis Steyerm 
56412 Palicourea canaguensis Ste 57298 Baccharis venulosa var. cuspidibrac tea- 
5044 1 Psychotria canaguensis Standl. e" Stey- ta Steyerm. 
erm. 57309 Psychotria tamaensis Standl. & Stey- 
5044 1 Rudgea nebulicola iaa erm. 
56455 Turpinia multijuga Steyert 57310 E reticulata Steyerm. 
56502 Rudgea lucentifolia Standl. & Steyerm. 57321 Schefflera tamana Steyerm. 
56507 Llagunoa venezuelana Steyerm. 57335 s mplocos pittieriana Steyerm. 
56509 Maytenus coriacea p 57336 Schefflera cuatrecasasiana Steyerm. 
56513 Meriania steyermarkii Glea 57340 Symplocos tamana Steyerm. 
56516 Saal Ms ola oa. 57359 Conomorpha tamana Steyerm. 
56536 Drymaria firmula S Meye orm. 57367 Zarex tachirensis Steyerm. 
50537 Baccharis meridensis Steyerm. 57391 Rhynchospora tomentosa Steyerm. 
56539 Canciones Veris a 51401 Carex tamana Steyerm. 
56550 Rhynchospora paramora Steyerm. 57404 Rhynchospora tamana Sad erm. 
56551 Myrcia sanisidre rensis Se 57422 Fimbristylis tamaensis Stey 
56569 Dipterode endron vene lee Steyerm. 57428 auwolfia steyermarkii W cod 
56580 Zanthoxylum pallidum Steye 57435 | sulcata St 
56582 Ternstroemia acrodantha Kobucki & 57441 alicourea tachirensis Standl. 8 Stey- 
Steyerm erm. 
56589 Calyptrant mes meridensis Steyerm. 57455 ^en rad ed e A 
56591 Hillia microcarpa Steyerm. 57456 Conomorpha liriensis Steyer 
56592 Havet etia laurifolia var. venezuelana 57481 Ranunculus PEE f. iM 'grus 
teyerm. Steyerm. 
56594 B has venezuelensis Steyerm. 57519 Polymnia meridensis Steyerm. 
56601 Guettarda steyermarkii Standl. 57528 Pectis venezuelensis Steyerm. 
56014 sychotria dev! Steyerm. 57551 Dioscorea bolivarensis Ste yerm. 
56653 llex nervosa var. glabrata Steyerm. 57587 Rhyne 'hospora trichochaeta var. vene- 
56681 Psychotria Senden ina Steyerm. zuelensis Steyerm. 
56721 Anthurium monsterioides Steyer 57629 Bulbostylis svensoniana Steye 
56792 Eupatorium steyermarkianum Badillo 57649 E tia SEM j0sa var. NEAR 
56806 Croton larensis Steyerm. 
56813 Phyllanthus larensis Steyerm. 57688 Bolus boliv ipo Steyeri 
56821 roton deserticolus Steyerm 5713 Alseis trichocarpa Standl. & Steyerm 
56854 Phyllanthus croizatii Steyerm 57717 Chry sop! hyllum stey ali M lose 
56869 Beloperone steyermarkii Le 57749 Eupa atorium crassicaule Steyerm 
56979 Ternstroemia son var. minor 57807 Monotagma duidae Steyerm. 
Kobuski & Steye 57820 Exochogyne aa Gilly 
57005 Hieronyma croizatii Stey 1 57821 Xyris esmeraldae Steyerr 
57019 Picramnia caracasana f. pam 57835 Xyris capillaris Steyerm. 
Steyerm. 57835a Genlisea esmeraldae Steyer1 
57039 md meridensis Steye 57850 Drosera tenella var. peli Stey- 
57040 Ve ex collumanthus Bu em erm. 
57063 Hy no ibi viride Steyerm. 57853 Urospatha savannarum Steyerm. 
57066 Verbesina negrensis Steyerm 57879 Doliocarpus esmeraldae Steyerm. 
SLOT? Viola venezuelensis Steyerm 57888 Calathea duidae Steyerm 
57084 Eupatorium stevioides Steyerm 57891 Phyllanthus orinocensis Steyerm. 
57111 Zanthoxylum tachirense Steyerm. 57908 Bactris duidae Steyerm. 
57136 Arachnothrix calycophylla Steyerm 57919 Psychotria paupertina Standl. & Stey- 
57137 Castilleja steyermarkii Pennell erm. 
57149 Verbesina tachirensis Steyerm. 57920 Calathea lasseriana Steyerm. 
57162 Cuphea serpyllifolia var. tachirensis 57933 Bactris venezuelensis Steyerm. 
Steyerm. 57944 Desmoncus duidensis Steyerm. 
57170 Eugenia tachirensis e 57975 Tovomita duidae Steyerm. 
57172a Eugenia tamaensis Steye 57979 Lisianthius scabridulus Steyerm. 
57175 Fuchsia verrucosa var. tamaensis Stey- 57986 Conomorpha apiculata Steyerm. 
erm. 51989 Schefflera duidae Steyerm 
57192 Palicourea ensiformis Steyerm. 57994 Spathicarpa duidae Steyerm. 
57198 Oreobolus venezuelensis Steyerm. 58001 Axonopus steyermarkii Swallen 
57204 ugenia crebrifolia Steyerm. 58012 Navia steyermarkii L. B. Smith 
57212 Hypericum ericifolium Steyerm. 58021 Sauvagesia duidae Steyerm. 
57215 MID frigidum f. tamaense Stey- 58022 Ilex tateana Steyerm. 
erm. 58038 Ilex stenophylla Steye 
57217 Espe Ea steyermarkii Cuatr. 58040 Stegolepis a Sus erm. 
57246 Pleurothallis steyermarkii Schweinf. 58047 Oyedaea blakeana Steyerm. 
57253 a sessilianthera Steyerm. 58048 Psychotria duricoria Standl. & Stey- 
5171255 sychotria uberta Standl. & Steyerm. erm. 


768 Annals of the 
Missouri Botanical Garden 

58056 Psychotria avia Standl. & Steyerm. 59227 Psychotria kukenanensis Steyerm. 
58057 Smilax duidae Steyerm. 59331 Microlicia steyermarkii Gleason 
58061 pire MI m 59333 Utricularia oliveriana f. parva Steyerm. 
58099 nthurium duidae 59335 Ut ddnde arenicola var. kavanayena 
58117 Posta Sane bien Sd & Ste 

Steyerm. 59336 Xyris inuidia actea Steyerm. 
58123 Ilex retusa f. glabra Steyerm. 59342 Poecilandra pumila Steyerm. 
58125 Mikania auyantepuensis var. brevipedi- 59349 lboda macrostachya var. robustior 

cellata Steyerm. Steyerm. 
58127 llex retusa f. major Steyerm. 59351 (reir aureolimba Steyerm. 
58168 s scaridula Steyerm 59359 ein : pressa var. kavanayena 
5817 Tofieldia duidae Steyerm. 
58186 Bonnetia duidae Kobuski & Steyerm. 59361 E pusilla Steyerm. 
58191 e ii cretacea var. composita 59362 Nietneria paniculata Steyerm 

59363 agenocarpus steyermarkii Gilly 

58225 Philodendron duidae Steyern 59309 Paepalanthus steyermarkii Mol 
58229 icularia humboldtii f. cuneata Stey- 59394 Ilex subrotundifolia Steyerm 

erm. 59396 Clusia pusilla Steyerr 
58233 Xyris culmenicola Steyerm. 59403 Calea oliveri var. taeniotricha Steyerm. 
58242 Styrax duidae Steyerm 59404 Utricularia congesta $ teyerm. 
58254 Rhynchospora duidae Steyerm. 59408 Calea lucidivenia f. lanceolata Steyerm. 
58262 Miconia steyermarkii Gleason 59440 Psychotria ptariensis Standl. & Stey- 
58265 Conomorpha lepidota f. acutata Stey- erm. 

erm. 59452 Polygala s sanariapoana Steyerm. 
58270 Vellozia duidae Steyerm. 59482 Phyllanthus major Steyerm 
58284 Ouratea duidae di Ma 59483 Phoradendron ptarianum Ste 
58340 Ilex venezuelensis Steyert 59484 Ledothamnus steyermarkii A. C. Smith 
58349 Ilex retusa f. glaucescens Penna 59497 Grammadenia ptariensis Steyerm 
58394 Remijia steyermarkii Standl. 59504 Eupatorium tepuianum Steyerm. 
58437 Xyris stenostachya Steyerm. 59505 Eupatorium roupalifolium f. latifolium 
58442 Tocoyena orinocensis Standl. & Stey- Steyerm. 

erm. 59509 Anthurium ptarianum Steyerm. 
58445 Genlisea sanariapoana Steyerm. 511 Ravenia ruellioides var. ptariana Stey- 
58446 Utricularia oliveriana Steyerm. erm. 
58461 Pagamea anisophylla Standl. € Stey- 59514 Stegolepis parvipetala Steyerm. 

erm. 59517 Utricularia tepuiana Steyerm 
58466 Burmannia sanariap oana Steyerm. 5952] Psychotria speluncae Standl. & Stey- 
58467 Sauvagesia miniata Steyerm. erm. 
58469 Rhynchospora sanariapensis Steyerm. 59527 Mikania phelpsii Maguire & Steyerm. 
58472 rosera sanariapoana Steyerm. 59534 Aulomyrcia bolivariensis Steyerm. 
58488 Utricularia orinocensis Steyerm 59537 Drimys brasiliense var. ptaritepuiensis 
58504 Mandevilla steyermarkii Woodson Steyerm. 
58505 Utricularia amazonasana Steyerm 59547 Magnolia ptaritepuiana Steyerm. 
58520 Utricularia ayacuchae Steyerm 59548 Malanea ptariensis Steyerm. 
58558 alanea setulosa Steyerm 59549 Phthirusa ptariana Steyerm 
58606 Clusia roraimana Steyern 59570 Bonnetia steyermarkii Kobuski 
58652 Rhynchospora tepuia teyerm 59574 Ilex ptariana Steyerr 
58678 Oliganthes roraimensis Steyerm 59625a Utricularia venezuelana Steyerm. 
58759 Conomorpha roraimae Steyerm 59632 Conomorpha inline ailis m. 
58701 Symplocos roraimensis Steyerm 59034 Pagamea standleyana Steyerm. 
58848 Rapanea minima Steyerm 59640 Cephalocarpus and Gilly 
58868 Eriocaulon steyermarkii Mold 59642 Ouratea ptaritepuiensis Steyerm 
58870 C oraimensis Steyerm 59644 Genlisea nigrc is Steyern 
58943 Phthirusa roraimensis Steyerm 59655 Stegolepis ptaritepuiensis Steyerr 
58956 Eugenia rondonensis Steyerm 59058 Perama galioides var. intermedia Stey- 
58983 Rapanea reticulata St n erm. 
59000 Magnolia roraimae Steyerm 59660 Xyris setigera var. elongata Steyerm. 
59004 Mollinedia roraimensis Steyerm 59677 nopsis ptariana Steyerm. 
59065 Polygala balkeana Steyerm. 59682 Philodendron ptarianum o 
59072 Calea clematidifolia Steyerm. 59688 Phoradendron t ee eri Steyer 
59107 Cupania roraimae Steyerm. 59695 Smilax staminea tusata oa 
59118 idgea bolivariensis Steyerm. 59699 Clusia imbriata Steyerm 
59136 onomat pha T aed 59702 usia cerroana Steyerm. 
59182 C ia kukenanica Stey 59711 Wn densiflora var. tepuiensis 
59191 Utricularia boliv: ariana Stey m. Steyern 
59195 Utricularia congesta f. aues Stey- 59719 na p SHARM 

erm. 59724 ulomyrcia ptariensis Steyer 


Volume 76, Number 3 


Taylor 
Plants Described by 


769 


Julian A. Steyermark 


60015 


60021 


60090 


60094 
60102 
60119 
60121 
60124 
60125 


60175 


Xyris — Steyerm. 
agam eyermarkit Stan 
Utricularia campbelliana var. minor 
Steyerm. 
Weinmannia stey ermarku Cuatr. 


Gomidesia bonnetiasylvestris Steyerm. 
Leothrix steyermarkii Mo 

Everardia steyermarkii Gill 
Rhynchospora ptaritepuiana Steyerm 


n. 
Brunellia integrifolia var. ptariana Stey- 


erm. 
Utricularia aureomaculata Steyerm 
A ma steyermarkii Woodson 
Mikania stictophora Steyerm. 
pdbaltenos ptariense Steyerm 
Eupatorium roupalifolium var. subinte- 


Clusia hexacarpa var. ptaritepuiana 


oteyerm. 
Psychotria arenaria Standl. & Steyerm. 


xt paucifolius var. steyermarkii 
ryon 

sis ptariana a Steyer 

i cn roraimensis var. lios 


teyer 
Artrosis stey rain McClure 
s ptariensis Steye 
U inlaws wre dd oma CM 
narkii Bartram 
ld 


Moronobea ptaritepuiana Steyerm 


Protium ptar ianum var. os Sie 


erm. 
Panopsis tepuiana Steyerm. 
Protium ptarianum Steyerm. 


Piper steyermarkii Yuncker 
Coussapoa steyermarkii Cuatr. 
Mollinedia ptariensis Ste 
Mikania ADA var. pa 
Steyer 
Concev cier ptarianum Steyerm. 
Maguire 


yrcia sor oropanensis Seyen. 


erm. 
Diodia lipesi var. leioc arpa Stey- 
rm. 
Merograda sororopaniana Steyerm 


Mahurea sororo pan tepuiana Steyerm. 
erm 


j ens E Stand. & 
Stey 


60184 
60185 


60191 


60201 
60203 


60668 
60686 


607 í 02 


60789 


Chiococca lucens Standl. & Steyerm. 
Cephaélis bolivariensis Standl. € Stey- 


erm. 
Psychotria heterocarpa Standl. & Stey- 


erm. 
atayba sororopaniana eoa 

Matayba ptariana Steye 

Rhynchospora bolivariana e 

Aulomyrcia salticola Steyerm. 

Sauvagesia longipes Steyerm. 

Roupala minima Steyerm. 

Thesium epoko a rm 

um Stey erm. 
Tm 


a bolivariana Sie erm. 

cia tepuiensis Steyer 
Ps eda parvifolia c rm. 
Humiria pilosa Steyerm. 
Ilex tepuiana Steyerm. ex Edwin 
Blepharandra ptariana Steyerm. 
Trattinickia glabra Steyerm. 
Oxythece steye rmarkiana Monachino 


Remijia stenopetala Standl. & Sus. 
Matayba reducta Steyerm. 

Pier sm: lon venezuelense n iN 
Palicourea aculeifera Steye 

Psychotria cubitalis Standl. n Steyerm. 


> maculata Steyerm. 
Predum puberulenta Steyerm. 
Tovomita angustata Steye rm. 
Utricularia subpeltata Steyerm. 


Micropholis ste ee À Monachino 
Souroubea guianensis var. tomentella 
Steyerm 
Sterigmapetalum guianense Steyerm. 
Phoradendron karuaianum Steyerm. 
Ilex karuaiana Steyerm. 
Matayba venezuelana Steyerm. 
Pourouma steyermarkii Standl. & 
C 


„uatr. 

Sloanea ptariana Steyerr 

Hieronyma oblonga var. exited gla- 
bra Steyerm. 


PE ain rcia karuaiensis Steyer 
Caraipa longipedicellata Steyerm. 
Mikania lucida f. hirtic aulis Steyerm. 


Geonoma karuaiana Steyert m. 


770 Annals of the 
Missouri Botanical Garden 
60813 Blepharodon steyermarkii R. Holm 62436 Melochia humboldtiana Steyerm 
60820 Diomma fruticosa Steyerr 62437 Clibadium surinamense var. macrophyl- 
60822 Taralea steyermarkii Schery lum Steyer 
60824 Euplassa venezuelana Steyerm 62491 Echeveria bicolor var. turumiquirensis 
60829 Cladium costatum Steyerm. eyerm. 
60831 Rhynchospora sororopana Steyerm. 62532 Croton turumiquirensis Steyerm 
60834 Trimezia fosteriana Steyerm. 6253 led turumiquirensis Steyerm 
60847 Anthurium krauseanum Steyerm 62542 mannia costensis Standl. & Stey- 
60849 Senefelderopis croizatii Steyerm. 
60863 rain englerianum Steyerm. 62548 Eugenia dear hice Steyerm 
60868 Rapatea steyermarkii Maguire 62513 Gnaphalium caeruleocanum m Steyerm, 
60871 Bonnetia tepuiensis Kobuski & Stey- 6257 Hypericum caracasanum var. turumi- 
quirense Steyerm. 
60875 Malas microphylla Standl. & Stey- 62583 c charis erec folia Steyerm. 
62605 ex culmenicola Steyerm. 
60876 Podocarpus steyermarkii Buchh. & 62611 Plinia cola Steyerm. 
Gray 62628 yrtus me var. turumiquirensis 
60879 Myrcia A McVaugh (dedicat- Steyer 
'd to 62088 ede ai venezuelensis id 
60883 Byrsonima carraoana Stey 62689 Abutilon turumiquirense Steyer 
60888 Pagamea pauciflora Stand, & Steyerm. 62691 Viburnum tinoides var. roraimense f. 
60892 Moronobea ptaritepuiana f. rosea Stey- turumiquirense Steyerr 
62705 Carex turumiquirensis Steyerm. 
60898 ( 'onomorpha depressa Steyerm. 62711 Befaria steyermarkii C. Smith 
60912 Cupania kavanayena Steyerm 62739 Stellaria venezuelana Steyerm 
60914 ualea ferruginea Steyerm 62752 Utricularia turumiquirensis Steyerm 
60920 Drosera arenicola Steyerm 62800 Capparis guaguaensis Steyerm 
60968 Desmoncus multijugus St 62875 croton sucrensis Steyerm 
60993 Homalium anzoateguiensis Steyerni: 62880 Heteropterys quetepensis Steyerm 
61016 Sloanea onotillo Steyerm. 62963 xcremis coarcta lba Steyerm 
61039 Bactris bergantina Bion 65135 Aster anomalus f. albidus Steyerm 
61317 iis hnothri reflexa var. glandulifera 66615 Heuchera puberula f. glabrata Steyerm 
teyer 67440 Viola triloba f. albida Ste 
61373 Magia is Standl. & Steyerm. 3749€ cce canadensis f. Pb Stey- 
61424 Arachnothrix costanensis Steyerm. 
61443 Freziera steyermarkii Kobuski 67915 Hydrophyllum appendiculatum f. album 
61513 Diospyros anzoateguiensis Steyerm. 
61519 Abutilon pseudogiganteum ene 67235 Mertens aa ica f. rosea Steyerm. 
61523 Croizatia neotropica a Steyerm 68024 Scutellar rvosa f. alba 2 erm 
61524 ams a pittieriana Bum 68254 spora idu idae Steyerm. 
6157: Styrax costanus Steyerm. 68397 Lysimachia quadriflora f. aren 
61612 Oyedaca em var. glabrior Steyerm. 
Steyer 70292 Ruellia strepens f. alba Steyerm. 
61626 fue unilocularis Kobuski & 70126 Helianthus mollis f. flavida Steyerm. 
Steyerm. 72011 Rosa setigera f. alba Steyerm. 
61659 llex vesparum Steyerm. 73223 d canescens f. pallidum 
61665 Ternstroemia steyermarkii aaa Palmer & Steyerm. 
61726 Sloanea onotillo var. major Steyer 73886 Ledothamnus roan Maguire, Stey- 
61731 Maytenus guianensis f. crenulata Sr erm. & Lute 
erm. 74630 Palicourea tepuic uis Steye 
61836 Dioscorea lasseriana Steyern 14850 Anthurium steyermar rkii Bunting 
61874 eperomia steyermarkii Surius 14652 Smilax chimantensis Steyerm. & Ma- 
61914 Banisteria alternifo la Steyerm. guire 
61919 Hasseltia monagensis Steyerm. 14656 Matayba chimantensis Steyerm. & Ma- 
61950 Cinnamodendron venezuelense Steyerm. guire 
61959 uarari teyermarkii Cuatr. (4781 Catostemma hirsutulum Ste 
62017 Oreopanax venezuelense Steyerm. 74816 Struthuanthus chimantensis Sen & 
62095 Oy edaea verbesinoides var. hypomalaca aguire 
yerm. 74877 Pagamea uniflora a 
62102 Duranta steyermarkii Mold. 14883 Graffenrieda steyermarkii Wurd. 
62110 Rhynchospora culmenicola eum 74886 lo ias ane Maguire, Stey- 
62185a Xylobium steyermarkii Folda erm. & Lutey y 
62189 Phyllanthus a e um 74980b Dicranoloma steyermarkii Seni 
62215 Sloanea breviseta Steyerm. 75059 Phthirusa hippoerateoides Steyerm. & 
62216 Sloanea ena Steyerm Maguir 
62225 Palicourea pustulata Steyerr 75110 Stenopadus affinis Maguire, Steyerm. & 
62232 Alseis microcarpa Standl. & Steyerm. 
62375 Aulomyrcia vinacea Steyerm 15157 This reticulata Steyerm. & Maguire 


Volume 76, Number 3 
1989 


aylor 


Tay 
Plants Described by 


Julian A. Steyermark 


Malanea chimantensis Steye 
Dacryodes chimantensis ea & 


aguire 
Chiococca nitida var. chimantensis 
Steyerm. 
Phoradendron tenuiflorum Steyerm. & 
ies 
Piper to a oo 


arkit L B. Smith & B. 
. Se 
n ballotifolia Maguire, Steyerm. & 


Nothochlaena steyermarkii Ma 
Kotchubaea longiloba Steye 

Rapatea chimantensis er 

Hymeno hyllopsis steyermarki Vareshi 
e tepuiensis Ste eyerm. & Ma- 


Tepuia multiglandulosa Steyerm. & Ma- 


guire 
Didymopanax chimantensis Steyerm. & 
aguire 
Myrica rotundata Steyerm. & Maguire 
Psychotria aubletiana var. cacuminis f. 
tomentella Steyerm. 
Magnolia chimantensis Steyerm. & Ma- 


guir 

ua lanocaulis Maguire, Stey- 
erm. & Wurd. 

Rhamnus chimantensis Steyerm. & Ma- 
guire 

Clusia steyermarkii Ma 

Men viarum f. pilifera Paus & Stey- 


ee pilosella f. laevigata Palmer 


eyerm. 
Rubus pensilvanicus f. albinus Palmer 
& Steyerm. 
Calathea coccinea Standl. & Steyerm. 
Lobelia siphilitica f. purpurea Palmer & 
Steyerm 
Silphium perfora f. petiolatum 
almer 
Munhlenbergia brachyphyll f. aristata 
almer & Steyerm. 
ien dcin var. occidualis f. ro- 


Randia Mic var. dasyphylla Stey- 


erm. 
Pittierothamnus elineolatus Steyerm. 
Psychotria lupulina s 
var. maypurensis f. pubens Steyerm. 
pude haenkeana var. mirandensis 
Steyerm. 
Dicranostyles imatacensis Steyerm 
Stenospermation steyermarkii Bunting 


ote 
Sipanea pratensis var. e” f. 
breviflora Steyerm 


89464 
89491 
89543 
89565 
89577 
89698 
89802 
89813 


89875 
90199 


90428 
90429 
90467 
90647 
90798 


90823 
90883 


90919 


Siparuna sancheziana Steyerm. 

Swartzia steyermarkii Cowan 

Stenospermation ammiticum Bunting 

Piper Se Steyerm. 

Stegolepis minor Steyerm. 

Navia ida x B. Smith & Steyerm. 

Amaioua brevidentata Steyerm. 

Sicydium araguense Steyerm. & Trujil- 
lo 

Dichapetalum e Prance 

Navia navicularis L. B. Smith & Stey- 


erm. 

Navia connata L. B. Smith & Steyerm. 

Remijia densiflora var. minima Steyerm. 

etalum Dora subsp. ichu- 
m. 


er 
Remijia roraimae var. Ads a Stey- 
erm. 
Qualea apodocarpa s 
Henriettiella steyerm 
od hyssopifolia f. peras Sie: 


Smilax suraimensis Mi ls 

cs dunster um Steyerm., Ari- 
. & Wur d 

dr trinitense Steyerm. 

Macrolobium steyermarkii Cowan 

Meliosma nubicola Steyerm. & Lasser 

Cestrum pariense Steyerm. 


Malanea pariensis Steyerm. 
Piptocarpha steyermarkii E 
Rytidostylis brevisetosa DURAM 
Coussarea d Steyerm 


Psy chotria sanmartensis var. costanen- 
sis Steyerm. 

Paullinia naiguatensis Steyerm 

Connarus steyermarkii Prance 

Heisteria olivae Steyerm. 

Paullinia toxicodendroides Steyerm. 

Lacmellea costanensis Steyerm 

Chrysophyllum E Stey- 
erm. 

Struthanthus mucronatus a 

Dacryodes steyermarkii San 

Ochthocosmus grandifolius Eom 

Pouteria venamoensis Steyerm. 

Orthaea venamensis Maguire, Steyerm. 
& Luteyn 

Piper venamoense Ste 

Piper hippoerepiforme Stey 

roreticulata event 
Vochysia pauciflora Steyerm. 
Psychotria coussareoides var. ciliata 
erm. 


otey 
Cissus venezuelensis Steyer m. 


Remijia roraimae var. auyantepuiensis 
Steyerm. 

Schoepfia tepuiensis Steyerm 

Chrysophyllum ficoides RN 

Ocotea steyermarkiana C. K. Allen 

Notopora auyantepuiensis Steyerm. 

Achnopogon steyermarkii Aristeg. 


772 Annals of the 
Missouri Botanical Garden 
93564 Cephaëlis cacuminis Steyerm. 97071 Manettia coccocypseloides var. glabrior 
93571 Sloanea steyermarkii C n Steyerm. 
93652 Chiococca auyantepuiensis Steyerm 97090 Palicourea meridensis Steyer 
93712 Heliamphora heterodoxa var. exappen 97239 Manettia lindenii f. pli uam Stey- 
diculata f. glabella Steyerr erm. 
93737 Mandevilla subcarnosa var. angustata 97243 Palicourea nubicola Steyerm. 
eyerm. 97257 Piper canovillosum Steyerm. 
93808 Sténapaduk steyermarkii Aristeg. 97325 Anthurium a Bunting 
93837 Utricularia ste yermarkii P. Taylor 97475 Chiococca naiguatensis Steye 
93844 Philacra steyermarkii 97655 Psychotria icu Eum 
93889 Blepharodon auyantepuiense Steyerm. 97664 Peperomia chapensis Steyerm. 
93892 Cynanchum auyantepuiense Steyerm. 97688 Psychotria yaracuyensis St m. 
93895 Lindmania riparia L. B. Smith, Stey- 97811 spe eel DM bolivariana Bunting & 
erm. & H. Robinson Steyern 
93917 Pouteria auyantepuiensis Steyerm. 97837 Maguireothamnus speciosus subsp. 
93980 eun emis Steyerm. jauaensis Steyerm. 
93926 10panax multiramosum Steyerm. 97851 Euceraea sleumeriana Steyerm. & Ma- 
93974 paran x auyantepuiense Steyerm. guire 
94103 Malanea auyantepuiensis Steyerm. 97852 Psychotria cerronis Steyerm. 
94203 Qualea rubiginosa var. angustior Stey- 97854 Tibouchina stey a da 
erm. 97857 Malanea jauaensis Steye 
94382 Markea costanensis Steyer 97865 ees ite L. B. Smith & Stey- 
94518 Lennoa madreporoides di australis 
Steyerm. 97865a Navia p L. B. Smith & Stey- 
94897 Piper pseudobredemeyeri Steyerm. erm. 
94923 Trichipteris steyermarkii Tryon 97878 Ledothamnus steyermarkii subsp. longi- 
94956 Psychotria sucrensis Steye setus Maguire, Steyerm. & Luteyn 
94962 Besleria mortoniana Ste 97881 Navia incrassata L. B. Sk & Stey- 
95002 Piper TOPUR e oo erm. 
95019 Psychotri ensis Steyer 97895 Lycopodium steyermarkii Ollgaard 
95022 Guzmania sieut cu L. n. Smith & 97917 Stomatochaeta steyermarkii Aristeg. 
erm. 97918 Ledothamnus jauaensis Maguire, Stey- 
95075 Psychotria humensis Steyerm. erm. & Luteyn 
95152 Sarcorhachis venezuelana Steyerm. 97928 Pagamea jauaensis Steyerr 
95187 Boehmeria ramiflora var. vinacea Stey- 97944 Ledothamnus stenopetalus Musee 
erm. Steyerm. 
95215 Psittacanthus costanensis Steyerm. 97945 Cardonaea jus Aristeg., Maguire 
95246 XR di hyalina Steyerm. & Steye 
95314 Psyc hotria costanensis var. pubibractea 97958 Ta es LR sessilis Steyerm. & Ma- 
Steyerm. guire 
95320 Vriesea splendens var. oinochroma 97979 Byrsonima steyermarkii Anderson 
Steyerm. 98016 Sipapoa steyermarkii Maguire 
95355 Peperomia borburatensis Steyerm. 98049 Tyleria phelpsiana Maguire & Steyerm. 
95374 g Ss var. trichophylla 98054 Vaccinium puberulum var. jauaense 
Ste aguire, Steye utey 
95418 Selaginella gigantea Steyerm. & A. R. 98057 Psychotria glandulicalyx var. densipu- 
ens Steyerm. 
95456 Mars stellulata Steyerm. 98068 Lisianthius jauaensis Steyerm. & Ma- 
95463 estrum aristeguietae Steyerm. guire 
95934 Palicourea Deua var. pu- 98070 Psychotria phelpsiana Steyerm. 
ens Steyerm. 98072 Symplocos jauaensis Steyerm. & Ma- 
96039 di anceps var. robustior Stey- guire 
98073 Smilax jauaensis Steyerm. & Maguir 
96161 REA steyermarkii Bunting 98078 Schefflera reticulata subsp. jentensis 
96216 Peperomia pariensis oe rm. Maguire, Steyerm. & Frodin 
96342 Elvasia steyermarkii 98086 Aspidosperma decussata n jauaen- 
96472 Pitcairnia venezuelana L B. Smith & sis Steyerm. & Ma 
Steyerm. 98098 Psychotria jauaensis Si erm 
96653 Clidemia steyermarkii Wur 98206 Pochota gracilis subsp. bolivariensis 
96715 Campylocentrum ste iid Foldats Steyerm. 
96736 Manettia tachirensis Steyerm. 98298 Manettia tamaensis Steyern 
96816 Palicourea demissa var. chimo f. pu- 98345 la di Foldats 
bens Steyerm. 98381 Lepanthes steyermarkii Fo vue 
96951 un ceca L. B. Smith & Stey- 98384 Maxillaria steyermarkii Foldat 
98558 


96993 


inicium steyermarkii Foldats 


Arcytophyllum nitidum f. indui 


Volume 76, Number 3 
1989 


Taylor 


y 
Plants Described by 


773 


Julian A. Steyermark 


100311 


10037 la 


100884 
100940 
101030 
101055 
101900 
101939 
102035 
102360 
102431 


102603 


102939 
103062 
103153 


103185 


103752 


Telipogon steyermarkii Foldats 
Dendrophthora steyermarkii Rizz. 
Brachionidium ucc d, Foldats 
Stelis steyermarkii Foldat 


Psychotria aubletiana var. oe Stey- 


erm 
Palicoure ea dur ill Steyerm. 
Psychotria aunda Steyerm. 
Psychotria sanluisensis P 
Piper galicianum Steyer 
Lepanthopsis steyermarhis Foldats 


Sterigmapetalu um het ho den Steyerm. 
& 


iesner 
Piper Wird UE p 


A otria chape 


Dio nose costanensis m Y & 
Austin 

ini yaracuyense Henderson & 
Steyerm. 


eromia croizatiana Steyerm. 
Psychotria decurrens Xung 
Hoffmannia aroensis Stey 
Hoffmannia stenocarpa N 
Psychotria dunstervil m Steyerm. 
Palicourea Peru vs Steven. 
Bulbophyllum steyermarkii Foldats 
Gomphicus steyermarkii Foldats 
Piper crenulatum Steyerm. 
Palicourea tamaensis var. psilophylla 

Steyerm. 

Psychotria norae Steyerm. 
Palicourea nana Steyerm 


Arcytoph 


oe caraboboensis Steyerm. & 


Bun 
Philodendron steyermarkii Bunting 
ir 


Heliconia julianii Ba 
Leandra steyermarkii Wurd 
naa maroana S m 


ra yavitensis Stey 
Heliantho stylis como A C. Berg 
Psychotria yavitensis Steyern 
Duroia maguirel var. ac 
he stey erm. 
Monotagma yapacanensis Steyerm. & 
ntin 


unung 
Bonnetia tristyla subsp. nervosa Stey- 
erm. 
Symplocos yapancanensis oe 
Sloanea yapacanae Steyerm 


Psittacanthus julianus Rizz. 
~ papillata ee eli 7 Südi 


Schradera andina Steyerm 


103937 


104418 
104454 
104569 


104978 


104989 


105009 


105237 


105274 
105306 
105390 


105468 
105500 


106037 


106178 
106573 
106763 


108880 
108895 


108921 
108938 
108984 
109039 
109041 
109076 
109178 
109181 


109199 
109237 


109252 
109255 


109294 
109298 


109315 
109373 


Tapeinostemon oo var. australe 
Maguire & Steyer 

Burmannia carrenoi en 

Piper dunstervilleorum Stey 

Sterigmapetalum resinosum er & 
ies 

xd oa f. tomentosum Stey- 

rm. 


Piper nubigenum var. venezuelense 
Steyerm. 
Heteropterys steyermarkii Anderson 
m. 


ge) 
w 
e 
e 
>. 
o 
= 
"i 
m 
4 
o 
d 
p 
Q 
o 


erm. 

Froesia venezuelensis Steyerm. & Bunt- 
ing 

Distictis steyermarkii pal 

Peperomia buntingil Steyer 

var. cernuum T papilla- 


. & L. B. Smith 
aane Bunting 


Piper dilatatum f. longipilum Steyerm. 
is onde glandulicalyx f. bracteifera 


yer 
TUN diederichsanae Steyerm. 
Styrax wurdackiorum Ste eyerm 
Ds Smith 


Raveniopsis jauaensis Steyer 
Navia scirpiflora L. B. Smith. A 
& H. Robinson 
ic jauana L. B. Smith, Steyerm. & 
obinson 
ravia steyermarkii de Roon 
Poc bs coer eh Steyerm. 
Phainantha steyermarkii Wurd. 
Sloanea cavicola Steyerm. 
Liriosma tepuiensis Steyerm. 
Psammisia breweri Steyerm. & Maguire 
Virola steyermarkii Rodrigues, ined. 
Mycerinus viridiflorus Steyerm. & Ma- 
uire 


Daphnopsis nevlingiana Steyerm. 

Tepuianthus sarisarinamensis Maguire 
& Steyerm. 

Maytenus jauaensis Steyerm. 

Chorisepalum breweri Steyerm. € Ma- 
guire 

Ilex jauaensis Steyern 

Men carrenol "NN & Ma- 


Thibaudia breweri Steyerm. & de 
Schefflera gracillima Steyerm. & M 
guire 


774 Annals of the 
Missouri Botanical Garden 
109389 Paepalanthus meseticola Mold. & Stey- 117421 Bonnetia guaiquinimae Steyerm. 
erm. 117468 Idina berryi Cowan & Steyerm. 
109390 Xyris jauana L. B. Smith & Steyerm. 117473 Croton guaiquinimae Steyerm. 
109405 CERE Mu e venetifolius Mold. & Stey- 117498 m guianensis MA guaiquini- 
erm. mensis Steyerm. 
109428 Psychotria carrenoi Steyern 117569 Decem longipedicellatus Stey- 
109447 Ouratea propinqua Maguire & dm erm. & Luteyn 
109485 Piptocarpha jauaensis ius & S 117570 s ds attenuatus Steyerm. & 
e 
109535 Tapeiaostémdn jauaensis Steyerm. & 117615 Neea. su be labrata Steyerm. 
Maguire 117841 Davilla steyermarkii Kubitzki 
109574 Styrax Jauaens is Steyer 117920 Roupala griotii Steyerm 
109584 i-e jauaensis Beye rm. & Ma- 117971 eea gualquinimae Steyerm. 
117977 Daphnopsis guaiquinimae Steyerm. 
109596 Phyllanthus carrenoi — 117982 — atrorosea L. B. Smith, 
109633 Navia luzuloide des B. Smith, Steyerm. Steyerm. & H. Robinson 
& H. Re shins 118338 ride pseudojamesoniana Steyerm. 
109673 atayba a jauaensis s Steyerm: 118651 Peperomia liesneri ‘ n 
109689 Piper jauaense Steyerm. 118876 Sabicea liesneri Steyern 
109695 Sloanea jauaensis var. minor Steyerm. 119027 Bartlettina liesneri King & H. Robinson 
109699 Sloanea carrenoi Steyer m. 119958 Peperomia minarum 
109765 Maguireothamnus jauaensis var. breweri 119971 Piper pseudohastularum Steyerm. 
Steyerm. 120032 Sterigmapetalum tachirense Steyerm. & 
109841 Sloanea jauaensis Steyerm. iesner 
109842 Cecropia steyermarkii Cuatr. 121697 Piper tacariguense Steyern 
110148 Sloanea larensis Steyerm. 122132 Psychotria ee Steyerm, 
111182 Besleria steyermarkiorum Wiehler 122197 Psychotria davidsei Steyerm. 
111304 Blepharodon julianii Morillo 122215 pc cataniapensis Steyerm. 
111402 llex altiplana Steyerm 122318 Guettarda leiantha Steyerm. 
112181 Billbergia manarae Ste m. 122366 News pis pedum ulata Steyern 
112735 Phthirusa enema Rizz. 122478 Navia igneosicola L. B. Smith, — 
112797 Ouratea steyermarkii Sastre & "Robins son 
112829 Dulacia redmondii Steyerm 122730 Peperomia cyclophylla f. zulianensis 
113200 Dipterx phaeophylla Steyerm. Steyer 
113330 Croton subincanus var. guaiquinimensis 122876 Hillia rel Steyerm 
Steyerm. 123742 Psychotria trujilloi var. VD 
113438 Pakaraimaea Wo seis subsp. ni- Steyerm. 
tida a & Ste 123808 Vriesea capituligera Steyer 
113463 Swartzia cowa ii Steyern 123846 Navia geaster L. B. S Smith, en & 
113837 Ochthocosmus milione var. canari- H. in: 
s Steyerm. & Luteyn 124311 Bonnetia euryanthera Steyerm. 
113851 dom SR Maguire & Stey- 124326 Tyleria breweriana Steyerm. 
erm. 124383 ran foliosa n marahuacensis 
113856 Guapira ayacuchae Steyerm. uire & Steyer 
114221 Piper cernuum var. araguense f. pu- 124406 Sc hefflera uli cda var. marahu- 
bens Steyerm. acensis Maguire, Steyerm. & Fro 
115564 Phoradendron E li Rizz. 124477 aio: disc rri L. B. Smith, Stey- 
115598 Sipanea carrenoi Pu ke .& son, ined. ms. — 
115645 Bonnetia ptariensis Stey Steyerbromelia desler L. B. Smith 
115968 Aphanocarpus oe f. glabrior & H. Robinson 
Steyerm. 124526 Cottendorfia geniculata var. minor L. B. 
116093 oue auyantepuiensis Maguire Smith, Steyerm. & H. Robinson 
»t 
116166 Ravenous le d 124530 rec guianensis subsp. sipapoensis 
116180 Cottendorfia brachyphylla var. pen 124582 Brewearia duidensis L. B. Smith, Stey- 
. B. Smith, Steyerm. & H. Robin . & H. Robi 
un 7 erm . Robinson 
117209 Piper pavasense Steyerm. 124622 TENA integerrima var. pilosa Stey- 
117221 Lubaria szczerbanii Steyerm Suen — 
117276 les guaiquinimae Steyerm. 124771 "s di var. adpressipilum 
117298 Navia emergens L. B. Smith, Steyerm. Steyer 
H. Robin 124826 Piper marturetense f. pilosum alll 
117361 Navia ovoidea L. B. Smith, Steyerm. & 125483 Psychotria subimbricata Steyer 
. Robinson 125174 Perissocarpa steyermarkii bp. tachi- 
117393 Ouratea culmenicola Maguire & Stey- rensis Steyerm. & Maguire 
erm. 125654 Navia aliciae L. B. Smith, Steyerm. & 
117407 Stomatochaeta colveei Steyerm. . Robinson 


Volume 76, Number 3 
1989 


Taylor 
Plants Described by 
Julian A. Steyermark 


775 


125925 


125996 


126049 


126328 


126381 
126421 


126472 


126510 


128104 


128465 


128465 


128474 


128500 
128611 


128611 


128637 


128815 
128945 


128976a 
128992 
129063 
129067 
129067 


129197 
129203 


129391 


129456 
129512 


129596 
129646 
129650 


129798 
129816 
129924 
129932 
129962 
130072 


130112 


Oritrophium marahuacense Steyerm. & 
aguire 
Cottendorfia arachnoidea L. B. Smith, 


dteye . : : 

Rhamnus marahuacensis Steyerm. & 

Maguire 

M dead marahuacae L. B. Smith, 
Steyerm. & H. aes on 

Bs hows even Ste 


Navia albiflora L. B. Smith, Sievert. & 
: Robinéon 
Piper subsessilifolium var. morii Stey- 
erm. 


Pitcairnia steyermarkii L. B. Smith 
m 


Lindmania sessilis L. B. Smith, Stey- 
. & H. Robinson 

S huberi L. B. Smith, Stey- 
erm. . Robinson, ined. ms 

Lindmania buberi L. B. Smith, es 
erm. & H. Robinson 

Lindmania imitans L. B. Smith, Stey- 
erm. & H. Robinson 

Elaphoglossum steyermarkit Mickel 

Cottendorfia aurea L. B. Smith, Stey- 
erm. & H. Robinson, ined. ms. 

Lindmania aurea L. B. Smith, Steyerm. 


. Robinson 

Adenanthe bicarpellata var. chimanten- 
sis Steyerm. 

Chimantaea huberi Steyerm. 

Lindmania saxicola L. B. Smith, Stey- 
erm. & H. Robinson 

Vaccinium steyermarkii Lute 

Psychotria crassa f. ms Sena 

Navia pedemontana L. B. 5 mith, Stey- 
erm. & H. Robinson 

Cottendorfia terramarae L. B. Smith, 
Steyerm. & H. Robinson 

Lindmania terramarae L. B. Smith, 
Steyer obinson 

NM Ade terramarensis Steyerm. 

Cottendorfia delascioana L. B. Smith, 
Steyerm. & H. shinee 

ce calophyllus var. angustifolius 


ub ll Steyerm 
m julianii Carnevali & Ra- 


mire 

Guzmania terrestris L. B. Smith & 
Stey 

Navia n L. B. Smith, Steyerm. & 
H. Robinson 


Scaphyglottis michelangeliorum Carne- 
vali & Steyerm 


Rutaneblina pusilla Steyerm. & Luteyn 
Phyllanthus jablonskianus Steyerm. & 
Luteyn 


yn 
Acopanea ahogadoi Steyer m. 
:himantaea acopanensis Steyerm. 
Stegolepis PUE ER zs eyerm. 
St Mos nin: diffusa L. B. Smith, 
teyerm. & H. n 
Ilex Tue Steyerm. 


130135 Tillandsia NOM d L. B. Smith & 
Ste yerm 

130228 Ochthocosmus berryi Steyerm. 

130318a Navia berryana L. B. Smith, Steyerm. 
& H. Robinson 

130328 Phyllanthus vaccinifolius var. vinillaen- 
sis Steyerm. 

130339 Remijia delascioi Steyerm 

130410 


Navia delascionis L. B. Smith, Steyerm. 
& H. Robinson 
marahuacae Steyerm. 


130425 Ilex 


130440 Remijia marahuacensis Steyerm. 
130481 Ilex holstii Steyerm. 
130531 Ilex brevipedicellata Steyerm. 
130029 Gongylolepis terramarae Steyerm 
130637 Holstianthus barbigularis Steyerm 
130721 Schradera marahuacensis Steyerm. 
130742a Peperomia marahuacensis Steyerm 
130878 Catostemma marahuacensis Steyerm. 
130880 Lissocarpa stenocarpa Steyerm. 
130907 Ouratea marahuacensis Maguire & 
Steyerm. 
131659 Simira ignicola Steyerm 
131850 Navia = Steyerm. & Holst 
131868 Symplocos acananensis Steyerm. 
132006 oa ie humilis Steyerm 


SUKSDORF, W. N. (Wilhelm Nikolaus) 


Grindelia hirsutala var. brevisquama f. 
tomentulosa Steyerm. 


SWINK, F. A. 

2775 Vernonia missurica f. swinkii Steyerm. 
TAMAYO, F. (Francisco) 

3133 Duroia gransabanensis Steyern 

4509 Arcytophyllum nitidum or. cara 


sanum var. culmenicolum CUN 
TATE, G. H. H. (George Henry ipm 


Ilex c dip 


431 persa eun ta a subsp. al Ma- 
re, B erm. & Frod 

526 Ilex pep AM. Steyerm 

567 Pagamea duidana Standl. Es Steyerm. 

1147 Borreria oligodonta Steyer 

1365 Pagamea auyantepuiensis em. 


THOMAS, W. W. (William Wayt) 
Navia po : B. Smith, Steyerm. 
1. 


bir 
Navia polyglomerata L. B. Smith, Stey- 


3261 
erm. . Robinson 

3245 Navia a L. B. Smith, Steyerm. 
& H. Robinson 


THURN, I. (Everard Ferdinand Im) 
16 Schefflera S Maguire, Stey- 
erm. & F 
TILLETT, S. S. idis 
82 


oizatia naiguatensis Steyerm. 


738-366 Manette honigii Steyerm 
743-230 Peperomia atabapoensis Steyerm 
745-385 Sabicea tillettii Steyer 

746-501 Manettia tillettii Steyerm 
746-798 Peperomia honigii Steyerr 
752-329 Siphocampylus tillettii Steyerm 
752-348 Duidaea marahuacensis Steyerm. 


776 Annals of the 
Missouri Botanical Garden 
761-83 Peperomia tillettii Steyerm. 3642 Piper galicianum f. hirsutopetiolatum 
43934 Rudgea tillettii Steyerm Steyerm. 
43944 Merumea plicata Steyerm 5457 Faramea cazaderensis Steyerm. 
43946 Psychotria tillettii Steyerm. X 
z s ae VARADARAJAN, G. S. 
43954 Psychotria boyanii Steyerm . -— " 
43996 Psychotria merumensis Steyerm. Connellia varadarajanii L. B. Smith € 
44824 Retiniphyllum guianense Steyerm. Steyerm. 
44825 Rudgea merumensis Steyerm VARESCHI, V. (Volkmar) 
44835 Duroia merumensis Steyerm. Rudgea vareschii Steyerm. 
44837 Calycophyllum merumense Steyerm. 4966 Tepuia vareschii S 
44867 Orthaea merumensis Maguire, mE 6850 Arcytophyllum vareschii Steyer 
Micya 7790 Psychotria vareschii Steyerm. 
44992 Schradera ternata Steyerm. 
45007 Psychotria psittacina Steyerm VEILLON, J. P. 
45008 Psychotria urniformis St l Dendropanax veillonii Steyerm. 
45057 Ceratostema glandulifera Maguire, Stey- 14 Styrax andinus Steyerm. 
erm. & Luteyn UNER 
45059 Psychotria ayangannensis Steyerm. delata j . . 
45079 Psychotria multiramosa oe Palicourea flexiramea Steyerm. 
45081 Malanea ciliolata Steye VOGEL, S. (Stefan) 
45085 Ixora intropilosa Steyer sychotria caquetensis Steyerm 
45105 Psychotria amplinoda var. ayangannae Pos Psyehorria capitata f. trichophora Stey- 
teyerm. 

45180 Alibertia triloba Steye ; 

312 E lic olii St 
45184 Bi ea cds var. psilocarpa iii a aca 

VOGL, PADRE C. (Cornelio) 
45194 Retiniphyllum scabrum var. ayangan- Aspilia avilensis Aristeg. & Steyerm. 
Steyerm. l , À iot A 
45114 Sipsries. ayangannensis Steyerm. ii M do v impu MD Wolfgang) 
45117 dgea enervia Steyerm. 2137 Chione panamensis Steyerm. 
45178 Schefflera ayangannensis Maguire, Stey- VON TUERCKHEIM, H. or von Türckheim (Hans) 
erm. & Frodin 

l Struthanth ill Standl. & Stey- 
45182 Rudgea ayangannensis Steyerm. po SA pee "m 
proe Piper perstipulare Steyerm. 1617 Xylosma sessile Standl. € Stey 
45567 Calycophyllum intonsum Steyerm. 1728 Elaterium macrophyllum Stan dl 

TORRES LEZAMAE, A. Steyer 
203 Simira lezamae Steyerm. 1732 Polymnia x PE Standl. & Stey- 
8410 


TRACY, J. P. (Joseph Prince) 
69 Grindelia nana var. altissima Steyerm. 
TRELEASE, W. (William) 
s.n. Grindelia lanceolata f. latifolia Steyerm. 
TRUJILLO, B. (Baltasar) 
Psychotria fernandezii Steyerm. 
137 Psychotria avilensis Steyerm. 


1921 Psychotria trujilloi Steyerm 

1955 Psychotria terepaimensis Steyerm. 

2763 Faramea larensis Steyerm. 

6555 Rudgea trujilloi Steyerm 

7594 Hoffmannia bernardii var. aracalensis 

Steyerm. 

7919 Peperomia trujilloi Steyerm. 
TUCKER, J. M. 

979 Hyperbaena salvadorensis SS 

1368 Bouvardia salvadorensis Steyerm 


ULE, E. (Ernst Heinrich Georg) 
5386 Psychotria 
6795 


manausensis Steyerm. 
Joosia ulei Steyerm. 


VAN DER WERFF, H. (Henk) 
237 Peperomia ouabiane var. sanluisensis 
Steyerm. 

Piper arieianum var. yaracuyense f. fal- 
conense Steye 


548 


acia IET Standl. & Stey- 
ermi. 


WESSELS-BOER, J. G. 


962 Psychotria wessels-boeri Steyerm. 

1953 Schoenobiblus amazonicus Steyerm. 
WHITEHOUSE, E. (Eula) 

s.n. Selenia oinosepala Steyerm. 


WHITTON, B. A. 


120 Chiococca multipedunculata Steyerm. 
WILLIAMS L. (Llewelyn) 

10639 Utricularia veu e 

11505 Eugenia caurensis Ste 

11691 Myrciaria caurensis Ste; 


12073 m rigida var. hirtibacca Stey- 
12970 ios pungens Standl. & 
teyerm 

12992 Psychotria calvifora n 

1317 Carex standleyana Steye 

13329 Rondeletia orinocensis cam 

13650 Maytenus pittieriana Steyerm 

13853 mo macrocephala Standl. & Stey- 
ern 

13918 Picram mia tristamina Ste 

13922 


Philonotion williamsii m 
4026 Faramea yavitensis Steyerm. 


Volume 76, Number 3 
1989 


Taylor 


Plants Described by 
Julian A. Steyermark 


14140 


15532 


WILLIAMS, 
112 


13178 


WILSON, € 


175 
188 


329 
356 


Caraipa ferruginea Steyerm. 


Posoqueria williamsii Steyerm. 

Platy d decipiens Woodson & 
Steyern 

Rudgea cardlipá m & Steyerm. 

Clusia reducta erm. 

Eugenia llewelynii S m. 

iid iiie ventuariana Stand! & Stey- 


Pao galioides f. quaternata Steyerm. 


Calathea acuminata Steyerm. 


Tocoyena brevifolia Sum 

». (Louis Otho) 

icis aubletiana var. centro-ameri- 
na Steyerm. 

Carex la ana Steyerm. 


L. (Carl Louis) 


Triolena izabalensis Standl. & Steyerm. 
Tetrorchidium brevifolium Standl. & 


dteyerm. 
Euphorbia verapazensis Standl. & Stey- 
eri 


‘rm. 
Hybanthus sylvicola Standl. & Steyerm. 


Lisianthius calciphilus Standl. & Stey- 
erm 


WINGFIELD, R. (Robert) 


5495 


6571 


Piper diffamatum var. angustius Stey- 
erm. 
Piper pseudodivulgatum Steyerm. 


WOOTON, E. O. (Elmer Ottis) 


s.n. 
2568 


Grindelia acutifolia Steyern 
Grindelia arizonica var. Lon 
Steyerm. 


WURDACK, J. J. (John Julius) 
911 


961 


SORAA parvifolia (a aliji 


Perama wurdackii Steyern 


34225 


34262 


39784 


39810 
39823 


39858 


Rd 0 


43762 


UNCKER, T. G. 


Hippotis latifolia Steyerm. 

Hippotis wurdackii Steyerm. 

Pentagonia tw urdackü Suis 

Simira wurdackii Steye 

Pentagonia parvifolia Sdn 

Eupatorium ibaguense var. pam retum 

laguire, Steyerm. & Wurd. 

Sauvagesia iban subsp. "a 
tensis Maguire, Steyerm. & Wurd. 

Chimantaea eriocephala Maguire, Stey- 
erm. & Wurd. 

Min an heterodoxa var. exappen- 
diculata Maguire & Steyerm. 

Alibertia latifolia var. pargueniana Stey- 
erm. 

Gouania wurdackii Steye 

Sipanea pratensis var. dic e f. gla- 
rior Steyerm. 

Sunira erythroxylon var. 
erm. 

"duis s m 

Sloanea wurdac 

Toulicia aber [e erm. 

Randia venezuelensis Steyerm. 

Mitracarpus frigidus var. orinocensis 
Steyerm. 

Borreria confertifolia Steyerm. 

Ouratea wurdackii Maguire & Stey- 


saxicola Stey- 


erm. 
Rudgea wurdackii Steyerm. 


1. 
Sipanea ned var. diehiotems f. gla- 
briloba Steyerm. 
Borreria bun Steyerm. 
Coussarea revoluta Steyer 
Schefflera tamatamaensis TEN 
er 


Stey 

Schradera hillifolia voli 
Capirona wurdackit 
Perama galioides var. longiflora Stey- 

erm. 
Euphronia acuminatissima Steyerm. 
(Truman George) 
Erblichia standleyi Steyerm. 
ceu qa JAHR eum 


ndl. & Ste 


778 Annals of the 
Missouri Botanical Garden 


APPENDIX III. Family Index. Numbers of taxa described by Steyermark arranged alphabetically by family. 


Family Genera Species Subspecies Varieties Forms Miscellaneous 


9 T E = > 


an 
N 


Acanthaceae 
Actinidiaceae 


Amaranthaceae 
Amaryllidac eae 
Apocynaceae 
Aquifoliaceae 
Araliaceae 


recacea 
Aristolochiae eae 


2 

yn 

= 

= t 
4 

o 

£ 

en 


Bignoniaceae 
Bom e 


[d — 
N 
No) 
| 
_ 


Boraginaceae 
Brassicaceae 

Bromeliaceae 
Brunelliaceae 


haleel wl! 
| 
| 


NPE Bw 


Burmanniaceae 
Burseraceae 
Cactaceae 
Campanulaceae 


N 


Canellaceae 
Capparidaceae 
Caprifoliaceae 
Caryophyllaceae 
Cecropiaceae 
Celastraceae 


w -n AeA 0 —-—tb€:20 tr) 
| 
| 
| 
| 
| 


— 


Chrysobalanaceae 
Clethraceae 


e 
e 
£e 
[v] 
[c] 
poo] 
c 
— 
-— N 
— Oe 
| 
| 


Commelinaceae 


NOK Dm- wer A NR 


Connaraceae 
Convolvulaceae 
Crassulaceae 
Cucurbitaceae 
Cunoniaceae 
Cupressaceae 
Cyperaceae 


Dichapetalaceae 


— — c to € 
lowon 
| 
| 
| 


— 
a 
© 
Ww 


Dilleniaceae 


ON be 
| 
| 
| 
| 


Dioscoreaceae 
ipsacacea 


| | 
— | 

| - 
| | 


Dipterocarpaceae 
roseraceae 
Ebenaceae 
oe 
Elatina 


eo 


| ed 
N 

mee ww 
| 

sd] 
| 


— 
Ww 


Volume 76, Number 3 
1989 


aylor 


Tay 
Plants Described by 
Julian A. Steyermark 


779 


APPENDIX III. Continued. 

Family Genera Species Subspecies Varieties Forms Miscellaneous 
Eriocaulaceae 1 3 — — — — 
Erythroxylaceae l l — — — 
Euphorbiaceae 20 54 — 1 — 3 
Fabaceae 20 23 = = 4 1 
Fagaceae l — — - 1 3 
Flacourtiaceae 0 10 — - — l 
Garryaceae l l = — o 
Gentianaceae 10 27 — 1 1 
Gesneriaceae 2 2 — — — — 
Grossulariaceae 11 — — — — — 
Gunneraceae l l — — -— pu 
Heliconiaceae l l — — — -— 
Humiriaceae l l - — = 
Hydrophyllaceae 2 — — 2 n 
Icaciniaceae 2 2 — - — 
Iridaceae 2 2 — 3 l — 
Ixonanthaceae l 5 — 2 — 
Juglandaceae l = l — 
Lamiaceae l — — — l — 
Lauraceae 4 7 — l — — 
Lecythidaceae l ] — — — - 
Lenno e l l — = 
L fla 'eae 3 29 — 2 3 -— 
Liliaceae 6 4 — l 5 l 
Limnocharitaceae l l = = = — 
Lissocarpaceae l 1 — = = = 
Loasaceae 3 3 — — — -- 
Loganiaceae 2 5 ms E A — 
Loranthaceae 3 12 — - — 
Lythraceae l — = 1 — — 
Magnoliaceae l 3 = = — 
Malpighiaceae 9 12 -— l — l 

alvaceae 4 8 -— — = = 
Marantaceae 4 8 — c — - 
Marcgraviaceae 3 4 — l -= — 
Melastomataceae 6 10 — — = 
Meliaceae 2 9 — = = = 
Mendonciaceae l 2 - = 
Me iie eae l 2 — — — = 
Monimiaceae 2 7 — = 
Manus 2 2 — — — -e 

yricaceae ] l - — — — 
M iniri 5 12 = ] — 
Mvrtacea 10 39 — -— — — 
Nyc ena 4 30 - — — n 
Ochna 6 38 3 7 — 7 
aa 4 5 — -— — = 
Oleacea 3 5 — 10 = — 
Onagraceae 3 2 = 2 l — 
Orchidaceae l l — — — = 
Oxalidaceae l 2 — - -— = 
Passifloraceae 2 3 — — — 
Piperaceae 3 39 2 10 13 — 
Poaceae l — — l m 
Polemoniaceae l — — - l - 
Polygalaceae l 4 = l = — 


780 


Annals 


of the 
Missouri Botanical Garden 


APPENDIX III. Continued. 

Family Genera Species Subspecies Varieties Forms Miscellaneous 
Portulacaceae 2 3 — — — 
Primulaceae l — — l — 
Proteaceae 3 13 — l — — 
Quiinaceae l l — = 
Ranunculaceae 4 3 — — 3 — 
Rapateaceae 3 13 = - — — 
Rhamnaceae 3 9 — — — 3 
Rhizophoraceae l 5 l — 2 
Rosaceae 3 l — l 7 — 
Rubiaceae 93 555 29 113 46 49 
Rutaceae 11 16 — l 3 1 
Sabiaceae l 5 — — - 

Salicaceae l -- — 3 — 
talacea l l — — 
Sapindaceae 9 35 2 l — — 
Sapotaceae 2 5 — — 
Sarraceniaceae l l — l 2 — 
Saxifragaceae l = — = l — 
Scrophulariaceae 5 3 — — 2 -— 
Selaginellaceae l l = — 
Simaroubaceae l 2 — l — 
Smilacac l 6 — l — 
Solanaceae 8 27 — l — 
Staphyleaceae l l -— - — 
Sterculiaceae 2 2 — — - — 
Styracaceae l 2 — — 
Symplocaceae l 11 — 
Taxodiaceae l — l — 
* Tepuianthaceae l 6 1 — - — 
Theaceae 4 13 2 3 2 
Lion slaeaceae 2 7 — 
Tiliac l 2 = 1 
Trigoniac eae l 2 - = 
Turneraceae l l — l — 
Urticaceae 3 3 — l 
Valerianaceae l 2 — — 
Velloziaceae l l — -— 
Verbenaceae 3 4 — — 
Violace: 4 3 — 2 1 
Viscaceae l 9 — — - 1 
Vitaceae l l — E 
Vochysiaceae 3 4 — l E — 
Winteraceae l — l — 
Xyridaceae 3 11 = 2 z 
Zingiberaceae 2 3 = 


SYSTEMATICS OF ZAPOTECA Héctor M. Hernández? 
(LEGUMINOSAE)! 


ABSTRACT 


Zapoteca (Leguminosae: Mimosoideae) is a si ies well- Veces genus belonging to the tribe Ingeae 
distributed from the southwestern United States and northern Mexico to Piin Argentina and the West Indies. 
The highest concentration of taxa occurs in the dry ben of so aod n Mexico and, to a lesser extent, in Central 
America and the northern Andes. The predominance of allopatric distributions and the apparent low frequency of 
hybrids suggest that r in this genus has primarily arisen from genetic differentiation of a isolated 
populations. The recent climatic episodes of the dulce appear to m had a prominent role on the current 
distributional patterns of Zapotec a species. Zapoteca species have similar reproductive biologies. All probably have 
nocturnal anthesis and are pollinated by ad settling moths. Self-compatibility appears to be widespread in 
the genus, Z. tetragona being the only known self-incompatible species. Andromonoecy is another widespread 
component of the breeding system of Zapoteca species. This probably maximizes the use of reproductive energy 
without reducing pollinator attraction. The basic chromosome d of Zapoteca (x = 13) and some of the generalized 
characters of the polyads (i.e., acalymmate, discoid, 16-grained polyads) indicate a relationship between Zapoteca 
and the remaining genera of the Ingeae, with the exception of Calliandra sensu stricto, which is cytologically and 
morphologically isolated in this tribe. I recognize 17 species in Zapoteca, three of which (Z. alinae, Z. andina, and 

ana) are newly described. Eleven subspecies are recognized, of which Z. portoricensis subsp. pubicarpa is 
newly described. A subgeneric classification is proposed, where taxa are grouped into four morphologically distinct 
subgenera: Nervosa, Zapoteca, Aculeata, and Amazonica. The subgenera are based primarily on differences in 
vegetative characters: number of pairs of pinnae, number of leaflet pairs per pinna, leaflet size, leaflet texture, venation 
patterns, and stipule type. 


RESUMEN 


Zapoteca (Leguminosae: Mimosoideae) es un género relativamente bien delimitado que pertenece a la tribu Ingeae. 
Se distribuye desde el suroeste de los Estados Unidos y el norte de México hasta el norte de Argentina y las Antillas. 
La más alta concentración de taxa se da en los bosques tropicales secos del sur de México y, en menor medida, en 
Centro América y en la porción norte de los Andes. La predominancia de distribuciones alopátricas, en combinación 


' This study is part of my doctoral thesis at Saint Louis University, Saint Louis, Missouri. I thank Dr. Peter H. 
Raven, Director of the Missouri Botanical Garden, for directing o research and for support and bp DUE UA in 
all stages of the pa The Missouri Botanical Garden provide ce space and herbarium, library greenhouse 
facilities. The assistance of the staff and technicians of the Missouri Botanical Garden was Sanii pee is gratefully 

dged 


I also thank Drs. Rupert Barneby, Gerrit Davidse, John Dwyer, Alwyn Gentry, Mario Sousa, ME an i 
reviewer for reading the manusc ript and providing eel ent and editorial advice; Ph. Gui . Kea 
pt . Lowry, and B. Stein for critically reading parts of the n J. Dwyer for duse ia the lad 

aia of the new taxa; C. Beutelspacher for identifying the flower visitors; J. Myers for his excellent botanical 
illustrations; F. e for Figure 44; M. Veith and M. Reyes for Ae a in the use of the SEM at Washington 
University and Instituto de Geología, UNAM, respectively; and D. Thiel for typing part of the m Ia 
especially grateful to the following persons who provided seed paas and other botanical material: M. 
Delgado, C. Dodson, E. Forero, C. Hughes, C. Johnson, J. Lewis, E. Lott, J. Miller, D. Neill, J. Oakes, P. E 
D. Smith, W. Stevens, R. Torres, J. Vassal, T. Wendt, and T. Zanoni. I also thank the curators of the following 
he rbaria for providing specimens: A, BM, BR, CAS, CGE, ENCB, F, G, GH, GOET, K, LE, MA, MEXU, MICH, 

P, S, TEX, UC, US, WIS, and XA 

d studies would noi have been possible without the financial assistance of the Missouri Botanical Garden and 
World Wildlife Fund U.S. (Tropical Botany mia a I thank the following piis who provided assistance in 


the field: M. Arredondo, S. Arredondo, E Ch rchil, M. Correa, M. Grayum, P. Gra , L. Hernández, D. Janz 
P. Moreno, A. Ramirez, A. Shilom Ton, ^p. Stevens, P. Tenorio, and R. Torres. 

The five-year stay in Saint. Louis, Md was made possible for my family and me by a sc holarehip. os ided by 
the Instituto de Biologia, Universidad Nacional Autónoma de México, through the Departamento de Becas. I express 


sincere gratitude to Dr. José Sarukhán K., then director of the Instituto de Biología. and Dr. Mario tum S., former 
curator of the Herbario Nacional de México (MEXU) for their confidence and continuous support. Thanks are extended 
to the Missouri bind Garden for partial financial support during our last year in Saint Louis. dapi my deep 
E to my wife Alina and to Pablo for their love and emotional support Bu this long experie 

ssouri joe Garden, P.O. Box 299, Saint Louis, Missouri 63166, U.S.A. Present m Herbario 
Nac pe de México, Instituto de Biologia, une Nacional pus ola de México, Apartado Postal 70-367, 
Ciudad Universitaria, 04510, Mexico, D.F., Méx 


ANN. MISSOURI Bor. GARD. 76: 781-862. 1989. 


782 Annals of the 
Missouri Botanical Garden 


n la aparente baja frecuencia de hibridos, sugiere que la especiación en este género ha sido el resultado de u 
BOX de diferenciación genética de poblaciones aisladas geográficamente. Los episodios climáticos recientes i 
Pleistoceno posiblemente tuvieron un papel determinante sobre los patrones de distribución actuales de las especies 
de Zapoteca. Las especies de Zapoteca son bastante uniformes en los rasgos generales de su biología ra 
Todas tienen posiblemente flores con antesis nocturna y son polinizadas por palomillas no especializadas. Los sistemas 
de auto-compatibilidad genética parecen estar bién representados en el género, Z. tetragona siendo la única especie 
en donde la auto-incompatibilidad ha sido demostrada. La andromonoecia es otro componente común de los sistemas 


reproductivos de las especies de Zapoteca; este fenómeno se a como un medio para maximizar el uso de la 
energi ia reproductiva, sin re 'ducir la atracción a los polinizadores. El número cromosómico básico de Zapoteca (x = 


3), en asociación con algunos caracteres generalizados de n políades (ej., políades acalimadas, discoides, con 
granos), reflejan relaciones entre Zapoteca y el resto de los géneros de la tribu Ingeae, con la excepción de Calliandra 
sensu stricto, género que es fuertemente atípico desde el punto de vista citológico y morfológico. En este trabajo se 


para la cier ‘ T 
como nueva. En base a los patrones de variación en caracteres vegetativos (ej., número de pares de pinnas, nümero 
res de foliolos por pinna, tamano de los n textura de los foliolos, patrones de vanación de los foliolos y 
tipo de estípulas) se propone una clasificación subgenérica, en donde las especies se agrupan en cuatro subgéneros 
distintos: Nervosa, Zapoteca, Aculeata y Amazonica. 


o 


Zapoteca (Leguminosae: Mimosoideae) is a ge- Laetevirentes. Zapoteca and Calliandra are com- 
nus of 17 species in the relatively specialized tribe pared in Table 1. 
Ingeae. Its species are mostly erect or scandent I consider all the species included by Bentham 
shrubs, of small to moderate size, found in vege- (1875) in Calliandra ser. Laetevirentes to be Za- 
tation types ranging from arid, scrubby thorn for- poteca. When I originally proposed Zapoteca 
ests to wet, evergreen tropical forests, althoug (Hernández, 1986a), the possibility of including 
most species occur in areas originally covered by Calliandra amazonica and C. aculeata, treated 
tropical deciduous forests. Zapoteca ranges from by Bentham in ser. Macrophyllae, was discussed. 
southwestern United States and northern Mexico Since these two species share a number of repro- 
to the West Indies and southward through most of | ductive characters with Zapoteca (1.e., compact, 
Mexico, Central America, and part of South Amer- spherical, homomorphic capitula; 16-grained dis- 
ica, to northern Argentina. It attains its greatest coid polyads with lens-shaped thickenings; elasti- 
diversity in southern Mexico, although in some cally dehiscent fruits with relatively thin. valves), 
areas of Central America and the northern Andes they are transferred in the present work to Za- 


diversity is also considerable. poteca. The stipular spines of Z. aculeata, the 

Zapoteca was recently separated from the P compound pseudopanicles of Z. amazonica, and 
genus Calliandra Benth. (Hernández, 1986a the small number and larger size of the leaflets in 
tensive analyses of the polyad characteristics, Res both species are interpreted as advanced charac- 
ling morphology, chromosome numbers, and sev- teristics. These species should not be considered 


eral reproductive features in Calliandra sensu by any means as intermediate between Zapoteca 
Bentham revealed differential variation patterns and Calliandra; they are specialized species within 
among its neotropical species. These variation pat- Zapoteca, with divergent morphological charac- 
terns were associated with two distinct taxonomic ters. 

groups: the first, including the members of Ben- Zapoteca is defined by its compact, spherical, 
tham's series Laetevirentes, and the second, the  homomorphic capitula; acalymmate, 16-grained, 
remaining neotropical species, these belonging to discoid polyads with lens-shaped thickenings (ex- 
series Macrophyllae, Nitidae, Pedicellatae, and cluding Z. nervosa); elastically dehiscent fruits with 
Racemosae. The strongly divergent nature of these membranous or coriaceous valves; and basic chro- 
two groups, along with the absence of intermediate mosome number of x — 13. This combination of 
forms between them, suggested that Calliandra, characters makes Zapoteca a well-cireumscribed 
as originally conceived by its author, is polyphyletic entity; however, with the exception of the distinc- 
(Hernández, 1986a). In an accompanying paper tive circular thickenings in the polyads, each of 
Romeo (1986) provided supporting evidence based — these characters occurs independently in other gen- 
on differing patterns of nonprotein imino and sul- era of tribe Ingeae. The general characteristics of 
phur amino acids in seeds and leaves of Zapoteca the Zapoteca polyads, its basic chromosome num- 
and Calliandra. Consequently, the new genus Za- ber, and some of its morphological characters re- 
poteca was proposed to include the species of series — flect close relationships of this genus with some 


Volume 76, Number 3 Hernández 783 
Zapoteca 


TABLE 1. Comparison of Zapoteca and Calliandra (from Hernández, 19864). 


Zapoteca Calliandra 
Leaflets thin membranous, rarely coriaceous chartaceous to coriaceous 
Inflorescences spherical heads; homomorphic obconiform, racemose, rarely spherical: 
homomorphic or heteromorphic 
Polvads l6-grained, discoid 8-grained, bisymmetric, with viscid aj 
pendage in “basal” grain 
Stigmas cup-shaped, with narrow area of expanded: discoid, or capitate 
led receptivi 
Pods membranous to coriaceous more rigid; coriaceous, ligneous, rarely 
membranous 
Seedlings cotvledons ovate, sessile. foliaceous, cotyledons sagittate, petiolate, fleshy, 
ephemeral persistent 
Chromosome numbers = 13 =8,11 


genera of the Ingeae, perhaps with members of the mately 3-4 m tall with woody stems. In most cases 


Pithecellobium complex; however, to establish def- — the stems are rather thin (ca. 1-2 em in diameter), 
inite generic affinities of Zapoteca is a difficult task — although occasionally individuals of Z. tetragona 
due to our poor current understanding about ge- and Z. portoricensis develop stems up to 20 cm 


neric delimitation within this complex. What seems in diameter. A few species are scandent or sub- 


clear is that Zapoteca has closer affinities with 


some of these genera than with Calliandra sensu TABLE 2. Outline of the classification of Zapoteca. 
stricto. Several characters in Calliandra (e.g., 
polvad features. chromosome numbers, floral mor- I. Subgenus Nervosa 
phology) are atypical in the tribe Ingeae and in- LZ nervosa 
dicate that this genus is evolutionarily divergent YE Sorens oiu 
from the other genera of the tribe. 9 > nei 
This study provides an account of the mor- 3. Zal 
phology, biogeography, reproductive biology, cy- à. ic 
togenetics, and taxonomy of Zapoteca. The taxo- 5. Z. filipes 
nomic revision is based on over 4,000 herbarium 6. Z. portoricensis 
specimens from the major collections of the world a. subspecies portoricensis 


and on extensive field observations. Field studies b. subspecies pubicarpa 


were conducted in most of Mexico, Nicaragua, 
Costa Rica, and Panama during 1983-1985. In 
addition, individuals representing eight species and 
five subspecies of Zapoteca were grown from seed 


subspecies flavida 
7. Z. mollis 
8. Z. caracasana 
a. subspecies caracasana 


b. subspecies weberbaueri 


in a greenhouse at the Missouri Botanical Garden 9. Z detramona 

during 1982-1986; these plants provided infor- bón. X andina 

mation for the introductory chapters and served LL. Z. costaricensis 

to complement the field observations and herbar- 12. Z. formosa 

ium studies when taxonomic decisions had to be a. subspecies formosa 


made. Based on the critical examination of cor- b. subspecies salvadorensis 
related morphological characters, in combination c. subspecies gracilis 
. i d. subspecies rosei 

with eco-geographical factors, I recognize l7 species i l 
; : e. subspecies mollicula 


and 11 subspecies in Zapoteca (Table 2). The f i . esti 
. . H % s SM species SCHO 

species are classified into four subgenera, which 13. Z. scutellifera 

represent different evolutionary lines. VE mess 


d ie RE III. Subgenus deuleata 
THE PLANTS Ve 


Z. aculeata 
VEGETATIVE MORPHOLOGY IV. Subgenus Amazonica 
; en . f : l6. Z. microcephala 
Habit. The species of Zapoteca are relatively / 
uniform in habit. Most are erect shrubs approxi- 


17. Z. amazonica 


784 


Annals of the 
Missouri Botanical Garden 


scandent. Zapoteca portoricensis subsp. flavida 
and Z. amazonica grow in moist thickets supported 
by neighboring shrubs. Similarly, Z. caracasana 
branches near the base of the primary stems and 
has spreading branches that become prostrate or 
subprostrate. The scandent and prostrate habits 
are found exclusively in warm, moist habitats. 

Leaves. The leaves of all species of Zapoteca 
are stipulate and twice pinnate with opposite leaf- 
lets. The wide variation in leaf characteristics pro- 
vides an excellent basis for classifying members of 
the genus. Mature leaflets vary in length from about 
0.5 em in Z. alinae and Z. media to a maximum 
of 22 cm in Z. amazonica. In addition, there is a 
clear tendency towards a reduction in number of 
leaflets concomitant with an increase in leaflet size. 
In all species of subg. Zapoteca the pinnae have 
from several to many pairs of leaflets; by contrast, 
in Z. amazonica the leaves have only one pair of 
pinnae with a single pair of large leaflets. Fur- 
thermore, all members of subg. Zapoteca have 
relatively membranous leaflets, whereas in the re- 
maining species they are clearly chartaceous to 
coriaceous. 

Extrafloral nectaries are uniformly present in Z 
filipes, Z. scutellifera, and Z. nervosa. They are 
located between the insertion of the pinnae and, 
less frequently, between the distal pairs of leaflets. 
In Z. scutellifera and Z. filipes an additional nec- 
tary is located near the base of the petioles. The 
nectaries are more or less cup-shaped in Z. scu- 
tellifera and Z. nervosa and cylindrical in Z. fi- 
lipes. According to Zimmerman's classification 
1983), all of these fall into the "Hochnek- 
category (elevated nectaries). Extrafloral 
nectaries have long been considered to be the focal 


(Elias, 


tarien 


point of a mutually beneficial relationship between 
the plants that possess them and certain groups of 
ants (see summary in Bentley, 1977). The ants 
exploit the nectar produced by these glands, which 
is rich in sugars, amino acids, proteins, and other 
1983) and provide the plant 
varying degrees of antiherbivore protection. 
However, such a relationship has not yet been 
demonstrated in Zapoteca. 


compounds (Elias, 
with 


lenation patterns. Despite the fact that leaves 


have numerous taxonomically useful characters, 
monographers generally limit themselves to super- 
ficial descriptions of the most conspicuous features 
. | describe the 
venation pattern of the leaflets of several species 


of leaf architecture. In this section 
of Zapoteca, in order to obtain a broad picture of 
the variation of this important component of the 
leaf architecture. 


Mature leaflets of ten species of Zapoteca, rep- 
resenting all the subgenera (Table 3), were cleared 
following Dilcher (1974). The terminology follows 
Hickey (1973, 1977). 

Variation in leaflet venation patterns is highly 
consistent with the subgeneric classification in Za- 
poteca and provides important information for phy- 
logenetic interpretation. All members of the genus 
The 


secondary veins usually depart from the primary 


essentially have brochidodromous venation. 


vein at an acute to right angle, curving abruptly 
to join superadjacent secondaries to form a loop. 
In addition, loops are always enclosed by arches 
formed by higher-order veins. Zapoteca nervosa 
differs from the remaining examined species in 
having eucamptodromous venation in the basal half 
of each leaflet and brochidodromous venation in 
the upper half (Fig. 30). Furthermore, the tertiary 
domains in the leaflets of this species are elongate 
and parallel to the primary vein, and the free- 
ending veinlets are more abundant in the distal 
area and usually are once- or twice-branched. 

The marginal venation is looped in all Zapoteca 
species with the exception of subg. Amazonica, 
where it is clearly fimbriated. Similarly, all species 
of the genus have incompletely closed areoles, ex- 
cept Z. amazonica and Z. microcephala, which 
have imperfectly developed areoles. 

With the exception of Z. tehuana, all species 
of subg. Zapoteca are characterized by relatively 
fine, inconspicuous tertiary and higher-order veins 
that are difficult to discriminate even under a dis- 
leaflets of Z. te- 


huana and all species of the other subgenera (Table 


secting microscope. By contrast, 
3) have thicker and conspicuous tertiary and qua- 
The vena- 
tion patterns of Z. tehuana (subg. Zapoteca) and 
Z. aculeata (subg. 


ternary veins and free-ending veinlets. 


1culeata) are strikingly similar, 
reflecting taxonomic affinities between the two sub- 
genera. These two species share the basic brochi- 
dodromous venation pattern of the genus; however, 
they differ in their marginal venation and in the 
orientation of their high-order veins and veinlets. 
Although the marginal venation of these two species 
is clearly looped, occasional free-ending marginal 
veins are present. The veinlets are characteristi- 
cally oriented towards the admedial-basal area of 
the leaflets in these two species. 

Zapoteca amazonica and Z. microcephala have 
several characters in their leaves that depart from 
the rest of the genus. The tendency towards the 
development of larger leaflets concomitant with a 
reduction in their number in these species is prob- 
ably associated with an increase in the degree of 
organization of leaflet venation. Hickey (1977) pro- 


Volume 76, Number 3 
1989 


Hernández 785 


Zapoteca 


TABLE 3. Characteristics of leaflet venation in Zapoteca species. 
Relative 
Thickness 
of 
Number Secondary 
of anc 
Primary Marginal Tertiary Areole 
Taxon Venation Type Veins Venation Veins Development Veinlets 
Subgenus Vervosa  eucamptodromous l looped moderate incompletely ] -2 branched 
basally and broch- closed 
idodromous dis- 
tally 
Subgenus Zapote: brochidodromous | looped hairlike incompletely ] -2 branched 
ca (excluding closec 
tehuana) 
Z. tehuana brochidodromous l looped with free- moderate incompletely admedially 
ending veins closec ramified 
dendroid 
Subgenus Aculeata  brochidodromous l looped with free- moderate incompletely admedially 
ending veins closed ramified 
dendroid 
Subgenus 4mazon- brochidodromous 2 fimbriated moderate imperfect dendroid 
ica 


Material examined. Subgenus Nervosa: Z. nervosa, Liman 284. Subgenus Lapel eca; Le dors rosa, 11-959, H-2 10. 
Q4. qu 


11-230. H-217: Z. tehuana, H-950; Z. alinae, H-95 


tetragona, H-956. Subgenus Aculeata: Z. act 


posed a system of leaf classification based on in- 
creasing levels of regularity of venation (leaf ranks) 
which provides a useful scale here. The venation 
patterns of the two species of subg. dmazonica 
reflect an advanced condition because their tertiary 
veins have relatively regular courses (third rank; 
see Fig. 48). In contrast, the leaflets of the re- 
maining species of Zapoteca fall in the second rank 
because their tertiary and higher-order veins are 
random in course and are usually not well differ- 
entiated from the lower orders. Of particular in- 
terest is the presence of two co-primary veins (Fig. 
48) in the leaflets of Z. amazonica and Z. micro- 
cephala. This probably indicates leaflet. fusion. 
Hickey & Wolfe (1975) reported that leaflet fusion 
is a common tendency in the Leguminosae. 

Leaf pubescence. With few exceptions, pu- 
bescence type is not a reliable taxonomic character 
in Zapoteca, because in most of the species it is 
clearly variable within and between populations. 
Zapoteca nervosa, Z. aculeata, Z. amazonica, 
and Z. microcephala appear to be uniformly en- 
tirely glabrous or nearly glabrous, whereas Z. mol- 
lis is consistently pubescent throughout. In the 
remaining species of subg. Zapoteca, pubescence 


; Z. mollis, 
des s ata, . Ast 
Woytkowski 5943; Z. microc ephala, Haus hi 1711 (F). € óllec tions in MO sud MN 'rwise indicated. 


11-793; 


amas ONO a, 


Grayum 120 . H-948, 


599, Seby Daae Z. 


is variable, reaching an extreme of inconstancy in 
Z. formosa and Z. portoricensis. A microscopic 
survey of cleared leaflets of 13 of the species re- 
vealed three basic types of hairs (Figs. | Their 
taxonomic distribution is shown in Table 
minology following Leelavathi & Ramayya, 1982). 
1) are the 


most abundant and are found at the base of leaflets 


(ter- 
(1) Unicellular conical hairs (Fig. 


and on the rachillas of Z. aculeata, along the leaflet 
margins of Z. microcephala, and on all leaf parts 
of all the pubescent species of subg. Zapoteca. Á 
very similar type of hairs, unicellular clavate (Fig. 
2), is ANS found on the rachilla of Z. alinae 
Table 


(2) ns ellular macroform conical hairs (Fig. 


— 


3) occur in all species of subg. Zapoteca except 
They 


mixed with other tvpes of hair, primarily on leaflet 


Z. formosa and Z. alinae. are found, inter- 
margins and rachillas. 

(3) Filiform capitate hairs and filiform clavate 
hairs (Fig. 4) usually have a uniseriate cylindrical 
foot formed by a variable number of cells of varied 
lengths, and a multiseriate (or rarely uniseriate?) 
head. These are usually restricted to leaflet margins 
and rachillas, and are found in all species of subg. 


Zapoteca except Z. formosa. 


786 Annals of the 
Missouri Botanical Garden 
TABLE 4. Distribution of types of leaf trichomes in selected species of Zapoteca. la = unicellular conical, Ib = 


unicellular clavate; II = mu 


Iticellular macroform conical, HI = filiform capitate or filiform clavate. Relative abundance 


is indicated by: + = occasional, ++ = regular, + ++ = abundant. All collections in MO unless otherwise indicated. 
H = author’s collections. 
Type 
Taxon Collection la Ib Il IH 
Subgenus Nervosa 
Z. nervosa Ekman 284 
Subgenus Zapoteca 
Z. alinae H-949 + B + 
Z. formosa H-210 +++ 
Z. formosa 11-959 
Z. formosa H-217 +++ + 
Le ales H-230 oed 
Z. formos Sousa 25€ +++ 
Z. lambertiana A = 412 ++ - 4 
Z. media H-951 +++ ++ ++ 
Z. mollis Grayum 4201 +++ ++ ++ 
Z. portoricensis H-94 +++ ++ ++ 
Z. portoricensis subsp. flavida H-793 +++ + ++ 
Z. tehuana H-950 FHF + + 
Z. tetragona 1-956 ++ ++ ++ 
Subgenus Aculeata 
Z. aculeata 4costa 13599 i 
Subgenus amazonica 
Z. amazonica Il oytkowski 5943 
Haught 1711 (E) ++ 


Z. microcephala 


Stipules. The stipules are persistent, leafy, 
and possess a series of veins running parallel to 
the margins (Figs. 5, 6); in most species the stipules 
are conspicuous (e.g.. Z. amazonica, Z. tetra- 
gona), although in some they are very small (e.g., 
Z. tehuana). Exceptional in this respect is Z. acu- 
leata, which has spinescent stipules (Fig. 46), and 
Z. nervosa, whose stipules are reduced to small 
(< 1 mm)sc 


are adpressed and probably have a protective func- 


ales. Throughout the genus the stipules 


tion during the early phases of leaf and inflores- 
cence development. Zapoteca andina is excep- 
tional by having descending or depressed stipules 
(Fig. 41). Moreover all species have stipels at the 
base of the distal pairs of pinnae and at the distal 
tip of every pinna. 


INFLORESCENCES AND FLOWERS 


Inflorescences. The flowers are consistently 


arranged in compact, homomorphic, spherical, 


capitate inflorescences. In the basic plan of inflo- 
rescence organization, solitary or fasciculate pe- 
duncles are axillary and subtended by a leaf, as in 


Z. alinae, Z. media, and Z. aculeata (Fig. 7A). 


indication of the derived character of the 


There is, 
the progressive reduction of the subtending leaves, 


however, an evolutionary trend toward 


which reaches its extreme in the two species of 
subg. Amazonica, where the leaves are always 
vestigial, and the inflorescence therefore appears 
to be an elongate panicle (Fig. 7D, E). Transitional 
forms between the two extremes are found in 
several species of subg. Zapoteca (Fig. 7B). For 
instance, in Z. mollis, Z. portoricensis, and ad- 
ditional species, the subtending leaves start deve- 
loping during or after anthesis. Sometimes, how- 
ever, the subtending leaves never develop, as in 
some individuals of Z. caracasana (Fig. 7C) and 
Z. formosa, in which the capitula are apparently 
aggregated in paniclelike inflorescences 
The trend towards retardation and suppression 
of leaf development that results in differences in 
foliation among the species of Zapoteca has also 
produced various patterns of inflorescence orga- 
nization. | interpret these types as steps in a con- 
tinuum of inflorescence specialization rather than 
as discrete and fundamentally different types. An 
"panicu- 


late" inflorescences of Z. microcephala, Z. ama- 


Volume 76, Number 3 Hernández 787 
1989 Zapoteca 


FIGURES 1-4. Leaf trichomes of Zapateca l. Unicellular conical, Z. alinae (H. Hernández 952).— 2. 
K ellular clavate, Z. alinae (H. Hernández 952). 3. Multi :ellular macroform conical, Z. mollis (Grayum 4201). 
. Filiform capitate, Z. tetragona (H. Hernández 950). All collections at MEXU and MO. Scale bars = 100 um. 


ie In summary, there has been an evolutionary 


— 


zonica, and other species in subg. Zapoteca is 1 
presence of persistent paired stipules at every node trend of inflorescence modification in Zapoteca, 
where retardation and suppression of the subtend- 
ing leaves has played a prominent role in changing 
the overall aspect of the flowering system. The 


axillary capitula subtended by a developed leat 


along the flowering system. These inflorescence 
types, therefore, cannot be considered strictly pan- 
iculate, and for this reason the term "pseudopan- 


icle" will be used. 


FIGURES 5, 6. Stipules of Zapoteca. —5. Z. portoricensis subsp. portoricensis (H. Hernández 948). —6. Z. 
tetragona (H. Hernández 956). Collections at MENU and MO. Scale bars = 1 cm. 


Annals of the 
Missouri Botanical Garden 


NS | 
\ ? 3 — 
NA 
y Y e 
M ET | ri 
O N E 
bla]; > 
IDA 
Sf ze) 
2 | -oQ 
E JR 
O 
) Eg a 
Rey 
| > 
eN A D 
| Vee 
V 
| | / 
| 
/ 
| / 
| 
| 


D 


FIGURE 7. Patterns of inflorescence ee in dapotéca .—A. 
a well-developed leaf, Z. alinae (Sousa et al. 7846, MO). 

Z. portoricensis (Higgins 2578, MICH). —C 
15982, LL) 


e capitula at each node are subtended by 


stigial. C. Z. caracasana (Asplund 
. D. Z. microcephala (Mutis 3 3800, US). E. Z. amazonica (Sehunke 6913, MO). 


Volume 76, Number 3 Hernández 789 
1989 Zapoteca 
(leafv, axillary capitula) appear to represent the in all species of Zapoteca except Z. nervosa. In 


basic pattern. The paniclelike inflorescences of Z. 
microcephala and, especially, Z. amazonica (subg. 
Amazonica) are highly modified and are clearly 
derived from the basic type. Intermediate types 
exist in many species of subg. Zapoteca, where a 
single individual may have both leafy, axillary ca- 
pitula and flowering branches with vestigial or late- 
developing subtending leaves. 


Flowers. | Aside from variation in size and in 
the relative proportions of their parts, the flowers 


of Zapoteca species are extremely constant. De- 


tails of floral morphology and certain aspects of 


floral biologv are in the generic and specific. de- 
scriptions and in the section on reproductive bi- 
ology. 


POLLEN 


The diversity of pollen types in subfamily Mi- 
mosoideae has been studied extensively by several 
authors (Guinet, 1969, 1981— a review; Sorsa, 
1969; Niezgoda et al., 1983). One of the outstand- 
ing features in this subfamily is the high frequency 
of permanently compound grains, these either in 
tetrads or more commonly in polyads. As Guinet 
(1981) and others pointed out, the basic polyad 
type in the Mimosoideae is the acalymmate type, 
in which the individual grains are more or less free 
from one another. This form of polyad cohesion is 
in contrast to the calymmate type, where the ex- 
ternal layer of the exine is common to all cells. 
Calymmate polyads are found in Calliandra. 
1986a), 
several comprehensive surveys on the pollen char- 
acteristics of Calliandra (Guinet, 1965, 1969, 
1981; Guinet & Barth, 1969; Sorsa, 1969; Niez- 
goda et al., 1983) indicate the phyletic distinctness 


As discussed elsewhere (Hernandez, 


of Zapoteca. The basic plan of polyad organization 
here is the same as that in the other genera of 
tribe Ingeae, but unlike that of Calliandra sensu 
stricto. All the species in the genus have acalym- 
radially symmetrical, l6-grained 


mate, discoid, 


polyads (Figs. 8-13). The eight central grains, 
which are distributed in two rows, are morpholog- 
ically different from the peripheral grains (i.e., the 
pollen grains exhibit heteromorphy). The external 
layer of the exine has an undifferentiated granular 


structure, and the individual grains is al 
apertures (see fig. 7 83). 


The 


size of the polyads ranges from 76 to 2 um. 


in Niezgoda et al., 

One very striking diagnostic character, found no- 
where else in Mimosoideae, 
prominent lens-shaped structures in the. central 


cells on one side of the polyad. These are found 


is the occurrence of 


connection with this, Guinet (pers. comm., 1985) 
observed that the rounded lens-shaped areas occur 
on both sides of the polyads (on eight grains) in Z. 
aculeata and Z. amazonica; nevertheless, they 
are less pronounced on one side. Such areas some- 
times occur on some or all of the peripheral grains 


of Z. alinae polyads (Fig. 9 


PODS, SEEDS, AND SEED DISPERSAL 


Pods. 
throughout Zapoteca (Figs. 14, 15). The pods are 


Pod morphology is relatively. constant 


usually straight, linear, and flattened: they gen- 
erally have unconstricted margins, visible seed 


chambers, and thickened sutures (terminology fol- 
1984). Pod size varies from 3.5 x 


0.7 cm in Z. formosa subsp. mollis to 24 x 1.2 


lowing Gunn, 
em in Z. amazonica. There is some interspecific 
variation in the texture (density) of the valves, 
which is of taxonomic value. In this respect, the 
species may be arranged into three groups: 1) Le 
lambertiana and Z. filipes with thin, membranous 
valves; 2) the remaining species of subg. Zapoteca, 
along with Z. nervosa, with thick, membranous 
valves; and 3) Z. aculeata, Z. microcephala, and 
Z. amazonica, with coriaceous valves. [t is im- 
portant to mention that the terminology of pod 
texture used here is relative, and as such its use 
leaflets). 


The pods are glabrous except those of Z. cara- 


is not applicable to other structures (e.g., 
casana subsp. weberbauert, Z. mollis, Z. porto- 
ricensis subsp. pubicarpa, and Z. formosa subsp. 
salvadorensis, which are short-pubescent, densely 
villous, puberulent, and pilose to densely pilose 
respectively. 

Seeds. In all species of subg. Zapoteca, the 
seeds are widely ovoid to rhomboid and compressed 
and lack rupture lines, arils, or wings (Figs. 16, 
17). In most members of this subgenus the testa 
is mottled, or dichrome, in which case the area 
encompassed by the pleurogram (areola) has a 
lighter color than the outside area. There is con- 
siderable interpopulational variability in seed size 


and the coloration of the testa, especially in Z. 


formosa; by contrast, Z. tetragona appears to be 


consistent in having dark, monochrome, ovate seeds. 
Another seed feature is the pleurogram, common 
1984). Seeds of 
subg. Zapoteca usually have irregular, 90% pleu- 


in subfamily Mimosoideae (Gunn, 


rograms (sensu Gunn, 1984), although those : FA 
The seeds of Z. 


amazonica differ from those of subg. Puis in 


tetragona are always regular. 


being narrowly rhomboid to narrowly elliptic, larg- 
er, and in lacking the characteristic pleurogram. 


790 Annals of the 
Missouri Botanical Garden 


FIGURES 8-13. Unacetolvzed 16- grained polyads of Zapoteca species. —8. Z. media (R. Fernández 3). 
9. Z. alinae (H. Hernández 206); one of the periphe = grains has the le 2ns-shaped structure charac te ristic a the 
12 


central grains. — 10. Z. lambertiana (H. Hernández 14 11. Z. tehuana (H. Hernández 950). . tetragona 
(H. Hernández 956). — 13. Z. caracasana subsp. "M us (Johnson 2494, MO); both = n the polyad are 
shown. All collections at ME XU unless otherwise indicated. Scale bars = 10 um Figures 8 ; 50 um Figures 
11-13. 

Seed dispersal. | Zapoteca pods dehisce ex- species of Zapoteca over land and is probably 


plosively with the valves splitting along both sutures effective over larger distances as a result of ac- 
from apex to base (Fig. 15). This is probably the cumulative short-distance dispersal events over 


prevalent short-range die ecl mechanism for | many generations. 


Volume 76, Number 3 
1989 


Hernández 791 


Zapoteca 


FIGURES 14, 15. 
Scale bars = 2 cm 


This dispersal mechanism, however, does not 
explain the presence of species such as Z. formosa, 
Z. caracasana, and Z. portoricensis on islands, 
which they presumably reached by long-distance 
dispersal over water. Zapoteca formosa, for in- 
stance, occurs throughout the West Indies and on 
some islands off the Mexican Pacifice coast (Revilla- 
gigedo Islands). Populations of this variable species 
frequently grow in coastal areas. Although exper- 
iments have shown that seeds and mature pods of 
Z. formosa subsp. formosa taken from a coastal 
population of the Yucatán Peninsula floated for 
only two to four days in seawater, the seeds are 
highly resistant to saline water. As shown in Figure 
18, seeds from this population (Hernández 940) 
germinated after soaking in saline. water (3.4% 
sodium chloride) for up to 70 days. Most of the 
seeds germinated normally when they were me- 
chanically scarified and transferred to petri dishes 
in a medium moistened with fresh water. Ridley 
(1930) presented evidence that seeds and pods may 


Pods of Zapoteca tetragona (H. Hernández 956). —14. Indehisced pod. — 15. Dehisced pod. 


wai for considerable distances floating on de- 


E 


. The great resistance to salinity. of the seeds 
of Z formosa, and presumably of Z. caracasana 
and Z. portoricensis as well, suggest that these 
species dispersed by flotation from the continental 
mainland. Further discussion about the distribution 
and dispersal of Z. formosa is provided below. 


SEEDLINGS 


The taxonomic importance of seedling charac- 
ters has been recognized by legume researchers 
for a long time (De Candolle, 1825; Compton, 

1912). Recently, such characters have been ap- 
plied | successfully in the construction of generic 
classifications in Leguminosae 
(Léonard, 1957; Vassal, 1969; 
der Schiff, 1971; Kupicha, 1977; Guinet et al., 

980: and others). The distinctive seedlings of the 


Robbertse & van 


species formerly assigned to ser. Laetevirentes in- 
dicated the heterogeneous nature of Calliandra 


Annals 
Meus a Garden 


FIGURES eds 
Norico du La ; 948 í 
500 um. 


(sensu Bentham) and provided evidence for rec- 
ognition of Zapoteca (Hernández, 19802). 

See d of 19 populations representing eight 
X Zapoteca, all belonging to subg. Za- 
poteca, were studied (Table 5). About 10 me- 
chanically scarified seeds per sample were germi- 
nated in petri dishes. After one to three days, when 
the radicles hzd emerged, the germinated seeds 
were transferred to individual pots. The seedlings 


were described following Duke & Polhill (1981). 


spe C les 


or Vue species showing the irregular pleurograms. 
17 


. Z. formosa (Torres 


S 
= 100 
[^^] 
o e 
o 
o 
16) 
o 60 
c 
2 
S 
= 
= 20 
h 
o 
D 
20 40 60 
days 
IGURE 18. Resistance of ra formosa subsp. 


is seeds to saline water (3.4% sodium chloride). 
e bars indicate percent of p germinated seeds. Ten seeds 


were tested at each time interval. 


portoricensis subsp. 


A 
4063). Collections at — 2 MO. Scale bars — 


The principal characters used in the descriptions 
were: 1) position (whether epigeal, geal, 
geal), morphology, texture, and degree of persis- 
2) type (whether pin- 


or hy po- 


tence of the cotyledons, and 
nate or bipinnate) and sequence (whether opposite 
or alternate) of the eophylls. 

These samples were remarkably uniform, the 
seedlings of each species examined being essentially 
identical. Diagnostic characters at the species level, 
such as the square stems of Z. tetragona or the 
ovate leaflets of Z. formosa, became manifest above 
the third eophyll, as the first metaphylls arose. A 
brief description of the seedlings listed in Table 5 
follows (Figs. 19, 20) 
Germination phaneroepigeal; cataphylls absent. 
Cotyledons sessile, elliptic to elliptic-ovate, folia- 
ceous, ephemeral (remaining functional for ca. one 
week). Leaves stipulate, glabrous; first two eophylls 
opposite and pinnate, the pinnae with 12 lan- 
ceolate to narrowly oblong leaflets; third eophyll 
alternate, bipinnate, with | pair of pinnae; pe 
eophyll alternate, bipinnate, with 1, rarely 2 pairs 
of pinnae. 
although the seedlings of subg. 
Vervosa remain un- 


In summary, 
Aculeata, Amazonica, and 
known, the seedlings that have been observed so 
far, all in subg. Zapoteca, display a high degree 
of inter- and intraspecific morphological homoge- 


neity. 


Volume 76, Number 3 Hernández 793 
1989 Zapoteca 


FIGURES 19, 20. Seedlings of Zapoteca. —19. Five-day-old seedling of Z. tetragona (H. Hernández 956) 
20. Three-week-old p vog of Z. portoricensis subsp. portoricensis (H. Hernández 948). Collections at MEXU and 
MO. Scale bars = m. 


PATTERNS OR MORPHOLOGICAI lines, which correspond to the four subgenera pro- 
DIFFERENTIATION posed in the systematic treatment. 
There are several morphological features com- Subgenus Nervosa comprises Z. nervosa, which 


mon to all Zapoteca species. Among the most appears to be morphologically the most isolated 
important are: brochidodromous venation patterns species in the genus. Superficially, this species ap- 
in leaflets; persistent, leafy stipules (excluding Z. pears out of place within Zapoteca; however, the 
aculeata), homomorphic, spherical, capitate inflo- morphology and the type of dehiscence of its pods 
rescences; cup-shaped stigmas; 16-grained polyads and the overall features of its inflorescences justify 
with prominent lens-shaped thickenings (excluding its placement within this genus. 

Z. nervosa), and dry, linear, flattened pods with Zapoteca nervosa exhibits several characters 
thickened margins and showing a particular mode unique in the genus, the most important of which 
of dehiscence. are: stems with fissured bark; large, coriaceous, 

The distribution of morphological features among grayish leaflets with a peculiar eucamptodromous 

the species of Zapoteca reflects four evolutionary brochidodromous venation; and 16-grained polyads 


TABLE 5. Species of Zapoteca in which seedlings were studied. H = author's collections. 

Taxon Collection Data Distribution 
Z. alinae Torres 3947 (MO, MEX U) Mexico, Oaxaca 
Z. caracasana subsp. weberbaueri Johnson 2494-82 (MO) Colombia, Valle 
Z. formosa subsp. formosa Johnson 483-78 (MO) Mexico, Yucatán 
Z. formosa subsp. formosa Torres 4262 (MO, MEXU) Mexico, Oaxaca 
Z. formosa subsp. formosa Johnson 490-78 (MO) Mexico, Yucatán 
Z. formosa subsp. roset Johnson 1270-79 (MO) Mexico, Guerrero 
Z. formosa subsp. rosei Torres 6612 (MEXU) Mexico, Oaxaca 
ZL. formosa subsp. schottil Starr & Birgy 169 (MEXU) Mexico, Sonora 
Z. formosa subsp. schottii Felger 84-124 (MEXU) Mexico, Sonora 
Z. lambertiana 4costa 134 (MO, XAL) Mexico, Veracruz 
Z. lambertiana Martínez O. 284 (MEXU) Mexico, Tamaulipas 
Z. media Delgado 53 (MEXU) Mexico, Hidalgo 
Z. portoricensis subsp. portoricensis H-154 (MO, MEXU) Mexico, Veracruz 
Z. tehuana Torres 4167 (MO, MEXU) Mexico, Oaxaca 
Z. tetragona H-167 (MO, MEXU) Mexico, Veracruz 
Z. tetragona H-675 (MO, MEX U) Costa Rica, Cartago 
Z. tetragona H-800 (MO, MEXU) Mexico, Veracruz 
Z. tetragona H-827 (MO, MEXU) Mexico, Veracruz 
Z. tetragona Johnson 1582 (MO) Mexico, Veracruz 


794 


Annals of the 
Missouri Botanical Garden 


without lens-shaped structures. In this connection, 
it is reasonable to hypothesize that Z. nervosa 
represents an Efe line that separated from 
the main stock early in the evolution of the genus, 
and its peculiar morphological features would be 
explained by divergent evolution. The lack of are- 
olate, lens-shaped thickenings in the polyads of Z. 
nervosa, a character also absent in the remaining 
genera of the Ingeae, appear to represent a prim- 
itive condition within Zapoteca and suggests that 
this species is phylogenetically basal in this genus. 
Additional information on this critical species is 
needed in order to learn more about its phyloge- 
netic relationships. 


Subgenus Zapoteca is the most widespread and 
diversified subgenus in Zapoteca. Yet a number 
of morphological features common to all the species 
All the 


species have pinnae with several to many pairs of 


make this subgenus easily recognizable. 


thin, membranous leaflets, which show character- 
istic usually inconspicuous and simplified venation 
patterns. The highest diversity of hair types (three 
types) of the genus is found in subg. Zapoteca; 
the species of the remaining subgenera are char- 
acterized by having only the unicellular conical 
type (Table 4). Further, the valves in the pods in 
subg. Zapoteca, along with Z. nervosa, are con- 
sistently thick-membranous, whereas those of subg. 
Aculeata and Amazonica are coriaceous. 
The pattern of leaflet venation Z. tehuana 
differs from those of the other species in subg. 
Zapoteca and shows strong similarity with those 
Aculeata). 


of Z. aculeata (subg. This may indicate 


a phylogenetic link between the two subgenera. 


Subgenus Aculeata contains only Zapoteca 
aculeata, which is characterized primarily by spi- 
nescent stipules (a characteristic that has clearly 
been derived within the genus), by an apparent 
arborescent habit, and by pinnae having fewer pairs 


of chartaceous-coriaceous leaflets than those of 


subg. Zapoteca. The coriaceous pods of Z. acu- 


leata resemble those of subg. Amazonica. 
Subgenus Amazonica is made up of Z. ama- 
zonica and Z. microcephala, which are charac- 
terized by having the most obvious combination of 
apomorphic features in Zapoteca. Their leaves are 
reduced to two or three large, coriaceous leaflets 
in each pinna. This is the result of an evolutionary 
trend toward reduction in the number of leaflets 
associated with increased leaflet size. This trend is 
also seen in other genera of Mimosoideae (e.g., 
Moreover, the leaflets in 


Calliandra, Mimosa). 


these species have a number of common architec- 


tural features reflecting an increased level of or- 
ganization, which is also interpreted as an advanced 
condition. A progressive trend toward suppression 
of leaves subtending the inflorescence in Zapoteca 
has resulted in the highly specialized, elongate 
pseudopanicles of these species. Among other clearly 
apomorphic features of subg. Amazonica are the 
elongate seeds of Z. amazonica, which lack the 
characteristic pleurogram and are the largest in 
the genus. 


BIOGEOGRAPHY 


Zapoteca ranges from the southwestern United 
States (southern Arizona and southwestern Texas) 
and northern Mexico, from about latitude 32°30'N, 
southward through most of Mexico, Central Amer- 
ica, and much of South America (Venezuela, Co- 
lombia, Ecuador, Peru, Bolivia, Guyana, Brazil, 
Paraguay) to Argentina (ca. 
2:7°30' "si Zapoteca also occurs on most of the 


northern latitude 
West Indies and in the PLE Islands (1.e., 
Isla Socorro and Isla Clarión 

Zapoteca species occur in a variety of environ- 
ments, ranging from open, arid or semiarid, scrub- 
by vegetation types (e.g.. Z. media, Z. alinae, Z. 


formosa subsp. mollicula, and Z. formosa subsp. 


schottii) to the wet evergreen forests of Amazonian 
Peru (Z. amazonica). Most populations of Z. por- 
flavida and Z. 


found in relatively wet, lowland habitats, generally 


toricensis subsp. tetragona are 


on disturbed sites. The greatest degree of diver- 
sification has occurred in areas covered by sea- 
sonal, tropical deciduous forests (sensu Breedlove, 
1981). In addition to the species found in clearly 
arid regions, most populations of eight species of 
Zapoteca (Z. nervosa, Z. lambertiana, Z. filipes, 
Z. mollis, Z. caracasana, Z. andina, Z. formosa, 
and Z. tehuana) occur in tropical deciduous for- 
ests. In addition, Z. portoricensis subsp. portori- 
censis and subsp. pubicarpa primarily occur in 
oak, 
montane moist forests. 

The occurrence of three of the four subgenera 


middle-elevation pine-oak, or subtropical 


of Zapoteca in South America (Zapoteca, Acu- 
leata, and Amazonica), compared with only one 
in Mexico and Central America, is probably a con- 
sequence of the more ancient existence of Zapo- 
teca in South America. Although only subg. Za- 
poteca is represented in Mexico and Central 
America, extensive diversification has taken place 
at the specific and subspecific levels, especially in 
Mexico. For instance, there are seven subspecific 
taxa in Mexico (i.e., Z. portoricensis subsp. por- 
toricensis, Z. portoricensis subsp. pubicarpa, Z. 


Volume 76, Number 3 Hernández 795 
1989 Zapoteca 
portoricensis subsp. flavida, Z. formosa subsp. casana, Z. tetragona, Z. costaricensis, Z. for- 


formosa, Z. formosa subsp. rosei, Z. formosa subsp. 


mollicula, schottii), four « 


In contrast, 


Z. formosa subsp. 


them restricted to this country. only 


four subspecies of subg. Zapoteca occur in all of 


South America (Z. portoricensis subsp. flavida, 
Z. caracasana subsp. caracasana, Z. caracasana 
subsp. weberbaueri, and Z. formosa subsp. for- 
mosa), and only one of these, Z. caracasana subsp. 
weberbaueri, is endemic 

Viewed from a global perspective, there are no 
clear centers of species diversity in Zapoteca. The 
total number of species and the number of endemic 
species are roughly correlated with the size of each 
region (i.e., Mexico, 7 species/4 endemics; Central 
America, 6/2; West Indies, 4 
10/6. 


the areas of species concentration and the distri- 


l; and South Amer- 
ica, However, a more detailed analysis of 
bution of endemic taxa ee several regions 
with high diversity (Fig. 

Southern Mexico ee a -— Chiapas, includ- 
ing southern Puebla) has the largest concentration 
of species and subspecies of Zapoteca. This rela- 
tively small region, characterized by its remarkable 
climatic and topographic diversity, contains seven 
species, and if subspecies are taken into account, 
the figure rises to eleven taxa (Z. media, Z. alinae, 
Z. lambertiana, Z. portoricensis subsp. portori- 
censis, Z. portoricensis subsp. pubicarpa, Z. por- 
toricensis subsp. flavida, Z. tetragona, Z. formosa 
subsp. formosa, Z. formosa subsp. rosei, Z. for- 
mosa subsp. mollicula, and Z. tehuana). Four of 
these are endemics (Z. alinae, Z. portoricensis 
subsp. pubicarpa. Z. formosa subsp. mollicula, 
and Z. tehuana). especially to dry areas of Oaxaca 
and the arid valleys of the Pueblan- Oaxacan bor- 
der region. The number of species and subspecies 
of Zapoteca suddenly decreases from this center 
Fig. 21). On the Pacific slope of 


Mexico, from Guerrero northwards, there are four 


— 


in all directions 


taxa (Z. tetragona, Z. formosa subsp. formosa, 
Z. formosa subsp. rosei, and Z. formosa subsp. 
schottii), decreasing to one in the northern portion. 
Zapoteca media and Z. formosa subsp. formosa 
are the only taxa that occur in the Central Plateau. 


Along the entire Atlantic slope, from Tabasco to 
Tamaulipas, together with the Yucatán Peninsula, 
there are only five taxa (Z. lambertiana, Z. por- 
toricensis subsp. portorte ensis, Z. portoric ensis 
subsp. flavida, Z. tetragona, and Z. formosa subsp. 
formosa). Zapoteca lambertiana is the only one 
of these that is essentially endemic to this region. 

In Central America, there are a total of eight 
taxa (Zapoteca portoricensts subsp. portoricensis, 
Z. portoricensis subsp. flavida, Z. mollis, Z. cara- 


© 
— 


mosa subsp. formosa, and Z. formosa subsp. sal- 
vadorensis). Diversity is quite uniform among the 
different countries (Fig. 21). Zapoteca mollis and 
Z. costaricensis are essentially endemic to Costa 
Rica, although a disjunct population of the former 
is known from Panama. Zapoteca formosa subsp. 
salvadorensis is restricted to southern Guatemala 
and El Salvador. 

Six taxa of Zapoteca occur in the West Indies 
(Z. nervosa, Z. portoricensis subsp. portoricensis, 
flavida, Z. 


subsp. caracasana, Z. formosa subsp. formosa, 


Z. portoricensis subsp. caracasana 


and Z. formosa subsp. gracilis). Generally only a 
single species occurs on a particular island; how- 
ever, in Hispaniola the four species, including the 
endemic and morphologically distinct Z. nervosa, 
occur together. Howard (1973) pointed out that 
Hispaniola and Cuba are the most floristically di- 
verse islands in the West Indies and that they show 
great habitat diversity and highest percentage of 
endemism (Hispaniola with 33% of endemic gen- 
era). Zapoteca formosa subsp. gracilts is the sec- 
ond endemic entity in the West Indies. It is pri- 
: the 
Bahamas, Cayman Islands, and northwestern Haiti. 


marily restricted to coastal areas in Cuba, 

Eleven taxa of Zapoteca occur in South Amer- 
ica. The greatest diversity occurs in the tropical 
Andean countries, Venezuela, Colombia, Ecuador. 
Peru, and Bolivia, where populations are primarily 
found in the foothills of the Andes, in the inter- 
Andean valleys, and less commonly in the Amazon 
Basin. As illustrated in Figure 21, the number of 
taxa in each of these countries ranges from four 
to six, with Colombia and Ecuador most diverse. 
Perhaps the most remarkable biogeographical fea- 
ture of this whole area is the occurrence of five 
endemic taxa, which, excluding Z. caracasana 
subsp. weberbaueri and Z. andina, comprise the 
most highly advanced species in Zapoteca, each 
with a large number of derived morphological char- 
acters (i.e. Z. Andes; Z. 


the Magdalena 


aculeata, Ecuadorean 


microc suada, Median Valley of 
River; and Z. amazonica, Amazonian Pe ru). Out- 


side this area, diversity is much lower, with one or 


two taxa oceurring within Guyana, Brazil, Para- 
guay, and northern Argentina (Fig. 21). Within 


however, there are two extra- 
£. filipes, restricted to 
the Brazilian states of Goiás and Minas 
Z. scutellifera. found in the states of Amazonas, 


this large region, 
Andean endemic species: 


Gerais, and 


Para and Rondonia, Brazil, and in the departments 
of El Beni and Santa Cruz, Bolivia. 

Zapoteca species are predominantly segregated 
geographically and/or ecologically, although a 


796 


Annals of the 
Missouri Botanical Garden 


Q 1, | - 
1 la 3 -» 
/ Y A A 
o | a NS a 
1 o 3 . o EN P Li 
Y 1 1 o m 
5 
T| 3 : : 
` 55 o 
ee A ss y i 
SS T 
: Ba 4 7 
9 64 - - j i d 
Pi ' f Ut 
y Wr 
S i 
«i a E LS, {| y Y 
Se, IR M 
p A ah pas 1 N ) : 
pos J > ~ ) 2: f 
i Vi AP M / / y) 
| r 2 1 
^ l » | 
{| N 
e 2 ` | 
EN 2 
t | 
^ $ | H | ; 2 
1000 Km : 5 [ Y AM 
A E | PON 
E a 3 T 


FIGURE 21. 


Number of Zapoteca species and subspecies per country. 


Mexican regions and Brazilian states are 


treated as separate units. Total number of taxa/endemic taxa. 


number of cases of gross geographical overlap are 
known. However, only a few records of true sym- 
patry exist: in the District of Tlacolula, Oaxaca, 
plants of Z. alinae (H. Hernández & Ramírez 
942) have been found intermixed with plants of Z. 
formosa subsp. rosei (H. Hernández & Ramírez 
943); in this same area of Oaxaca, Z. formosa 
subsp. formosa has occasionally been found grow- 
ing alongside Z. formosa subsp. rosei (H. Her- 
nández & Torres 216, 217 and Magallanes 111, 
113). Also, in the tropical deciduous forest of El 
Chorreadero, Chiapas, Z. lambertiana (Breedlove 
39694, H. Hernández & Ramírez 891) and Z. 
portoricensis subsp. portoricensis (Breedlove 
11889) have been collected; however, it is unclear 
whether these are narrowly sympatric. 

There are some presumed hybrids. For example, 
in southern Guatemala and in southern Mexico, a 
number of collections provide evidence for exten- 
sive hybridization between Z. portoricensis subsp. 
portoricensis and Z. tetragona; in southern Puebla 
and northern Oaxaca, some plants with characters 
intermediate between Z. media and Z. formosa 
likewise suggest hybridization. In Guerrero popu- 


lations of Z. formosa subsp. formosa and Z. for- 


mosa subsp. rosei come into close proximity, and 
there are a number of collections of probable hybrid 
origin. 

The predominance of allopatric distributions and 
the apparent rarity of hybrids in Zapoteca suggest 
that hybridization has played an insignificant role 
in the evolution of the genus. The few cases of 
strict sympatry may represent secondary contact 
of previously differentiated populations. It could be 
hypothesized that speciation in Zapoteca has pri- 
marily resulted from genetic differentiation of geo- 
graphically isolated populations. 

There are wide disjunctions in the distribution 
of Z. formosa subsp. formosa in South America, 
from northern Colombia and Venezuela to Bolivia, 
southeastern Brazil, and Paraguay (Fig. 44). As 
elaborated below, its distribution was probably con- 
tinuous across the Amazon Basin during the cool- 
dry phases of the Pleistocene, when a drier and 
presumably more suitable environment for this sub- 
species was widespread (Ab'Sáber, 1982; Absy, 
1982; Bigarella & Andrade-Lima, 1982; van der 
Hammen, 1982; Prance, 1982). Zapoteca scu- 


Volume 76, Number 3 Hernández 797 
1989 Zapoteca 
tellifera is closely related and is known from a few — et al., 1980), and also by the wide distribution of 


disjunct populations in the Amazon Basin (Fig. 44). 
Specialized characters in Z. scutellifera (e.g., scu- 
telliform extrafloral nectaries, scandent habit) and 
its presumed edaphic specialization suggest that it 
was probably derived from Z. formosa subsp. for- 
mosa. Zapoteca scutellifera could have been de- 
rived from populations of Z. formosa subsp. for- 
mosa that became isolated after the return to the 
present-day, warm-wet climatic conditions. 

This pattern. along with the patchy distributions 
of dry-forest species like Z. E Tllana and 
mollis, supports Gentry's (1979, 2) theory of 
reverse refuges. This theory assumes that the pres- 
ent discontinuities in the distribution of the dry 
forests (dry forest refuges) coupled with the exis- 
tence of geographically restricted, morphologically 
differentiated dry-area populations, result from a 
shift towards more mesic conditions after the cli- 
matic deterioration in the Quaternary. 

A second example of possible allopatric specia- 
tion in Zapoteca involves the two members of the 
highly specialized subg. Amazonica: Z. microceph- 
ala and Z. amazonica. Zapoteca microcephala 
is restricted to the Middle Magdalena River Valley, 
whereas F amazonica occurs in Amazonian Peru 
(Fig. 47 


closely ies probably differentiated as a con- 


These two species, which are obviously 


sequence of a vicariance event caused by the uplift 
of the northern Andes, which reached their present 
elevation at the beginning of the Pleistocene some 
three million vears ago (van der Hammen, 19 
Additional evidence suggesting an enini 
model of speciation for Zapoteca is the relatively 
common occurrence of closely related taxa dis- 
playing contiguous or nearly contiguous distribu- 
tions. This distribution pattern is shown by closely 
related species, such as Z. tetragona and Z. an- 


dina (Figs. 40. 


species in the genus (Figs. 36, 39 


42). and by virtually all the sub- 


REPRODUCTIVE BIOLOGY 


During the past few years a considerable body 
of data has accumulated on the diverse reproduc- 
tive mechanisms of the Mimosoideae (Simpson et 
al., 1977; Kenrick & Knox, 1982; Koptur, 1983, 
1984; Knox & Kenrick, 1983; Hopkins, 1984; 
1983). Entomophily was probably the 
original mode of pollination in the subfamily, as 


— 


Bernhardt. 


indicated by the presence of apparently insect, 
probably hymenopteran, pollinated mimosoid inflo- 
the Middle For- 


mation, Tennessee (Crepet & Dilcher, 1977), and 


rescences in Focene Claiborne 


in the Oligocene of the Texas Gulf Coast (Daghlian 


this pollination system in the subfamily. It has been 
suggested (Raven & Polhill, 1981; Polhill et al., 
1981) that Mimosoideae were already 
tiated from the other subfamilies by 
B.P.) 


spread in tribe Mimoseae (e.g 


differen- 
the start of 
the Eocene (54 m.y. ). Entomophily is wide- 

. Desmanthus, Leu- 
caena, Mimosa, Prosopis), aiee is generally con- 
sidered to be the basal tribe of the Mimosoideae. 
Furthermore, most species of Acacia (Acacieae) 
are insect-pollinated (Bernhardt, 1983; pers. obs.); 
however, bird pollination has been derived second- 
arily in some Australian species of this genus (Ford 
& Forde, 1976; Bernhardt, 1983; Kenrick et al., 
1983). In contrast, Parkia (Parkieae) is primarily 


bat-pollinated (Baker & Harris, 1957; Hopkins, 
1984), although a few species are pollinated by 

g | I ) 
insects (see discussion in Hopkins, 1984) 


High-energy pollination systems such as those 
involving hawkmoths. birds, and bats are concen- 
trated among the genera of the more advanced 
tribe Ingeae (Cruden et al., 1976; Arroyo, 1981; 
1983; Hernández, 1986a; unpublished 


obs.). In the Ingeae, the transition from low-energy 


Koptur, 


insect pollination to high-energy systems, which 
demand greater allocation to nectar and pollen, is 
associated with inflorescence modifications. For in- 
stance, connation of the staminal filaments to form 
a tube and increase in number and length of the 
filaments made it possible for the flowers to hold 
larger quantities of nectar necessary to satisfy the 
energy requirements of larger, homothermic pol- 
linators. Similarly, changes in periodicities of nectar 
and aroma production, nectar quality, and timing 
of anthesis have enabled the plants to adjust to the 
physiological and behavioral characteristics of par- 
ticular pollinators. 

Mimosoideae have a number of additional 
productive traits that make them extremely inter- 
esting from an ecological and evolutionary point 
Guinet (1969, 1981) 
that the most conspicuous 


pollen is the occurrence of permanent units of 


of view. discussed in detail 


feature of mimosoid 
dispersal in the form of tetrads, octads, and polyads 
throughout the subfamily. Such units of pollination 
presumably provide the plants with a selective ad- 
vantage in reproduction. As has been demonstrated 
for Acacia species (Kenrick & Knox, 1982: Knox 
& Kenrick, 1983 


fertilize all the af] in the ovary. 


. a single pollen unit is able to 
Another wide- 
spread feature of the mimosoid inflorescences that 
is important in the breeding systems of the species 
is the occurrence of female sterile flowers. Viewed 
in an evolutionary perspective, these reproductive 
phenomena have great potential as characters on 


798 


Annals of the 
Missouri Botanical Garden 


which to base phylogenetic interpretations. Pub- 
lished reports concerning the reproductive biology 
of the species here assigned to Zapoteca are vir- 
tually nonexistent, and the present chapter is in- 
tended to provide an account, based on my own 
observations, of some of the reproductive features 


of these plants. 
METHODS 


Nectar volumes were measured with calibrated 
micropipets, and sugar concentrations were deter- 
mined using a temperature-compensated Erma hand 
refractometer. In a few cases, when the flowers 
were very small, a syringe was used to extract the 
nectar from the staminal tubes. All measurements 
were made in the greenhouse early in the morning 
in order to estimate the total amount of nectar 
produced per flower. Between four and ten indi- 
viduals per population were examined. It is quite 
possible that the results in Table 6 might be lower 
volumes than would be found in the wild, since it 
has been shown that nectar volumes and sugar 
concentrations may be significantly lower in plants 
growing in greenhouse pots compared with wild 
plants (Cruden & Hermann, 1983). However, they 
allow species comparisons. 

Field observations on flower visitors were carried 
out continuously during the first three to five hours 
of anthesis. Since all Zapoteca species flower at 
night, the behavior of flower visitors was observed 
with the aid of a head lamp. In most cases, visitors 
were successfully collected with insect nets. Insect 
voucher specimens are deposited in the Entomo- 
logical Collection at the Instituto de Biologia, 

The number of ovules per gynoecium was de- 
termined by squashing fixed ovaries in a drop of 
Alexander's stain (Alexander, 1969); the ovaries 
were examined with a compound light microscope 
within 15-20 minutes after staining. Sex expres- 
sion was characterized for each population by dis- 
secting fixed flowers and counting the proportion 
of hermaphrodite and female-sterile flowers in each 
inflorescence. Self-compatibility was assessed on 
plants growing in the greenhouse using the tech- 
niques outlined by Bawa (1974). Pollinations were 
accomplished using a fine brush within four hours 
of the beginning of anthesis, once the anthers were 
dehiscent. Anther dehiscence was determined by 
touching the anthers with a clean microscope slide; 
a large number of polyads on the slide revealed 
open thecae. Given the small size of the flowers, 
the inflorescence was taken as a unit in the polli- 
nation treatments. In every population, a deter- 
mined number of inflorescences were artificially 


eross- and self-pollinated, and several additional 
inflorescences were left untreated (open) to test for 
autogamy. Due to the difficulty involved in emas- 
culating the delicate flowers of Zapoteca, the pos- 
sibility of apomixis was not tested. For the cross- 
pollinations, mixtures of polyads from several in- 
dividuals were used whenever possible. For the 
self-pollinated inflorescences, the pollen donor was 
taken from the same inflorescence or from other 
inflorescences in the same individual; therefore, a 
great number of these flowers were geitonoga- 
mously pollinated. Treated inflorescences were 
marked with hanging labels. As the greenhouse was 
insect-free, inflorescence manipulation was mini- 
mized by avoiding the need to bag inflorescences. 
Sample size varied from two to nine individuals per 
population. After each treatment, brushes and hands 
were sterilized with ethanol. Since pod abortion 
occurs in Zapoteca species at several stages of 
pod development, the resulting pods were counted 
Seed set was estimated 
calculating the average 


only if they matured fully. 
in every population by 
number of fully developed seeds in mature pods. 


FLOWERING PHENOLOGY 


Although it is difficult to characterize the phe- 
nological behavior of a given species throughout 
its range, I shall attempt to draw some generali- 
zations about the flowering patterns in Zapoteca. 
This discussion is based primarily on species re- 
stricted to Mexico and Central America. In wide- 
ranging species such as Z. tetragona, flowering 
times vary from one region to another. This species, 
which usually grows in secondary vegetation de- 
rived from lowland tropical wet forests, flowers 
mostly during the dry season; nevertheless, in the 
Mexican state of Morelos, where a more seasonal 
climate exists, the populations flower primarily dur- 
ing the rainy season. Zapotec a portoric ensis subsp. 
flavida, which grows in wet habitats, also flowers 
during the dry season. By contrast, the species 
growing in drier habitats (thorn forests, tropical 
deciduous forests, and tropical semideciduous for- 
ests) tend to flower predominantly during the sec- 
ond half of the rainy season (e.g., Z. formosa, Z. 
lambertiana, Z. media, Z. alinae). The species 
from drier areas have shorter, more synchronized 
flowering periods than species living in wetter hab- 
itats (e.g., Z. amazonica and Z. tetragona), which 
continue to produce flowers for longer periods. 
Finally, Z. portoricensis subsp. portoricensis, which 
generally grows in montane mesophyllous forests 
(terminology following Rzedowski, 1978), also flow- 
ers in the wet season. 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


799 


24 


rescences of Zapoteca species at anthesis; all photographs were 


Figures 22-26. LH 


formosa subsp 


. formosa (right) and Z. formosa subs} 
portoric ensis subsp. abv ensis 
26. Z 


UH. Hernández 95 


A possible explanation for this differential flow- 
ering pattern is that in wet areas excessive rains 
may have destructive effects on the delicate inflo- 
rescences of Zapoteca and may reduce the activity 


of potential pollinators (Janzen, 1967). | observed 


the devastating effects of rain on inflorescences of 


several Zapoteca species in. wet and dry forests. 
These effects are clearly reduced in areas of less 
precipitation. 

s far as I have observed, species of Zapoteca 
display two distinct flowering strategies. Most species 
(e.g.. Z. formosa, Z. media, Z. portoricensis, 
tetragona) have a very generalized flowering be- 
havior. Here regular numbers of inflorescences open 
By contrast, Z. 
Indi- 


every night over several months. 
alinae exhibits a more specialized strategy. 
viduals of this species intermittently produce rel- 


atively large numbers of inflorescences on scattered 


¡A rosel 
Hernández dag). 24 
i oum (H. Hernández 950). 


taken at aed 
i ap (H. Hernández 216, 217, respec tively 

. Z. tetragona (H. Hernández 956). 25. * media 
All collections at MEXU and MO. 


NN 
wh 
NN 


nights throughout its flowering period, from July 


to September. This specialization is similar to the 
"multiple-bang" strategy described by Gentry 
(1974) for tropical Bignoniaceae. It differs, how- 
ever, in that the flowering is not synchronized among 
the different individuals that make up a given pop- 
ulation of Z. 


of this strategy is that these flowering bursts pro- 


alinae. One possible adaptive value 


duce conspicuous visual and olfactory stimuli, thus 


increasing the chances of pollinator attraction. 


THE INFLORESCENCE AS THI 
POLLINATOR ATTRACTION 


UNIT OF 


Species of Zapoteca always have their flowers 


arranged compact. homomorphic, capitate in- 


florescences, which act as units of attraction (Figs. 
22-26). 


In each flower, the numerous elongate 


800 Annals of the 
Missouri Botanical Garden 
TABLE 6. Volume and sugar density of nectar in selected species of Zapoteca. All collections in MO. H = author's 


collections. 


Volume (pl) Sugar density (%) 
C (CF S.D.) D.) 


Taxon Collection Data x (S 
Z. alinae Mexico, Oxaca, 11-949 < 1] 17.27 
Z. formosa Mexico, Jalisco, H-230 «] 13.9 
Z. lambertiana Mexico, Chiapas, /7-891 1 
Z. portoricensis Mexico, Tamaulipas, H- 146 epi 
Z. portoricensis Mexico, Veracruz, H-948 1.04(0.8) 18.43(3.4) 
Z. tetragona Mexico, Veracruz, 11-956 3.05(2.0) 19.83(2.5) 
Z. tetragona Mexico, Veracruz, 11-960 5.12(2.0) 20.32(3.2) 


filaments, which give the inflorescences their at- 
tractive powder puff aspect, are fused basally, 


forming a tube in which nectar accumulates. 


Anthesis and the process of flower presenta- 
tion. Field and greenhouse observations have 
shown that species of Zapoteca display predictable 
flowering mechanisms. All members of the genus 
exhibit nocturnal anthesis and a remarkable syn- 
chrony in floral behavior during flower presenta- 
tion. Usually the filaments of all the flowers in a 
head become erect in a springlike fashion late in 
the evening (ca. 17:00-18:00 hours), during the 
last two hours of daylight. Subsequently, when the 
19:00 


At about the same 


filaments are completely erect (ca. 18:00 
hours), the anthers dehisce. 
time, nectar secretion begins, accompanied by pro- 
duction of a sweet aroma. Each head remains func- 
tional for only one night and the filaments. wilt 
within approximately the first two hours of daylight. 
With 


few exceptions, the color of the staminal filaments 


Attractants: filament color and aroma. 


is constant in a given species of Zapoteca and thus 
is of taxonomic value. The filaments may be white 
(e.g., Z. alinae, Z. portoricensis, Z. tetragona), 
red-purple (Z. lambertiana), or a combination of 
white in the basal half and pink or red-purple in 
the distal half (e.g., Z. amazonica, Z. caracasana, 
Z. media). The widespread Z. formosa shows some 
geographical variation in filament color, and the 
recognition of subspecies within this species is based 
in part on this feature. This character is nearly 
always stable at the population level in Z. formosa; 
however, in a population observed in eastern Oa- 
xaca a mixture of plants with white (11. Hernández 
& Torres 216) and white pink filaments (/7. Her- 
nández & Torres 217) turned up. Similarly, in- 
dividual plants of Z. tehuana with either white or 
white/pink filaments were observed in a 
house population (/7. Hernández 950). The flowers 


green- 


of all species of Zapoteca produce a delicate, sweet 


fragrance, which presumably functions as does 
the color of the stamens in attracting pollinators 


(Brantjes, 1978 


Rewards: nectar production. The amount of 
nectar produced by individual flowers varies among 
the species of Zapoteca, perhaps as a function of 
the size of the nectaries and the physical capacity 
all the 
tetragona produce 


As shown in Table 6, 
Z: 


quantities of nectar below or slightly above one 


of the staminal tube. 


species analyzed except 


microliter (ul) per flower. In the two populations 
of Z. tetragona (the species with essentially the 


most massive flowers in the genus), the nectar 


volumes were substantially higher (Table 6). The 
nectar of Zapoteca species has rather low sugar 
concentrations, ranging from 13.9 to 20.3%. The 


nectar volumes and sugar concentrations in the 
flowers of Zapoteca species reported here agree 
with those reported by Opler (1983) and Baker & 
Baker (1983) for settling moth-pollinated flowers. 


FLOWER VISITORS 


Inflorescences Zapoteca species are well 
adapted for pollination by moths. Field observations 
conducted in ten different populations representing 
five species of Zapoteca clearly showed that the 
flowers are consistently visited and pollinated by a 
variety of settling moths, mostly of families Noc- 
The 


moths were usually observed to visit the inflores- 


tuidae, Pyralidae, and Geometridae (Table 7). 


cences after some nectar had accumulated in the 
staminal tube and after floral fragance was detected 
by observers. Moths usually landed in the upper 
part of the inflorescence, where the underside of 
their bodies contacted the anthers and the stigmatic 
surfaces (Fig. 27). Moths generally moved from 
one part of the inflorescence to another in search 
of nectar, and the length of time they remained 
on an inflorescence was often up to 20-30 minutes. 


Aside from the species in Table 7, a substantial 


Volume 76, Number 3 


Hernández 
Zapoteca 


number of species of smaller moths were excluded 
because they were considered to be ineffective pol- 
linators. 

Many species of moths were observed to exploit 
the nectar of Z. tetragona (population J) illegiti- 
mately. They extracted nectar by standing on the 
peduncles without touching the anthers and stig- 
mas. However, five species of moths were observed 
to visit the flowers of this species legitimately; among 
these, Palpita flegia (Fig. 27) and Diaphania 
jairusalis may be considered the primary polli- 
nators on the basis of their abundance (Table 7). 
The color of the wings of these two species is 
strikingly similar to that of the white filaments of 
Z. tetragona. In contrast, the nectar robbers, along 
with the few other occasional pollinators, have wings 
with a variety of color patterns. 

Although inflorescences of Zapoteca species dis- 
play the typical syndrome of characteristics asso- 
ciated with hawkmoth pollination (Faegri & van 
der Pijl, 1979), hawkmoths were rarely observed 
as visitors. During the course of my observations, 
only two hawkmoths were observed, one visiting 
the flowers of Z. formosa (population C) and one 
on Z. lambertiana (population J). A possible ex- 
planation for this is that the individual flowers pro- 
duce rather small amounts of nectar, making the 
visits energetically unprofitable to hawkmoths. Ac- 
cording to Opler (1983) and Haber & Frankie (in 
hawkmoth-pollinated flowers usually pro- 
duce much greater amounts of nectar (maximum 
available nectar = 130.5 ul; 1983) 


compared with the relatively smaller amounts found 


press), 


Opler, 


1 


in the flowers of Zapoteca (Table 6). 

Two unidentified species of crab spiders (Ara- 
neida: Thomisidae) were observed preying on moths 
visiting the inflorescences of Z. formosa subsp. 
rosei and Z. portoricensis subsp. portoricensts. 
These light-colored spiders would be inconspicuous 
on the inflorescences of Zapoteca species, where 
their cryptic coloration permits them to prey on 
flower visitors. Given that the spiders occurred in 
very low numbers, their negative impact on the 
reproductive potential of the plants is probably 
Such an association between crab 
spiders and. flowers has been reported by other 
authors (Wickler, 1968; Kevan, 1972; Gertsch, 


1979, and others). 


unimportant. 


POLLINATION RELATIONSHIPS 

MI species of Zapoteca have very delicate, cup- 
shaped stigmas, and as indicated. elsewhere (Her- 
nández, 1986a), the concave area of receptivity is 


so narrow that only a single 16-grained polyad can 


FIGURE 27. Palpita, flegia visiting the flowers of e 
in a e near Catemaco, Ver: 
“XU, MO) 


H. He rnåndez 7 . MI 


poteca tetragona 


pæn, 


cruz 


be held (see fig. 11 in Hernández, 1986a). Con- 
sidering that the likelihood of a pollen unit reaching 
the stigma is directly proportional to the receptive 
(Cruden & Miller- Ward, 1981), 


one might anticipate that the chances of suecessful 


area of the stigma 


pollination in species of Zapoteca are rather low. 
As pointed out earlier, however, the settling moths 
that were recorded visiting species of Zapoteca 
normally remain on the inflorescences for consid- 
erable periods of time. The chances of successful 
pollination are also affected by the time the body 
of a particular pollinator makes contact with the 
stigma; therefore, the relatively long visits by set- 
tling moths compared with hawkmoths might be 
understood as a compensating factor to the intrinsic 


reaching the stigma in 


low chances of a polyac 
Zapoteca. 
The average number o 


ovules ovary in 


per 
Zapoteca species is highly correlated with the num- 
ber of pollen grains in the polyads. As indicated in 
Table 8, 
15 and never exceeds tl 
polyad (16). 
polyad to the number of ovules is close to one, 


the number of ovules ranges from 11 to 
ie number of grains in the 
ratio of grains 


Therefore. the per 


Annals of the 
Missouri Botanical Garden 


TABLE 7. 
numbers of individuals collected from inflore »scences. FO 
TETRA 


Z. media, PORTO 


— 
£ 
= 


= £. portoricensis, 


Pollinators of Zapoteca species. Capital letters designate populations and the figures in parenthesis the 
) 


7. formosa, LAMB = Z. lambertiana, MEDIA 


= Z tetragona. 


FORMO 


Pollinator 


LAMB MEDIA PORTO TETRA 


Noctuidae 


Rhizagrotis malefida Guenee 
Heliothis 


dutographa biloba us 


zea Boddie 


Walker 


uenee 


Pseudoplusia includens 
Prodenia ornithogalli € 
Gonitis edetrix Guenee 
Leucania sp. 
unidentified spp. 
Pyralidae 
Pilocrocis tripunctata Fabr. Bl) 
Conchylodes platinalis Guenee B(3) 
C. alcifera Hampson B(4) 
Diaphania jairusalis Walker 
Palpita flegia Cramer 
Lineodes mesodonta Dyar 
unidentified spp. 
Geometridae 
Mecoceras aulis Druce 
{metris mitocris Cramer 
4picia medusa Druce 
unidentified spp. 
Arctiidae 
dgaraea longicornis H. Sch. 
Ctenuchiidae 
Psilopleura vittata Walker 


H(1) 


I(1) 


J(4) 
I(1) J(35) 
H,1(10) 


G(1) 


G(1) 


ae 
Chiapas (11-8 
Veracruz (/T- f ie T Mexico, Veracruz (H- 165 


ranging from 1.04 in Z. formosa to 1.45 in Z. 
alinae. An almost identical situation has been re- 
ported by Kenrick & Knox (1982) and Knox & 
Kenrick (1983) for species of Acacia. In this ge- 
nus, however, the range of variation in the ratio 


of number of grains to number of ovules is some- 


what broader than in Zapoteca, but the species of 


Acacia with 16-grained polyads also have ratios 
approaching one (Kenrick & Knox, 1982). In Cal- 
liandra (sensu Hernández, 1986a) ratios also vary 
around one; however, in several species of the 
highly advanced ser. Racemosae, 
ovules greatly exceeds the eight grains found in 
the polyads (Nevling & Elias, 1971; Cruden, 1977; 
Hernández, 1984, and unpublished results). This 
reduction in the ratio of number of grains to number 
of ovules is probably associated with the highly 
modified fungiform, discoid, or capitate stigmas 
found in Calliandra, which have a greater area of 


rus Michoacán (H- 185): B. Mexico, Jalisco (H-230); C. 
. Mexico, Hidalgo (11-133); F. Mexico, Hidalgo (11-122); G. Mexicc 


the number of 


Mexico, Oaxaca (H-207); D. Mexico, 
», Hidalgo (H- 136); H. Mexico, 


X J. Mexico, Veracruz (11-799). ( ollections i in MEXU and MO. 


receptivity than the representatives of the other 
genera mentioned (Hernández, 1980a) 


BREEDING SYSTEMS 


Sex expression. In species of Zapoteca, sex 
expression and genetic self-incompatibility systems 
are the primary components of the breeding system 
that affect pollination and fertilization. Analysis of 
the inflorescences of several species has shown that 
morphologically expressed andromonoecy resulting 
from abortion of the gynoecium is widespread in 
the genus (Table 9). Functionally male flowers with 
an aborted gynoecium are irregularly distributed 
among populations. Generally, individual plants have 
inflorescences with varying proportions of her- 
maphroditic and functionally male flowers; how- 
ever, individuals with inflorescences composed en- 
tirely of hermpahroditic flowers are not uncommon. 


Volume 76, Number 3 


Hernández 803 


Zapoteca 


TABLE 8. Average number of ovules in ovaries of Zapoteca species. Collections in MO. H = author's collections. 
Number of Ovules Polvad 
Species Collection Y S.D. (N) Ovule Ratio 

Z. alinae H-2060 11.0 + 1.02 (46) io 

Z. formosa subsp. formosa H-954 13.6 + 0.71 (37) 1-2 

Z. formosa subsp. rosei H-165 13,3: 0:77 (32 1.2 

Z. formosa subsp. rosei H-209 15.4 + 0.50 (11) 1.0 

Z. formosa subsp. rosei H.217 12.3 + 0.80 (30) 1:2 

Z. formosa subsp. roset H-230 13.7 + 0.65 (40) 2 

Z. formosa subsp. mollicula 11-218 11.2 + 0.94 (27) 1.4 

Z. lambertiana H-147 14.1 + 0.86 (33) m 

Z. media H-192 12.5 + 0.68 (38) 1.3 

Z. portoricensis subsp. portoricensts H-122 14.2 + 0.62 (30) 1.1 

Z. portoricensis subsp. portoricensis H-165 12.1 + 0.70 (35) 1.3 

Z. tetragona Veill 5366 13.4 + 0.71 (32) 1:2 


The ratios of hermaphroditic to female-sterile 
(functionally male) flowers change during the course 
of the flowering season in individual populations. 
Therefore. the figures provided in Table 9, which 
correspond to a single stage of the flowering cycle 
in each population for which data are enumerated, 
do not reflect the actual proportions of the two 
flower tvpes. 

1 


interspecific variation in sex expression among the 


'able 9 indicates a great amount of intra- and 


species of Zapoteca. The ratios of hermaphroditic 
to female-sterile flowers varied from 0.4 to 77.5 
in the studied populations. Thus far, I have been 
unable to establish any correlation between the 
nature of sex expression and factors such as alti- 
tude, frequency of pollinator visitation, and incom- 
patibility systems. Nevertheless, a planned, detailed 
analysis of temporal variation in the ratios of her- 
maphroditic to female-sterile flowers in natural pop- 
ulations would probably contribute significantly to 
understanding this reproductive phenomenon. 
Incompatibility systems. The results of the 
incompatibility experiments show that of the five 


species studied, only Z. tetragona is clearly ge- 
netically self-incompatible, as indicated by the in- 
ability of the flowers to set pod following the self- 
and open-pollinated treatments in the two popu- 
lations (Table 10). In contrast, plants of the two 
populations of Z. formosa that were studied proved 
to be genetically self-compatible, and those from 
one additional population (H. Hernández 959) were 
f 


relatively high proportion of the untreated inflo- 


partially autogamous, judging from the fact that a 
rescences (20%) set pods (Table 10). 
Incompatibility studies on the remaining species 
(Z. media, Z. alinae, Z. portoricensis) were in- 
conclusive but suggested that these species are 
genetically self-compatible, at least in part. In both 
Z. media and Z. alinae, pod set was significantly 
higher in untreated (open) inflorescences compared 
Table 10). 


although this may have been partly the result of 


with the self-pollinated inflorescences 


— 


mechanical damage to floral parts during the hand 
pollinations. In Z. portoricensis, the difference in 
pod set between the untreated and the self-polli- 
nated inflorescences was less pronounced: however, 


the few pods that were derived from the hand self- 


Tagle 9. Sex expression in flowers of Zapoteca species. Collections in MO. H = author's collections. 
Percent of 
Taxon Collection à Flowers (N) g à Ratio 

Z. alinae H-206 27.0 (211) P 
Z. formosa subsp. roset 1-209 12.2 (54) 0.4 
Z. formosa subsp. roset H-217 1.4 (144) 71.0 
Z. formosa subsp. rose 11-230 14.0 (107) 6.1 
Z. formosa subsp. mollicula H-21€ 1.3 (157) TRO 
Z. lambertiana H-165 3.1] (143) 16.7 
Z. lambertiana 11-89 1 1.5(133) 66.5 
Z. media 11-192 2.9(175) 2349 
Z. portoricensis subsp. portoricensis H- 105 6.0 (181) 15.6 
Z. tetragona Veill 5388 11.0 (100) 8.1 


804 Annals of the 
Missouri Botanical Garden 
TABLE 10. Results of incompatibility experiments in Zapoteca species. Pod set (%) is given as the percent of 


treated inflorescences (N) that developed 
seeds per pod. SI — genetically self- -incompatible; S 


collections. 


at least one pod, and seed set as the average number of fully developed 
— genetically self-compatible. Collections in MO. H = author's 


Average 
: Number of 
^5 
Collec- Indi- Treat- Fon Se Pods Per In- Seed Con- 
Taxon tion viduals ment % florescence Set clusion 
Z. alinae H-949 9 Open 8.5 71 2:3 9.3 Ss 
Self 0.4 239 1.0 1.0 
Cross 6.5 186 1.3 7.8 
Z. formosa H-954 4 Open 0 98 5 
Self 5.6 53 3 1.5 
Cross 18.2 11 2.0 3:8 
Z. formosa 11.959 3 Open 20.0 20 1.0 5.0 S¢ 
Self 5.3 19 1.0 2.0 
Cross 66.7 6 1.7 6.5 
Z. media H-951 4 Open 15.6 167 1.4 6.9 Ss 
Self 0 80 
Cross 18.9 58 LS 5.9 
Z. portoricensis H-948 7 Open 2.7 221 2.8 SC(?) 
Self 1.6 122 2.0 0 
Cross 13.2 128 2.2 4.7 
Z. tetragona H-956 2 Open 0 24 SI 
Self 0 43 
Cross 33.3 12 1.8 11.2 
Z. tetragona 11-960 4 Open 0 18 SI 
Self 0 14 
Cross 100 l l 8 


pollinated inflorescences developed no mature seeds. 
In contrast, the pods set in untreated inflorescences 
produced a relatively low average number of seeds 
per pod (2.8), which indicates some degree of ge- 
netic self-compatibility in Z. portoricensis. Pre- 


suming that there was no accidental transfer of 


polyads among the untreated plants, the individuals 
of all three species tested were at least in part 
genetically self-compatible. 


CONCLUDING REMARKS 


Zapoteca species, are relatively consistent in 
several important features of reproductive biology. 
All the observed species exhibit nocturnal anthesis. 
The small flowers are organized in compact, cap- 
itate inflorescences, and there is remarkable syn- 
chrony among the flowers of a given capitulum in 
all the stages of flower presentation and in the 
patterns of aroma and nectar production. This sug- 
gests that ecologically the spherical capitula act as 
units of attraction. The particular combination of 
morphological and behavioral features in the inflo- 


rescences of species of Zapoteca is well adapted 
for pollination by settling moths. Moreover, the 
remarkable morphological correlation between the 
stigmas and the polyads in Zapoteca, along with 
the specific behavior of its pollinators, reflects a 
highly organized reproductive system. Reduction 
in the number of ovules in each ovary increases 
the reproductive efficiency of these species, giving 
a single polyad the potential of fertilizing all the 
ovules in an ovary. In other words, the high cor- 
relation between the number of ovules in an ovary 
and the number of pollen grains in a polyad is a 
reflection of the high probability of the ovules being 
fertilized by the pollen grains contained in a single 
polya 

According to Arroyo (1981), about half of the 
genera of Mimosoideae are andromonoecious; this 
condition occurs especially in taxa with capitate 
inflorescences. The results presented in Table 9 
indicate that andromonoecy is widespread in Za- 
poteca, and the presence of a vestigial gynoecium 
in functionally male flowers strongly suggests that 
andromonoecy is derived from hermaphroditism in 


Volume 76, Number 3 Hernández 805 
1989 

this genus. Several models have been developed to numbers of x = 13, with the exception of the 
explain the selective basis for the evolution of an- highly specialized Calliandra, in which n = 11 
dromonoecy (e.g., Heithaus et al., 1974; Ruiz & and — 8 have been reported (see review in 
Arroyo, 1978; Primack & Lloyd, 1980; Dulberger Hernández, 1986a). As discussed earlier. differ- 


1981; Bertin, 1982). The following discus- 


sion will be centered on the possible factors playimg 


et al., 


a selective role in the evolution and maintenance 


of female sterility in Zapoteca. 


— 


In species of Zapoteca, as in the majority of 
1981). 


flowers produced by individual plants is significantly 


plant species (Stephenson, the number of 
larger than the number of pods reaching maturity. 
In natural populations, the vast majority of the 
capitula abscise after anthesis. Even in the re- 
maining capitula, only a fraction of the fertile ova- 
ries develop into pods. For instance, in a population 
of Z. tetragona near Catemaco, Veracruz (H. Her- 
nández 600), it was estimated rn each capitulum 
.3; N = 425) 


mature pods. comparative ly D in relation to the 


produced an average of 2.04 


average number of hermaphroditic flowers per head 
(17.8 + 2.7; 100) in the same population. 
Thus, most of the flowers produced in Zapoteca 
serve only as attractive elements in the inflores- 
cence and as pollen donors (Stephenson, 1981, 
and references therein) 

Rather than being limited by pollinator activity, 
the 
probably affected by energetic, 


inflorescences of the Zapoteca species are 
and certainly by 
physical architectural constraints, which set the 
upper limit for maximum pod production. In this 
connection, Stephenson (1981) reviewed the avail- 
able evidence and suggested that “an upper limit 
to the number of pods that set is usually determined 
by resources rather than the number of morpho- 


logically female flowers, and . . . natural levels of 


pollination exceed pod set in many species." In 
this context, the adaptive nature of female sterility 
can be understood: given the lower pod set in 
Zapoteca populations, a reduction in the number 
of functional gvnoecia in an inflorescence would 
not reduce pod set. Vestigial gynoecia must be 


energetically less expensive to produce than the 


arger, functional ones. Thus, andromonoecy should 
be understood as a system that makes more efficient 
use of reproductive energy without compromising 
pollinator attraction (Ruiz & Arroyo, 1978; Pri 
mack & Lloyd, 1980; Arroyo, 1981; Bertin, 1982 


and as a means to increase the pollen ovule/ ratios 


— 


and, consequently, the chances of pollination. 


CY TOGENETICS 


According to Goldblatt (1981), the genera of 


the Ingeae consistently have basic chromosome 


ential patterns in chromosome numbers and in mor- 
phological characteristics provided the justification 
for the segregation of Zapoteca from Calliandra 
sensu Bentham. 


METHODS 


Somatic chromosome counts were obtained from 
actively growing root tips taken from germinating 
seeds or mature plants in the Experimental Green- 
house at the Missouri Botanical Garden. The meth- 
ods employed were those of Goldblatt & Gentry 
979). Root tips were pretreated in 0.003 M 
hydroxyquinoline for about seven hours at refrig- 
erator temperatures, 2:1] 
ethanol: glacial acetic acid for one minute. The 


— 
jæi 


then fixed ı absolute 


roots were hydrolyzed in 10% HCI for six minutes, 
then squashed and heated in formic-lacto-propionic 


orcein. 


RESULTS 

Chromosome numbers in Zapoteca were re- 
ported in Hernández (1986a). Counts for two ad- 
All ey- 


of Zapoteca have 


ditional taxa are included here (Table 11). 
tologically investigated species 

= 13 (Figs. 28, 29), which is 
basic numbers reported for the remaining genera 
of Ingeae (Goldblatt, 1981). Chromosome numbers 


consistent with the 


are now known for seven of the 17 species rec- 
ognized in this monograph, all of which are mem- 
bers of subg. Zapoteca. The species of the re- 
maining subgenera remain unknown cytologically. 


SYSTEMATIC. TREATMENT 


TAXONOMIC HISTORY OF ZAPOTEC AL 


Analysis of several critical characters of the 
neotropical species of Calliandra, a large genus 
currently undergoing revision (Forero, 1984; Her- 
nández, 1984), showed that Calliandra sensu Ben- 
tham apparently is polyphyletic. Consequently, the 
species of Bentham’s Calliandra ser. Laetevi- 
rentes were recognized as generically distinct (Her- 
nández, 1986a). For this natural group | proposed 
the name Zapoteca. 

Mimosa portoricensis Jacq. (— Zapoteca por- 
toricensis) and M. caracasana Jacq. (= Z. caraca- 
sana) were the first species to be published of what 
is now Zapoteca. They were described in 1791 on 
the basis of material collected by the Austrian bo- 


Annals of the 


806 
Missouri Botanical Garden 
TABLE 11. Chromosome numbers in Zapoteca. 
Taxon 2n Reference or Collection 
Z. alinae 26 Mexico, Oaxaca: (Hernández, 1986a)! 
Z. caracasana subsp. weberbaueri 26 Colombia, Valle: Johnson 2494-82 (MO) 
Z. formosa subsp. formosa 26 Mexico, Oaxaca: (Hernández, 1986a) 
Z. formosa subsp. formosa 26 Mexico, Yucatán: (Hernández, 19862) 
Z. media 26 Mexico, Hidalgo: (Hernández, 19862) 
Z. portoricensis subsp. flavida 26 Mexico, Veracruz: 11-793 (MO, MEXU) 
Z. portoricensis subsp. portoricensis 26 Mexico, Veracruz: (Hernández, 1986a) 
Z. tehuana 26 Mexico, Oaxaca: (Hernández, 1986a) 
Z. tetragona 26 Mexico, Veracruz: (Hernández, 19862) 
Z. tetragona 26 Mexico, Veracruz: (Hernández, 1986a) 
Z. tetragona 26 Costa Rica, Cartago: (Hernández, 1986a) 


Referred to as sp. nov. (1 


Referred to as sp. nov. (2) 


tanical expeditions to the West Indies and Vene- 
zuela during the second half of the eighteenth cen- 
tury. These species, as well as a few that have 


been described in Acacia, were treated by Bentham 
as Calliandra ser. Laetevirentes. Bentham (1875) 
recognized 12 species in ser. Laetevirentes, five 
of which he described in previous papers (Bentham, 
1840, 1844). Seven of the 12 species included by 
Bentham in ser. 
the present treatment. 

The species in Bentham's ser. Laetevirentes were 
treated by Britton & Rose (1928) 
group Portoricenses and group Formosae. They 


Laetevirentes are maintained in 


Anneslia 


recognized 25 species within these groups, of which 
11 were published as new. Ten of these 25 species 
are maintained in the present treatment, including 
two of the new species. Three additional species, 
all of them clearly recognizable as Zapoteca, were 
arbitrarily included by Britton & Rose under An- 
neslia group Molliculae. Britton & Rose’s treat- 


FIGURES 28, 29. 
950). — 29. Z. tetragona, 
5 um. 


ais gui wales of Zapoteca. 
n 


28. Z. 
> (metaphase) (H. He ndn 075). Collections at MEXU and MO. Scale bars 


ment is characterized by excessively narrow defi- 
nitions of taxa. and their subdivisions are artificial. 
Britton & Rose studied only a fraction of the over 
4,000 herbarium specimens that were available for 
the present study. Calliandra was conserved over 
Anneslia (Bunting, 1967; Hernández, 1986b) and, 
as a consequence, a number of names recognized 
by Britton € Rose required new combinations un- 
der Calliandra in recent regional floras. Since Brit- 
ton & Rose's treatment, only five new species of 


Zapoteca have been published. 


Zapoteca H. Hern., Ann. Missouri Bot. Gard. 73: 
755-763. 1986. TYPE: 
(Willd.) H. Hern. 


Zapoteca tetragona 


Ramose, erect, scandent, prostrate or subpros- 
trate, glabrous or hairy, unarmed, shrubs, rarely 
small trees, rarely with ed ent stipules, the 


young branches terete or 4-angled. Phyllotaxy dis- 


tehuana, 2n — 26 (prometaphase) (H. Hernández 


Volume 76, Number 3 Hernández 807 
1989 Zapoteca 


tichous. Leaves bipinnate; petiole rarely with nec- ceous, linear, straight, rarely slightly curved, plano- 
tariferous glands, usually with à conspicuous adax- compressed, with the margins thickened, usually 
ial channel; leaflets | to numerous pairs per pinna, with constrictions in interseminal areas, the valves 
opposite, sessile, usually membranous, rarely char- dehiscing elastically from apex to base. Seeds in | 
taceous or coriaceous, usually glaucous beneath, series, hard, ovoid to rhomboid, rarely ellipsoid, 


glabrous or hairy. Stipules usually conspicuous, — nonarillate, nonalate, usually with irregular or reg- 
leafy, rarely spinescent, persistent. Inflorescences ular 90% pleurogram. Seedlings phaneroepigeal: 


capitate, pedunculate, densely flowered, axillary or cotyledons ephemeral, foliaceous, sessile, elliptic to 

organized in simple or compound pseudopanicles, — elliptic-ovate: first and second eophylls opposite, 

with the peduncles solitary or fasciculate at distant the third and fourth alternate; leaflets thin, mem- 
j 


nodes, homomorphic, heterogamic or homogamic. branous. x = 13. 
Flowers bracteate. sessile, actinomorphic, (4-)5 —— 
i Distribution. Usually abundant in ope n sites 
(-6)-merous; calyx cup-shaped, dentate or dentic- 
i derived from tropical deciduous forest, in arid 


ulate, glabrous or hairy; corolla campanulate or 
or semiarid, scrubby vegetation and wet evergreen 
infundibuliform, membranous, with the petals val- i . - o 
forests. from southwestern United States and 

vate in bud, usually revolute at anthesis; nectar- . : Fun 
5 f northern Mexico, from about latitude 32°30'N, 
iferous disk present in all flowers; stamens ca. 30- : . : 
: southward through most of Mexico, Central Amer- 
60; filaments long-exserted, ca. 19-43 mm long, . li . d 
D ica, West Indies, and part of South America, to 

r a combination of two 


~ 


white, pink, red-purple, . f a Lia 
keene northern Argentina at about latitude 27230'5. Col- 
colors, the staminal tube included; anthers dithecal, . | E 
pe m lections come from altitudes of 02,850 m. 
dorsifixed, eglandular. each anther containing 8 ñ x » 
HM Zapoteca consists of four subgenera, 17 species, 
polvads: polyads 16-grained, discoid, heteromor- oe ce <li a . . 
: den i . . and 11 subspecies. The following systematic treat- 
phic, usually with eccentric lens-shaped thicken- SA » 
ment includes all the known taxa in the genus. The 
ings on the central cells; ovary 1, sessile or shortly . 
E E ^ sequence of the taxa reflects approximate. rela- 

stipitate, with ca. 10-15 ovules: style filiform, 


Ae . 
i MEO tionships. 
15-56 mm long in fertile pistils; stigma cup-shaped. 


Pods usually pendent, dry, membranous to coria- 


KEY TO THE SUBGENERA AND SPECIES OF Z APOTECA 


la. Pinnae 1-, rarely 2-jugate; leaflets usually larger, (1.5-) 2.2-22 cm long, chartaceous to coriaceous, 1-7 


pairs per pinna, usually glabrous or ine te; leaflet venation patterns str ongly conspicuous on both surfaces 

2a. Leaves with cup- rs ed glands at the insertions of the pinnae and sometimes between the distal pairs 
of leaflets; leaflets grayish, e -green or brown; polyads without lens-sha uh E hi gine pods i kly 
membranous; Blond iola ius Nervosa (1l. Z. nervosa) 


2b. Leaves eglandular; pe usually green or olive-green; polyads idi ae mie ved alas: pods 
coriaceous; Colombia, Ecuador, and Peru. 
Stipules spinescent; leaflets 3-7 pairs per pinna, ovate-lanceolate, 1.5-6.3 cm long; « capitu 
| ) IH 15 


axillary; Ecuador y hes (e Aculeata A acule 'ata) 
3b. Stipules leafy; leaflets 1 - 1/2 pairs per pinna, falcately lanceolate or falcately ovate- iren 
22 cm long: capitula in simple or compound, elongate sicud les; Colombia and Per 
IV. Subgenus poe onica 
4a. Leaflets 112 pairs per pinna, ciliate; stipules to 2 mm long; pseudopanicles simple; filaments 
white; erect Hen Colombia 16. Z. ils sa 
4b. Leaflets 1 pair per pinna glabro us Or glabrata: PNA te: 22 mm long; Saag el nen 
compound; sinn nue in tbe basal half and red-pur y; scandent shrubs; Amaz onian 
17. Z. amazonica 
lb. Pinnae ls 8-jugate; leaflets usually smaller, 0.4-4(-5.9) em d ad rarely thinly chartaceus 
3-07 pai s per pinna, glabrous to densely pubescent; leaflet venation mostly inconspicuous (except Z 
te huana); Z filipes and Z. scutellifera with somewhat Senado. veins abaxia y H. Subgenus Zapoteca 


~ 


Leaflets oblong- obovate to very widely obovate (except Z. formosa subsp. gracilis and occasionally 

subsp. rosei with elliptic to elliptic- -oblong leaflets), nii: 3-17 pairs per pinr 

6a. Leaves without glands; stipules lanceolate to ovate- lanceolate, or linear- orien to 13 mm long: 

filaments white, red-purple, white in the basal half and pink distally, or white basally and red- 

purple distally; essentially erect shrubs; southern United States and northern Mexico to uo: n 

Argentina and West Indies 12. Z. formosa 
eaves with scutelliform glands at the petioles, at the insertions of the pinnae, and sometimes 

between the distal pairs of leaflets; stipules linear-triangular, to 4 mm long; filaments white basally 

and pink distally; scandent shrubs; Brazil and Bolivia 13. Z: por ra 

5b. Leaflets narrowly oblong to elliptie or lanceolate, oblong-ovate or oblong-lanceolate excepi Z. ta- 

ricensis with the distal leaflets oblong-obovate), usually 5-67 pairs per pinna. 


e) 
— 
ej 


— 
~ 


808 


Annals of the 
Missouri Botanical Garden 


7b. 


(a. Leaflets 2 


Leaflets 0.4-2 « 


2-2.7 em long; stipules ovate to widely ovate, rounded to cordate at the base; Costa Ric 
l. 


^. costaricensis 
n long; nal EE to shallowly triangular, triangular-lanceolate, rarely 
na 


ovate; Mexico to pe Arg 


8a. 


8b. 


I. ZAPOTECA SUBG. 
NOV. TYPE: Zapoteca nervosa (Urban) H. Hern. 


Caules dense lignosi cortice fissurato. Foliorum pinnae 


unij ugae, nectaria 
in pinna utraque 


rum manifeste conspicua; stipulae foliosae. Capitula ax 
Pollinia sine tumorib 


illaria. 


crasso-membranacea glabra 


Stems densely wooded; the bark fissured. Pinnae 


: nae ne 


Branchlets conspicuously y on at least in the younger par 
9a. Filaments white; stipules triangular-linear to triangular- TM a 
basal auricles, to 2.5 cm long; seeds with regular pleurograms; Mexico to Ec 


bes | 


B tetragona 
9b. Filaments white basally and red-purple distally; stipules triangular, descending or de- 
pressed, with 2 basal auricles, to 0.6 cm long; seeds with irregular pleurograms; "m dor, 


Z. andina 
Branchlets terete or angled, but never as above (except some populations of Z. N E 
subsp. portoricensis). 
10a. Leaves with ui AIDE cylindrical glands on the petiole, at the points of sen of 

the pinnae, and usually between the leaflets; Brazil . filipes 
10b. Leaves without di (except Z. lambertiana very ca with minute glands ore. 
the proximal pairs of pinnae); Mexico to Brazil and Per 
lle l re 1% 2? 


, and 3? veins conspicuous PP. the broc gk etn 
venation pattern, mostly thinly chartaceous; stipules to 2 mm long; south 
axaca 


stern 
de hna 
Leaflets with the venation patterns mostly inconspicuous, essentially inem bre ede 
stipules usually larger than above, to 14(-17) mm long; Mexico to Brazil a 

eru. 
] 2a. 


llb. 


m predominantly 1-2(-3)-jugat 
3a. Filaments white basally i red. -purple distally; leaves glabrous, pu- 
bescent or pilose; stems usually well branched near the base; sumer m 
Texas ud northern Mexico to Puebla and Oaxaca Z. media 
ua ah white; leaves glabrous or nearly so; stems usually Tuis hed 
ar above the base; enl 3. 
Pinnae edemas 3- 7-jugate, rarely 1-22 
a. Pods membranous, with the as fra dile saan fil- 
aments red-purple; northeastern to southeastern Mex 


13b. 


alinae 


]2b. 


. lambertiana 
h the 


1 4b. Poda lead fore rigid than above (thick membranous), with 
at, glabrous, puberulent, densely v 

or pubescent filaments white, or 

Mexi 

15a. 


E 


, canescent 
white basally and red- Duis distally; 


iens white basally and red-purple distally; leaflets 0.4- 
0.1-0.3 cm; capitula commonly arranged in pseudo- 
ds sometimes axillary; subprostrate or prostrate shrubs 
/. caracasana 
15b. Filaments white; leaflets 0.4-2 x 0.1- cm; capitula com- 
Md axillary, sometimes arranged in pseudopanicles; erect 
shrubs. 


andent 
a. “Pods glabrous or oe D dps Mexico 
South America, and We 6. Z. portoricensis 
l6b. Pods densely villous sath Das tawny hairs; Costa ica 
and Panama . Z. mollis 


NERVOSA H. HERN., SUBG. — lens-shaped thickenings. Pods thickly membra- 


nous, glabrous. 

This subgenus comprises a single species, Z. 
glabra vel sub-glabra; foliola 3 
ordinatione vena- 


nervosa, which has a combination of characters 
dis unlike that of any other known species of the genus. 


us lenticularibus. Legumina 


— 


poteca nervosa (Urban) H. Hern., comb. 
nov. o nervosum Urban, Symb. 


l-jugate, with nectaries, glabrous or nearly so; Ant. 2: 257. 1900. Calliandra nervosa (Ur- 
leaflets 3-4 pairs per pinna, moderately large, co- ban) l libn & E. Ekman, Ark. Bot. 22A(8): 


riaceous, with a very conspicuous venation pattern; 


stipules leafy. Capitula axillary. Polyads without 


27. 1928. Jupunba (?) nervosa (Urban) Brit- 
ton & Rose, N. Amer. Fl. 23: 27. 1928. TYPE: 


Volume 76, Number 3 
1989 


Hernández 809 
Zapoteca 


E iN ¡És 
| | | | | ‘igs T um 


LE. 
Late | 


FIGURE 30. Zapoteca nervosa. — A. Branchlet with inflorescences at anthesis, and dehisced pods. — B. Detail of 


the primary stem showing fissured ba 


rk. — C. Detail of leaflet venation at leaflet apex. — D. Petiolar gland. — E. Flower. 


sho ss 
(From Ekman 8959 (US) and Ekman 8914 (MO); stem and flowers from Mejia et al. 35509 (MO).) 


Haiti: ad Corail, 1.200 m, Dec. 1894, Pi- 
carda 1362 (lectotype, here designated, NY 
(fragment)). Aside from the fragment at NY, 
which consists of several leaflets clearly iden- 
tifiable as this species, there appear to be no 
additional extant duplicates of Picarda's col- 
lection in any of the herbaria where his col- 


lections are presumably deposited (B, BR, L, 


and S). Figure 30. 


Shrubs to 3 m high, with slender stems ca. 2 
cm diam., the bark fissured; branchlets terete, 
grayish-brown, smooth, glabrous. Pinnae consis- 
tently |-jugate; petioles terete, with a conspicuous 


810 


Annals of the 
Missouri Botanical Garden 


100 Km 


FIGURE 31. 


channel in the adaxial surface, 5-13 mm long, 
glabrous; rachillae 2.2-7.2 cm long, glabrous, 
sometimes sparsely villous in the extremes between 
the leaflets; extrafloral nectaries located at the in- 
sertion of the pinnae and occasionally between the 
distal pairs of leaflets, elevated, cup-shaped, ca. 1 
mm diam. or less; leaflets 3-4 pairs per pinna, 
with the proximal anterior one reduced in size, 
ovate to lanceolate, sometimes rhombic, the base 
oblique, the apex obtuse to rounded, sometimes 
usually mucronulate, the 


.9-3.8 em, the proximal 


acute, rarely truncate, 
distal leaflets 2.7-6.8 x 0 
ones smaller, all coriaceous, usually grayish adax- 
ially and olive-green to brown abaxially, glabrous, 
occasionally floccose abaxially at the base, ciliate 
when developing: primary, secondary, and higher- 
order veins very conspicuous on both surfaces; 
stipules reduced to small scales, deltate, | mm long 
or less, glabrous, rarely villous. Capitula always 

axillary; peduncles solitary or fasciculate, filiform, 
(1.6-)3.7-5.5 em long at anthesis, glabrous, oc- 
casionally sparsely pubescent. Calyx denticulate to 
subtruncate, 1 mm long, glabrous or sparsely pi- 
lose, the teeth very shallowly triangular, ciliate: 
corolla infundibuliform, 4-5 mm long, glabrous, 
the lobes ovate-lanceolate; filaments purple, the 
staminal tube ca. 2 mm long; ovary ca. 1.5 mm 
iid glabrous, the stipe ca. 1 mm long. Pods ca. 


3.5 cm long, rounded at apex, rostellate, 


a membranous, Proin 
Distribution (Fig. 31). 


iola, where it has been collected at several scattered 


Endemic to Hispan- 


Su» 189 N 


68% w 


Distribution of Zapoteca nervosa, Hispaniola. 


localities, mainly on karst limestone, including lle 
Grande Cayemite, Haiti; 0-250 m 


Common name. 


Pti-gaiac (Haiti). 
Additional no ns examined. HISPANIOLA ISLAND. 


Harri: Grande Cayemite, Anse à Macon, Ekman 8914 
e Grande Cayemite, Anse du Nord, Elnan 8959 (F, 


Domingo. Coti 
tises, Boca del Infie erno, go ii 542. 3 (A, GH, US); Los 
Haitises, near the ee of Barracote River, S of Samaná 

, Liogier 14510 (NY, US); Pilancón, Bayaguana, 
e ogier & Liogier 18868 (F, NY); El Cl s Cabarete, 
Puerta Plata, Liogier & Liogier 20501 (MO. Y); Prov. 
Seibo, Parque Nacional Los Haitises, Mejía et s 35509 
(MO). 

This species, the only member of subg. Vervosa, 
is highly isolated morphologically and geographi- 
cally from the rest of the genus. Its general features 
and the manner of dehiscence of the pods are 
identical to those of subg. Zapoteca. These char- 
acters, as well as the overall characteristics of its 
inflorescences and flowers, justify its inclusion with- 
in Zapoteca. It is readily recognized by having 
fissured bark, extrafloral nectaries, conspicuous eu- 
camptododromous / brochidodromous leaflet vena- 
tion patterns, large, coriaceous, grayish leaflets, 
and denticulate to subtruncate calyces, all of which 
are unique in the genus except the extrafloral nec- 
taries. The stems of Z. nervosa, which are char- 
acterized by having dense hardwood, differ from 
those of subg. Zapoteca, in which the wood usually 
is comparatively softer and lighter. Of particular 


Volume 76, Number 3 


Hernández 811 


Zapoteca 


interest, however, is the absence of the jun 
lens-shaped structures in the central cells 
16-celled polyads (Guinet, pers. comm.), ou E as 
pointed out above. are important diagnostically in 
the genus. This plesiomorphic character state in 
Z. nervosa, in association with its distinctive mor- 
phology, suggests that this species has been sep- 
arated from the main Zapoteca stock since early 
in the evolution of the genus. 


II. Zapoteca subg. Zapoteca. TYPE: 
tetragona (Willd.) H. Hern. 


Zapoteca 


Benth., 


Trans. Linn. Soc. 


London J. 
London 


eii be ser. Laetevirentes 
. 1844; Benth., 

: 542. 1875. 
culiando Benth. sect. Eucalliandra Griseb., Fl. Brit. 
Pei 224. 1860; Cat. Pl. Cub. 83. 1866, pro 


— 


Brit. 


Pseudacacia Griseb., F 


Celtis nde Benth. sect. 
W. Ind. 224. 1860. 

Anneslia Salisb "group ue Britton & Rose, N. 
mer. Fl. 23: 49 28. 

omisi Salish ‘group Mol ulae” 
mer. Fl. 23: 49. 

dimesiia Salisb. * Lu Portorcones 

1928. 


N. Amer. Fl. 23 


Britton & Rose, N. 


Britton & Rose, 


Caules RA molliter lignosi, cortice fere laevi. 
Pinnae (1-)2-8-jugae, petiolo plerumque sine nectariis 


glabris vel dense pubescentibus, foliolis (2- )3- 07 -jugis is 
n- 


conspicua; stipulae foliosae. Capitula ax 
dopaniculis simplicibus vel Rond disposi Pollinia 
tumoribus lenticularibus praedita. Legumina crasso-mem- 
branacea, glabra vel dense iri Semina pleuro- 
gramate praedita 

Stems usually with soft wood; bark essentially 
smooth. Leaves glabrous to densely hairy, the pin- 
nae (1-)2-8-jugate, the leaf-stalk usually without 
nectaries, leaflets (2-)3-67 pairs per pinna, usually 
small, occasionally moderately large, membranous, 
rarely thinly chartaceous; venation usually incon- 
spicuous; stipules leafy. Capitula axillary or orga- 
nized in simple or compound pseudopanicles. Po- 
Pods thick- 


membranous, glabrous to densely pubescent. Seeds 


lyads with lens-shaped thickenings. 
with pleurogram. 


This subgenus is the most diversified evolution- 
ary line within Zapoteca, comprising 13 species 
and ll subspecies. It ranges from the southern 
United States and northern Mexico to northern 
Argentina. 


2. Zapoteca media (M. Martens & Galeotti) H. 
Gard. 73: 755- 


Acacia media M. Martens & 


Hern., Ann. Missouri Bot. 
163. 1986. 


Acad. Roy. Sci. Bruxelles 10: 
316. 1843. Anneslia media (M. doped & 
Galeotti) Britton & , N. Amer. Fl. 23 

66. 1928. Calliandra ds (M. Mans. A 
Galeotti) Standley, Publ. Field E Mus.. 

Bot. Ser 1929. : Mexico. Hi- 
lalgo: Hacienda de e pres Mextitlán, 
5,000 ft., June-Oct. 1840, Galeotti 3302 
pro parte (lec oe BR; Hernandez, Ann. 
Missouri Bot. 
are four specimens labeled Galeotti 3302, but 


Galeotti, Bull. 


Gard. 73: 757. 1986). There 
they are not duplicates of the specimen at BR 
and consequently are not treated as isolecto- 
types. The specimen at K is from "ravines de 
Regla," whereas the ones at G and P are from 
“ravines de Real del Monte”; a fragment at 
F lacks locality information. As MeVaugh 
(1978) pointed out. evidence indicates that 
Galeotti's Mexican collections were sorted sys- 
tematically and then arbitrarily numbered af- 
ter they arrived in Europe. In this way, three 
collections made on different dates were 
brought together under number 3302 after 


they were interpreted to represent the same 


species. 
Calliandra angelica Benth., London J. Bot. 3: 100. 
1 an th., Trans. Linn. Soc. London 30: 544. 


1875. Feuilleea angelica (Benth.) Kuntze, Revis. 
Gen. Pl. YPE: Mexico. Hidalgo: ravines 
Re E ee 3362 pro parte [lectotype here 
a "signated, K. The m where Bentham 
e located of Galeotti 
3362 with the loc ality formation specified by Ben- 
tham. As noted above, other sheets with this e 
(at BR, G, and P fuse different locality data 

Calliandra ws S. Watson, Proc. Amer 
s : 3902 882. dnneslia coulteri (S. Watson) Brit- 
1 Amer. Fl. 23: 66. 
Meno: C volui Soledad 25 mi. "pede of Mon- 
v . 1880, Palmer x 2 otype he 
nated, im isc a ectotypes K, S fragment) 
adici x oaxacana Rose, Contr. uc Natl. Herb. 
189 Anneslia oaxacana (Rose) Britton & 

2! 


= 
a 


ua N ones Fl. 23: 66. 1928. TYPE: Mexico. 
Oaxaca: granitic ledges, Tomellin Canyon, 4,000 
10 July 1897, Pringle 0:34 (lectotype here 


designated, US no. 316706; isolectotypes, A, BM, 
GH, K, LE, MEXU, MO, NY, S, UC, US). 


Shrubs 0.2-1.5(-3 


usually branching at the base; branchlets terete, 


— 


m tall, the stems slender, 
costate under magnification, glabrous to densely 
pubescent or pilose. Pinnae 1-2(+3)-jugate: peti- 
oles eglandular, terete, with a conspicuous channel 
in the adaxial surface, 0.7-4 cm long, glabrous to 
sparsely pubescent; rachis of 2-jugate pinnae (1-) 


2-12 mm long; rachilla |—4.5(—7) em long; leaflets 


812 Annals of the 
Missouri Botanical Garden 
\ o 
- ` EN Y 
. > me 
à > N MÀ 
à E T N 
4 E e a \ 
/ . o % n = 
4 V B ny 
A D 3 V 
2s ^ NS j 
\ | 
S 
d N $ o 
b > . \ 
e? 5 N iod 
A 1 t : 
8 4 f 
y hat í f 7 
~ MJ * p = il e 
e s | J 
|. | Y 
ee N F. 
` à E me 
15°N 
= | 
$ o 300 
Km já: 
FIGURE 32. Distribution of Zapoteca media. Populations with essentially glabrous leaflets = dots; populations 


with pilose or pubescent leaflets = triangles. 


(5-)7-17(-22) pairs per pinna, oblong to elliptic, 
oblique at the base, ae proximal pairs 
cuneate at base, acute to rounded at the apex, 

4-13 


branous, glabrous to pubescent or pilose on both 


xcept 


sometimes rostellate, 5.5 mm, mem- 


surfaces; leaflet venation usually inconspicuous; sti- 
pules leafy, lanceolate to ovate-lanceolate, rarely 


widely deltate, striate, (1—)2- 7 mm long, glabrous 
to pubescent or pilose. Capitula axillary, rarely in 
short, terminal pseudopanicles; peduncles solitary 
fasciculate at distant nodes, 


or, more commonly, 
(1-)1.6-7(-10.5) em long at anthesis, glabrous, 
sometimes pilose. Calyx ca. 1.52 mm long, gla- 


brous or finely ciliate, rarely sparsely pilose, the 
teeth narrowly oblong, acute at apex; corolla cam- 
panulate, ca. 3-3.5 mm long, glabrous, the lobes 
narrowly elliptic, acute at apex; filaments ca. 20 
mm long, white in the basal half, red-purple in the 
distal half, staminal tube ca. 1 1.5 mm long; ova- 
ries ca. 1 mm long when fertile, occasionally 2 per 
flower, glabrous, shortly stipitate. Pods to 10 x 
0.6 cm when mature, apex rounded to acute or 
truncate, rostellate, thickly membranous, glabrous, 
reticulate-veined. Seeds widely rhomboid to widely 
x 3 


ovoid, flattened, mm; pleurogram ir- 


E 


regular. Somatic RR number 2n 


— 


Distribution (Fig. 32). 
primarily on dry rocky calcareous slopes and along 


This species occurs 


the margins of seasonal streams in thorn and trop- 
ical dry forests, and in arid scrublands. It ranges 
from the Chinati Mountains (Presidio County) of 
Texas south over the Mexican Plateau and the arid 
slopes of the Sierra Madre Oriental and, to a lesser 
extent, the Sierra Madre Occidental, where it has 
been collected at scattered localities. 
limit is in the districts of Teposcolula, Nochixtlán, 
and Cuicatlán of Oaxaca, about 1 7?30'N. Collec- 
tions come from elevations of 1,000-2,150 m. 


Its southern 


Common name. — Cabellito de angel (Hidalgo). 

Additional examined. U.S.A. TEXAS: 
Presidio Co., 5 Chinati Mesquite Ranch, Tinaja 
Prieta Canyon, Lott 64 n MEXICO. AGUASCALIENTES: 
9 km NE Calvillo, sobre carretera a Aguascalientes, Aze- 
dowski - MeVaugh 1245 (ENCB, MICH); road to Cal- 
villo, W of Aguascalientes, near km 40, Mclaugh 18303 
(MICH). CHIHUAHUA: vicinity of E 4 km SE Rancho 
Enc illas; wart 773 (F, GH, LL, MEXU ). COAHUILA: 


4 (air) mi. E Em air) mi. 


fiat 


103°42'W), He nrickson 6876 
ea Canyon of Sage Blanca, central mass o 
lo as Cruces, W iens M ies, Johnston & Mueller 
| 


208 (GH, LL, 


Canyon Bonanza, SE CM ra Purísima, Johnston et al. 


| slope of 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


l'anón i. la Gavia, 5 Rane ho 


10300 (LL, MEXU, MO); ( 


de la Gavia, Johnston et al. 3 (LL, MEXU); 1. 
mi. N of Puente la Muralla He 57 (199 mi. 5 of 
Piedras ras) Lewis 82 (LL) Müzquiz, Santa na 


Canyon, ido 140 (F, GH, MENU, TEX); 5 

Mts., Marsh 1457 (F, GH, TEX); ae 25 mi. 
Monclova, "Palmer 2129 (GH, S) Canon del Indio 
Felipe, ca. 28?33'N, Sierra Hec a ‘eros, close to the Chi- 
huahuan boundary, Stewart 160 (GH); vicinity Rancho 
Tule, 24 km N Castillón, close to the Chihuahuan 
borane, Stewart 512 (LL, GH): Sierra de la Gloria, 
Cañón Oscuro C hiquillo, entering C. Chilpitin from S near 
El € -hilpitín (26°47'3 due 10121 7'50"W ), H endt & Ris- 
kind 1699 (ENCB, MEXU, MO). DURANGO: Mpio. 
Tepehuanes, Los Cedros al SW de Tabal iueto, camino 
ienda- Topia, al. 4207 


bahueto (al Catorce), 196 km W de Tepehuanes, Torres 
gi al. 3553 (MEXU). GUANAJUATO: near Xichú, Kenoyer 
2254 (A) HIDALGO: Mpio. Tasquillo, 3 km N Puente 
Tasquillo del Rio Tula, c arr. a a Zip, Delgado et al. 
e ; (ENCB, F, MEXU, 5 km Pina 4n. Ojo de 
Agua, González Q. 2751 P ); Paso del León, entre 
a el Grande y Los Venados, H. Hernandez & 
Torres 129 (MEXU, MO): 12.6 km de Los Venados por 
Barranca de Meztitlán, HM. Her- 
MO); Mpio. Zimapán, 8 


He rnández 


carretera a Meztitlán, 
nández & Torres 133 (MEXU 
km de Zimapán, en el Cañon de Tolimán, 7T. 
& Torres 153 (MEXU, MO): Mpio. Taxquillo, cuesta de 
Texquedo, R. Hernández 3644 (ENC B, (Us 5 km 
i s, Medrano et al. ); Mpio. Car- 
, Canada de La Pi bra a de To- 
pines Medrano et al. 9152 (ME NI MO); Barranca 
de Metztitlán, Medrano & Ortiz 106% 80 (E. NCB, MEXU 
km 276 on highway NE Jacala, Moore & Wood 
3980 (A, MEXU. MICH); Barranca de Tolimán, on road 
from Pues to Mina Loma del Toro, Moore & Il ood 
1383 (A, MEXU, MICH); near Denon, Rose et al. 
8928 a NY, US) cerca de Orizatlán, Rzedowski 
32210 (ENCB); Cerro del Junquillo, SW de Tasquillo, 
Tenorio & Romero 505 “XU, MO); 15 km N Ato- 
tonile o, por carr. a Tampic o, Hillaseñor et al. 128(N e 
NUEVO LEÓN: Huasteca E ca. 
(by road) SW pag IF ard 5724 (MIC H). OAXACA: 
Dist. Etla, El Parián, Conzatti 1558 (US); Distrito No- 
ahlin, El Parián. Conzatti 1929 (F), Dist. Uk LU 
de León, 2 km S de Reforma, carr. Acatlár A. 
García et al. 1454 (MENU, MO); cerros SE oe 
Miranda 4747 (MEXU); Lagunas, Orcutt 027 
Tomellin a i Pringle 6734 (ENCB, F, K 
, UC, US); Dist. Teposcolula, 5 km NE Chilapa de 
Ro edowski 34813 (ENCB); Totolapan, Rio Te- 
huantepec, Sousa 2666 (CAS). PUEBLA: highway 150 km 
259 SE Mexico City, Barr A al. 62 (MENU); Mpio. 
os de Osorio, 3 km de atlán por carr. a Tehuit- 
go, H. Hernández & bes 192 (ME XU, MO); Ani- 
cano, Miranda 2921 (MEXU); Pollatzin, Miranda 2948 
(MEXU) San Luis Tultitlanapa, Purpus 2007 (BM, 
an MO, NY, UC) Mpio. Cuicatlan, 15 km al 5 ds 
Dominguillo, sobre la carre R zedowski 


34940 (ENCB) Tehuitzingo, 9804 


HE 
— 
QE 
[e 
= 
O 
T: 


a 
t 


oX, 


Dia. 


Sou sa 


(ENCB, MEX U). QUERETARO: camino a | San Luis Potosí, 
km 10, Arguelles 1400 (ENCB, MENU); Mpio. Jalpan, 


a Ee e camino a Pinal de Amoles, Fernández 
3163 (MEXU): Rose & Rose 
(NY, ^ S). SAN LUIS POTOSÍ: camino Rio Verde 


ear Querétaro, 
San Ciro, 


Rzedowski 4511 (MEXU). TAMAULIPAS: Sierra de San 


Carlos, vicinity of San Miguel, La Tamaulipeca, Bartlett 

10590 (F, US) El Madroño, 14 m y 
. Victoria along X 101 ega Polis i et 

al. 42 ( 10); ia fex gd 


leno o 
Nogales, M^ ynski 489 (B. 
(US). ZACATECAS: San Juan € onem Rose 3550 US E 
8 km NE Jalpa, Rzedowski 14190 (ENCB); 5 mi. NE 


Jalpa, Me : iod 18485 (MICH). CurTIvATED: U.S.A 
Missouri, St. Louis, Missouri Botanical Garden, H. Hae T- 
nández 9. a (MEXU, MO). 

This a plastic. species is wide- 


spread in dry and semiarid regions of central and 
northern Mexico. It can be readily distinguished 
from its close relative Z. alinae primarily by having 
larger, white/ purple-red filaments, usually shorter 
stature, and copious branching. In addition, the 
leaflets of Z. media are usually broader than those 
of Z. 
species is discussed further under Z. alinae. 


As shown in Figure 32, 


alinae. The relationship between these two 


most of the known 
populations consist of plants with leaves glabrous 
or ciliate and very sparsely pilose on the rachis 
and rachilla. Plants with pubescent or pilose leaflets 
have been collected primarily in restricted areas 
in the southern part of its range (Oaxaca, Puebla, 
San Luis Potosi, and Ta- 


Hidalgo, 


maulipas). Most of the collections with hairy leaflets 


Guanajuato, 
are concentrated west of Hidalgo (Fig. 32), where 
With the exception of 
a population from the Barranca de Tolimán, Mpio. 


Galeotti collected the type. 


Zimapán, Hidalgo. where plants with hairy and 


. with glabrous leaflets have been collected (41. Her- 


nández & Torres 15%, Moore 1383), populations 
of plants with hairy leaflets seem to be geograph- 
ically segregated from the glabrous ones. 

The distributions of Z. media and Z. formosa 
approach one another closely in some areas of 
northern Oaxaca and southern Puebla, but indi- 
viduals from these species are not known to grow 
together. There is evidence of hybridization, which 
results in morphological intergradation between the 
two. Plants collected in the area of Acatlán de 
Osorio and Tehuitzingo, Puebla (11. Hernandez & 
Torres 192, 2921, 294 
9801). and Cuicatlán and Nochixtlán. 
(Conzatti 1929, 1747 


hairy leaflets, fewer pairs of leaflets per 


Miranda 8. Sousa et al. 
Oaxaca 
Miranda ) have character- 
istics (e.g., 
pinna, and broader leaflets) that differ from those 
typical of Z. media and tend to approximate those 
of Z. formosa subsp. mollicula or subsp. roset. 
3. Zapoteca alinae H. Hern., sp. nov. TYPE: 
Mexico. Oaxaca: 6 km NE 
de Mitla por camino a Albarradas, 1.090 m, 


veg. 


Distrito Tlacolula. 


matorral xerófito muy alterado, | Aug. 


814 


Annals of the 
Missouri Botanical Garden 


7 E 33: 


Zapote ca alinae. 
B. Le —C. Stipules. — D. Flower. 
MO): s from Hu He UR 952 (MEXU, 


1983, H. Hernández & R. Torres 206 (ho- 
lotype, MEXU; isotype, MO). Additional iso- 


types to be distributed. Figure 33 


Frutices recti caulibus valde grac sn omnino Ls 
vel fere glabris. Pinnae 1-jugae, interdum 2- vel 3 
foliola 16 19-juga in pinna utraque ue bee el 
lanceolata, venatione inconspicua. Capitula axillaria raro 
in pseudopaniculis brevibus simplicibusque disposita. Flo- 
res filamentis albis praediti. Legumina crassa, membrana- 


A. Branchlet with inflorescences at anthesis, and dehisced and indehisced pods. — 


E. SR F. Seeds. 
0).) 


(Drawn from living plant H. Hernández 942 (MEXU, 


cea glabra. Numerus chromosomatum somaticus 2n = 
20. 

Shrubs to 4 m high; stems very slender, to ca. 
1.5 em diam. at the base, the bark smooth; branch- 
lets terete, costate under magnification, dark, more 
or less lustrous, glabrous. Pinnae 1(-3)-jugate; pet- 
iole eglandular, terete, 0.9 3.7 cm long, glabrous; 
rachis 3-10 mm long in leaves with 2 or more 
pairs of pinnae, glabrous; rachillae 1.7-4 cm long 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


815 


FIGURE 34. 


with star). 


in well-developed leaves; leaflets 16-19 pairs per 
pinna, narrowly oblong to lanceolate, at base 
strongly oblique, at ix x rounded or acute, usually 
mucronulate, 4-12 2.5 mm, membranous, 
glabrous, sometimes enne leaflet venation in- 
conspicuous; stipules leafy, narrowly triangular to 
lanceolate, 2-3 mm long. glabrous. ee ieee ax- 
illary, rarely also in simple pseudopanicles 1.5 

cm long: peduncles solitary or fasciculate, filiform, 
2.5-6.5 em long at anthesis. Flowers ca. 15-25 
per capitulum; the calyx ca. 2 mm long, the teeth 
ovate-lanceolate, acute at apex, mucronulate, gla- 
brous; corolla campanulate, ca. 4 min long, the 
lobes ovate to narrowly ovate, acute at apex, gla- 
brous: filaments 15-20 mm long, white, the stam- 
inal tube ca. 1 mm long; ovaries ca. 1.2 mm long, 
on occasion 2 in a single flower, glabrous; style ca. 
16 mm long. Pods to 10 x 0.8 cm when mature, 
thick-membranous, 


at apex rounded, rostellate, 


glabrous. Seeds widely rhomboid to widely ovoid, 
flattened, grayish or darker, ca. 5 X 4 mm; pleu- 
rogram irregular, usually demarcated by a dark 


line. Somatic chromosome number 2n = 26 


Distribution (Fig. 34). 
central Oaxaca, 


Essentially endemic to 


in the districts of Tlacolula, Yau- 


Ls (ENCB, U 


Map of Oaxaca, Mexico, showing the distribution of Zapoteca alinae (dots) and Z. tehuana (dot 


tepec, Centro, Miahuatlán. and Teposcolula. Za- 
poteca alinae grows mainly on steep. rocky slopes 
in tropical deciduous and thorn forests and in sec- 
ondary vegetation derived from these vegetation 
types; it has also been collected along seasonal 
streams. Collections come from elevations of 1,050 


1.800 m 


Common name. Barba de chivo. 


Additional specimens examined. — MEXICO. OAXACA: 
nue t Centro, Cerro del Tule, Conzattt 5474 (NY); 5 
Miahuatlán, 1007 (MEXU); Dist. Tla- 
Sa a 7 km NE de Mitla por carr. a Tamazula yan, 
He rnández & Torres 27 1 (MEXU, MO); Dist. 
de Totolapan por carr. a Tehuantepec, 
He rnández & Torres 280 (MENU MO): Dist. Tlacolúla, 
33 km de San José de Gracia sobr carr. Tehuantepec 
Oaxaca, entre San José de Gracia y San Juan Guegoyache, 
Ramírez 939 (MEXU, Dist. 
W de Totolapan por y a Oaxaca, 
2 (ME . MO); arroyo 
abajo de Cueva — Se hoen niwetter 
JS); E d canón Ri 
Sousa 2689 (CAS, GH, MEXU, US); 
km NE San ip Albarradas, 19 a E-NE 
7846 (MEXU, N Tlac B 
«e Téllez 850 ) (€ 
Yautepec, 1 km 5 El a 


Garcia 


a 


Tine solula, 


-— 


> Tlac HY 


Sousa et al. 
kmi E-NE Totolapan, Sousa 
ENCB, MEXU, MICH); Dist. 


e 7 
— 


816 


Annals of the 
Missouri Botanical Garden 


rr. Est. Microndas, Sousa & Téllez 8611 (MEXU, MO); 
Dist. Tepossnnus, Las Pilas, a 4 km al E de Tamazulapan, 
Sousa et al. 899] (MEXU, MO); Dist. "He 16 km 
N Totolapan, Sousa et al. 9424 (ENCB, MEXU); Dist. 
Tlacolula, 6 km N de Mitla, carr. a a Torres 
et al. 3947 (MEXU, MO); Dist. Yautepec, Torre de 
zy onde San Cristobal, al W de Portillo Nejapa, Torres 

: Martinez 7449 (MEXU). 


This endemic species, which is relatively con- 
sistent morphologically, is closely allied to Z. media 
by its l- to 3-jugate pinnae, small, numerous leaf- 
lets, and habitat preferences. It can be readily 
distinguished from that species by the usually more 
slender stems, branchlets, and peduncles, narrower 
and consistently glabrous leaflets, and primarily by 
its flowers with shorter, white filaments. Zapoteca 
alinae is allopatric with Z. media. Its distribution 
overlaps at the east of its range (H. Hernández & 
Torres 942) near Totolapan, Dist. Tlacolula, with 
that of Z. formosa subsp. rosei (H. Hernández & 
Torres 943), but no evidence of hybridization was 
observed there. 


4. Zapoteca lambertiana (G. Don) H. Hern., 
Ann. Missouri Bot. Gard. 73: 755-703. 1980. 
Acacia lambertiana G. Don, Edward’s Bot. 
Reg. 9: t. 721. 1823. 
tiana (G. Don) Benth.. 


Calliandra lamber- 
London J. Bot. 3: 100. 
1844. Benth., Trans. Linn. Soc. London 30: 
543. 1875. Feuilleea lambertiana (G. Don) 
Kuntze, Revis. Gen. PI. 188. 1891. Anneslia 
lambertiana (G. Don) Britton & Rose, N 
Amer. Fl. 23: 66. 1928. TYPE: 
cality unknown, cultivated by George Don from 


Mexico: lo- 
material sent to him by Mr. Lambert from 
Boyton, where plants of this species had been 
raised from seeds collected in Mexico (lecto- 
type, K; Hernández, Ann. Missouri Bot. Gard. 
73: 757. 1980) 


Erect shrubs to 4 m tall; stems slender; branch- 
lets terete, costate under magnification, glabrous 
to villous or pilose. Pinnae (1-)2- 4-jugate, rachis 
usually eglandular, rarely with small (ca. 0.3 mm 
diam.), cup-shaped, slightly elevated glands be- 
tween the basal pairs of pinnae, the proximal pairs 
of pinnae rarely alternate; petiole terete, with a 
conspicuous adaxial channel, (1.2-)1.5-4.6 cm 


long, glabrous, rarely villous or pilose; rachis of 


2-jugate pinnae (0.2-)0.4-1.8 cm, much longer 
when 3- or 4-jugate, glabrous to villous; rachillae 
2.4-6.4(-7 1 7(-22) pairs 


per pinna, narrowly oblong to oblong or oblong- 


1.5) em long; leaflets (6 —)7 


ovate, occasionally lanceolate, at base oblique, at 


apex acute, the median ones 8 14(-16) x 2.5- 


4(-5) m membra- 


nous, all glabrous to villous or puberulent on both 


m, the proximal ones smaller, 


surfaces, occasionally hairy only abaxially, some- 
times ciliate; leaflet venation visible superficially; 
stipules leafy, triangular-lanceolate to ovate, acute 
at apex, striate, 2-8 mm long, glabrous to sparsely 
villous. Capitula axillary, or more commonly ar- 
ranged in compound or most commonly simple 
pseudopanicles 3.5-9.5 cm long: peduncles fas- 
ciculate at distant nodes, (0.5)1.5-6.5 cm long at 
anthesis, glabrous. Calyx ca. 2 mm long, glabrous, 
sometimes ciliolate, the teeth narrowly oblong or 
triangular, acute at apex; corolla campanulate, 3- 
4 mm long. glabrous, the lobes narrowly elliptic, 
acute at apex; filaments ca. 2.5 cm long, red- 
purple, the staminal tube 1-1.5 mm long; ovary 
sessile, glabrous. Pods to 9 x | cm when mature, 
rounded, retuse or acute at apex, rostellate, mem- 
branous, glabrous. Seeds quadrate-rhomboid to very 
dark brown, 


widely ovoid, 3.5 3-3.5 mm, 


the areola Does pleurogram irregular. 
Distribution (Fig. 35). 


tiana has been collected primarily in tropical dry 


Zapoteca lamber- 


forests and less commonly in tropical thorn, tropical 
subdeciduous, or tropical evergreen forests, where 
it usually is associated with secondary vegetation. 
It is distributed discontinuously from northeastern 
to southeastern Mexico, where a series of disjunc- 
tions occur. Most collections have been made in 
the following regions: 1) Sierra de Tamaulipas, 
2) Region of Villa Juarez and Huas- 
3) Coastal Plain of the Gulf 
of Mexico, east and northeast of Xalapa, Veracruz; 


Tamaulipas; 
teca, San Luis Potosi; 


4) Region of Los Tuxtlas, Veracruz; and 5) Slopes 
of the Central Depression of Chiapas. Collections 
come from altitudes of 190-900 m 


Common names. Barba de chivo (Tamauli- 


— 


pas), Usak xiixit (Huasteca 


Additional dp ce e us d. | MEXICO. CHIAPAS: 
Mpio. Cintalapa, Rizo de Breedlove 36732 (DS, 
MEXU, MICH, NY); Mpio. T ón. above Chicoasén, 
Breedlove 3725. e MEXU, TEX); Mpio. Acalá, 45 
km from Tuxtla, P resa La Angostura, Breedlove 37412 

JS, MEXU); Mpio. pon de iron El Chorreadero, 
Breedlove 39694 (DS, MEXU, 
nández & Ramírez 891 eem 
cona-5. Fernando (NW Tuxtla Cutiérre x ). Miranda 6462 
(MEXU); 5.6 mi. SE de Chiapa de Corzo, El Chorreadero, 
Ton 2942 (DS, ENCB, LL, MO, NY, US); Mpio. Venustia- 
no Carranza, above Finca Carmen, along the road from 
Acalá to Pujiltik, Ton 3042 (DS, MO, NY, US). san LUIS 
porosi: S of Villa Juárez, Clark 7418 (MO); Mpio. San 
Antonio, Lejem, Alcorn 1756 (MEXU). TAMAULIPAS: Mpio. 
Aldama, cas i ba iu) ne bu 40 km NNW of 
Aldama, Los € a Borrada pud 
98°10'W), ee paras ns Mpio. Aldama, 17 


Volume 76, Number 3 


Hernández 
Zapoteca 


817 


d 
\ 
s A 
Y 
x 
" 
E 
Y 25°N 
! = 
T o 
[to] 
v e o 
2 | í 
ne 
«e ENS 
2 gi | 4 
e D 
$e `N. | 
> peed 
400 Km £ 7 
f / 
ba PEE 


FIGURE 35. Distribution of Zapoteca lambertiana. 


km W de la carr. Aldama-Soto la Marina, por camino a 
Rancho Las Yucas, 38 km NW de Aldama, H. i n 
& Torres 147 (MEXU, MO); Chamal, Ocampo rd., 

oyer & Crum 35960 (A, MIC H): n Los Alados, Cerro 
de Metate, Martínez O. 264 (MENU, MO). VERACRUZ: 
Mpio. Actopan, Sierra de Manuel Dos po 134 (XAL, 
MO); Mpio. Vega de Alatorre, Canada de Mesillas, y 
por Santa Ana, Calzada 7724 (XAL) Cerro Gordo, 
km de Plan del Rio, carr. Xal: pa Veracruz (19°2 SM 
96°39'W), Castillo & Tapia 1223 (XAL); Mpio. Vega 
de Alatorre, El Centenario, a 20 k cm de Santa Gertrudis, 
Castillo & Venavides 2003 (XAL); region of San Andrés 
Tuxtla, along river draining Lagi ma de Catemaco, Dres- 
sler & Jones 226 (BM, GH, MEXU, MICH, MO, NY, 
UC); Mpio. San Andrés Tuxtla, Salto de Eyipantla, H. 
Hernández & T orres 780 (MEXU, MO); rim of the 
Barranca Zacuapán, Purpus 10770 (US); 
292 e entre El Carrizal y Los 
y > la Laguna Nixtama- 
lapan, d 2201 MENU x a M del Lago Catemaco, 
AS, GH, ME ; Mpio. Zapata, La Laja, 
entre ( Sd D y B a 900 m de carr. Jalapa- 
Veracruz, 2 or 16 km SE Jalapa, Sousa & Ramos 
1757 (MEXU); Dos Rios, Plan del Rio, Fentura 4197 
(ENCB, ua. Mpio. Dos Ríos, Coral Falso, Ventura 


T 


Souse sa 


5807 (ENCB, MEXU, MICH); Dos Rios, Plan del Rio, 
Ventura 10594 (CAS, ENCB, MEXU) 


Zapoteca lambertiana can be readily distin- 
guished from other species of subg. Zapoteca pri- 
marily by its much thinner pods with undulate 
surfaces and thinner margins, and specifically from 
Z. portoricensis by its red-purple filaments. Extra- 
floral nectaries between the pinnae in a single spec- 
imen (Dressler 2006, GH) and presumed similar- 
ities in the texture of the pods suggest a bu 
close relationship between this species and Z. 
lipes. Leaf indument is very variable in Z. "a 
bertiana: it varies at both intra- and interpopu- 


lation levels and does not seem to follow a 


geographical pattern. 

There is no morphological differentiation among 
the disjunct populations that occur over the ex- 
tensive geographical range of Z. lambertiana. It 
is likely that such discontinuities are of recent 


origin. Perhaps they resulted from tropical dry 


818 


Annals of the 
Missouri Botanical Garden 


forest fragmentations that occurred in the tropics 
subsequent to the Pleistocene glaciation, after the 
present wetter conditions originated disjunctions in 
the formerly extensive and continuous areas cov- 


ered by tropical dry forests. 

5. Zapoteca filipes (Benth.) H. Hern., 
nov. meet Benth., J. Bot. (H 
er) 2: ; Benth., Lond. J. Bot. 
99. C D Trans. Linn. Soc. 
30: 542. 1875; Benth. in C. Martius, Fl. Bras. 
15(2): 412. 1876; J. F. Macbr., Field Mus. 


Nat. Hist., Ser. Bot. 13: 71. 1943; L. Cár- 
denas, Revista Fac. Agron. (Maracay) 7: 131. 
1974. Feuilleea dimidiata Kuntze, Revis. 


Gen. Pl. 185. 1891. (This was published as a 
nomen novum because the name F. filipes 
was preoccupied by a species based on /nga 
TYPE: Brazil: Pohl 1458 (lectotype, 


felipes.) 
K, here designated; isolectotype, 

Erect shrubs to 2 m tall; branchlets terete, cos- 
tate under magmineation, sparsely villous, becom- 
ing glabrous. Pinnae 4—5-jugate; petioles terete to 
subtetragonal, with a conspicuous adaxial channel, 
2-3.5 em long, glabrous to sparsely villous; rachis 
2.5-4.5(-7.5) em long; rachillae 2.5-5.5(-7) cm 
long; with. glands near the base of the petiole, at 
the insertions of the pinnae, and usually at the 
insertions of the leaflets, these cv'indrical, concave 
at apex, to ca. 1 cm long but usually smaller; 
leaflets (9-)1 1 16 pairs per pinna, mostly falcately 
oblong-lanceolate, oblique at the base, at apex 
rounded to acute, 8-14 x 2.5-5 


mm, membranous, glabrous or at most sparsely 


mucronulate, 


ciliate; primary and secondary leaflet veins con- 
spicuous, especially abaxially; stipules leafy, lan- 
ceolate, parallel-veined, usually 3-5 mm long, gla- 
brous. Capitula axillary; peduncles fasciculate, 
(3.5-)4.5-8 em long at anthesis, glabrous. Flowers 
basically pentamerous but the number of flower 
parts variable: calyx ca. 2 mm long, glabrous, the 
teeth irregular, triangular or narrowly oblong, acute 
at apex; corolla campanulate, ca. 4 mm long, gla- 
filaments white in the 
basal half and red-purple in the distal half, the 
staminal tube ca. 2 mm long; ovary 1.5-2 mm 
long, sessile, glabrous. 


brous, the lobes lanceolate; 


Pods ca. 7.5 x 7 em 
when mature, at apex truncate, 


branous, glabrous. 


rostellate, mem- 


Distribution. This species has been collected 
at a few localities in the Brazilian states of Goiás 
and Minas Gerais, where it is commonly associated 
with cerrado vegetation at 950 m elevation. It was 
incorrectly reported by Macbride (1943) from Peru 
and by Cárdenas (1974) from Venezuela. 


comb. 


L dos 


Additional specimens examined. 
GERAIS: Sierra do ee tana ca. a km N 
road to Salinas, Irwin et al. 2 
810-811, Pabst & Pereira E 1 (NY 
Pohl 3492 (F, NY). 


BRAZIL. MINAS 


— 


GOlÁS: Salgado, 


Zapoteca filipes is probably closely related to 
Z. lambertiana as judged by the vegetative char- 
acteristics of these species, especially by the com- 
mon possession of membranous pods. Extrafloral 
nectaries, universal in Z. filipes, are occasional in 
Z. lambertiana. Zapoteca filipes can be readily 
distinguished from the remaining species of subg. 
Zapoteca by the cylindrical glands on the petiole, 
rachis, and rachillae. 


Hern., 
163. 1986. 


Collectanea 4: 


6. Zapoteca e eru H. 
Ann. Missouri Bot. Gard. 73: 755- 
Mimosa portoricensis RN 


143. 91; Jacq., Icon. Pl. Rar. 3: 20, t. 
633. 1793. Acacia portoricensis (Jacq.) 
Willd., Spec. 4: 1069. 1806; DC., Prodr. 2: 
467.1825;Schldl. & Cham., Linnaea 5: 595. 
1830: G. Don, Gen. Hist. 2: 419. 1832; 
Schldl., Linnaea 12: 566. 1838. Calliandra 


portoricensis (Jacq.) Benth., 
3: 99. 1844; Benth., 


"giga Bot. 
London J. ved £102. 


1846; Griseb., Fl. Brit. W. Ind. 24. pr 
Benth., Trans. Linn. Soc. Lenis 30: 543, 
1875; Benth. in C. Martius, Fl. Bras. 15: 411. 
1876; Standley, Publ. Field Mus. Nat. Hist., 
Bot. Ser. 18: 492. 1937; Standley & Calde- 
ron, Fl. Salvadoreña 125. 1941; J. ee 


Field Mus. Nat. Hist., Bot. Ser. 13: 72. 1943, 
pro parte: Standley & Steyerm., doo 
Bot. 24: 25. 1946; L. Cárdenas, Revista Fac. 
Agron. (Maracay) 7: 131. 1974. Feuilleea 
portoricensis (Jacq.) Kuntze, Revis. Gen. Pl. 
184. 1891. Anneslia p (Jacq.) J. 
D. Smith, Enum. PI. Guat. 2: 18. 1891. TYPE: 
Puerto Rico: Bredema yer 16 (neotype, B-W, 
here designated, seen on microfiche). Although 
there is no direct evidence, it is likely that the 
Bredemayer specimen collected in Puerto Rico, 
and here designated as the neotype was used 
by Jacquin as a basis for his description of 
Mimosa portoricensis. Bredemayer partici- 
pated in two of the Austrian botanical expe- 
ditions to the West Indies in 17 1785 and 
1785-1788, which provided teta for 
Jacquin to use in his Collectanea and other 
works. In connection with this, D'Arcy (1970) 
pointed out that “it is likely that Jacquin based 
many of his descriptions of new species on 
plants brought in by the official Austrian bo- 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


tanical expeditions and grown in Austrian gar- 


dens. 


7) m 


to 3( 


cm diam. at 


Erect or scandent shrubs usually 


tall: stems slender. sometimes to 12 


base: branchlets terete, occasionally 4-angled or 
nearly so, glabrous to very densely villous with 
white or tawny hairs, usually becoming glabrate. 
Pinnae. (1-)3-6(-8)-jugate: 
usually angulate or 4-angled, (1-)2-5 cm long, 
villous to densely villous, rarely glabrous; rachis 
0.9-5.5 10) em long; rachilla (2-)3-9.5(- 11) em 
long; leaflets (8—)1 5-67 pairs per pinna, narrowly 
oblong to lanceolate or oblong to oblong-obovate, 


petioles eglandular, 


occasionally subfalcate, oblique at base, acute to 
rounded at apex, usually mucronulate, the median 
leaflets 4-20 x 1-6 
glabrous, villous, puberulent, or sericeous on both 


mm, membranous, either 


surfaces, usually ciliate: leaflet venation inconspic- 
uous; stipules leafy, adpressed, triangular to tri- 
angular-lanceolate, occasionally linear-triangular, 
sometimes slightly curved, to 11(-17) mm long, 
glabrous to densely villous. Capitula usually axil- 
lary, occasionally arranged in pseudopanicles to 20 
-15) 


em long at anthesis. Flowe S pentamerous, rarely 


em long: peduncles fasciculate, (1.3-)3 1 2(- 


tetramerous: calyx cup-shaped, 1-30 3.5) mm long, 


glabrous to ciliate or densely villous around the 


teeth, these usually triangular to deltate, sometimes 


anceolate to linear-lanceolate or narrowly oblong, 
3-8(-10) mm long. 


the lobes 


acute; corolla campanulate, 
glabrous, occasionally villous apically, 
long, white, the 


lanceolate: filaments ca. 2.5 cm 


staminal tube ca. 2 mm long; ovary ca. 1.5 mm 
long when fertile, glabrous, sessile. Pods linear, 
rarely slightly curved, the apex rounded to trun- 
cate, rostellate, to 16.5 cm long when mature, 
thickly membranous, glabrous or pube rule nt. t. Seeds 
widely rhomboid to very widely ovoid, (X 

5 mm, brown. dark brown, or black, i areola 
sometimes mottled: pleurogram irregular, some- 


times regular. Somatic chromosome number 2n = 


Distribution (Fig. 36). Frequent in open sites 
such as forest clearings, forest edges, 


river banks. and wet thickets; in general in sec- 


roadsides, 


ondary habitats derived from a variety of vege- 
tation types. Collections come from elevations of 

2.100 m. Standley (1930) incorrectly included 
eh species in the Flora of Yucatan. 

Zapoteca portoricensis as circ umscribed here, 
consists of three rather variable subspecies: 
toricensis, pubicarpa, and flavida. Each has a 
few distinguishing, variable characters (Table 12), 
although they intergrade and overlap. Consequent- 


ly, these subspecies are defined on the basis of 
combinations of characters. Viewed in a local con- 
text, each subspecies might be accorded specific 
rank; however, a synthetic analysis of all available 
material reflects the absence of strong morpholog- 
ical discontinuities distinguishing them. 

The three subspecies of Z. portoricensis have 
distinct habitat preferences: subsp. portoricensis 
occurs primarily in relatively wet, mid- to high- 
altitude habitats; subsp. flavida is found wet, 
lowland areas: and subsp. pubicarpa occurs in dry, 
in the West Indies, 


subsp. portoricensts usually occurs in much drier, 


mid-altitude areas. However, 


low, sometimes coastal, habitats. 


ZAPOTEC A PORTORICENSIS 


la. ae scandent; leaflet 2-6 mm; 
rolla 3-4 mm long; mature m to 16.5 c 
n € farida 
lb. Shrubs erect; M 4-12( -15 Xx pex 
rolla 4-8(-10) mm long; mature pods to 12 
(14.5) cm long. 
2a. Pods glabrous, rarely hairy; peduncles 
(1.8-)3.5-12(-15) cm long at anthesis; 


KEY TO THE SUBSPECIES OF 


calyx glabrous to ciliate, the teeth rarely 
sparsely villous; corolla 4-8 mm long . 

ya. subsp. 
Pads puberulent; pedunc les 1.3- 
cm long at anthesis; calyx teeth pars 
tently villous; corolla 7-8(-10) mm long 

6b. subsp. pubicarpa 


Dm 
5(- 6) 


bo 
5 


6a. Zapoteca portoricensis (Jacq.) H. Hern. 


subsp. portoricensis. 


Beskr. Guin. PI. 


Mimosa guineensis Schum. & Thonn., 

323. 1827. 

feacia linearis Desv. ex Ham., Prodr. Pl. Ind. Occid. 
59. 1825. 


4. hamiltonii Desv. ex Ham., Prodr. Pl. Ind. Occid. 59 
l 


1826. 
1833. 


. alba Colla, Hort. Ripul. Appe DE e 339. 


. Bot. . 183. 


1. 

Lysiloma marc Boa Chisels: Fl. Brit. W.I. 223. 1860. 
TYPE: Jamaica, kh s.n. (holotype, GOET (not 

seen); 1 , GH, NY 
Calliandra 1 esol guensis En 
Jahrb. Syst. 23: 124. 1897. 4nneslia e 
] & pw ex Loes.) Britton & I N. 
Amer. Fl. 23: 63. 1928. TYPE: sb bur. 
galpa: Can Eu 1,000 m, 4 Aug. 1843, 
Rorhachah 204 (lectotype, (s IS no. 1266103 (frag- 

ment and photo), here designated; d totype, NY 
nt). ine holotype, which was at B, was de- 

1943 (P. Hiepko, Say comm., ee 


ds 
, MEXU, MO, NY, 


& Loes. ex Loes., Bot. 


photos P u. specimen are at F 
TEX, 


crie imd Var. Md i m Bot. Mag. 
133: t. 8129. 1907. Type: Cult. in Hort. Bot. [^ 
Raw. pes 1906 lens here ia ad, 


specimen was presumably collected by a 


820 Annals of the 


Missouri Botanical Garden 


e A -— T T = 
n a LAA S | 
H `A S 
~ «f A 9] < y 
f qa y Y M 
o l i - “SS | 
| SS P x 
vj o ] N 
g [] "ut 
= R 
7 Fe 0 
*. E 9 
Te ya " 4 
| Yo, > 2 
| i x i e Y ] 
^ v 
r xd 7 ] i 
o Í 
f | : 
= h af 
1 yo JN 
e N | 
Do o. x NR OR N 
` ` XA e 7. S [4 ( 
| | AS EFE ff A) 
aif $ A is 
A | T 
e af 4 ( {| J `y ) ? 
è f j A CN | : ; ( 
N \ ] | | a) ( 
t .- | ? ? / f t t ( * 
e. A C] ] / c4 
Jee Ya 4) ^V] 
| N LET 
x ] | 
AUS .t 
DE: 
1000 Km 4 i } , 
; Y L 
X ' Z T 4 
CN ON CN 
| N yas | 
0 \ ‘ Y 1 | 


FIGURE 36. 
pubicarpa (stars), and Z. portoricensis subsp. flavida (dots). 


Sprague mentioned in his discussion that ** 
plants were purchased in 1906 


the Kew 
from a nursery-man 


in Hyéres e ‘Inga alba” and flowered in the Tem- MEXU, MO, 
erate House in June and July of that year.’ 
Mii "pes Britton & Rose, N. Amer. Fl. 23: made by “Schiede 6% 
928 Medida spraguet (Britton & Rose) Lun, 


dell Lloydia 2: 89. 1939. TYPE: Mexico. Veracruz: 


TABLE 12. Comparison of the subspecies of Zapoteca portoricensis. 


LE and MO are labeled “Schiede and Deppe 6 
The specimens from BM and G are labeled 


Distribution of Zapoteca portoricensis subsp. portoricensis (open squares), Z. portoricensis subsp. 


Xalapa, Schiede 690 bius here designated, 
NY; isolectotypes, BM, CAS, CGE, G, GH, K, L 

UC, US— fragment). The specimens 
at NY and K are the only ones labeled as collections 


" whereas the specimens at 
90.” 


“Schiede 


Subspecies portoricensis Subspecies pubicarpa 


Subspecies flavida 


Habit 
Pairs of pinnae 


Leaflet 


erect shrubs 


(1-32 7(-8) 


erect shrubs 
2-4 


Shape narrowly oblong to lanceolate narrowly oblong to lanceolate 
lx w 5-15 1-3 mm 4-7 x 1-2 mm 
Pairs, pinna (12-)16-67 15-30 
Stipule length 100617) mm 11 mm 
Peduncle length (1.8-)3.5-12(-15) em 1.3-3.5(-6) cm 
alyx 
Length 1-3 mm 2-3(-3.5) mm 
Indument glabrous to ciliate villous to densely villous 
Corolla length 4-8 mm 7-8(-10) mm 
Po 
Length 12(-14.5) cm 6-7 em 
Indument glabrous puberulent 


scandent 
(2-)3-5( 


oblong to oblong-lanceolate 
6 mm 


l-2 mm 
glabrous 
3-4 mm 


9.5 cm 
glabrous 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


P a " (s.n.), and the one at ee is labeled 
ede" (s.n.). The specimens at CAS, CGE, GH, 
vee do not have any a eee tó o 
although oe feom the last herbarium make ref 
n as collected by Schiede. W ith 
E pui of na (BM, d and U de all 
the collections are hand annotated as “Acacia por- 
toricensis Willd.," and judging from the handwrit. 
ing, the other data provided, and the aspect of the 
plants, all these Weg cu almost certainly are du- 


cet thi 1 


plicates of the s e Schiede 690 collection: The 
specimens at G, GE , and NY hav mark, presum 
ably made a posteriori, with the name “Schle 

nevertheless, it is l known that Schlechtendal 


R. 
T comm.), Schlechtendal distributed Schiede’ s 


plar 
Calliandra nogale nsis Lundell, PR 2: 88. : 
exico. Veracruz: Nogales, E. Matuda 1153, 
2 May 1937 (holotype, MIC E isotypes, F, MEXU, 


Palliandra siltepensis Lundell, Phytologia 2: 1. E 
YPE: Mexico. Chiapas: Barranca Honda, ps ec, 
Oct. Nov. 


1940, E. Matuda 4040 (holotype, MICH: 
isotypes, A, F, MEXU, NY). 
Erect shrubs, 2-3(-6) m tall; stems slender, 
sometimes to 12 cm diam. at the base; branchlets 
terete, sometimes 4-angled, glabrous to densely 
villous with tawny hairs, often glabrate. Pinnae 
(1—3-7(-8)-jugate: leaflets (12-)16-67 pairs per 
inna, narrowly oblong to lanceolate, rarely oblan- 
ceolate, oblique at the base, acute to rounded at 
the apex, usually mucronulate, 5-15 x 1-3 mm, 
glabrous to villous, puberulent, or sericeous on both 
surfaces, usually ciliate; stipules triangular to nar- 
rowly triangular, sometimes slightly curved, 10 
(-17) mm long, glabrous to villous. Capitula axil- 
lary; peduncles (1.8-)3.5-12(-15) cm long at an- 
thesis, glabrous to densely villous. Calyx 1-3 mm 
long, the teeth usually triangular, sometimes lan- 
ceolate to linear-lanceolate, usually glabrous or cil- 
iate, sometimes with a few hairs in the area of the 
teeth; corolla 4-8 mm long, glabrous. Pods to 12(- 
14.5) x 1(-1.2) em, straight or MA curve. 
glabrous. Seeds widely rhomboid to ovoid, 
3-5 mm, brown to black, the areola sometimes 
mottled; pleurogram usually irregular, sometimes 
regular. Somatic chromosome number 2n = 


Distribution (Fig. 36). 
species has been collected in several relatively lo- 
calized areas: 1) Atlantic slopes of the Sierra Madre 
Oriental, in the states of Hidalgo, Puebla, and San 
Luis Potosi, and the eastern portion of the Trans- 
verse Volcanic Belt in Veracruz; 2) Sierra Madre 
del Sur, including parts of the Mixteca Region in 
Oaxaca; 3) Sierra Madre of Chiapas, and the Chia- 


This widespread sub- 


pas and Guatemala Highlands; and 4) W est Indies, 
in Jamaica, Hispaniola, and Puerto Rico (including 
some of the Virgin Islands). Disjunctions occur in 
the Sierra de Tamaulipas, Tamaulipas; Temascal- 
tepec, State of México; and in Honduras, Nica- 
ragua, Costa Rica, 
lection from Nova Friburgo, Rio de Janeiro, Brazil 


and Panamá. There is a col- 
(Curran 655). which was probably obtained from 
cultivated plants. This subspecies has usually been 
collected in mountain areas of moderate to rela- 
tively high altitude (800-2,100 m), in plant com- 
munities such as oak, pine, pine-oak, temperate 
deciduous, wet montane cloud, and montane rain 
forests, especially in secondary habitats. Never- 
theless, in some areas such as the Huasteca in the 
states of Hidalgo and San Luis Potosi, Mexico, 
populations usually occur at lower elevations, 
reaching as low as 165 m. Similarly, plants from 
the West Indies usually inhabit low elevation areas, 


down to sea level, with a much drier climate. 


Common names. Barba de chivo (Nicaragua), 
carboncillo blanco (Costa Rica), granolino (Domin- 
ican Republic), guacamaya montés (El Salvador), 
guaje cimarrón (Veracruz), guajillo (Veracruz), 
guajillo blanco (Veracruz), night-flowering acacia 
(Jamaica), pelo de angel (Puebla, San Luis Potosi), 
tamarindo de monte (Guatemala), timbrillo (Vera- 
cruz), white tamarind (Guatemala, Jamaica). 


Additional AE ns examined. — MEXICO. CHIAPAS: 
21 km al este de las Lagunas de Monte Bello, Cabrera 
3690 (MEXU, MO); El Chorreadero, 5.6 mi. east of 
Chiapa de Corzo, Breedlove 11689 (C AS, LL, MICH, 
MO, US); Mpio. of Motozintla de rwr SW side of 


Cerro Mozotal, Breedlove 25917 , MEXU, MICH, 
MO, NY); Mpio. of PEE Pus Granadilla, 
Laughlin 1069 (CAS, MICH, US) Ovando, Matuda 
17923 (F, MEXU, NY); Escuintla, El Triunfo, Matuda 


18098 (CAS, MEXU, NY); Siltepec, Cascada, Matuda 
18666 ee MEXU, NY). HipaLco: Mpio. de Tlanchinol, 
a 0.5 k SE de Tlanchinol, Flores 233 (CAS, ! , 
MICH, NY): Rancho Viejo, Jacala, Fisher 461 20 (C AS, 
F) 13 km al sur de Tepemaxac, pd 591 (MEXU); 
BR de Pun de Doria, 10 km NE of Tenango de 
Doria, H. Hernández & Torres 1 22 (MEXU. MO); Mpio. 
Meztitlán. ] jen de la desviación a Santa Mónica, H. 
Hernández & Torres 134 (MEXU, MO); Mpio. de Tam 
guistengo, 5 km NW Tianguistengo por carretera a Mo- 
lango, H. a & Torres 135 (MEXU, MO); Mpio. 
de Xochicoatlán, 16 km NW de Tianguistengo, H. Her- 
nández & Torres a (MENU. MO); Mpio. Molango, 1 
km N de Molango, H. Hernández & Torre s 138 (MEXU, 
MO); Mpio. de Chapulhuacán, 5 km 5 de la frontera 
SLP-Hgo., H. Hernández & Torres 149 (MEXU, MO); 
Mpio. Chapulhuacán, 9.6 km SW de Chapulhuacán por 
carr. a Jacala, H. Hernández & Torres 150 pa : 
MO); 29 km SW Chapulhuacán poe E: a Jacala, 
H. Hernández & Torres 151 (MEZ "MO. Mpio. de 
Tianguistengo, 4 km al E de A eer R. Her 


dez 3963 (GH, MEXU, WIS); Mpio. de Thane hinol. 


822 


Annals of the 
Missouri Botanical Garden 


km al S de Saree hacia Apantlaso, R. Hernández 


5372 (MEXU); Mpio. de Molango, cle 2 km al 
N de yaa R. ee see 9880 (MEXU, MO); Mpio. 
de Juare , Itztac oyotla, a Ni le lua ez, R. Her- 


nandez 610 5 (MEXU, van wa 301 on highway between 
Jacala and Santa Ana, Moore 2680 (GH); camino entre 
To : y Tenango de Doria, entre 16 y 31.5 km al 
NE de Tulancingo, s h7 da (MEXU); El € s 6 
in del entronque de a a Pisaflor res con la 
rretera Jacala- T: mazunchale, p norio & iie 2381 
(MO) 42 mi. N of rto Ignacio Isidro Díaz, N of 
Zimapán, Wunderlin et ne" 1119 (MO). MÉXICO: i ns 
Temascaltepec, Bejucos, Hinton 2019 (A, F, G, MEXU, 
JS). OAXACA: Dy Eu Cerro Espino, Conzatti 3184 
US); 5 km al N a Galera, carr. Pochutla-Oax. 
Sierra Pi vi Sousa et al. 5610 (CAS, MEXU); 
Jistr. de Juxtlahuaca, a 2 km al SE de Copala, Sousa el 
al. e 9 (MEXU, TEX); Distr. de Putla, a uu al 
de Putla de Guerrero, Sousa et al. 7688 (MEXU); sur 
del Distr. Cuicatlán, a 9 km N de Nacate Sousa et 
al. 10446 (BM, MEXU, MO); Distr. de Juquila 3 km 
al W de Pueblo Viejo y a 10 de San Juan Lacháo, Sousa 
et al. 12603 (MO); 6.1 km al N de San Gabriel Mixtepe "C. 
carr. Puerto Escondido Sola de V ega, Torres & Martínez 
6612 (MO). PUEBLA: 7 km al NE de pepper 
Basurto & Durán 91 (CAS, ENCB); de al 5 
Pahuatlán, Basurto & Durán 107 (CA S, MO); 8 bie al 
NE de Honey, o & Durán 324 (BM, MEXU); 
Mpio. de Venustiano Carranza, 3 km al W de Villa Lázaro 


Cárdenas, Basurto & Durán 366 (MEXU, MO); 2 km 
al W de Chic one uautla, Basurto & Durán 483 (CA 
MEXL uente Apulco, Boege 750 (MEXU) 


alrededores n. Xicotepec de Juárez (Villa Juárez), Quin- 
tero 763 (CAS, MICH, TEX); Zoactepan, Ibarra et al. 
75 (ENCB, M i» SAN LU S POTOSÍ: Mpio. Huehutlan, Tzi- 
neja, Alcorn 2868 (MEXU); 21 mi. N of Morelos, Ken- 
oyer & ers 4089 (; "w near Tancanhuitz, Nelson 4367 
(GH, NY, US); Las Canóas, Pringle 3752 (BM, s E 
G, GH, MEXU, MO, NY, UC, US» Tamasopo Can 
Pringle 3722 (A, BM, BR, F, G, GH, MEXU, MO, Us S). 
Mpio. de Xilitla, 3 km al NE de i de Xilitlilla, R ze- 
dowski 10506 (MEXU, MICH, 2 km al SE de 
Ahuacatlán, Rzedowski 10933 È v TEX); 
Valles, alrededores de Cd. Valles, Torres s.n. (MEXU); 
along hwy. 85, 7.4 mi. N of Tamazunchale, 3.7 mi. S E 
Matlapa, Dziekanowski et al. 3500 (ENCB, NY). T: 
MAULIPAS: Mpio. Gómez Farias, 1 km SE Rancho del 


Cielo, H. Hernández & e 's 146 (MEXU, MO); g 
de Guatemala, Sullivan NCB, NY, TEX). 
RACRUZ: Mpio. Tonayán, es OI is 'gación de Ixtepan, 2 


zada 2113 (MEXU); Mpio. Tantima, Sierra de Tantima, 
Calzada 5539 (MEXU); Mpio. Huiloapan, Cerro de San 
Cristobal, Calzada 8622 (WIS): Mpio. Yecuatla, Barranca 
del Cedral, Cházaro & Gutiérrez 3665 (WIS); Texolo 3 
km al SW de Xico, Dorantes 85 (F); 2 km despues de 
2 El Recreo E Rubeo a Palma Sola, Dorantes et al. 
7 (F, MEXU ) Cerro del ade de Zongolica, Galicia 
s.n. (TEX); Tlaltetela, camino € osautlán Ixhuacán de los 
Reyes, Ixhuacán de los Reyes, C. Hernández et al. 136 
(ENCB, MEXU); 6.8 km N de Xalapa por carr. a Naolin 
co, H. Hernández & Torres 154 Visi MO); Mpio. 
Naolinco, 1.8 km de Naolinco por carr. a Misantla, H. 
Hernández & Torres 155 (MEXU, MO): Mpio. Rio Blan- 
co, N de Río Blanco, por camino a Tenango, H. Her- 
nández & Torres 165 (MEXU, MO); Mpio. Rio Blanco, 

a l km de Río Blanco por camino al Cerro a Cristo, H. 


He rnández & Chacón 465 (MEXU, MO); ca. 5 mi. E 


~ 


of Huatusco along road to Cardel, Higgins 2578 (ENCB, 
MICH); Mpio. Ixhuacán, El Chorro, Luna et al. 141 
MEXU); Pastizal de Olmo, Chalatla, Luna et al. 279 
(MEXU); Tierra Colorada, arriba de Andrés Tenexapa, 
Medrano 1593 (MEXU); Mpio. Choc amán, 8.5 km by 
road W of Chocamán, Vee 2325 XU); Al E de P lan 
de Las Hayas, Vevling e Gomes: Pompa 1005 (GH 
MEX U); ps San Mig 2 km al W de Orizaba, 
Vevling & Gómez- Pompe 2256 (CAS, F, GH, 
Xalapa, Pringle 8091 (G, GH, LL, MEXU, MO, NY, 
UC, US); orillas del Rio Matzinga, near Orizaba, Ramos 
301 (CAS, MEXU, NY); El Esquilón, cerca de Xilotepec, 
Ramos 348(CAS, MEXU, US, WIS: El Naranjito, carre- 
tera "ir Zongolica, nau. , 1460 An JAS, 
MEXU, US) Mpio. Zozocolo, 2.5 km A 
Tenorio et al. uud (MEXU); e Magdalene Mag- 
dalena, Licea 11 y Mpio. Nogales, Nogales, Licea 
158 (F); Mpio. € hw. amán, Chocamán, `); 
Mpio. de Xico, Temimi, Ventura 862 (CAS, MICH, WIS); 
Mpio. de Tlac uilán. Tlacuilan, | entura 8857 (MEXU); 
Mpio. de Banderilla, Banderia; Ventura 9949 (LL, 
MICH); Mpio. de Xalapa, La Mesa de Chiveras, ia d 
10339 (MEXU): Mpio. de Xilotepec, San Marth Pr 
tura 13931 (MEXU). GUATEMALA. ALTA VERAPAZ: er 
San Pedro Carchá y Sacoyoú, Molina & Molina 1 2100 
4; carretera San Juan Chamelco, Molina & 
Molina 12287 (CAS, F, LL, NY); 5 km N of San Pedro 
2'N, 90°15’W), W illiams et al. 40220 (F, 


oe 


of Camotan, Ste halt 31680 
ston 1033 (F); Pacaya, Johnston 1. 331 (F). ESCUINTLA: 
between San Vice a and Calderas, o & 
Molina 27668 (BM. F, MICH, a 5). GUATEM Arra- 
zola, Heyde & Lux s.n. (GH, K, l 7? mi. E of Guatemala 
1 2340 (MO). HUEHUETENANGO: bet 


Steyermark 49476 (F). JUTIAPa: vicinity of Jutiapa, Stan- 
dley 70311 (F, US). QUEZALTENANGO: along old road 
between Finca Pirineos and Pa tzulin, Aot 865 574 
(F); vicinity of Santa Maria de Jesús, Steyermark 34256 
(F); Cotza l, Finca San Francisco, Shute h 1849 (A, F, 
NY, US). sanra rosa: Cocan Jumaytepeque, Heyde & 
Lux 4463 (G, GH, NY, US). HONDURAS. LEMPIRA: Mon- 
tana ae entre Guatán y Cuábanos, Molina 12942 (F, 
US). yoro: ca. 16 km from Yarucha on Quebrada 
de Oro in Cerro Büfalo, Spin s 4380 (NY). 
SAN SALVADOR: Volcán de S 
GH, NY, US). 5 
mi. NE of Metapan along ro 
12340 (MO); La Majada a zen Hacienda p n 
and Los Planes, Molina et al. 16966 (F, NY, US). Nic. 
RAGUA, ESTELÍ: Las Cruces MO); Laguna 
de Miraflores (13?16'N, 86?16'W), Moreno 8272 (MO); 
La Guayabita (1321 4/N, 86214 W), Moreno 17549 (MO); 
La Cabana (13°13'N, 86°13'W), Moreno 17569 (MO). 
JINOTEGA: Sa nta A between Matagalpa and Lino: 
2 (MO); Cordillera Central de 


— 
— 


pm 


= 


ma y 


a a € oie 
ac icd on e to 
, WIS); 3 mi. N 
Maie gu 5739 e sli of the upper 
Rio D Viejo, White 342 (MO, DOMIN 

REPUBLIC. SANTIAGO: Navarrete, Abbott iR (GH, US) 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


823 


4 aw: dllard 13564 (NY, US). azua: Hispan- 
iola, Valle de San Juan, on road to El Cercado, Ekman 
133; Boo F, GH, US); Santiago, Cordillera Septentrional, 
Las Lagunas, at Ariana nd e man 16086 (G 
GH, LL, NY, US). BARAHONA: EL I , Fuertes 1040 
(A, F, G, GH, K, NY, US): Boca C nd as yon 1623 
Y), Guayac anes, Lavastre 1858 (A, NY); San Cris- 
tobal: cerca de Najayo, Liogier & Jiménez 6221 (NY); 
Santo Domingo City, near Los Tres Brazos, Liogier 11823 


x 


(NY, US); E of Higuey, near Boca de Yuma, Liogier 
12263 (F, GH, NY, US); Jina eae Higue) y, lente T 
12349 (F, MEXU, NY, US); ni. E Santiago, 


Jaiqui Picado, Liogier 15168 E p 10 mi. E of 
Santo Domingo City, Guayacanes, Liogier 1 5642 (NY, 
Las Mercedes, Pedernales, joue r 160768 (F, 
GH, NY, US); farallón de Cumayasa, Liogier 19163 
NY); Seibo, Bavajibe, Ta vlor 188 (F, NY). Harri. DEPT. 
DI NORD: St. Michel de l Atalaye, Leonard 8020 (F 
S); Leogane, Pride 320 (WIS E pvc Keon 
Constant Spring, Britton 938 (NY); S Andre W Parish, 
Barkley 38491 (GH); Gordon Town, 
Blue Mountains, Hitchcock s.n. (Mt 
Howard & Proctor 13681 (A); S of Gutters, p & 
Mid 13865 (A); Mount Diablo, Maxon & Killip 161 
A, , NY, US); near Green River, between Cinchona 
a ga Hall, Maxon & Killip 1350 (A, F, GH, 
Y, US); Port Royal Mountains, Flamstead, Maxon 8728 
a H, US); Hollymount, "e m 10404 (NY, US); Bala- 
claya, Orcutt 1469 (MO, ; Troy, o 1347 US 
G HC Christiana, Holmwood, P. octor 10265 (A, NY, 
"Y ish, Craig Hill, Port Royal Vina 
Proctor 23828 (A, F, IL. MICH, NY, US); Portland 
rar ish, Stony River Gorge, near Macungo River junction, 
28. 38 34 Nau Pi E Rico. Island of Culebra, 
US) be twe en Aibonito and 


e 


` 


— 


— 


Coamo, Pd et a E (NY) Ceiba, icr & 
Britton 7811 (NY): Loiza, Britton » An 89 )(G, NY, 
); Puente Alfonzo, Goll et al. CS near 


Laguna de Tortuguero, Liogier 101. 50 (GH, NY, 
Inter Fajardo et Cabeza de San Juan, "s nis 16042 (U P 
Cayey, fiam Cedro, Sintenis 209 , NY) bein a, 
Sintenis 3544 (G, GH, US): a Sintenis vie 
(US); Maunabo in Montis Malapascua, Sintenis 9234 
GH Lares, Barrio Piletas, Síntenis 6021 (G, MO bs 
US); Yanco, Underwood & Griggs 4 em NY, US). V IRGIN 
ISLANDS. SAINT THOMAS: yenit 2n GH, NY). TORTOLA: 
Long Look, D'4rev 1448(A); Edney's, Fishlock 4 16 1(NY 
a to ea Bush, Britton & Shafi 'r 717 (E; NY, US). 
Cl MISSOURI: Saint Louis, conte Bo- 
tanic al Carde rnández 948 (MENU, MO). caL- 
IFORNIA: Santa Barbara. Hall 8132 (UC). Mixi O. DISTRITO 
FEDERAL: Ciudad Universitaria Facultad de Ciencias, Rico 
245 (MEXU). MORELOS: (GH). Brazil. Rio 
de Janeiro, Nova Friburgo, Curran 055 (F, GH, US 
DOMINICAN Rr PUBLIC. 
s (GH). Jav 
Warburg 2 2109 (NY). 


Cuernavaca 


Santo Domingo, Melo 49 (NY). 
AVA. Horto Botanico Bogoriensi, 


As shown above, there are several names that 
have been included among the s of synonyms of 


75; Britton & 


I have ri unable to 


Z. portoricensis (e.g . Bentham, 
Rose, 2 ic r. 
associate any relevant specimens with these epi- 
thets. 


1052. 


Acacia venusta Willd. (Enum. Hort. Berol. 
1809) has been traditionally. included 


synonymy under Z. (Calliandra) portoricensis, 
which is probably incorrect. The only material | 
have been able to relate to this name is a specimen 
at the Willdenow Herbarium (Cat. No. 19143), 
Bot. Berol. W.." as 


indicated by a handwritten note in the lower right 


which comes from "Hort. 


margin. The brief description provided in the orig- 
inal publication of this species is a replica of the 
specimen label, which probably was handwritten 
by Willdenow himself, judging by the style of hand- 
writing (Burdet, 19). This specimen consists of 
two leaves, with the pinnae widely separate from 
each other, and rather few pairs of leaflets per 
pinnae. Because of the lack of flowers or pods, it 
is not possible to relate this specimen to Z. por- 
toricensts. 

Zapoteca portoricensis subsp. portoricensis, as 
delimited here, is an extremely variable, broadly 
defined subspecies, the most variable within the 
species. It can be distinguished from closely related 
subsp. pubicarpa by its glabrous pods, usually 
smaller flowers, glabrous to ciliate calyx teeth, and 
12). It differs from 
subsp. flavida by its erect habit, usually smaller 


usually longer peduncles (Table 


and more numerous leaflets, usually larger flower 
parts, and smaller pods (Table 12). 


Perhaps the most highly variable character 1 


subsp. portoricensis is the degree of pubescence 
of its vegetative structures. Plants range from en- 
tirely glabrous or glabrate to having branchlets and 
leaf parts densely villous with long tawny hairs. 
This tvpe of variation occurs at both inter- and 
intrapopulational levels, and, consequently, is of 
little taxonomic value. There are geographical pat- 
terns in the expression of this character. One ex- 
treme occurs in Veracruz, and to a lesser extent 
in Hidalgo and Chiapas, where there is a prevalence 
of collections with densely villous branchlets and 
leaves. The other extreme occurs in San Luis Po- 
tosi, Tamaulipas, and Central America, where gla- 
brous or sparsely villous leaves and stems predom- 
inate. In the West Indian plants, the leaflets are 
consistently. glabrous, but the branchlets may be 
villous or glabrous. 

Other variable characters in Z. portoricensis 
subsp. portoricensis are the size and number of 
pairs of leaflets per pinna. Most plants from the 
Sierra Mixteca, Sierra Madre del Sur, and Sierra 
Madre de Chiapas, in the states of Oaxaca and 
Chiapas, Mexico, are characterized by pinnae with 
smaller, more numerous (to 67) pairs of leaflets 
usually densely distributed along the rachilla, giving 
the pinnae a feathery aspect. Plants with small, 
numerous pairs of leaflets per pinna are also found 
in Central America. In Oaxaca, plants with leaves 


824 


Annals of the 
Missouri Botanical Garden 


of this sort also tend to have larger flowers and 


usually larger stipules. However, these characters 
have been found in numerous collections from dif- 
ferent regions (e.g.. Veracruz, Puebla, Hidalgo), 
where they occur in plants that do not have feath- 
ery leaves like the ones in Oaxaca, so that there 
is no definite geographical pattern and no basis for 
recognizing additional infraspecific taxa. In con- 
trast, many of the plants from the West Indies 
have larger and less numerous leaflets, thereby 
resembling subsp. flavida. 

e vast majority of collections of Z. portori- 
censis subsp. portoricensis have terete branchlets; 
however, plants with four-angled branchlets, a 
characteristic diagnostic for Z. tetragona, are not 
uncommon in some regions. Some of these plants 
may be the result of hybridization between subsp. 
portoricensis and Z. tetragona. These two taxa 
are generally ecologically isolated in Central Amer- 
ica and southern Mexico, where their geographical 
ranges overlap. In these regions, populations of 
subsp. portoricensis are found primarily in mon- 
tane cloud forest or similar habitats, whereas those 
of Z. tetragona occur in warmer lowland habitats. 
Populations with four-angled branchlets are most 
frequent in Guatemala, where these two taxa come 
into close proximity and in some cases are probably 
strict sympatric. A number of Guatemalan collec- 
tions of subsp. portoricensis have four-angled 
branchlets, although sometimes this characteristic 
is expressed only distally, or is less marked than 
in typical. Z. tetragona (e.g., Molina & Molina 
12287, Standley 86574, 87132, Steyermark 
34256, Williams et al. 12100, and others). Sig- 
nificantly, these plants are derived from areas where 
Z. portoricensis subsp. portoricensis and Z. tetra- 
gona approach each other, as for example, in the 
areas of San Juan Chamelco-Cobán-San Pedro 
Corchá, Departamento de Alta Verapaz; Colomba 
Santa Maria de Jesús, Departamento de 
lenango; 


Ipala-Comatán, Departamento de Chi- 
quimula; and Escuintla-San Vicente Pacayas, 
Departamento de Escuintla. Individuals with four- 
iden branchlets also occur in some other regions 
(e.g.. Orizaba-Cordoba, Veracruz, Chiapas), and 
even in areas where Z. tetragona does not occur, 
such as Haiti. 

West Indian populations often differ from those 
on the mainland in some characters, possibly the 
result of the smaller gene pools in insular popu- 
lations and the selective effects of local climates. 
In many of the plants collected in the West Indies, 
the leaflets are comparatively larger, approximat- 
ing those of Z. portoricensis subsp. flavida. Ja- 
maican specimens are characterized by smaller 


(4.5-6 cm long) pods and by usually smaller, black 
seeds with regular pleurograms, as compared with 
the mainland populations. In addition, some of the 
plants from this island have | -2-jugate pinnae with 
fewer pairs of leaflets. 

Zapoteca portoricensis subsp. portoricensts and 
its close relative Z. caracasana subsp. caracasana 
co-occur on Hispaniola, although it is unclear 
whether they really grow together. Here, as else- 
where in the West Indies, most plants of subsp. 
portoricensis differ from mainland populations in 
having calyces with the teeth lanceolate to linear 
lanceolate, a characteristic of Z. caracasana. In 
some cases, the identification of herbarium speci- 
mens is troublesome if information concerning habit 
and filament color is not provided. However, there 
are several diagnostic characters for each taxon 
(Table 13, while comparing the two species, applies 
to the two subspecies discussed here). Zapoteca 
caracasana subsp. caracasana differs from Z. 
portoricen SIS subsp. portoricen sis in its prostrate/ 
scandent habit, smaller flowers, white /red-purple 
filaments, and by the frequent occurrence of pseu- 
dopaniculate inflorescences with more slender pe- 
duncles. The relationships between these two taxa 
are discussed further under Z. caracasana. 


6b. Zapoteca portoricensis subsp. pubicarpa 
. Hern., TYPE: Mexico. Oaxaca: 

Dist. Huajuapan de León, a 6 km NW de San 
Jerónimo Silacayoapilla, casi en la desviación 


subsp. nov. 


a Ciénega Zahuatlán, veg. alterada de Bursera 
con Lysiloma, 1,900 m, 4 Dec. 1978, Sousa 
& Zárate 9822 (holotype, MEXU; isotype, 
CAS). 


1 2-4-juga; foliola 15-30] juga in pinna utraque 
Teu re vel lanceolata. Calyx 2-3(-3.5) mm lon- 
gus, dentibus villosis saepe forte villosis; corolla 7-8(-10) 
mm Aai saepe apice villosa. Legumina S S 


o 
=~ 


Erect shrubs, to 3 m tall; branchlets villous to 
densely villous with white-gray hairs. Pinnae 2-4- 
jugate; leaflets 15-30 pairs per pinna, narrowly 
oblong to lanceolate, oblique at base, acute at apex, 
x 1-2 mm, 
ciliate, rarely sparsely villous; pal narrowly 


mucronulate, the median ones 4- 
triangular, to 11 mm long, usually densely villous. 
Capitula axillary; peduncles costate, 1.3-3.5(-6) 
cm long at anthesis, glabrous to densely villous. 
Calyx 2-3(-3.5) mm long, the teeth triangular, 
the area of the teeth villous to densely villous; 
corolla 7-8(-10) mm long, usually villous apically; 
Pods ca. 6-7 


ovary glabrous. cm long, brown, 


puberulent. 


Volume 76, Number 3 
1989 


Hernández 825 


Endemic to a restricted 


Distribution (Fig. 36). 
area in southern Puebla and northern Oaxaca, 
where it has been collected in disturbed places in 
the transition zone between oak and tropical de- 
ciduous forests and in thorn forests. Collections 
come from 1.850-2,120 m. 


species grow in a generally drier environment than 


lants of this sub- 
other mainland populations of the species. 
MEXICO. 


Additional specimens examined. PUEBLA: 


259 km S exico City, Barr et al. 62-709 (MEX U); 
vicinity of San | uis, Sep ind ‘a de Tlacuilosto, Purpus s.n. 
(BM, F. GH, MO, , UC, US); Cerro de la Yerba, San 


is Titia con Mixteca, Pun 2008 (G, UC); 

.1. mi. SW « [ Ac 'atepec, Schaffner et al. (4-046 BAM 5 
17 kmalS e Acatepec, Sousa 7740 ( 
de Zapotlán de is Salas, a 8 kmal 5 

OUSA ico 1032 s 2) El Coro, 10 km ] 
, Te e 6659 (MEXU); 
Mpio. Caltepec, Rincón Fn y Hierl rba, p al W 
de Caltepec, Tenorio & Romero 7604 y XU 


Zapoteca portoricensis subsp. pubicarpa is 
closely related to subsp. portoricensis, from which 
it can be distinguished primarily by having puber- 
ulent pods, villous calyx teeth, larger flowers, and 
usually shorter peduncles (see Table 12). The ova- 
ries are glabrous, but pubescence becomes evident 
soon after the pods start to develop. The charac- 
teristics used to define this subspecies are not dis- 
continuous with those of subsp. portoricensis. 
However, the particular combination of morpho- 
logical characters defining subsp. pubicarpa is dis- 
tinctive and appears to be relatively constant 
throughout its range. This and its. biogeographic 
consistency justify its recognition as a separate 
subspecies. 

The pubescence of the pods is the main diag- 


nostic character for this subspecies. Nevertheless, 


hairy pods have been found in two collections of 


Z. portoricensis subsp. portoricensis from Nica- 
ragua (Laguna 333, Moreno 8272). These plants, 
however, have smaller flowers and longer peduncles 
than are characteristic of subsp. pubicarpa:; they 
clearly fall within the limits of subsp. portoricensis. 
Flower size is variable in subsp. portoricensis, with 


large (calyx ^ 2-3 mm long. corolla — 6-8 mm 
long) flowers found in plants throughout the range 
of this subspecies: such flowers, however, are rarely 
as large as those of Z. portoricensis subsp. pubi- 
carpa, and the plants on which they occur never 
have the hairy pods or short peduncles character- 
istic of subsp. pubicarpa. 


6c. Zapoteca portoricensis subsp. flavida 
(Urban) H. Hern., comb. et stat. nov. Cal- 
liandra flavida Urban, Ark. Bot. 24A: 4 


1931; Britton & Killip, Ann. New York Acad. 


Sci. 35: 137. 1936. TYPE: Grenada, ad Bel- 
vedere, 1,600 ft., Dec. 1889, Eggers 6226 


(lectotype, US 938738, here designated; 
olectotvpe. GH). 


Scandent shrubs, 2-3(- 7) m tall; stems slender; 
branchlets terete, usually spreading and descend- 
ing, glabrous to sparsely villous, sometimes densely 


villous in the younger parts. Pinnae (2-)3-5(- 7)- 
ger |} 


jugate: leaflets 8- 18(-36) pairs per pinna, the me- 


dian ones oblong to oblong-obovate, sometimes 
subfalcate, oblique at base, rounded to acute at 
apex, mucronulate, 7-20 x 2 


6 mm, finely villous 
on both surfaces; stipules triangular-lanceolate, oc- 
casionally linear-triangular, to 11(-13) mm long, 
glabrous, ciliate. Capitula commonly axillary, oc- 
casionally in pu S les to 20 cm long: pe- 


duncles slender. 3-7.5 cm long at anthesis. Calyx 
1-2 mm long. ue the teeth triangular to 
deltate, rarely narrowly oblong, acute, occasionally 
irregular; corolla 3-4 mm long. Pods to 16.5 x 
1.2 cm at maturity, thickly membranous, usually 
black at maturity. Seeds ovoid to very widely ovoid, 
ca. 6 X 4-5 mm, brown to dark brown, the areola 
mottled, Somatic 


usually pleurogram irregular. 
> 


chromosome number 2n = 26. 


Distribution (Fig. 36). 
sis subsp. flavida primarily occurs in lowland trop- 


Zapoteca portoricen- 


ical forests, where it is commonly found along river 
banks, in inundated areas, and in wet thickets. Its 
ange extends from southeastern Veracruz and 
northern Oaxaca, Mexico to Venezuela and Peru, 
where it has been collected in several disjunct pop- 
ulations. Additional collections have been made in 
Grenada (Lesser Antilles), Guyana, in the states of 
Pará, Ceará, Pernambuco, Goiás. Mato Grosso, 
Brazil, and in the department of Santa Cruz, Bo- 
livia. In Mexico and Central America it is primarily 
a lowland subspecies, extending altitudinally from 
0 to 670 m elevation: in Venezuela it reaches 1,100 
m. This subspecies has been introduced to Africa, 
where it has been collected from naturalized pop- 
1011) and 


from cultivated individuals in Nigeria (Guile 701). 


ulations in Ghana (Enti 275, Ligne 


Additional specimens examined. MEXICO. CHIAPAS: 
Mpio. Palenque, a 6 del av a Palenque, 
H. Hernández & Torres ; 805 (MEXU, Ea 
3 (GH, MO, US) Mpio. P ocingo, 5 km 
Martínez 9120 (MEXU): Mpio, 
Ococingo, 2 km W Crucero Corozal, camino buie a 
Boca Lacantun, Martínez 10156 (MEXU); N . Oco- 
emge 4 km S Ejido Benemérito de las Americas, camino 
: r de Cacao, Martínez i ¿9 (MEXU) Mpio. A 
que, a 5 km al W de Catazaja, Sousa et 


Pal tal o 
(BM. MEXU: Mpio. de Ocosingo, ca. Bonfil, a 


Echeverria, 


826 


Annals of the 
Missouri Botanical Garden 


2291 (XAL). OAXACA: Chiltepec, Martínez A. 93 (XAL); 
i Martínez-Calderón 304 (A, LL, 
ABASCO: Tacotalpa, Rio Tacotalpa, 
MEXU); Majito, Rovirosa 652 (K, 
an un 41 km al W de Cárdenas, 
Sousa et al v ). VERACRUZ: Chacal- 
tianhuis, La iu a, López s.n. (NAL); Mpio. Catemaco, 
Rio Coxcoapa que desemboca en Laguna Sontecomapa 
H. Hernández & Torres 793 (MEXU, MO); Minatitlan. 
Vell s.n. (NY, US); Loon de Sontho omapan, sobre la 
ribera del Rio Coscoapan, Catemaco, Menéndez 131 (F, 


Cowan 2731 (LL 
JS); Huimanguillo, S 


— 
eae 


MEXU, MO). GUATEMALA. ALTA VERAPAZ: Laguna Sapalá, 
| mi. SW of Sibicté, Steyermark 44929 x bl 
FUEFCNMAEUR 7737 (GH, K, MEXU, NY, US). IZABAL: 


oH. aie 1075 (F, US); Nis Dartmouth 
s Lago lzabal, 
Steyermark 39103 (F, G); Lago 
Felipe, between San Felipe and mouth of Rio Juan Vicente, 
Stevermark 39685 (F). PETÉN: Lacandón, i El Caribal, 
about 6 km SSW, Contreras 3537 (F, LL, MEXU); San 
Pedro, Cadenas Road, Contreras 9480 (F, L L, MEXU). 
RETALHULEU: Retalhuleu, Be rnoulli & Cario 1257 (K). 
BELIZE: Moho River, Peck 752 (GH); Rio Grande, Schipp 
1106 (4, BM, F, 6, GH, K, MIC H, MO, N IC) Jacinto 
Creek, Schipp 1216 (A, BM, F, G, GH, "MICH. MO, 
NY, UC). HONDURAS. SANTA BARBARA: Limones, Standley 
& Lindelie 7540 (F). NICARAGUA. ZELAYA: Cerro La Glo- 
ria (13%40'10N, 84%50'25"W), Ortiz 1131(MO). PANA- 
MA: abri Rio Areti, Duke & Nickerson 14906 (MO). 
VENEZUELA. DELTA AMACURO: region of Morón and Temi- 
che to ees of Rio Cuyubini, 


Izabal, opposite San 


rm 


Steyermark 87702 (F 


MIC Y). CARABOBO: between Puerto € xo and San 
Felipe, "Pittier 9133 (GH). vaRACUY: Los Canizos, Pittier 
8759 (GH, NY, US). COLOMBIA. CHOCÓ: Rio Truandó, 


between Río Sucio and La Nueva, Duke 9815 (NY). META: 
p de la Macarena, Plaza Bonita, Philipson et al. 

8 (US). SANTANDER: Carare, André 294 (F, GH, K, 
Na US). ECUADOR. GUAYAS: ue ira, between Guayaquil 
and Daule, Dodson 2 "Ge try 12579 (MO). Los RÍOS: 
between Quevedo and So o dnd & Gentry 
6293 (MO). PERU. HUANUCO: Pachitea, Schunke 2571 
(F, G, US). JUNÍN: Rio Pauc artambo “Valley, Killip «€ 
Smith 25272 (NY, US); Puerto Yessup, Killip & Smith, 
26320 (NY, US) Puerto Bermüdez, Killip & Smith, 
20435 (NY, US). LORETO: ees Ule 67 25 (F, G, 
K). SAN MARTIN: Chazuta, Rio Huallaga, Klug 40/4 (BM, 
CAS, F, GH, MO, NY, UC, US); oh n Alto Rio 
a Klug 4298 (A, BM, F, , UC, US); 


'áceres, Tocache 


k 378 (US). « GOlÁs: Maurilándia, 
dos Bois, Hatschbach 34270 (NY); Serra do Caiapó, 30 
km S of Caiapônia, Irwin et al. 10993 (NY). MATO GROSSO: 
Aripuana, Projoto Juína, Silva & Rosário: 1811(F). PARÁ: 
Serra de Baterrité, Ducke 1517 (BM, US); Monte Alegre, 
Colonia Itahuajury, Ducke 16051 (BM, US); Monte Ale- 
gre, Serro do Luna, Fróes 30295 (US). PERNAMBUCO: 
Tapéra, Bento 2430 (F, GH, MICH, US, WIS); Vicencia, 
Mata do Eng. Jundia, Pedra da Negra, Tavares 7254 (NY). 

A CRUZ: Prov. is Buena Vista, Stein- 
bach 822 (F, GH, MO, NY, WIS); Prov. Del Sará, 
Cuenca del Surutú, Cantón oo Steinbach 2: 160 


(A 


This subspecies can be easily distinguished from 
subsp. portoricensis and pubicarpa by its scandent 


habit, larger, usually less numerous leaflets, and 


usually larger pods. In addition, subsp. flavida 
primarily grows in habitats associated with lowland 
tropical forests, whereas the two other subspecies 
have been collected chiefly in subtropical, mid- to 
relatively high-altitude habitats. Zapoteca porto- 
ricensis subsp. flavida can be distinguished from 
^. caracasana by having larger leaflets, flower 
buds constantly glabrous, deltate-triangular calyx 
teeth, white filaments, and larger pods. The capit- 
ula are arranged in pseudopanicles only occasion- 
ally in Z. portoricensis subsp. flavida; this is a 
very common condition in Z. caracasana. 
Zapoteca portoricensis subsp. flavida is rela- 
tively constant morphologically throughout its wide 
range; however, a number of collections from South 
America have a combination of characters that 
differ from those of the Mexican and Central Amer- 
ican plants. Characteristics of a collection from 
Colombia (Philipson et al. 1418) suggest mor- 
phological intergradation between Z. portoricensis 
subsp. flavida and Z. caracasana: narrowly tri- 
angular to linear-triangular stipules, leaflets 8 X 
2 mm, numerous (to 36) pairs of leaflets per pinna, 
capitula arranged in elongate pseudopanicles, calyx 
with linear-lanceolate teeth, and white filaments. 
An almost identical situation is displayed by most 
of the Brazilian collections of Z. portoricensis subsp. 
flavida (Andrade-Lima 70-5777, Ducke 1517, 
16051, Fróes 30295, Guedes 378, Hatschbach 
34270, Pickel 2430, Tavares 754). Here, several 
of the vegetative and reproductive characters di- 
agnostic of either Z. portoricensis subsp. flavida 
or Z. caracasana intergrade in particular popu- 


ations. 

Although it is not known whether Z. portori- 
censis subsp. flavida and Z. caracasana ever ac- 
tually grow together in specific habitats, they have 
been collected in comparable microhabitats (e.g., 
river banks, shady sites, and thickets), which raises 
the possibility of hybridization. Populations of Z. 
caracasana, however, occur more commonly in 
drier habitats than do those of Z. portoricensis 
subsp. flavida. The distributions of these two taxa 
overlap only in the area of the Andes, where there 
is much local microhabitat differentiation. 

Collections from Bolivia have some characters 
that do not correspond with those commonly found 
in individuals of Z. portoricensis subsp. flavida. 
For instance, the filaments are reported to be white- 
pink, instead of white, in Steinbach 822. The calyx 
in almost all the Bolivian collections has lanceolate 
teeth (Steinbach 5412, Weddell 4237, Williams 
207). In addition, the mature pods are always 
smaller than typical for this entity (to 10 cm long) 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


FIGURE 37. 


and the valves are thicker (e.g., Pierce s.n., Troll 

1738, Williams 244). 

7. Zapoteca mollis (Standley) H. Hern., Ann. 
Missouri Bot. Gard. 73: 755-703. 1986. Cal- 
liandra mollis Standley, Contr. U.S. Natl. 
Herb. 17: 431. 1914; Standley, Publ. Field 
Mus. Nat. Hist., Bot. Ser. 18: 492. 1937. 
Anneslia mollis (Standley) Britton & Rose, 
N. Amer. Fl. 23: 65. 1928. TYPE: Costa Rica: 


1900, Tonduz 13536 
(holotype, US no. 578114; isotypes, BM (2), 
F, G (3), GH, K (2, LE, NY (3), US (2)). 


buissons à a Nicoya, Jan. 


Erect shrubs to 3 m tall; branchlets terete, cos- 
tate under magnification, abundantly villous with 
tawny hairs. Pinnae (3—)-4+ 6-jugate; petioles eglan- 


dular, terete, with a conspicuous channel in the 


1.2-4.5 
9.5 em long, abundantly villous; 


rachillae 4.5-12.5 em long in well-developed leaves; 


adaxial surface, em long, abundantly vil- 


lous: rachis 3.5 

leaflets 15-26 pairs per pinna, narrowly oblong, 

oblique at base. acute to rounded and mucronulate 
1 

18 x 2-5 mm, the 


proximal ones smaller, all membranous, drying a 


at apex, the median leaflets 7 


light green in the abaxial surface, densely pilose 
with white hairs on both surfaces; leaflet venation 
inconspicuous; stipules leafy, triangular-lanceolate, 
acute, 


ca. 6-14 mm long, parallel-veined, pilose, 


ciliate. Capitula in simple, rarely compound, pseu- 
dopanicles 3.5- 15 em long: peduncles usually fas- 
ciculate, rarely solitary, 2.5 4 em long at anthesis, 
usually densely villous with tawny hairs. Calyx ca. 
2-3 mm long. glabrous, the teeth lanceolate, acute, 
ca. 3-4 


2 mm long: corolla infundibuliform, ca 


Distribution of Zapoteca mollis (dots) and Z. costaricensis 


— 


triangles), Costa Rica and Panama. 


mm long. glabrous, the lobes ovate-lanceolate, acute: 
filaments about 2.5 cm long, the basal half white 
and the distal half ochroleucous, the staminal tube 
ca. 2 mm long; ovary ca. 2-3 mm long when 
fertile, glabrous, sometimes with fine translucent 
sessile. Pods to 9 x 0.7 


with the apex rounded, rarely truncate, rostellate, 


hairs, cm when mature, 
thickly membranous, densely villous with long tawny 
, flattened, ca. 4 X 
3 mm, dark colored, the areola lighter, pleurogram 


hairs. Seeds widely rhomboid 


irregular. 


Distribution (Vig. 37). 


collected in tropical dry forests near Nicoya (Pen- 


This species has been 


insula of Nicoya), in San Lucas Island (Gulf of 


Nicoya), and in Rio Lagarto, all in Prov. Puntare- 
nas, Costa Rica. Plants from a presumably disjunct 
population have been collected in Santa Cruz, near 


Panama City, Panama. 


Additional specimens examined. Costs Rica. 


PUNTARENAS: Island of E Lucas, Gulf of Nicoya, Bar- 
clay 2750 (BM, MO, ; Isla de San Lucas, Golfo de 
Nicoya, along rds Hacien dá Vieja, 9°57'N, 
84954'W, Grayum et al. 4164 (MEXU, MO); Isla de 
San Lucas, Golfo de Nicoya, between Playa us TX 
and Punta Barrigona, Grayum 4201 (ME XU, ; Isla 


de San Lucas, Golfo de Nicoya, dus irail to Thes A "rial 

Gravum 4290 (MENU. MO): Río Loris. entronque : 

Monteverde y Guac imal, Téllez 4307 (MEXU). PANAMA: 
(MO) 


Santa Cruz, Arroyo 34 


This species closely resembles Z. portoricensis 
subsp. flavida in general characteristics ~ leaves 


and inflorescences, but it can be dist ished readily , 
y having an erect habit, which onto with the 
scandent habit of the subspecies, 


and by being 


villous or pilose throughout, including the pods, 


828 


Annals of the 
Missouri Botanical Garden 


which are densely covered with long tawny hairs. 
Further, the filaments are white in Z. portoricensis 
subsp. flavida, but white in the basal half and 
ochroleucous in the distal half in Z. mollis (Grayum 
1162, 4290). (It would be desirable to check more 
living individuals for this feature.) Contrasted with 
Z. portoricensis subsp. flavida, which is commonly 
found in thickets in wet lowland habitats (e.g., along 

ater courses and near ponds), populations of Z. 
mollis usually occur in relatively dry habitats. With 
the exceptions of Z. formosa subsp. salvadorensis, 
Z. caracasana subsp. weberbaueri, and Z. por- 
toricensis subsp. pubicarpa, this is the only species 
in the genus with hairy pods. 

In addition to the few collections of this species 
from Costa Rica, there is a collection from Santa 
Cruz, near Panama City (Arroyo 37), which in- 
dicates a disjunction in its range. This collection, 
with immature pods, matches the plants collected 
in Costa Rica well. Recent attempts (1984) to re- 
collect from this population have been unsuccess- 
ful, presumably due to widespread destruction of 
natural habitats in this part of Panama. 


8. Zapoteca caracasana (Jacq.) H. d Ann. 
Missouri Bot. Gard. 73: 755-763. 1 la 
mosa caracasana Jacq., Collectanea H 142. 


cacia caracasana (Jacq.) Willd., 
Spec. t 1068. 1800; uis Prodr. 2: 467. 
1825; G. Don, Gen. Hist. 2: 419. 1832. Cal- 
i. caracasana ee Benth., London 
J. Bot. 3: 99, 1844; Benth., Trans. Linn. Soc. 
London 30: 543. 1875; Britton & Killip, Ann. 


New York Acad. Sci. 35: 137. 1936; J. F. 
Macbr., Fieldiana, Bot. 13: 69. 1943; W pos 
son & Schery, Ann. Missouri Bot. Gard. 


260. 1950; L. Cárdenas, use Fac. 
(Maracay) 7: 131. 


sana (Jacq.) Kuntze, 


n 
. Feuilleea caraca- 


Gen. Pl. 184. 


Ah 


1891. 4nneslia caracasana (Jacq.) Britton 
& Rose, N. Amer. Fl. : 65. 1928. TYPE 
Jacq., Icon. Pl. Rar. 3: 2n t. 632. 1793 


(neotype here designated, copies at MO, NY, 
U). I have designated this excellent illustration 
as neotype (Fig. 38) because no other element 
on which the description of Mimosa caraca- 


sana was based appears to have survived. 


Scandent, subprostrate or prostrate shrubs; stems 


Y 


usually branching shortly above soil level, ca. 


cm diam.; branchlets flexible, elongate, usually to 
3(-5) m long, terete, sometimes angulate, costate, 
glabrous to densely villous. Pinnae (1—)2- 7-jugate; 
leaf primordia usually densely villous or pilose; 
petioles eglandular, terete, with a conspicuous 
channel adaxially, 0.9-5 cm long, usually villous 
or pilose; rachis (0.4-)1-7.5 em long; rachillae 2- 
6(-7.5) em long; leaflets 10-26(-37) pairs per 
pinna, narrowly oblong, oblique at base, rounded 
to acute at apex, the median ones (4-)5- 10(-11) 
X (1-)1.5-2.5(-3) mm, the proximal ones smaller, 
all membranous, glabrous to short-villous, usually 
ciliate; leaflet venation usually conspicuous abax- 
ially, the primary and secondary veins prominent; 
stipules leafy, narrowly triangular to linear-trian- 
gular, sometimes subauriculate at base, acuminate, 
parallel veined, to 7(-12) mm long, glabrous to 
villous, usually ciliate. Capitula axillary or in simple 

less frequently, compound pseudopanicles to 
20(-26) cm long; peduncles fasciculate at distant 
cm long at anthesis. Flowers pen- 


nodes, 1.5-6.5 
tamerous, sometimes hexamerous; calyx 


shaped, deeply divided to the base, 1.5-2 mm long, 


cup- 


glabrous to densely covered with relatively long 
wavy hairs in bud, glabrous, villous or ciliate at 
anthesis, the teeth lanceolate to linear-lanceolate; 
corolla campanulate, 3-4 mm long, glabrous, the 
lobes lanceolate; filaments ca. 2.5 cm long, white 
in the basal half, purple-red in the distal half, the 
staminal tube ca. 1.5 mm long; ovary ca. 1 mm 
long when fertile, glabrous or occasionally with a 
few hyaline hairs, shortly stipitate. Pods straight, 
sometimes slightly curved, apex usually curvedly 
rostrate, 10(- 11) x 0.7(-0.8) em, thickly mem- 
branous, glabrous, glabrate, or canescent, becom- 
ing densely short-pubescent. Seeds ovoid to rhom- 
boid-ovoid, 4-6 x 3 
pleurogram irregular. Somatic chromosome num- 


4 mm, mottled or brown; 


26 


ber 2n — 


Distribution (Fig. 39). 
species occur primarily in disturbed habitats de- 


Populations of this 


rived from a wide variety of vegetation types, rang- 
ing from dry scrubby vegetation to wet evergreen 
forests. Its range extends from northern Venezuela 
(including Isla Margarita) southwest to Colombia, 
Ecuador, and Peru, where it has been collected in 
scattered localities. It has also been found in His- 
paniola and in a distant disjunct population in Hon- 
duras. Perhaps this is the species within Zapoteca 


FIGURE 38. 
3: 20, t. 632. 17 


— 


“Mimosa caracasana" (= Zapoteca caracasana subsp. caracasana), from Jacquin's Icon. Pl. Rar. 
793 


Volume 76, Number 3 
1989 


Hernández 


829 


830 


Annals of the 
Missouri Botanical Garden 


Ne i 


1000 Km 


g -y 
y, 
D aye 
X 
N PES 
D a 
SN 
G 
0 
H 
n y 
2 
3 
J 
| 
Ey 
as 
Y, ore 
de TY 
i 4 
* 6 
at 
\ 4 y ? t-— 
oe MM! 
XN å 
* i 
* * l 
E 
R 
Y 
+ 
N 
( > ae 


FIGURE 39. 


of which populations occupy the widest altitudinal 


range: 0-2,500 m. Woodson & Schery (1950) 


incorrectly included this species in the Flora of 


Panama. 


Zapoteca caracasana is closely related to Z. 
portoricensis, and these two species have been 
merged at times. For instance, Bentham (1844 
considered Z. (Calliandra) caracasana as a syn- 


— 


onym of C. portoricensis, although subsequently, 
1875), 
he recognized them as distinct species. Later, 
Woodson & Schery (1950) treated Calliandra 
portoricensis as a synonym of C. caracasana. The 


in his revision of Mimosoideae (Bentham, 


most important characters distinguishing these two 
Table 13. Although iden- 


tification of herbarium specimens is admittedly dif- 


species are outlined in 
ficult in some cases, Z. caracasana can be distin- 
guished from Z. portoricensis by its presumably 
constant scandent or prostrate habit, usually small- 
er leaflets, and capitula with more slender pedun- 
cles, which are usually arranged in pseudopani- 
culate inflorescences, a condition rarely found in 
Z. portoricensis. Observations on cultivated ma- 
terial and information from herbarium specimens 
suggest that the scandent/prostrate habit of Z. 


Distribution of Zapoteca caracasana subsp. caracasana (dots), Z. caracasana subsp. weberbaueri 
(stars), and Z. caracasana of unknown subspecific affinity (open square 


— 


caracasana is widespread throughout its range; 
however, it is uncertain whether it is characteristic 
of all the populations or if erect plants occasionally 
occur, as indicated by label information on a few 
specimens. Within Z. portoricensis, the scandent 
habit occurs only in subsp. /lavida. There are, 
however, important differences in growth habit be- 
tween that entity and Z. caracasana: cultivated 
specimens of of Z. caracasana (H. Hernández 
961) had extremely short primary stems, usually 
ds rise to drooping, very elongate branchlets 
10 cm above the soil level, while those of Z. 
Mises subsp. flavida develop very long, 
flexible primary stems, sometimes reaching several 
meters before giving rise to secondary branches. 
The flowers of Z. caracasana are generally 
smaller than those of Z. portoricensis, and the 
calyces are consistently deeply divided to the base, 
with the teeth lanceolate to linear-lanceolate. In 
addition, the two species are easily distinguished 
by their filament colors (Table 13) and by the 
usually smaller pods in Z. caracasana. Pod size, 
however, is highly variable in Z. portoricensis. 
Although the degree of pubescence of stems and 
leaves is extremely variable in Z. caracasana and 
in Z. portoricensis, the branchlets of the former 


Volume 76, Number 3 Hernández 831 
1989 Zapoteca 
TABLE 13. Comparison of Zapoteca caracasana with Z. portoricensis. 
Z. caracasana Z. portoricensis 

Habit scandent, prostrate scandent, erect 

Leaflet | x w 4-11 x 1-3 mm 1-20 x 1-6 mm 

Capitula commonly arranged in pseudopanicles, or usually axillary, rarely in pseudopani- 
axillary cles 

Calyx deeply divided with the teeth lanceolate to teeth triangular or deltate, rarely lan- 
linear-lanceolate ceolate to linear-lanceolate 

Corolla length 3-4 mm 3-8(-10) mm 

Filament color white. red-purple white 


Maximum pod length to Ll em 


to 16.5 em 


are never as densely villous as commonly in the 
latter. 
highly variable in Z. caracasana, most of the col- 


Similarly, the indumentum of the calyx is 


lections with completely glabrous calyces being 
concentrated in Ecuador. However, the peculiar 
vestiture observed in the flower buds of this species 
has never been observed in Z. portoricensis. Fur- 
ther discussion of the relationships between these 
two species is under Z. portoriensis. 

Zapoteca caracasana consists of two closely 
related subspecies: caracasana (with glabrous pods) 
and weberbaueri (with canescent or pubescent 
pods). As in the other taxa of Zapoteca with pu- 
bescent pods (Z. mollis. Z. portoricensis subsp. 
pubicarpa, Z. formosa subsp. salvadorensis), the 
ovaries of Z. caracasana subsp. weberbaueri are 
glabrous, and the hairs of the pods presumably 
develop after fertilization. However, the ovaries in 
this subspecies occasionally have a few hairs. 


KEY TO THE SUBSPECIES OF ZAPOTECA CARACAS ANA 


la. Pods glabrous or nearly so 


"EE oo. Ba. subsp. caracasana 
lb. Pods canescent or pubescent .. 
8b. subsp. PA 


8a. Zapoteca caracasana (Jacq.) H. Hern. 


subsp. caracasana. 


Branchlets villous to glabrate, usually glabrate, 
rarely densely villous. Calyx densely covered with 


usually wavy, long hairs or glabrous in bud, the 
teeth usually ciliate at anthesis; ovary glabrous. 
Mature pods to 10( 11) x 0.7 


rarely glabrate. Seeds 5-6 x 4 mm. 


(-0.8) em, glabrous, 


Distribution (Fig. 39). Relatively 
along roadsides, in cultivated fields and thickets, 


common 
and in shady sites. In the West Indies it occurs on 
Hispaniola. On the mainland, it has been collected 
in northern Venezuela (including Isla Margarita), 
in a few presumably disjunct localities in the de- 


partments of Magdalena and Meta, Colombia, and 
the Department of Ayacucho, Peru. Zapoteca ca- 
racasana subsp. caracasana appears to be allo- 
patric with Z. caracasana subsp. weberbaueri, 
although their ranges come into proximity in Co- 
lombia and Peru. Collections come from altitudes 
of 200-1.300(- 2.500) m. This subspecies has also 
been collected from cultivated individuals at the 
2078). 


Given the fact that the only collection from Hon- 


Botanical Garden of Pará, Brazil (uber 
duras lacks pods, it is not possible to assign it to 
a subspecies. 


Common name. — Cujicillo (Dominican Repub- 


~ 


Additional specime ns examined. HAITI: n f des 
Cahos, Les Gonaives, a an 9081 (GH, K, MO, Ss, 
US). DOMINICAN REPI . MONTE CRISTI: Santo Domine, 
Cordillera Centr al, Monein, Ekman 12612 
Rio del . Ekman 13733 (GH, S, US). B CO: 
between El i suacate and Sapotén, Liogier 19782 
NY). saNTIAGO: Hato del Yaqua, Valle del Cibar, Pian 
16527 (A, F, G, GH, a vos Cerro de la Culebra, i AS nez 
9 [0 (US); Constanza. El Chorrito, Liogier 12230 (NY) 
near Las Mercedes, e nales. ower 10087 (NY); 
near Sapotén, Liogier 16190 (NY). VENEZUELA. ARAGUA: 
Carmen, Williams 10411 (F, US). DISTRITO FEDERAL: 
Naiquetia, Gentry & Morillo 10326 ( MO): near Caracas, 
Lower Cotiza, Pittier 7307 (G, GH, P, US; L 
5:2 (A, S, UC, US): between Sabana Grande and Baruta, 
W yere 10592 (F, US). MERIDA: near D initas (8°4.5' 
70°40'W), Breteler 3149 a NY, 5): Tycoporo Forest 
E 'N, 70°45'W) ipi 3445 (US) ). NUEVA 
ESPARTA: El Valle, Miller & Johnston 26 1 (E). PORTUGUESA: 
Guanare, 17 km NW of Tucupido, Davidse et al. 21420 
(MO). COLOMBIA. META: V illavicencio, Cuatrecasas 4691 


osa, Tan nayo 


Perijá foothills, Cuatrecasas & Ca 
PERU. AYAC U CHO: Estrella, ear Hu anta and Rio Apu- 
rimac, Killip & Smith 23091 (NY, US). HONDURAS: 
»AN(?): One N of Copan, Poole & Watson 1034 


8b. Zapoteca caracasana subsp. weber- 


baueri (Harms) H. Hern., comb. et stat. nov. 


Annals of the 
Missouri Botanical Garden 


Calliandra weberbaueri Harms, Repert. Spec. 
Nov. Regni Veg. 17: 88. 1921. TYPE: 
Huancavelica: layacaja, Valley of the 
Mantaro, northeast of Pampas, 1,600 m, Mar. 

3, Weberbauer 651 1 (lectotype, GH, here 
F, G (fragm.), S, 


Peru. 


Prov . 


designated; 
US) 


isolectotypes, 


Branchlets densely villous to usually. glabrate. 
Calyx very densely covered with usually wavy, long 
hairs, or glabrous in bud, the teeth ciliate or gla- 
brous at anthesis; ovary glabrous, very rarely with 
a few, translucent hairs. Mature pods to 10 x 0.7 
cm, usually canescent when Ts densely short- 


pubescent at maturity. Seeds 4-5 -3.5 mm. 


Somatic chromosome number A = 26. 


Distribution (Fig. 39). 


primarily in relatively dry, scrubby vegetation. It 


This subspecies occurs 


has been collected from scattered localities in Co- 
lombia, in the departments of Antioquia, Cauca, 
Cundinamarca, Santander, Tolima, and Valle; in 
Ecuador, in the provinces of Chimborazo, Guayas 
and Loja; and in Peru, in the departments of Apu- 
and 


Cuzco, Huancavélica, Lambayeque, 


Collections come from elevations of 0 


Additional specimens examined. COLOMBIA. 
ANTIOQUIA: vicinity of Medellin, Toro 787 (NY, US). 
CUNDINAMARCA: alrededores de Puente de San Antonio de 
Cuatrecasas 8284 (F, US); La Esperanza, García 
IS). Santandercito, Uribe 467 (US). SANTANDER: 
Río Suratá Pur a El ws illo e Urata, Killip 
& Smith 16433 (GH, IS); between El Roble and 
Tona, Killip da xem TEM (A. GH, NY, US). TOLIMA: 
Chicoral, Haught 6442 (US). VALLE: 30 35 km E Pal- 
mira, pm. 2494-82 (MO). ECUADOR: CHIMBORAZO: 
between Huigra and Naranjapata, Asplund 15503 (G, 
e Rio € thane po near Huigra, 


Camp E-3107 (F, GH, wo. pit ntura, pene « 


Tena, 
467: > 


~ 
— 


and Ceresita, 
Asplund ieee (B, G, LI NY. P, S, US); km 19 Wi 
Pasquales on Guayaquil- Daule road, Dadon et al. 13596 
(MO); ru km 21, Guayaquil a Daule, Dodson & 
aged 13862 (MO). Loja: 55 km W of Loja, between 
San Pedro ind Chinchas, Espinosa es (US). PERU. 
\PURIMAC: Casinchihua in Rio B ae Valley, 33 km 
SW of Abancay, c et al. 6 (NY, WIS); Abancay, 
O): Haci je "E Lucre, iones Valley, 

, G, UC o: Prov. 
Alfaro. 36 1 (MO); Prov. Convencion, Echara- 
te, Var ras 1045 > (F, 6, GH, UC); Prov. Convención, 
alrededores de Quill, largas 249 (F); Prov. Con- 
vención, Chauca Mayo, Pare as 5471 (MO). 
f ea Haschem 'olpa, Pede u 
US) Tayac aja, between Chejyaj a 


Convención, Ro- 


Tovar 4145 
cubamba, Tovar 4579 (US 
Olmos, between pera and Jaen, Hutchison & W right 
3411 (NY, UC). TUMBES: between Quebrada Seca y Ma- 
tapalo, Ferreyra et al. 10685 (MO, US). 


Asplund a ; (B. NY, S Guages ful. 
if 


5). LAMBAYEQUE: km 98 E of 


This epithet has traditionally been associated 
with Z. portoricensis (Macbride, 1943), perhaps 
because of the superficial similarities between this 
iod ies and Z. caracasana. The type material of 
“Calliandra weberbaueri, however, has a num- 
ber of features suggesting its inclusion. within. Z. 
caracasana. For instance, the calyx is deeply di- 
vided, and the filaments are "white; pinkish-red 
as indicated in Weberbauer's collection. In addi- 
tion, the "prominent" leaflet veins in the type 
collection, as seen under magnification, are like 
those observed in many other collections of Z. 
caracasana. 

Acacia fragrans Ten. nom. nud., Cat. Ort. Nap. 
| was apparently never described. There is, how- 
ever, one specimen at K, clearly identified as Z. 
caracasana subsp. weberbauert, which was pre- 
sumably collected by Michele Tenore in the Orto 
Botanico di Napoli; its original source is unknown. 


9. Zapoteca tetragona (Willd.) H. Hern., Ann. 
Missouri Bot. 73: 755-763. 1986. 
Acacia tetragona Willd., Sp. Pl. . 4(2): 1069. 
1806. Calliandra Bis udo (Willd.) Benth., 
J. Bot. ain 2:15 ww Benth., 


Gard. 


Lon- 


don J. 3: 99 ; Benth., London J. 
Bot. 5: ps 1846; a. Trans. Linn. Soc. 
London 30: 543. 1875; S. Watson, Proc. 


Amer. Acad. Arts 22: 410. 1887; Britton € 
Killip, Ann. New York Acad. Sci. 35: 137. 
1936; Standley, Publ. Field Mus. Nat. 
Bot. Ser. 18: 493. 1937; Standley & S 
Salvadorena 125. 1941; Standley & 
Steyerm., Fieldiana, Bot. 24: 27. 1946 
Woodson & Schery, Ann. Missouri Bot. Gard. 
37: 261. 1950; L. Cárdenas, Revista Fac. 
Agron. (Maracay) 7: 131. 1974. Feuilleea 
ud (Willd. Kuntze, Revis. Gen. 
189 . Anneslia d es illd.) J. p. 
Smith, Lum Pl. Guat. 1: 89. Britton 
& Rose, N. Amer. Fl. 2 23; 2 on Mimosa 
quadrangularis Poiret in Lam., Encycl. Suppl. 
810, nom. illeg. 
epithet was based on W illdenow's Acacia te- 


ron, Fl. 


po x 


— 
w 
— 


(This superfluous 


tragona.) TYPE: Venezuela. Caracas: in ripis 
arenosis fluvii Guairito, Bredemayer 17 (lec- 
totype designated by Hernández (19862), B- 
W cat. no. 19147, not seen; photos at MEXU, 


MO). 


ie cb portoricensis var. multijuga M. pie de pol. 
z. (C rawfordsville) 20: 285. 1895. : Guate- 
Rosa: San Juan Utapa, 4, 500 ft. Oct. 
1893, Heyde & Lux 6098 ( ecd here desig- 

ted, G; isolectotypes, GH, 2)). 
Calliandra toroana Britton & re in pem & Killip, 


Volume 76, Number 3 Hernández 833 
1989 
1000 Km 
Figure 40. Distribution of Zapoteca tetragona. 
Ann. New York Acad. Sci. 35: 137. 1936. TYPE: glabrous. Seeds usually ovoid, 5-6 x 3-5 mm, 


rd Antioquia: vicinity of Medellin, 18 Aug. 
. Toro 465 ( (holotype, NY, phe o, G). 


Mostly shrubs to 5 m tall, rarely trees to 12 m 
tall; stems 2-4(-207) em diam.; 
spicuously 4-angled, sometimes herbaceous in the 


branchlets con- 


— 


younger parts, with the surface costate under mag- 
nification, mostly densely villous with tawny hairs 
when young, becoming glabrate. Pinnae (3 )4- 7- 
jugate: with a 


petioles eglandular, conspicuous 


adaxial channel, angled, 1.7 © cm long, glabrous 
to densely villous; rachis (1.5~)3-9(- 11.5) em hea 
glabrous to densely villous: rachillae (3.5~)4. 

10(-13) em long: leaflets 10- 
narrowly oblong. oblique at base, rounded to acute 


at apex, the median leaflets (5-)8-18 x | 


the proximal smaller, all membranous, glabrous or 


35 pairs per pinna, 
4 mm, 


ciliate marginally, always glabrous on both sur- 
faces; leaflet venation inconspicuous; stipules leafy, 


adpressed, triangular-linear to triangular-lanceo- 


ate, acute at apex, straight or curved, parallel- 
veined, (0.5-)0.8- 2.5 cm long, ciliate, villous or 
glabrous. Capitula axillary, rarely arranged in pseu- 
dopanicles 6-22 cm long; peduncles fasciculate, 
(2.5-)5-12 cm long at anthesis. Calyx 2-3 mm 
long, the teeth deltate, ciliate; corolla funnelform, 
(5-)8 mm long. 


glabrous, the lobes lanceolate, 


sometimes with tufts of hairs apically; filaments 
40 mm long, white, the staminal tube 


3-4 mm long; ovary 2- 2.5 mm long, glabrous, 


usuallv ca. 


shortly stipitate. Pods to 16.5 x 1.4 em, at apex 
truncate to obtuse, rostellate, thickly membranous, 


mostly black or dark brown; pleurogram regular, 


obscure. Somatic chromosome number 2n 


Distribution (Fig. 40). 


has been collected primarily in moist thickets, along 


This widespread species 


streams or river banks, and, in general, in ruderal 
habitats derived from tropical and subtropical ev- 
Mexico 


(Nayarit and northern Veracruz) southeast through 


ergreen forests. Its range extends from 
Central America along the Andean Cordillera im 
which it reaches southern Ecuador. Populations 
develop primarily at moderate and less frequently 
at low altitudes (0- 1,500 m), although im some 
parts of Central and South America they can reach 
altitudes of 2.300 m. 
Common names. 
(Puebla), 


co (Oaxaca), 


Angel (Veracruz), atzalaquio 
barba de chivo (Veracruz), cabellito blan- 
cabellito de angel (Michoacán, Guer- 
rero, Puebla), cabeza de viejo (Oaxaca, Veracruz). 
(Costa 


carbonero blanco (Colombia), carboncillo 


Rica). carboncillo de flor blanca (Costa Rica), cola 
de iguana (Jalisco). guaje silvestre (Oaxaca), gua- 
l lig tión (Oa- 
xaca), palo costillo (Guerrero), pelo de angel (Pueb- 
(Jalisco), 


saqi c"alib (Quecchi name, Guatemala). 


jillo (Jalisco), guajillo blanco (Oaxaca). 


— 


la), potosina quebracho de montana 


(Honduras). 


MENICO. 
Mpio. Te a wo Paraje Mahben Chauk, Breedlove 
(Ds, ENC .H, US); Mpio. 
Oc siue along the rd. to Malpaso, Breed 
Thorne 2099. 5 (DS, LL, MEXU, MICH, dn 


CHIAPAS: 
601 
Ocozocoautla, 18 inia n 


4dditional specimens examined. 


> 


& 
Mi 10. 


Annals of the 
Missouri Botanical Garden 


Pc along Rio Testecapa, 10 km SE Mapastepec, 
iis e & Thorne 30739 (MEXU); Mpio. Frontera 

"m apa, along rd. to Ciudad Cuauhtemoc, Breedlove 
Pine (DS, MEXU); Mpio. Peltaleingo, Ahk'ulbal Nab 
above Peltal Icingo, Breedlove 50102 (MEXU); El Sal- 


Azules, García et al. 2 

WM Ux Ixhuatán-N. Mac Dougall 76 (NY, 5); 
cuintla, Turquia, Matuda 17073 (DS, F, MEXU, NY); 
Guamuchil, SE de Mapastepec, Miranda 6770 (MEXU); 
near Yajalon, is 3411 (NY, US); near Tapachula, 
Velson 3838 (NY, US); alrededores de Rayón, Ramos 
& Funk 501 (MO); Mpio. Venustiano Ca za, above 
Soyatitán, along rd. from Las be to Pi sei, Ton. 31. 30 ) 
(DS, ENCB, F, LL, MICH). coLima: Mpio. Comala, 

km al N de El Naranjal, Ma MUR "s qn iu 2620 red 
MICH). GUERRERO: Dist. A 'ana, Alparosa, Hinton 10847 
(BM, F, G, GH, MO, NY, US); Dist. er Plan de 
Carrizo, res 11059 (DS, F, GH, , US) Dist. 
Montes de Oca, Vallecitos, Hinton 117 "80 DS, GH, NY, 
US) Atoyac, Kerber 169 (BM, BR, K, MICH, Us). 
JALISCO: W slopes of Cerro San Miguel, central segment 
of Sierra de Manantlán, /ltis et al. 470 (MICH); Mpio. 
bare El Agostadero camino a Pihuamo, Martínez et 
al. t (MEXU); 16 mi. SW Autlán, Mel Pe et al. 
198 ea H); Guadalajara, Palmer 635 (BM, GH, NY, 

US). MÉXICO: Dist. Temascaltepec, Bejucos, Hinton 2019 
(F, ME XU); Dist. ars 'altepec, Tejupilco, Hinton 2207 
A, BM, ; Temascaltepec, Acatitlan, Hinton 
T101 (A, BM, F, G. MICH, US). MICHOACÁN: Dist. Coal- 
coman, Zarzamora, Hinton 12274 (DS, GH, MICH, NY, 
US); Dist. Coalcomán, Coalcomán, Hinton 12 23. 31 (DS, 
F, GH, MICH, NY, T, US); Dist. € , Sierra 
Torrecillas, Hinton 15278 (GH, US); Puente Alto, 
km S Ario de Rosales, Sousa & Soto 8008 (CAS). MORELOS: 
Pa nar 17 (MEXU); Cuautla, Las Tazas, Levy 
. ; Huajintlán, Lyonnet ae (BM, MO, US 
Pring gle 9727 (F, G, ¢ , MO, NY, US). 
paraa Puerto i NUR ione Hoe 28 
(ENCB); San Blas, Rudd 3029 (MEXU, US). Oaxaca: 
Mpio. Pochutla, 2 km NE Cha calapilla, o séa 3 km Puente 
de San Juan, Delgado 694 (CAS, MO, UC); Dist. Juchi- 
tán, Mpio. Sta. María Chimalapa, Matías Ror omero, Co- 
lonia C Son rA. "ii a 923 (ENCB, F, MEXU, dpi 
NY); Dist. Put > km NE de Putla por car 
Tlaxiaco, H. Her bud & Torres 453 (MEXU, o; 
Sierra de San Pedro Nolasco, Talea, Jurgensen 792 I, 
GGE, G; y Tuxtepec, € 2 ec ris vicinity, "on 
tínez- Calderón 297 (A, LL, UC, US); Choapam, Yaveo, 
near oe Yave Mexia 9216 (B. CAS, T G, GH, MO, 
Y, 


N IC, US): Ranchería Toltepec, camino a Pluma 


~ 


z 


~ 
c 


Hidalgo, p km N Pochutla, pos et al. 5338 (CAS, 
US, W Dist. Juquila, 8 km NW de San Gabriel Mix- 
tepec, Sousa et al. 6375 (MEXU, MICH, MO); Dist. 
Peotitlán, 2 km al N de Puente de Fierro, en la carr. a 


km al NE de Huautla, 
M); Dist. Jamiltepec, 11 kra SE de 
San Sebastián bi Pu Sousa 8450 (ENCB, MEX U); Dist. 
Putla, 2 km Putla de Guerrero, Sousa et al. 8480 (ENCB, 
MEXU, MICH); Dist. Juchitán, 10 km NW de Palomares, 
Sousa & Téllez 8791 (MO); Dist. Tuxtepec, 
Bonita, San Felipe il ur be 9 km NW 
Oaxaca-Veracruz, car s Romero-Tuxtepec, Sousa 
« Téllez 8806 (MO); Dist. hie Monte Soria, 5 km 
NE de Valle Nacional, Sousa & Rico 9265 (CAS, ENCB); 
Dist. Jamiltepec, 1.2 km N de desviación al Vivero Guapi- 
nol hacia San Agustín Chayuco, Torres et al. 1636 (MO). 


PUEBLA: Mpio. Panter ec, 1 km N de Mecapalapa, Basurto 
& Durán 118 (CAS, ENCB, MO); Mpio. Xilotepec de 
Juárez, 1 km S de Villa Avila Camac de Ho & Durán 
498 (CAS, ENCB, MO); SE Alpuye Miranda 4108 
XU). VERACRUZ: km 6 carr. Miza al -Naolinco, Aven- 
daño 111 ENCB, F, NY); Valle de Córdoba, roit 
S); 1 km N Sihuapán, Calzada 


~ 
— 


895 ie Te F, MO. NY); San Andrés Tuxtla, 
Rancho Palzoquiapan de Don Aristeo (18°27'N, 95?13'W), 


b ono 4220(F, WIS); 1 mi. E de Papantla, Crutchfield 
& Johnston 5223 (TEX); Mpio. de la Palma, La Palma, 
Potrero, Delgado et al. 896 (CAS, ENCB, MO); km 10 
del camino C 'edillo- La Laguna, Hidalgotitlán, idis 's et 
a 4079 (F, MO, NY); Mizantla, camino a Tenochtitlán, 

Hernández et al. 203 (F, '); Mpio. anne l 
E de Sontecomapan por camino a Coxcoapan, H. Her- 
nández & Torres 167 (MEXU, MO); Mpio. Cabadas, 
12 2 6 km NW de Santiago Tuxtla por carr. a Lerdo de 
orres 776 (MEXU, MO); 
e Catemaco 
por camino a Montepio, entre la Estación de Biología Los 
Tuxtlas y Sontecomapan, H. o & Torres 799 
¿A i e Catemaco por 
camino a es epio, H. Hernández & Torres 800 (MEXU 
MO); Mpio. Córdoba, sobre la autop e Fortin-Córdoba, 
H. aa & Torres 827 (MEXU, MO); La Higuera, 
entre San José Acateno y Huaytamalco, Márquez et al. 
651 (F); Ignacio de la Llave, Martínez-Calderón 1549 
(A, BM, CAS LL, MO, NY, US); Mpio. d 
ip Azul, Martínez -C 'alderőn 1803 (A, CAS 

MO, NY, US); Mpi 
ne Hwy. 5. 
(F); 5 km E of Angel Cabada, Veill 5388 (MO); Barranca 
del Fortin, near San Martin Tlacotepec, Purpus 16439 
(A, F, US); Mizantla and Papantla, Schiede & Dep 
691 (BM, G, GH, K, MO, NY); Zongolica, camino kid d 
a ver Nacaxtla (18940' N, 96°5 /) Vazquez 
192 (ENCB, F); Río Solosúchil, entre Cedillo y La Es- 
cuadra, Lu (17?16'N, 94°37'W), Våzquez et 
al. 1314 (BM, MO); URS pers Loma de San Ra- 
tael, Ventura 4565 (CAS, E 3, LL); Amatlán, Rio Seco, 
Ventura 4976 (CAS, ENC ^ rue H, TEX). GUATEMALA. 


— 


ALTA VERAPAZ: Cobán, Turckheim 691 (GH, P, US); near 
San Juan Chamelco, Wilson 4094 (F). € die IMULA: Vol- 
cán Ipala, near Amatillo, Steyermark 30585 (F). PETÉN: 


Dolores, river bank of Rio Mapas uo. 10584 (F, 
LL, MEXU, S). EscUINTLA: San Juan Mixtan, Donnell 
Smith 2282 (K, US) near Escuintla, Standley 63903 
(F); La Trinidad, rd. between Escuintla and Finca Zapote, 
Standley 64916 (A, F); near Rio Sinacapa SE Escuintla, 
Standley 89266 (F). IZABAL: El Estor, Contreras 11112 
] SXU, S). QUEZAI TENANGO: between Colomba 
idle °). RETALHULEU: Re- 
ulli & Cario 1155 (K); Las Victorias, 
las, S perd niaii, 
a . 


Standley | | Porve nir, 
Potrero m a Tajamulco, Ste Sale 37048 (F). 
SANTA ROSA: Casillas, Heyde & Lux 414 , GH, US); 
uci of Platanares, between Taxico ai nd a vn 
REY 79083 (F, G); near El Molino, Standley 783: 
U S). SUCHITEPEQUEZ: r Mazater bee Standley 
009 87 (F): San Francisco Zapotitlán, Steyermark 5214 
(F). BELIZE. EL cayo: El Cayo, Grano de Oro, pt to id 
La Flor vicinity lumber mill, Dw yer 10893 (F, NY); Vaca, 
Gentle 2183 (A, MICH); 37 mi. section, Hummingbird 
highway, Gentle 8998 (F, LL, MEXU, MICH, S); 6 km 


pem 
— 


Volume 76, Number 3 


Hernández 835 


Zapoteca 


E Cave's Brand, Sousa et al. 12189 ecos Macal 
River bottoms at the end of Kinloch Rd., 
(MO); Barton Creek, Whiteford 2897 (BM). EL S. 
DOR. AHUACHAPÁN: vicinity of Ahuachapan, Standley 19814 
(GH, NY, US); near Salto Atehuecia, Stan lley & Padilla 
2873 (F, NY). L4 LIBERTAD: n Andres, 
Clausen & Kovar 6208 (MEX í 

vicinity of San Salvador, Due Y "197 206 (GH, NY, US). 


SANTA ANA: S of Santa Ana, Allen & Armour 7009 (F, 
LL, NY, US). Honpt Ras. \TI Hei vicinity of Tela, 
Standley 55164 (A " US). COMA 3 km to 


Molina & Molina T " 2 ^F, MO). 
COPÁN: between Copán ruins and Santa Rita, Molina & 
Molina. 24661 (F, MO, NY, US). CORTÉS: d Lake 
Yojoa, : illiams & Williams 18798 (F, GH, US). SANTA 
BARBARA: Río Sisicarpa, Molina 3738 (F). ot ANCHO: be- 
tween Tutic salpa and PET ae Rio 
Blackmore & Heath 1957 (BM, 
Culmi, near Las Marias, Blac ime £ ie ath 1665 (BM); 
vicinity of Catacamas, Standley 6 (F). YORO: Pijol, 
eddie, von Hagen & von i. gen nO (F, . NI- 
CARAGUA. JINOTEGA: entre Wiwili y El Carmen, márgenes 
Rio Coco - o 85?50'W), Araquistain & Moreno 
MO): ZELAYA: Bluefield Cukra, Long 100 
(E); La omen 5 km W de Waslala, carretera Waslala 
; carretera nueva a 
Wani, El Hormiguero, Ortiz 1215 (MO); Amparo, Ortiz 
1304 (MO); Mpio. Siuna, Siunawas arriba, Ortiz 1513 
(MO); sector de Negro Was, entre El Empalme y Rosita 
ics B4995"W. Ortiz 2184 (MO); El along 
Pis Pis and surrounding hills ca. (14°04'N, 84°38'W), 
Pipoly 3633 (MO); Cano Corozal, near i Kun, Ste- 
vens & Moreno 197 11 (MO). Costa Rica ei 
Pedro de San Ramón, Brenes 14974 1 (NY); Tor Edel Rio 
Burio, menes 1769 (F, NY, US); Alfaro Ruiz, j^ e 
Smith 1292 (F. NY). CARTAGO: vicinity of Turrialba, 
Godfrey 66 192 ? (MO): Finca Perla, entre Reserva de Vida 


San José Comay Brus. 


— 


Silvestre Tapanti y Purisil, H. Hernández et al. 675 
(MEXU, MO); 2 km de Cervantes por carr. a San Jose 
H. Hernández et al. 730 (MENU, MO). GUANACASTE 


Finca la Pacifica, Opler 858 (F); entre Liberia y Quebrada 
Grande, Poveda 1137 (F, MO); ); Quebrada Zopilote, Par- 
» Nacional Rincón de la Vieja, al. 12750 
$ vicinity of Tilarán, 


: Barba, 


piles, Carlsen 3449 (F, MEXU, US). SAN JOSÉ: 
rradura, tributary Rio C post del Pacific o, NW Canaan, 
General M alley (9°29'N, 
MO, NY, a 
(BM, BR, G US): Rio A 
F, GH, MO NY, US) along Rio Pus 
Quizarra, Williams et al. 26404 (F, NY, 
CHIRIQUÍ: Cerro Horqueta, Blum & P yer 2010 
Dist. Boquete, abov i 
Dist. Tm Bajo. Mo: 520 (A, 
ao Highway to Conce pei ión at Cuesta de Piedra, 
5308 (TEX, UC, US); en Hato del Jobo ind Cerro 
Vaca, Pittier 5: 423 (US); 3 mi. S El Volcán, Tyson 873 
MO). cocré: El Valle, upper Rio Antón, Allen 3819 (G, 
a MO, S); along road 2 2 mi. above El Valle on road to 
a Mesa, Croat 1329 O, NY). paniEN: Rio Pucuro, 
ds en Cerro Mali nA Cerro "d Gentry & Mori 
nd O). VERAGUAS: 5 mi. W a Fe, on rd. past 
“scue ih Agricola Alto Piedra, Croat 2 2 23106 (CAS, F, MO, 
1 ). VENEZUELA. DISTRITO FEDERAL: Cordillera de Avila, 
de south-facing slopes above Caracas, Steyermark 55 14 


E» José, Tonduz 


Tonduz 9800 (BM, BR, 


. PANAMA. 
LL): 


~ 
~ 


, Davidson . 


p 


alrededores de Tovar 
arriba de 


(F, MO). MÉRIDA: Depto. Tovar, 
Badillo 5862 (F). MIRANDA: Los Guayabitos, 
Baruta, Aristeguieta 5932 (NY); Petare and vicinity, 
; Pittier 7471 (GH, hae I ACHIR 

' Lobatera, La Llanada, Cárdenas 
(F); between Las pir and Praxulte. along Rio Tachi- 


A: entre 


ra, Steyermark 57420 (F); along highway, N edge of 
Delicias, Stevermark ^ Liesner 118%. >6 (MO). TRUJILLO: 


Depto Carache, entre El Paramito y El Bale , Be nites 
de Rojas 1989 (MO); San Jacinto, 7« 
nacimientos del Río Momboy, La P . Tamayo ix 
(Us). Cc O ANTIOQUIA: Itiribe, vicinity of Mede 
Toro 577 (NY, US). Boyaca: Cordillera Oriental, vertiente 
oriental, entre márgenes del Río 
Súnuba, Cuatrecasas 9701 (F, US). CALDAS: el 
tera Chinc bina; Duque-Jaramillo 4017 NY, US). 
| Lehmann. 5366 (F, GH, S, U p ioca 
Valley, Pittier 926 (US). CUNDIN 
MARCA: entre Chipaque y Caqueza, alrededores de Puente 
Serviez, Gar cía- Barrige ra & Jaramillo 20536 (F). MAG- 
DALENA: Santa Marta, rcg La 
Sirena (10°59'N, 7 ), Kirkbride 2163 (US NORTE 
R: vicinity of Chinácota, 
5) E wy iar ey, 
of Toledo, Killip . & Smith 2 i (A, GH, NY 
TOLIMA: El Libano, Garcia- os 12227 (US). VALLE: 
bids Oc cidental, vertiente occide nn Hova del de San- 
quinini, La Laguna, Cuatrecasas 1504 : Cor- 
dillera De cide a Hova del Rio Alban, El Zane Cua- 
trecasas 22400 (F, S, US); Bitaco, Johnson 2466 (MO). 
ECUADOR. AZUAY: ass Rio Patol, between Hacie nda Yuba 


i. que y Guayata, 


~ 
£z 
e 
a 
Jy 
- 
a 
c & 
S 
O 
& 
c 
E 
= 
w 


and Hacienda San José de Coimotan, in region of ud 
guin, Stevermark 52757 i MO). BOLÍVAR: Cord. Oc- 
cidental, Limón, Acosta > (F). CARCHI: trail e. 


Chical to Maldonado, e 7 San Juan, Gentry & 
RIO 26611 (MO). CHIMBORAZO: vicinity of Ventura, 
Rose 23524 (GH, NY, US). COTOPAXI: Quevedo- 
qe una road, areas along Rio Pilalo (79*10' W, 09535) 
Holm-Nielsen et al. 3075 (F, MO, NY). CULTIVATED: 
U.S.A. MISSOURI, Saint Louis, Missouri Botanical Garder 
H. Hernández 956 (MENU, MO). Java. Mountaingarden 
Tjibodas (Mt. Gede), van E. 13707 (BR). 


Zapoteca tetragona is closely allied to Z. por- 
toricensis but can be readily distinguished pri- 
marily by having conspicuously tetragonal branch- 
lets, larger corollas, and longer filaments. Despite 
the opinion of Woodson & Schery (1950), who 
suggested that Z. tetragona (Calliandra tetra- 
gona) is distinguished “unnaturally” by its tetra- 
gonal branchlets, implying that it is not differen- 
they call the 
this entity is maintained 


tiated from what “caracasana 
portoricensis complex." 
here as a separate species. Discussion of the re- 
lationship between Z. tetragona and Z. portori- 
censis is provided under the latter species. 
Zapoteca tetragona is probably also related to 
can readily be distin- 


Z. andina, from which it 


guished by having adpressed, usually larger, tri- 


angular-linear to triangular-lanceolate stipules, de 


teeth, and totally white filaments. 


Discussion of the relationship between these two 


species is provided under Z. andina. 


836 


Annals of the 
Missouri Botanical Garden 


Acacia quadrang eularis Link (Enum. Hort. Be- 
rol. Alt 5 

cluded in the list of synonyms of Calliandra tetra- 
but 
synonymy. 


822.) has traditionally been in- 
gona, | have been unable to confirm this 
Link did not cite Willdenow’s Acacia 
tetragona, nor Poiret's Mimosa quadrangularis, 
and given the fact that almost all of Link's personal 
herbarium was destroyed during World War H, 
there is no material available with which to evaluate 
this hypothesis. 


10. Zapoteca andina H. Hern., sp. nov. TYPE: 
Ecuador. Azuay: valley of the Rio Tarqui, a 
ew km south of Cuenca, 2,500 m, 1-2 Feb 


1945, M. 4. Guiler 47 (holotype, US no. 
2106703; isotype, NY). Figure 41. 


Frutices ramulis praese rtim juvenilibus c onspic ue tetra- 
gonis. Pir 3-6-jugae; foliolis 15-32-jugis in pinna 
utraque, angusto- -oblongis; stipulae vadoso-triangulares basi 
auriculis duabt od vel depressis praeditae. 
Calyx ca. 2 mm longus, « entibus angusto-oblongis; fila- 
menta basi alba ad apicem ism ea. 


mae 


15 desc cen 


Shrubs to 2 


4-angled, especially the younger ones, costate un- 


m tall; branchlets conspicuously 


der magnification, glabrous to sparsely villous. Pin- 
nae 3-6-jugate; petioles eglandular, with a con- 
spicuous adaxial channel, 1.3-4 cm long, glabrous 
to villous; rachis 2-4 cm long; rachillae to 8 cm 
long; leaflets 15-32 pairs per pinna, narrowly ob- 
long, oblique at base, acute at apex, the median 
leaflets 5-10 


brous or ciliate; leaflet venation inconspicuous; sti- 


1-3 mm, all membranous, gla- 


pules leafy, shallowly triangular, with 2 basal au- 
ricles, descending or depressed, parallel-veined, to 
0.6 em long, glabrous or ciliate. Capitula axillary; 


peduncles fasciculate, 3.5-7 cm long at anthesis. 
calyx ca. 2 mm long, the teeth narrowly oblong, 
corolla funnelform, ca. 


acute; 4.5 mm long, gla- 


brous, the lobes lanceolate; filaments white in the 
basal half, red-purple in the distal half, the staminal 
tube ca. 2 mm long; ovary glabrous. Pods ca. 7 
X l cm, acute at apex, rostellate, thickly mem- 
branous, glabrous, reticulate-veined. Seeds ovoid, 
ca. 5 X 3 mm, mottled; pleurogram irregular. 


Distribution (Fig. 42). 
endemic to restricted areas of the Andes, 


Zapoteca andina is 
in the 
provinces of Azuay and Loja, Ecuador; the de- 
partments of Cajamarca, La Libertad, and Piura 
in northern Peru; and the department of La Paz, 
Bolivia. This species is allopatric with Z. tetragona; 
taken together, the two species occupy a nearly 
continuous area of the northern Andes (Figs. 40, 
42); 1,400-2,650 m. 


Additional iur examined. ECUADOR. LOJA 
Hac. Huaico-Pamba, 55 km S de Loja 
(MICH). PERU. CONTUMAZÁ: 
tana. (Guzmango-Contumazá), 
(MO); ao Guzmango, Sagástegui 10. 321 (MO); Depto. 
Cajamarca, Quebrada Honda ane Yumal), Sagás- 
legul & López 10627 (MO). La LIBERTAD: Prov. Otuzco, 
lee hal, aie a al. 26: 18 (US). PIURA: Prov. Can- 
d Ferreyra < 3 (MO); entre Yu y Faique, 
cerca a Cancha ique, pons 10920 (US). BOLIVIA. LA 
PAZ: So rata, Holway « Holway 25 GH. US); Sorata, 
prope Espada, Mandon 759 (BM BR, GH, K, NY); 
without laesit: Bang 1812 (BM, F, GH, MO, NY, US). 


Zapoteca andina resembles Z. tetragona in 
althoug 
branchlets are sometimes visibly four- miris only 


most of its vegetative features, the 
in the youngest parts. It differs from Z. tetragona 
primarily by having smaller, triangular, descending 
or depressed, auriculate stipules and by having 
narrowly oblong calyx teeth, smaller floral dimen- 
sions, and white/red-purple filaments. Available 
collections of Z. andina do not reveal the habit, 
and it is uncertain whether plants of Z. andina 
are erect shrubs or low, subprostrate or prostrate 
shrubs, as suggested by the aspect of some of the 
. Bang 1612, Mandon 


Bolivian collections (e.g 


Ll. Zapoteca costaricensis (Britton & Rose) 
H. Hern., comb. nov. Anneslia costaricensis 
Britton & Rose, N. Amer. Fl. 23 . 1928. 
Calliandra costaricensis Pli & Rose) 
Standley, Publ. Field Mus. Nat. Hist., Bot. 

: 492. 1937. TYPE: Costa Rica: bords 
du Rio avala: Jan. 1897, Pittier 10643 
(lectotype, US no. 577350 here designated; 
isolectotypes, BR (2), US (2)). 


Shrubs; branchlets slender, terete, smooth, gla- 
brous. Pinnae (2-)3-jugate; petioles very slender, 
terete to tetragonal, with a conspicuous adaxial 
1.3-7 cm long, glabrous; rachis 2-2.5 
4.3-6.2 
glabrous or glabrate; rachillae 7- 


channel, 
cm long in 2-jugate leaves, cm long in 
3-jugate ones, 
13 cm long; leaflets 10-13 pairs per pinna, the 
median ones narrowly oblong, the distal ones ob- 
long-obovate, oblique at base, acute at apex, 2- 
2.4 A 0.5-0.7 


narrower, all olive-green adaxially, pale-green 


cm, the proximal pairs smaller and 


abaxially, membranous, glabrous; leaflet venation 
inconspicuous; stipules leafy, ovate to widely ovate, 
rounded to cordate at base, acute at apex, to 13 
X 8 mm, glabrous, palmately veined. Capitula ax- 
illary; peduncles 4.5-6.5 cm long in fruit. Pods to 
ca. 15 cm long when mature, acute, rostrate, thick- 
ly membranous, glabrous. Seeds ovoid, 7 X 4 mm, 
brown: pleurogram irregular. 


Volume 76, Number 3 Hernández 837 
1989 Zapoteca 


loa 


FIGURE 41. Zapoteca andina. — A. Branchlet with inflorescences at anthesis, and dehisced pod. — B. Leaflet. 
C. Stipule. — D. Flower. — E. Polyad. — F. Seeds. (From Espinosa 2329 (MICH); pod from Sagástegui et al. 10321 
(MO).) 


Distribution (Fig. 3?) | Zapoteca costaricen- Additional specimen examined. — Costa RICA. LIMÓN: 
TER bord de la voie ferrée de l'Atlantique à Siquirres, 19 Nov. 
R). 


sis is endemic to Costa Rica, where it is known 3 VOLG LGI 
1890, Pittier 3154 


[- 


from only two collections, one from Valle del Rio 
Diquis (Río Convento) near San José and the other The relationships of this species with other mem- 
from near Siquirres, Province of Limón. bers of Zapoteca are difficult to establish, espe- 


838 


Annals of the 
Missouri Botanical Garden 


y | 
ies ON 
e ) yu Aa 
+ 
$ 
LI 
ze 
^ 


FIGURE 42. Distribution of Zapoteca andina. 

cially due to the paucity of material. On the basis 
of leaflet morphology and obscure leaflet venation 
patterns, Z. costaricensis is probably related to Z. 
portoricensis subsp. flavida. Zapoteca costari- 
censis can be easily distinguished from that entity, 
however, by its much larger leaflets and. charac- 
teristic ovate/ widely ovate, cordate stipules. The 
narrowly oblong leaflets of Z. costaricensis con- 
trast with those of Z. formosa, which are almost 


always obovate. 


12. Zapoteca formosa (Kunth) H. Hern., Ann. 
Missouri Bot. Gard. 73: 755-763. mee 
store ta formosa Kunth, Mimose os E52 


822; Kunth, Nov. Gen. Sp. 6: 278. Med 
b Prodr. 2: 466. 1825. bui for- 
mosa (Kunth) Benth., London a Bot. 3: 98. 


. London J. Bot. 5: 102. 


Benth., Trans. Linn. Soc. ion 30: 542. 
18 Benth. in C. Martius, Fl. Bras. 5: 411. 
1876; León, Contr. Ocas. Mus. Hist. Nat. 


Colegio “De la Salle" 10: 237. 


1846; 


1951. Feuil- 


leea formosa (Kunth) Kuntze, Revis. Gen. Pl. 
187. 1891. 4nneslia formosa (Kunth) Britton 
& Millsp.. Bahama Fl. 159. 1920; Britton & 
Rose, N. Amer. Fl. 23: 67. 1928. TYPE: Mex- 
ico, Guanajuato: Bonpland 4288 (holotype, 
P, seen in microfiche; isotype, P). The label 
information on these specimens does not in- 
dicate the locality, but according to the de- 
scription, it was collected in Guanajuato 1,068 
hex (1.922 m) elevation. The holotype is ex- 
tremely similar to and. presumably served as 
the model for the illustration in Kunth's work 
(Fig. 43). 

Erect shrubs to 5 m tall; stems erect, sometimes 
drooping, usually slender, sometimes to 5 em diam.; 
branchlets terete to angular. Leaves eglandular, 
the pinnae ] -3(-4)-jugate; poe terete, with a 
conspicuous adaxial channel, 0.3-7.8 cm long, 
glabrous to villous; rachis 0.3- B 8 cm long; rach- 
illae 0.6-12.5 cm long; leaflets (2-)3-15 
(=l 7) pairs per pinna, oblong to oblong-obovate or 
very widely obovate, the proximal ones sometimes 


FIGURE 'acia formosa 
plate was ka Pied on Bonpland 4288, 


-— 


”(= Vi ra a formosa subsp. formosa), from ~ s Mimoses t. 32. 1822. This 
collection made in Guanajuato, Mexi 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


839 


Jf yp v4 P d ORINA 


Annals of the 
Missouri Botanical Garden 


FIGURE 44. 


salvadorensis (solid squares), Z. formos 


ia subsp. g 


becoming elliptic or widely obovate, all oblique at 
the base or the proximal ones sometimes becoming 
acute, rounded to acute at apex, usually mucro- 
the distal leaflets 0.4-5.9 0.2-3.6 cm, 
the proximal ones smaller, mostly membranous, 


nate, 


glabrous to pubescent, pilose or villous; leaflet ve- 
nation mostl y INCONSPICUOUS, primary and second- 
ary veins occasionally conspicuous; stipules leafy, 
triangular to linear-triangular, or lanceolate to ovate- 
lanceolate, to 13 mm long, glabrous to villous. 
Capitula axillary or in simple pseudopaniculate in- 
florescences; peduncles usually fasciculate at dis- 
tant nodes, 

pentamerous; 


1-11 cm long at anthesis. Flowers 
calyx cup-shaped, 1.5-4 mm long, 
glabrous, the teeth triangular to oblong-lanceolate; 
corolla campanulate, 3-5 mm long, glabrous, the 
lobes elliptic to lanceolate; filaments ca. 2.5-3 cm 
long, white, greenish white, red-purple, or white in 
the basal half and red-purple or pink in the distal 
half, 


ca. l- 


the staminal tube ca. 1-3 mm long; ovary 
3 mm long, glabrous. Pods to 13(-15) em 
long when mature, at apex truncate to rounded or 
acute, usually rostrate, thickly membranous, gla- 
brous, glabrate or densely pilose. Seeds rhomboid- 


rien of Zapoteca se da ra (stars), Z. eee subsp. formosa (dots), Z. formosa subsp. 
racilis (open triangles), Z. 
formosa subsp. mollic a (solid triangles), and Z. cnn subsp. o (open squares 


ormosa subsp. rosei (open circles mA 


2 


4-8 x 3-0 mm, 
brown or gray, the areola usually lighter; pleuro- 


ovoid, sometimes very widely so, 


gram irregular, sometimes semiregular. Somatic 
chromosome number 2n = 


Distribution (Fig. 44). Usually common in 
disturbed or undisturbed habitats derived from arid 
scrub, tropical deciduous forests, and less frequent- 
ly semievergreen forests. This species ranges from 
Arizona to Argentina and into the West Indies. 
Collections come from 0-2,000 m. 


Zapoteca formosa as circumscribed here con- 
sists of six highly variable subspecies: formosa, 
salvadorensis, and 
schottii. 
primarily on leaf and flower characters (Table 14). 


gracilis, rosei, mollicula, 


This infraspecific classification is based 


Varying degrees of morphological intergradation 
occur between different pairs of subspecies. A total 
of 24 specific epithets have been published for 
plants that fall within Z. formosa. Most of these 
names really designate regional variations in leaflet 


— 


characters. In light of the enormous degree o 
morphological variability within this widespread 


species and the relatively common occurrence of 


841 


Hernández 
Zapoteca 


Volume 76, Number 3 


1989 


sno4qe[5 snoJqe[a snodqe[s sno4qe[z aso[rd sno4qe[5 juauinpu] 
un ) un cr) wo 6 uo 11 Ul 6 ui (ciet yidua] aura] uinurxe[« 
a[dand-poa 
“ud /a]tq« 
xutd/ajtw — a[dand-poa ayy 's[dund-poa / o1tq« 31M ayy” oy 10[09 yuəurej g 
aje[norued aje[norued aje[norued 
AIP[[IXE A1e[pxe -opnasd *Aie[[uxe AiP[[xe -opnasd *AJe[[Ixe -opnasd *'Aie[pgxe uoneziuezio a2ua2salogu] 
uiui c une wu 6 uui c wu ZT wuu (CT-)O0I uisue| əmdys urnurixe[q 
AJIPu. snoiqeys Aney- snoJqe[a AJITtey - snoJqe[3 sno.qe|3 AJtey Adley snoJqe[s JUSTINPUT 
unu (2-)9-7 x (PI )c[ p Wu 6-2 x plp uugpc-t€ x [£-g uu [[-C£ x 22.6 "m6]|-,x097. 91 Wu ge € x 6£- TI Mx | 
(1-)9-t(-€) (9)6- £C c) (LIZ L-(- €) G1-1(=8) 9-t (11-)8-€ euurd ‘sated 
Ulo jogeo'T 


ui o9'c-g'e(-g' 


wo (31-18-97 


Yau] e[uqoew 


ur C'r-6'[I ui g'[-9'0 (€ TI-)€9-6'I ui CCIT-GE 
un cE€-Z1 uo TICO uo (c-)t-I uo CCE] wo crt-pe(-g D uo) g'-cc yi3ua] epouaq 
Gal (3) €-I I-e G t-6 (*-)€-c(- D aeuutd jo sued 
1110128 pjnoigoutr 19804 sipiopas sisuo40ppa]ps psousof 
sai adsqng satadsqny satadsqny satadsqny satyadsqny satoadsqny 


“psoulof D3910dn7 jo satbadsqns ayy jo uostreduro:) 


tI rruv[ 


Annals of the 
Missouri Botanical Garden 


intermediates, a conservative approach has been 
chosen in this treatment. The following sequence 
of the subspecies reflects their putative relation- 
ships. 


KEY TO THE SUBSPECIES OF ZAPOTEC A FORMOSA 


la. Pods pilose; plants hairy Vp bua s 


. salvadorensis 
plants glabrous 


2b. subsp 
Lb. Pods glabrous, rarely eee 
to ha 
2a. Filaments white or greenish white 
3a. Leaflets oblong, ane ie stipules nar- 
rowly triangular, to 5 mm long ...... 
2c. subsp. gracilis 


3b. Leaflets oblong- obovate, glabrous to 

villous; stipule lanceolate to ovate-lan- 

ceolate, to 13 mm ong Dr UE 
subsp. formosa 


2b. Filaments either red- purple or uui s ally 


t 


and red- el Or dur tally. 
4a. Leaflets 2- i mm; capitula 
ri i white pun 


and 
either pink « or re y 


Petioles 


purple dis 
3- il mm TA - 
1 2e. a molida 
5b. Petioles 12-35 mm long ae 
12f. subsp. sc ichot 
4b. Leaflets 8 3l x 3 21 mm; capitula 
axillary or pseudopaniculate; fila- 
ments either wholly red-purple or white 
basally and red- pue to B distally 
12d. subsp. rosel 


12a. Zapoteca formosa (Kunth) H. Hern. 
subsp. formosa 

Calliandra capillata Benth., London; Bot. 3: 98. 1844; 
Be ‘rans. Linn. Soc. London 30: 542. 1875. 


Anneslia yra (Benth. ) Britton & Rose, N. Amer. 
Fl. 23: 68. 1928. rype: Mexico. Oaxaca: ad radices 
Ves ri Felipe, Julio, Andrieux 404 (lectotype, 

, here designated; isolectotypes, F (fragment), G, 


K, 
Calliandra malacophylla Benth., ia Bot. 3: 100. 
; Benth., Trans. Linn. Soc. London 30: 544. 
Lo 5. TYPE: Mexico. Dara ad radices nds ad 
Felipe, Julio, 4ndrieux 401 (lectotype, K, her 
ignated; isolectotypes, F (fragment), G, K P, 
ragment)). 
Calliandra viridiflora Benth., "E J. Bot. 3: 98. 
844; Benth., Trans. Linn. . London 30: 542. 
Brazil: Tweedie s.n. Lie K, here 
designated; isolectotype 
C nara gracilis Klotzsc il m Baker, Refug. Bot. 5: 
294. 1873, non Griseb., , nec Standley, 1929; 
nec Renvaiae, 1981; , Trans: Linn. S 
don 30: 542. 187! i 
(Maracay) 7: 131. 1974. Calliandra formosa var. 
gracilis Riia :h ex Ph r) J. F. Macbr. 
Gray Herb. 59: 4 19. ~ moritz iana 
ardenas, de 23: ‘his name 
was published as a nomen novum for Calliandra 
gracilis Klotzsch ex he designation of 


Moritz 887 (BM) as s ea made by Cár- 


— 


1875. TYPE: 


oc. Lon- 


denas is incorrect; the type collection is Otto 404, 
which Klotzsch explicitly cited in the original de- 
scription of C. gracilis.) TYPE: 
(lectotype here deal LE). ' 
presumably was at B, was ci destroyed in 
1943 (P. Hiepko, pers. comm. 86); additional 


duplic ates of this collection were si found at GOET 


nor 
d penduliflora Rose, Contr. U.S. Natl. Herb. 
1899. Anneslia per ee (Rose) Britton 
& aaa N. ie Fl. 23 . 1928. TYPE: Mexico. 
Jalisco: Bolanos, 10 19 Sent 1897, N. Rose 
CHA Mn totype here designated, US no. 301875; 
isolec pes, GH, K, NY). 
Cilia marinate Grin b. ex R. O. Williams, Fl. 
rin. & Tob. 1: 2 1931: "Britton & er Ann. 
New York Acad. E 35: 137. 1936 des i ms as, 
Revista Fac. Agron. (Maracay) 7: 13 An- 
neslia marginata (Griseb. ex R. O. Wins un 
Revista Sudamer. Bot. 6: 177. 1940 : British 
West Indies, Trinidad, Crueger ds dicte here 
designated, K; isolectotypes, NY, . The ds 
m was = ovided by p. in his Fl. Br 
IJ. 710. 54. 
a ge ntryi Standley, Field Mus. Nat. Hist., 
Ser. 17: 261. 
Wash. 527 
Guasaremos, 
Sonora, 


Bot. 
937; gue Fup. oe e 
2:122. s YPE: Mex Chihuahu 
Rio Mayo, ' Canyon Chanate, oM 
5 Aug. 19 36, X 5. Gentry 2303 (holotype, 
E; isc es, A, ME , MO, S, UC, US). 
4nneslia chapade roana rai & Roce, N. Amer. FI. 
23: 69. 1928. TYPE: s p mala. Santa Rosa: Cha- 
padero, 5,000 ft. 892, He yde & Lux 3740 
Ae here e US no. 942577; isolec- 
toty 
fanno pol Britton & Rose, N. Amer. Fl. 23: 68. 
. Calliandra cookii (Britton & Rose) Standley, 
hii Field Columbian Mus., Bot. Ser. 3: 277. 1930 
: Guatemala. Petén: between Yaxmuxan and 
Ya xhá. 9 Apr 1922, Cook & Martin 209 (lecto- 
type, here designated, US no. 108466 ud 
dr izalcoensis Britton & Rose, N. A . Fl. 23: 
. 1928; León, Contr. s Mus. Hist, ‘Nat. Cole- 
gio "De la Salle" 10: 237. 1951. rype: El Salvador. 
Sonsonate: vicinity of a in thicket, 19-24 Mar. 
1922, Standley 21818 oe here designated, 
isolectotypes, GH, 
Annestia Sene Britton ^ a N. Amer. Fl. 23: 
. 1928. Calliandra orie naa (Britton & Rose) 


" a. Se 


Santiago, 27 Aug. 1906. N. Taylor 4 (holotype, 
NY). 
N. Amer. Fl. 


dos suborbicularis Britton & Rose, 
69. xico. Veracruz: Banos del 


28. TYPE: 
ae neal: Aug. 1912, Purpus 6064 (lectotype here 
designated, NY; isolectotypes, F, GH, p S). 

4nneslia a itton, N. Amer. Fl. 23: . 1928 

Calliandra voroensis (Britton) Standley, a Field 

Columbian Mus., Bot. Ser. 4: 309. 1929. TYPE: 

Honduras. Yoro: near P rogres 30 r m, wet thicket, 

24 Jan. 1928, Standley 55095. 3 5 (lectotype here des- 
ignated, NY; isolectotypes, A, F, US). 


Erect shrubs to 3( 
der, sometimes reaching 5 


5) m tall; stems usually slen- 


cm diam.; branchlets 


Volume 76, Number 3 Hernández 843 
1989 Zapoteca 

generally ascending. sometimes drooping. Pinnae 4dditional besa examined. MEXICO. CAM- 
(1-)2-3(-4)-jugate: petiole 2 7.8 cm long; rach- : PECHE: alrededores de las Grutas de Xtacumbilxunam, a 


is 1.7-6.8 cm long; rachillae 2.0 8(- 12) cm long; 
leaflets 3-8(-1 1) pairs per pinna, the distal leaflets 
1.1-5.9 x 0.5 
widely obovate, 


3.6 em, oblong-obovate to very 
sometimes falcate, the proximal 
ones elliptic to widely obovate, the distal ones oblique 
at base, usually becoming acute proximally, round- 
ed to acute at apex, usually mucronate, the prox- 
imal leaflets smaller, all glabrous to short-pubescent 
or finely villous: stipules lanceolate to ovate-lan- 
13) mm 


long. glabrous to pubescent or villous. Capitula 


ceolate, sometimes slightly curved, to 10( 
axillary or in pseudopanicles to 22 em long; pe- 
duncles 2.1-11 cm long at anthesis. Calyx 2 
mm long, the teeth triangular to oblong-lanceolate, 
sometimes deeply divided; corolla 4-6 mm long; 
filaments white, sometimes greenish white. Pods to 
19-15) x 


occasionally villous or sparsely villous in early de- 


1.2) em, glabrous when mature, but 


velopmental stages. Seeds rhomboid-ovoid to widely 
so, 6-7 3-6 mm, brown, the areola usually 


lighter; pleurogram strongly irregular. Somatic 


chromosome number 2n — 2 
Distribution (Fig. 44). 


widest distribution within Z. formosa. 


This subspecies has the 
Its range 
extends from northern Mexico, in the states of 
Tamaulipas. Chihuahua, and Sinaloa, south and 
southeast through Central America to central Costa 
Rica. In South America it occurs in northern Ven- 
ezuela and northern Colombia, and also in Brazil 


(Minas Gerais), Bolivia, Paraguay, and northern 
Argentina. It has been collected on Cuba, Trinidad, 
This 


subspecies has not been collected in Panama, most 


and other smaller islands in the West Indies. 


of Colombia, Ecuador, or Peru. It is relatively 
common in open sites in seasonal, dry environments 
such as tropical dry forests, coastal thickets, and 
arid thorn serub, although occasionally it is found 
in wetter disturbed or undisturbed habitats, in semi- 
evergreen or evergreen tropical or subtropical for- 
ests. It has also been collected occasionally in pine 
and pine-oak forests in Mexico. Collections come 
2.000 m. Cultivated individuals 
in Guanajuato, Mexico yielded the collection Duges 


from altitudes of O 


s.n. 


Common names. — Bellotica (Atlántico, Colom- 
bia), (Yucatán), 


chipilin de monte (Guatemala), chapuli (Guerrero), 


caba pixoy cabellera (Oaxaca), 
claveyino (Chihuahua), barbón montanés (El Sal- 
vador), huaje (Sinaloa), ibche (Yucatán), taman- 
ché (Yucatán), tepehuaje (Guanajuato), veranero 
(Bolivar, Colombia), yag civ (Oaxaca), yaque (Su- 
cre, Venezuela). 


3 km W de Bolonchén de Rejón, Cabrera & Ce 
95 14 (MEXU); 8 km SE de Conhuas sobre el camino al 
Centro Regional Calakmul, en km 99 
cega-Chetumal, Cabrera & Cabrera 10973 (MEXU); 
Reforma, Lundell 83. GH, de H, Mo XY. 

IS); Tuxpeña, Lundell 1073 (F). CHIAPAS: 
Mpio. ui Tuxtla Gutiérrez, 15 km N of ed uxtla Gutiérrez 
i road to El Sumidero Breedlove 10035 (LL, MO, 

S); Mpio. of Chiapa de Corzo, Mexican hwy. 190 in the 
eee paraje of Muktajok, Breedlove 30531 (MO, 
NY); Mpio. of Ocozocoautla de Espinosa, 13-18 km S 
of Ocozocoautla, Breedlove 37844 (MEXU, MO); Mpio. 

i Trinitaria, © km al S de La 
Frontera Comalapa, H. 
(MENU, MO 
Flores, Miranda 5977 (MEXU); road between Tenejapa 
and Yajalón, Velson 3244 (GH, US); Mpio. of Ocosingo, 
near Ranel 10 Mumuntik near Ocosingo, Jon 3491 (LL, 
MO, NY, US). cuintanta: Wisiego, Rio Mayo, Gentry 
VR , MO): Mpio. s entre Kirare y la Bufa, 
Tenorio & Romero 6585 (MEXU). COLIMA: SW foothills 
of the Nev ado de Colima, l- i: y mi. 5 of Hacienda San 
As laugh 1611. 13 (MIC H). DURANGO: Rose 2332 

JUATO: Mpio. Guanajuato, Canada de Pastita, 
delata s.n. (ENCB). 10 km SW of León, road to San 
Francisco del ii) ón, Mc laugh 24272 (MICH). 
GUERRERO: 4 mi. rrapato, Rowell 107 (MICH); 
2.5 mi. N of E Herold & Clark 449 (MEXU, 
MICH); Distr. Montes de Oca, Vallecitos, Hinton 11026 
CAS, GH, MEXU, MICH, NY, US); Tecpan- El Verde, 
Hinton 14335 (CAS, GH, MEXU, MICH, NY, UC, US); 
Mazatlán, Kruse ; (CB): La Corréa 
271 (F, G, GH, 
Sur, N of Río Balsas 
Mexia 8942 (B, CAS, F, G, GH, K, LL, MO, NY, UC, 
US); near Iguala, et et al. 9417 (US); Mpio. de Tecpan 
de Galeana, El Pusulmiche 3 km del entronque de la 
brec in a Corinto con la Rods Tecpan de od Zi- 
nejo, Tenorio et al. 7 (MO); Mpio. de Zihuata- 


e 17 km al NE c ees por la carr. hoe 


ibrera 


) de la carr. Escar- 


Trinitaria por carr. a 
Hernández Chacón 


Rancho Lindavista, 24 km E de 


~ 


, Langlassé 


tanejo- Cd. Altamirano, Tenori io et al. 398 (MO): 1.5 km 
al N de Chilpancingo, Torres et al. 1205 (MO): s km 
al N de Zumpango del Río hacia Iguala, Torres 1247 
(MO). JaLisco: above Lake Chapala, Barkley et al. 2615 
(F, TEX) o de la Cruz, La Cruz, rue r 
5485 (ME and W facing Wm ] 12-1 


of Autlán, P 4 augh & Koelz 883 
of Autlán, Mel ins) 1985; (MIC He N si La 
Cue asta below the pass Talpa de Allende, boe d 
¿ (MICH): ne e Sir T duo Hé (364 

(G H. US): near Etzatlan, Ro. : Pai 524 (NY, 
US). MICHOACAN: vicinity ‘of Vota ius ne. : 59061 " BM, 
G, GH, MO, NY, Us); Aquila, La Mina, Guerrero et al. 
494 (MEXU); Distr. Coale a Sierra Maier Pe Hinton 
et al. 15131 (GH, LL, NY, US) bet 1 San Juan de 
an Ana), "Me 1 augh 17950 
uitzeo, oe a s S i 
ud HH, MO ; ias , US); Mon- 
teleón, Pringle E 25 (F, GH, ce s Ed e» 17 
mi. E of d T 10404 (US). MORELOS: 6 
mi. W of 15 3 (MEXU, MICH); Cuer- 
navaca, Ros i NAYARIT: 
Santa María del Ors. ndi 9084 (MEXU) 
e atlán on the road to Barranca de Oro and dames 
McVaugh & Koelz 259 (MICH). oaxaca: Distr. 


los Plátanos and Bs 


844 


Annals of the 
Missouri Botanical Garden 


Tlacolula, enfrente de la Virgen Juovila, 12 km E de 
Totolapan, H. Hernández & Torres 216 (MEXU, MO), 
Dist. Pochutla, a ^ km NE de Chacalapa por camino a 
Finca Monte Cristo, H. Hernández & Torres 394 (MEXU 
MO); Dist. Puta, 20 km NE de Putla por carr. a ns. 
H. Hernández & Torres 143 (MEXU, MO); Teotitlán 
del Valle, 10 mi. E of city, Kenoyer 1629 (A); vicinity 
of Mitla, Messer 247 (MICH); vicinity of Yalalag, Nelson 
945 (US); near Oaxaca, Pringle 4801 (A, BM, BR, F, 
E GH, MEXU, MICH, MO, NY, P, UC, US); a 10 km 
; de Teotitlan del Camino, Rico et al. 251 (I 
MEXU. MO); Monte Albán, Smith 115 (GH, US); Mpio. 
de e 27 km a e Rio Verde, Sousa et al. 
ah (MEXU); Distr. de PA ee Mpio. de Loma Boni- 
, San Agustin, 4 km al 5 de Mano Marquez, Sousa et 
al /288 (MO); Desv. : Cruz, a 5 km al 
N de Santa Cruz, Sousa et al. 7599 (CAS, MEXU); Distr. 
Tuxtepec, a 3 km al N de San Matéo Yetla, ids 7955 
(CAS, MEXU); Distr. de Tlacolula, Santa Rosa, a 3 km 
al E de Totolapan, Sousa & Tellez z 6561 (MO); Distr. de 


uz Mixtepec, Sousa et al. 104 
(CAS, MEXU, TEX): l i al SW de San José de Gracia, 
Téllez & Magallanes 111 (CAS, MEXU, MO, NY); Dist. 
Tehuantepec, ladera W del Cerro Guingola, L. Torres 
395 (MEXU). PUEBLA: Acatlán, Miranda 3025 (MEX U), 
behind Cerro Colorado between Nacozcalco and San An- 
tonio : vir along Ws Barranc a de los gie 5mith 
tal. 3 (G, GH, M SX, US). QUERÉTARO: 
,"21 e N of Querétaro on di to San 
Anderson & Anderson 5041 (ENCB, GH, 
MICH, NY); Hacienda Rivera, Arsène 10061 (A, BM, 
F, MO, NY, US); El Batán, Arguelles 6 15 (CAS, MEXU); 
Magallanes 2672 (MEXU). QUINTANA ROO: 
Mpio. Carrillo Puerto, Rancho El Ramonal, Durán & 
Olmsted 694 (MEXU); 22 km al riis id Felipe € arrillo 


, camino a S. Chico, 


~ 


x 


de ll 7846 n TH IS Coa inel: km 17 da la 
carr. costera W, s rez e Ramos 1003 (LL, MEXU, 
MO), 18 km al S de las Ruinas de Tulum, Quero & 
Grether 924 (MEXU, MO); 3 km al S de Cancún, Sousa 
et al. 10804 (MEXU, NY); Lázaro Cárdenas, en el camino 
blanco de Kiun rumbo San Ars di Ucan & Chan 
2926 (XAL). SAN LUIS POTOSI: 13 mi. W of Nuevo Mo- 
relos, Kenoyer & Crum 3953 (A). SINALOA: Imala, Gentry 
1959 (CAS, GH, MEXU, MICH, MO, NY. UC); Las 
Mesas, Sierra Surutato, Ge ntry 0150 (GH, MICH, ipn. 
NY). TABASC 10: San ues Balancán, límite N de la zona 
al, e resa Garei ía & Palma 
AMAULIPA : Mpio. Gomez Farias 
las por camino corto al Banc ho 
; z Torres 145 (MEXU, MO); 
10 km NW of El Progreso which is 18 km NW of en 
(23°N, 99930" W), Stanford et al. 1095 (CAS, GH, MEX 
MO, NY, UC). VERACRUZ: Buena Vista, Dorantes et i 
1430 (LL, MEXU); Mpio. de Jalcomulco, alrededor de 
Jale omuleo, Gandara & Dorantes 100 (MEX 
pan, Purpus 8401 (GH, UC); Mpi 


o 


EXU). YUCATÁN: en los alrede- 
de Balancanché, a 36 km W de 
Cabrera & pri 9,33 (MEXU); Canta- 
km después de mayec i 0, 
Chan 1814 (XAL); d pa 12 carr. Hunuema a 


dore Elo uic 


Valladolid, 


Sisal, Chan 2827 (XAL), Chikindzonot, M as Chan 
a s 5 km E of Motul, Conway & Johnson 490- 
8 (MO, ) Chankom, Enríquez 690 (MEXU, US); 

R, CA ;H 


o, 11; ha . SF G 
UC 


MA, MICH, MO, NY, , US); Suitún, Gaumer e 
23410 (F, G, GH, MA, ), US); Kantunil, Gaumer 
et al. 23431 (A, F, G, US); 2 km al NW de Ticul, Sier 
de Ticul, Grether 523 (MEXU); 7 km al N de Colonia 
Yucatán, brecha de Cc lonia Yucatán a El Cuyo, Grether 
557 (MEXU) 5 km al de Chimax, Grether 1333 
d MO); 3 km W of Valladolid, JoAnsun & Conway 
E 


8 (MO); Oxkutzc ub, Xul, 
tza, bs ds 1438 (BM, F, MEX 
(MICH); Muna, Steere 2152 eae D 
235 (GH); Valla ue Pixoy, 


A a 
LO 


. ALTA VERAPAZ: between San C d 
Verapaz and Chixoy, Steye eibi 43911 - 
MALTENANGO: Alame s Johnston 925 (F); Tm: Martin, 
Morales 1302 (F). € B 
elow Mor t ina 


Rio Negro, 
30954 (F). GUATEMALA: 2.5 km S of pcr dum 
& Dwyer 2987 (F, ER LL, MO, US). JALAPA: 
vicinity of Jalapa, Standley 77380 (F, AEN PETÉN: «Tikal 
entrada de de Santa Fe, Contreras 191 (F, LL, 
MEXU); Dos Lagunas, on Ixcanrio Road, 3 km NE of 
the village, Contreras 1544 (F, LL, MEX A Uaxactún, 
Contreras 3594 (F, LL, MEXU); Milpa Grande, Ma- 
canché, Contreras diui ks " on NY); G 4 


PE ME Laguna 
MEX 


PEE oe 
1 r 


“I N 
~ 


O). SACATEPEQ 
renzo, Standle M 50866 (A, F). SANTA ROSA: near Cuilapa 
Standley 78533 (F); between Santa Ana Hui 
Sierra de los ( 
ermark 51357a (F). BELIZE. 
bal, Arnason & Lambert 17882 (MO); El Cayo, Chanek 
108(MEXU?). HONDURAS. cer AGUA: near Lake Yojoa, 
Hazlett 1079 (MO). corTés: 3 km de San Pedro Sula, 
Montana La Zona, canon del Rio Piedras, Molina 3435 
F, GH, US); 


m Mua 


= 


AZAN: near San 


(F. G T LL. MENU. 
San Francisco La Paz, Mantin Ù v 3i / 
LL, NY, US). EL SALVADOR. AHUACHAPÁN: Padilla 88 
(US). CHALATENANGO: La Reina, Calderón 2408 (F, US). 
LA LIBERTAD: Santa Tecla, slopes of is án San Sig 
Williams & Molina 15229 (GH). NICARAGUA. BOA 
Pitas (12°28'N, 85%35'W), Moreno rope 2 (MO) Santa 
Cruz (12%24'N, 85°49'W), Moreno 186 19 (MO). c ARAZO: 
Oriente, 1 km de la carr. S (11°49 
86?08'W), Moreno 10710 (MO); Comarca El Aguacate, 
4 km S of Jinotepe, Neill 1058 (MO). CHINANDEGA: SE 
del Cosiguina, Fonseca. 146 (MO); Volcán San Cristóbal, 
Finca Los Rojos, a 20 km de Londen (12°42'N, 
87°02’W), H. Hernández et al. 582 ? (MEXU, MO); falda 
SE del Volcán Casita, 1 ki y 
860?57"W), Moreno 1572 (MO) 
de € hinandeg 'ga (1 2°41'N, 87°00'W), Moreno 25040 (MO); 
ule, camino al Volcan Chonco 
, Sandino pud (1 'ALE 
Busto Monota N of Acoy acia (11?58'N, 
vens 10942 (MO); Hda. San Na artin, near pies of 
Rio El Jordán and Rio La Pradera (1221 7'N, 85%15'W), 
Stevens 22620 (MO). ESTELÍ: i: Salto de Estanzuela, 5 km 


t 

Ur 

= 

fs 

- 
E 
lon 
Z 
=> 


HS 
D 
w 
zz 
oc 
- 
E 
E: 


Volume 76, Number 3 
1989 


Hernández 


S Ciudad de Esteli, H. Hernández et al. 034 (MEXU, 

MO); 6 km de Pueblo Nuevo carr. a Limay Cerro San 
Ramon (13%21'N, 86°31'W), Moreno 2140 (MO); La 
Gavilana (13903'N, 86°19'W), Moreno 21883 (MO), 
Mpio. de Condega, 5 km al NE de Condega (13?22'N, 
86°20'W), Moreno 25333 (MO). GRANADA: Isla Apater 
Cerro El Llano (falda nente) _Gryale a 808 (MO); ' 
Agustín, camino a Cutirre, a 2.5 km de Granada (85%58'N 
11*555'W), Moreno 2632 (MO); Volcán Mombacho, su- 
biendo por la finca Las Delicias (LIPS'N, 85°58'W), 
Moreno 4089 (MO); 1 km al N de Casa de Teja (11°50’N, 
80°01" W). 5404 C MO) 


— 


Moreno 


Ju 
W), lega & Robleto 106 (MO). LEON: carr. 
vieja a L eón, km 6 h debajo del puente del Rio Tamarindo 
(12214'N, 86°43 Guzmán et al. 1025 (MO); Ca- 
rretera a. El Tránsito San Pablo (12?05'N, 86°41'W), 
Moreno 4841 (MO). Mapriz: Cerro Quisuca (13°31'N, 
86?3]'W), Stevens 22205 (MO). MANAGUA: Cuesta del 
Coyol (12230'N, 86903 W), Araquistain 36 14 (MOX San 
Isidro (12905-06'N, ior 'W), Ara iquistain . 3654 (MO); 
Cuenca sur (12%00-03'N, 85?15-I Araquistain 
3749 (MO); Managua, Punta Chiltepe, costado W de 
Laguna Apoyeque, i icd a et al. 3104 (MO); carr. entre 
Las Cone ehitas y ) 1961 


(MO); 5 


Franca, Sandino 349a (MO); along hwy. 8 ca. 2.4 km 


SW of intersection He hwy. 2, km 28 (11?57'N, 
86?20'W), Stev 394 (MO). MATAGALPA: Ermita c 
pan 20 pt es de El Tuma (1 FEQ AUN 


85°30 yw), m a & Grijalva 1366 (MO); 
tera Ab cana, esta El Venado, 4 km al E Las 
Delicias (12°41'N, 86°03 W). be dicis (MO) I 
km al E de la Hacienda Los Ange les (1321 2'N, 85°4.2'W), 
Moreno 19004 (MO); carr. vieja a fori Las Trancas 
(12°59'N, 85°58'W), Moreno 22927 (MO); El Tempis- 
que, 3 km al E de Puertas Viejas (12°35 » 86?00'W ), 
24831 (MO); along hwy. itrance to La 
Cruz, ca. 1.8 km SE of Las Calabazas (12939'N, 86°04' W), 
Stevens & Montiel 18611 (MO); Macizos de Pelias Blan- 
cas, SE side, drainage of Quebrada El bp slopes 
N and W of Hda. San Sebastián (1321 4 15'N, 85°38' W), 
Stevens et al. 21271 )) NUEVA SEGOVIA: carr. a Di- 
pilto, 3 km al N de Ocotal Santa Marta (13°39’N 
86?28'W), Moreno 25253 (M ZELAYA: La Luz 
(l 3944'N, 84?47'W), Ortiz 1589 MO) Suina, Seymour 
3200 (MO). Costa Rica. GUANA : Bahia. El Coco, 
md Playa Hermosa (10%32N, "an 59. Ht W), Burger & 
Baker 9942 (F), Parque Meis Santa Rosa. H. ne T 
nández et al. 6098 (MENU, MO). PUNTARENas: Monte 
verde, Haber . 392 (MO). SAN JOSE: i Rio Virilla, Son 
José, Brenes 20565 (NY); Planta Hidroe léc trica Brazil- 
bus Colón, Por eda 934 (F); La Caja, 


CErce 


CUBA. SANTIAGO DE CUBA: Santiago Bay, Clemente 4279 
(US). Sardinero, Clemente p (GH, US); Finca El Sa- 
lado, Clemente 6268 (G S); Finca Caimanes, E 


mente 0340 E H); Loma € ed C : "nente 7030 (CH): 
, Clemente 7439 (GH, NY); Bayate, Ek- 
man 6432 (NY "i en la zona entre el Puente de la Mina 
a Estación de Radio, carr. de Punta Gorda, López 217 
it S); bordes del camino en Santa Maria de Loreto, López 
& Acuna 346 (US) VILLA om Santos Espir iritus 
to Loma de Cantú, ya 5. 3501 NY, in | TRINIDAD: Is pe 
of Hues Frier 5 (NY). VENEZUELA. ANZOATE( 
Isla C nds Seg s N of Guanta (10°18 N, 64°36’ Y . 


Steyermark & Manara 107998 (US). ARAGUA: detrás 
del Instituto de Botánic a Agric ola, Ms ultad de Agronomía 
as 1979 (F), Magdaleno, H il- 
liams 10303 (F, G, . BOLÍVAR: headwaters of Hio 
Saca, Wurdack «€ upp 60 (F, NY, US). CARABOBO: 
alrededores de Mariara, 2s 1051 (F); Pto. Cabello, 
Curran & Haman 114 1 (A, F, GH, MO, NY, Us): 

>] Cura, near San reo Pittier 7902 (F, 


£ 
= 
^ 
< 
cS 
`) 
E. 
= 
= 


haci ie "nda de 


A La Cabrera, on Lake Valencia, Pittier 12212 
(A, ( , US). a dri RAL: between Caracas and 
cn pude to La Guaira, Pittier 11619 (A, F, MO, 
NY, US). MÉRIDA: Peninsula Paraguaná, La Ciénaga, 


Nuevo and Santa Ana, Breteler 4373 
(F, MO, NY, UC, US); Chama Valley between Estanques 
and La Pietaria,  Birateler 4502 (F, MO, UC, US). mi- 
RANDA: Parque Nacional Guatopo, 22 km N of Altagracia 
de Orituco, N of the border with Estado de Guárico, Vee 
17935 (NY): Guarenas Valley, Pittier 11903 (A, NY, 
US). NUEVA ESPARTA: Isl 
SUCRE: Puerto de Hierro, 


be tween. Pueblo 


jio Aristeguieta 1719 (NY, UC, US); Peninsula de 
Araya, 20 km NW of Cariaco, 2 km W of corner where 
road inan Cariaco to Chocopata mee r ocean (10°38'N, 

1340'"W), Liesner & Gonz les z 12037 (MO). TACHIRA: 
: of La Mulata, near Venezuelan E ia do 
(79550'30"N, 72?27'3"W), Steyermark et al. 120203 
(MO). Yaracuy: Hacienda Iboa, near Guama, | to 


11160 (GH, , US). ZuLta: San Martín, 
Palmar, Pier 105. 31 is H, U x COLOMBIA. 


Canacoima, Dugand 676 (F); Los Pendales, 


Riodulce, Dugand & Jaramillo 1104 (US); Costa del 
Caribe, S de Salgar, Dugand 6421 (US); Costa del 

Caribe, erto a m 5, Dugand 60574 
(US); Barranq il Elías 314 (N SOLÍVAR: La 


ena, Killip de HM 14008 (A. BM, 
s; Torrec illa, near Turbaco, Killip & 
A 14388 (A. GH, NY : 
tecita and Carraipia, Rio Cesar, Cuatrecasas & € 
neda 25525 (US). MAGDALENA: Becerril, Haught poe 
(A, US); trail pm Pueblito, Parque Nacional Tayrona 
(11219, 73358 W), Kirkbride 2513 (F, MO, NY, US); 
0.5 km E of Aeropuerto Simón Bolivar and 14 km Sc 
Santa Marta (11907'N, 74°15'W), Kirkbride 2606 (MO. 
NY, US); Mpio. de Santa en Don Jaca, Romero 
10595 (F, MO, NY) NORTE DE pa AN 
Peralonso, en los alrededores de 
Barkley 18 (US). BRAZIL. N 
ert 755 (K). Be 


Popa, near Cartas 


denas 4775 (US): Alto Parapeti, de Michel 
ie AGUAY. Fuerte Olimpo-Chaco, Rojas 

: Zwischen. Rio va und Rio Aquila Estrella, 
E 4304 (BM, GH). ARGENTINA. CHACO: Las Brenas, 
Femi 10198 (A). JUJUY: Dique La Ci T Krapovt 
ckas & Cristóbal 17540 (LL, UC, WIS), SALTA: jesus 
González, Aguilar 252 (NY); Dept. de Orán, Carapari, 
Bridarolli 3262 (GH); Cerro 20 He Febrero, Filipovich 
255 (US); 10 km E de Tartagal, camino a buco Kra- 
povickas & Schinini 39282 (WIS); 14 km E de Em- 
barcación, Krapovickas & Schinini Po 1 (G); Depto. 
San Martín, Gral. Ballivin, pus El Saladillo, Logname- 
Medina & Cuezzo 10144 (WIS); Pasaje del tio ura- 
mento, Lorentz & Hieronymus 225 (BR, G, ! JC); 
Depto. Orán, Ruta 50, Quebrada El Cebilar, 
et al 504 (MO, P): Depto. Caldera, Mojotoro, 
35760 ire NY): El Tabacal a Rio Santa Maria, 
4521 (UC); Depto. Oran, Urindel a Rio San Francisco, 


Marunak 
: 


Annals of the 
Missouri Botanical Garden 


Meyer 8413 (A, NY); Depto. Orán, Bella Vista, Pierotti 


/ , , Alen mal ma, 
Venturi 9812 (A, BM, GH, MO, NY, 
Cerro Negro, Venturi 10. 390 (BM, x 
i pp 


SANTIAGO DEL 


Estero, ins Jovi E 390 (MEXU) 


to. Burruyaco, 
MO, NY, UC, US): Depto. Trancas, Tapia, ert 5824 


(A, CAS, US); ao Burruyaco, Cerro del Campo, Fen- 
uri 7525 i CAS, GH, US). CULTIVATED. U.S.A. 
MISSOURI: St lis, M Botanic al Garden, H. Her- 
nández 946, 953, 1 (MEXU, MO). Mexico, Guana- 


juato, Garden of d eds: Dugès 1 (GH, NY, US). 


vecimens not assigned to subspecific status, prob- 
ubl intermediate between Z. eas subsp. formosa 


and Z. formosa subsp. rosei. EXICO: GU d Distr. 
Coyuca, Pungarabato, dues 087 BA, , F, GH,LL, 
EXU, NY, US); Distr. Mina, den eres Camerún, 


Hinton 9191 (ENCB, K, iL. a NY, UC); 13 km NE 
of Atoyac, al 1270-80 E NY); Mpio. de Zum- 
pango del Río, 8 km al N de Milpillas, carr. Iguala 
REM Ad. d et al. 120 eee MO); Mpio. de 

Chilpanei i F NS ravos, sobre la desv. a Chichihualco, 
a 8 km al NW Gilneas ae Ladd et al. 146 (MEXU, 
MO); Distrito inn N of Rio alsas, Temisco, Barranca 
de la Julia, Mexia 31 (CAS, GH, MO, NY, UC, 
US); 4 mi. W de Zumpango del Rio, Rico & Funk 201 
(CAS, MEXU). 


Zapoteca formosa subsp. formosa is the most 
widespread and morphologically variable subspe- 
cies within the species. Fourteen names have been 
published that fall within the boundaries of this 
subspecies as | have defined it; most of them have 
been based primarily on regional variations in leaf- 
let morphology and vestiture. These characters, 
along with the size of the flower parts and pods, 
are highly variable at an inter- and intrapopulation 
level, especially in the Mexican and Central Amer- 
ican populations, and thus they are not useful tax- 
onomically. Perhaps the most variable character 
is leaflet size, but the variation does not correspond 
to a geographical pattern. Á similar situation occurs 
with variation in leaflet vestiture; plants with short- 
pubescent or villous leaflets occur in the same areas 
as plants with glabrous leaflets throughout most of 
the range. In addition there is continuous variation 
in leaflet apex morphology, ranging from rounded 
to acute. Plants from the southern limits of this 
subspecies are comparatively consistent. morpho- 
logically. In this area, they usually have medium 
to relatively small leaflets, and the number of pairs 
of leaflets per pinna is quite regular. 

The entity most closely related to Z. formosa 
subsp. formosa appears to be the Central American 
subsp. salvadorensis, which is distinguished by its 
densely pilose pods. Zapoteca formosa subsp. for- 


mosa is consistent in having glabrous pods, al- 
though in El Salvador and Venezuela, plants with 
glabrate pods have been collected. In a few col- 
lections of subsp. formosa from Oaxaca, Jalisco, 
Honduras, Venezuela, and Cuba, the pods are pi- 
lose or sparsely pilose when very young, but soon 
become glabrou 
Zapoteca Pm subsp. formosa also appears 
closely allied to subsp. rosei, from which it differs 
by having white or white-green filaments (Table 
4). Plants of subsp. formosa also usually have 
larger leaflets and are generally more robust than 
those of subsp. rosei. In a number of instances, 
however, the identification of these two subspecies 
is troublesome if flowers are not available or if 
filament color is not known. In some areas of the 
Mexican Pacific slope, especially in the state of 
Guerrero, where these two subspecies presumably 
occur sympatrically, hybridization probably takes 
place, resulting in morphological intergradation. In 
this area, many collections of subsp. rosei have 
larger leaflets than normal in this entity, thus re- 
sembling those of subsp. formosa. Such areas of 
intergradation are in Tecpan de Galeana (e.g., Hin- 
ton 14335, 10894, Ladd et al. 206) and Zihuata- 
398, Ladd et al. 216, 


Sousa 6159). Similarly, in some localities of the 


nejo (e.g., Tenorio et al. 
District of Yautepec in Oaxaca, plants from the 
two subspecies have been found growing together 
215, 217; Telles 
13). The collections belonging 


— 


e.g., H. eie & Torres 
Magallanes 111, 
either to Z. formosa subsp. formosa or to subsp. 
rosei, but of ambiguous identity, are listed sepa- 
rately under Z. formosa subsp. formosa. 

Zapoteca formosa subsp. formosa is distinguish- 
able from the West Indian subsp. gracilis by hav- 
ing oblong-obovate leaflets, usually larger, lanceo- 
late to ovate-lanceolate stipules, and glabrous, 
pubescent, or villous leaflets, which contrast with 
the consistently glabrous ones of subsp. gracilis. 
Subspecies formosa differs from subsp. mollicula 
and subsp. schottii by its more robust habit, white 
filaments, and larger Pers leaflets, stipules, flower 
Table 1 

ere is a wide rude tion in the South Amer- 
ican distribution of Z. formosa subsp. formosa, 


— 


parts, and pods 


from northern Colombia and Venezuela to Bolivia, 
southeastern Brazil and Paraguay (Fig. 44). Mor- 
phologically, the populations from the southern 
region clearly belong to the subspecies. Perhaps 
this geographical fragmentation is related to the 
current scarcity of relatively dry, seasonal, lowland 
habitats in the intervening region. As reviewed by 
Haffer (1982, 


fluctuations during the Pleistocene had a great 


and references therein), climatic 


Volume 76, Number 3 
1989 


Hernández 


effect in tropical areas. In lowland areas of the 
Neotropics, alternating periods of cooler and drier 
climates probably caused extinction or displace- 
ment of species adapted to the warm-wet forest 
conditions during the interglacial periods, and only 
a few areas (forest refugia) remain relatively un- 
changed. Consequently, great expanses of wet for- 
ests were replaced by drier, more open vegetation 
in the Amazon Basin (Ab? 1982; Absy, 1982; 
Bigarella & Andrade-Lima, 1982; van der Ham- 
men. 19 1982; and others). In light 
of this body of ideas, it can be postulated that the 
populations of Z. formosa subsp. formosa found a 
more suitable environment during the dry phases 
of the and then were continuously 
distributed across the Amazon Basin; and that the 
America is the 


Saber, 


Prance, 


Pleistocene, 


present-day disjunction in South 
result of fragmentation of a formerly more contin- 
uous range after the warm, wet present-day con- 


ditions were established. 


12b. Zapoteca formosa subsp. salvadorensis 
(Britton & Rose) H. Hern., comb. et stat. nov. 
Anneslia salvadorensis Britton & Rose, N. 
Amer. Fl. 23: € 1928. TYPE: El Salvador: 
vicinity of San Salvador, 650-850 m, 2-7 
Feb. 1922, Standley 20152 (lectotype, NY, 
here designated: isolectotypes, GH, MO, US). 


Erect shrubs to 4 m tall, usually finely to densely 
villous throughout, except for the flowers. Pinnae 


— 


2-3-jugate: petioles (1.8-)3.1-4.5 em long; rachis 
(0.9-)1.6-4 cm long: rachillae (1.8-)3.8-5.0 cm 
long; leaflets 4-6 pairs per pinna, oblong-obovate, 
oblique at base, rounded to acute at apex, usually 
mucronate, the distal leaflets 1.6-2.6 x 0.7-1.9 
cm, the proximal ones smaller: stipules lanceolate, 
sometimes curved, to 1.2 em long. Capitula usually 
axillary, occasionally in simple pseudopanicles to 
7 em long: peduncles 2.2 
Calyx 3-4 mm long. the teeth oblong-lanceolate, 
glabrous; corolla ca. 5 mm long. glabrous; filaments 
white. Pods to 9 x 0.8 cm, pilose to densely pe 


8.2 cm long at anthesis. 


with white hairs. Seeds rhomboid-ovoid, 
4 mm. sometimes red when immature, Sdn when 


mature, the areola lighter: pleurogram irregular. 


Distribution (Fig. 44). Endemic to southern 


Guatemala and El Salvador, 
collected in the departments of El Progreso, Chi- 
quimula, Quiché. Sololá. and Zacapa, Guatemala, 
It has been 


where it has been 


and near San Salvador. El Salvador. 
collected primarily in disturbed, brushy habitats at 
altitudes of 300—1.800 m. 


4dditional examined. 


spe € iT 
CHIQUIMULA: between Chiquimula and Sta. Roa. Moli. 


nens 


na & Molina 25136 (F, MO, US); near divide on road 
from Zacapa to bee m Standley 73743 (F). EL 
ds. RESO: Motagua \ kon & Breckon 2118 

MEXU, NY). QUICHÉ: near village of Rio Blanco, 
ams e ~ 22456 (BM, F, GH, NY, US). 5 OLOLÁ: 
villag line Volcán Sun Podio: north 
“lopes d Lage 
ZACAPA: Rio Hondo, A of Sierra de las Minas, Stands y 
i 7397 9 (F) Río Hondo, Steyermark 29494 (F). 


i 


— 
> 
£5. 

2 5 
E 
T 0 


Zapoteca formosa subsp. salvadorensis is closely 
related to subsp. formosa, from which it can be 
distinguished by having pilose or densely pilose pods 
and by being villous on all vegetative parts. By 
contrast, presence or absence of vestiture in subsp. 


formosa is expressed irregularly throughout its 


range. Geographically, these two subspecies are 


allopatric. 

12c. Zapoteca formosa subsp. gracilis 
(Griseb.) H. Hern., comb. et stat. nov. Cal- 
liandra gracilis Griseb., Mem. Amer. Acad. 
Arts 8: 180. 1861, non Klotzsch, 1871, nec 
Standley, 1929, nec Renvoize, 1981. Annes- 


lia grisebachii Britton & Rose, N. Amer. Fl. 
23: 67. 1928. This was published as a nomen 
novum, Anneslia gracilis 


was preoccupied by Britton € Rose for a 


because the name 


species based on Acacia gracilis M. Martens 
€ Galeotti (see synonymy under Z. formosa 
subsp. rosei). Calliandra grise bachu (Britton 
& Rose) Standley, Publ. Field Columbian Mus., 


Bot. Ser. 3: 277. 1930. C. grisebachii (Brit- 
ton & Rose) León, Contr. Ocas. Mus. Hist. 
Nat. Colegio "De La Salle" 10: 237. 1951; 
Liogier, Fl. Cuba (Supl.). 72. 1909. TYPE: 


Prope Cuba Orientali, Sep. 1859 Jan. 1800, 
Wright 151 pro parte (lectotype here desig- 
nated, GH: isolectotypes, CGE, F, G, LE, MO). 
As León (1951: see synonymy below) pointed 
out, Wright 151 includes two elements under 
the same number. In addition to the specimens 
considered here to represent the type of this 
there are five collections, also num- 
at G. GH, LE, NY (fragment), 


came from another 


taxon, 
bered as 151. 
and S, that presumably 
locality and belong to Z. formosa subsp. for- 
mosa. One additional specimen at G includes 


material from both. 


Calliandra comosa Griseb., . PL Cub. 83. 1866, non 


(OUS formosa var. cubensis J. 


enth., 
Macbr., a Gray Herb. 59: 4. 1919. (Based 
on the same pe as Callic ae. comosa.) Anneslia 
(J. F. Macbr.) Britton & Rose, Amer. 


* to 
uw 
~ 
O 
^ 
o 


e Calliandra cubensis (J. F. Macht) 
Lai on, ‘Came. Ocas. Mus. Hist. 

Salle" 9: 7. 1950; León, Contr. 
“De La Salle" 10: 23 


Nat. Colegio T iogier, 


Annals of the 
Missouri Botanical Garden 


m Cuba (Supl.). 72. 1969; Proctor, Kew Bull. 11 
81. 1984. TYPE: CUE: Wright 2406 (lectotype 
De designated, GH) 


Erect shrubs to 2(-5) m tall, glabrous through- 


out. Pinnae 2-3(-4)-jugate; petioles 1.3-5.5 em 
long: rachis 0.7-5 cm long; rachillae 3.5-12.5 cm 
long; leaflets (5—)7—15 pairs per pinna, the median 
ones elliptic to elliptic-oblong, the distal ones oblique 
at the base, the proximal ones becoming acute, 
rounded to subacute, rarely acute at apex, some- 
times mucronate, the median leaflets 9-22 x : 

ll mm, all usually drying light green; stipules 
linear-triangular, to 5 mm long. Capitula axillary; 


peduncles 3.5-10.5 em long at anthesis; calyx 
2.5-3 mm long: corolla 5-6 mm long; filaments 
white. Pods to 11 x 1.2 em, glabrous. Seeds ovoid, 
5-6 x 3-4 mm, usually mottled, pleurogram se- 


miregular. 


Distribution (Fig. 44). Restricted to several 
islands of the West Indies: Cuba, Bahamas, Cay- 
man Islands, and Haiti, where it primarily occurs 
in rocky, calcareous areas, in dense thickets, and, 
in general, in secondary, arid habitats at elevations 
of 0-600 m. 


Additional specimens examined. BAHAMA ARCHI- 
PELAGO. ABACO: Eight Mile Bay, Brace 1854 dubi. 
NGUILLA ISLAND: hacia Wilson AT NY). ANDR 


Brace 6661 (F, NY); Be rires 
s South Roca S of Little 
k, Correll 4348 1 (NY): 2 mi. E of Andros Town 
Hill 3003 (NY); Mangrove Cay, near Lisbon 

NY 


airport, 


Creek, Small & Carter 8439 (F, GH, US). BERRY 
ISLAND: Whale Cay, Britton & Mills paugh 2183 (F, 
NY); Goat Cay, Britton & Vileooagh 2288 (F, NY) 


Anderson Cay, Great Harbour Cay, Correll end (E, 

NY). BIMINI ISLANDS: South yon Correll 42 

NY); Bimini, Cat Cay, Correll 456 14 (NY); E Bimini 

NE corner of Island, Gillis a. (F). CAICOS ISLA 

Millspaug eh & Millspaugh 92 35 (F. 
Millspaugh ra oa 

adcos, Hilson 7690 (F, GH, 


Window to Harbor Island, Britton & ca 5393 
(F, NY); North Eleuthera, near Watkins Hill, 
£1060 (MO); 6 mi. N of James Cistern, Corre 
15234 (NY). EXUMA CHAIN: | B : iay, Britton & 
Millspaugh 2843 (F, GH, NY, : Great Ina- 
ES Rocky Point, Correll 47471 nra - LONG ISLAND: 
Clarence Town, Britton & Milspangh 6291 e: NY). 
NEW PROVIDENCE: near Nassau, Curtis 63 (A, F, G, GH, 
MO, NY, US); Blue Hills, between Lake C pn ues and 
Lake Killarney oe 25°03.5'N), Webster et al. 
S) Hog Island, H ilson 8272 ). 
: Great Ragged Island, Hilson 7806 (F, 
NY). ROSE ISLAND: Britton & Millspaugh 2149 (F, NY). 
s T Proctor 29355 
(LL); 0.8-1 km inland from Spot Bay, Proctor 35189 
(US). CUBA: CAMAGUEY: Cayo Sabinal, between Corte Fi- 
cote and El Fuerte, Ekman 15539 (US) La Gloria, near 


Laguna Grande, xcd 602 (GH, NY, US); Santa Lucia, 


Shafer 974 (GH, NY, US); Cayos Canal Nuevo, Cayo 
Lagua, Shafer 2666 (F, NY ); Cayo Guajaba, hills 
NW end, Shafer 2855 ( l, A vicinity of 


Tiffini, Shafer 2901(F, NY, US). ORIENTE: 
Nipe Bay, Britton et al. 12471 (NY, US); Río del Medio, 
Jauco region, León 12426 (GH, US); Loma de Tabaco, 
Manatí Bay, León & Alain 18937 (GH, US); Holguin to 
ida er nd 1410 (NY, US) northward of Raimon, 
Nipe I , Shafer 1792 (NY). PINAR DEL RÍO: Pan de 
DER " Mulata, Barker 16513 (NY); Pan de Gua- 
jaibón, Alain & Acuña 2995 (NY); Sierra de Nipe, Rio 
Piloto, Ekman 5780 (G); Banes, prope ena Rico, Ek- 
man 6007 (NY). HAIT: lle de Nord 

“kman 2846 (NY, US): 
Leonard & Leonard 113 312 i GH, NY. 5) CULTI- 
VATED: U.S.A. FLORIDA: Miami, Fairchild Tropical Garden, 
Gillis 8672 (MO) 


Zapoteca formosa subsp. gracilis is relatively 
uniform morphologically compared with the other 
subspecies of Z. formosa. It is related to subsp. 


formosa, from which it is distinguished by having 


elliptic to elliptic-oblong leaflets and narrowly tri- 
angular stipules and by being glabrous (Table 14). 
The seeds of subsp. gracilis differ from those of 
the remaining subspecies of Z. formosa by having 
semiregular, instead of irregular, pleurograms. 
Subspecies gracilis and subsp. formosa are essen- 
tially allopatric, although there are a few collections 
Cuba. Their rela- 


of both from Oriente Province, 


tionship is discussed further under Z. formosa subsp. 


formosa. 


Although leaflet morphology is quite constant 
among populations of Z. formosa "E gracilis, 
a few plants from the Bahamas (e.g., Correll & 
Correll 45614, Curtis 82, Gillis 11 oe Hill 3003) 
have unusually narrow leaflets. These plants could 
be confused with Z. portoricensis but can be readi- 
ly distinguished by their smaller and narrower stip- 
ules and relatively thicker leaflets (also see Table 
14). In addition, Z. formosa and Z. portoricensis 


are strongly segregated geographically in the West 


— 


Indies. 


12d. 
gins) H. Hern., 


Zapoteca formosa subsp. rosei (Wig- 
comb. nov. Calliandra rosei 
Dudley Herb. 4: 17. 1950 
rosei (Wiggins) 
Acad. Sci. 6: 83 


Wiggins, Contr. 
Calliandra schottii subsp. 


Felger & Lowe, J. Ariz. 


1970. Acacia gracilis M. Martens & Gal. 
eotti, Bull. Acad. Roy. Sci. Bruxelles 10: 311. 
1843. Anneslia gracilis (M. Martens & Ga- 


leotti) Britton & Rose, N. Amer. Fl. 23 

1928. Calliandra gracilis (M. Martens & 
Galeotti) Standley, Publ. Field Columbian Mus., 
Bot. Ser. 4: 309. 1929, non Griseb., 1861, 
nec Klotzsch, 1871, nec 1981. H. 


Renvoize, 


Volume 76, Number 3 
9 


Hernández 849 


Zapoteca 


S. Gentry, Publ. Carnegie. Inst. Wash. 527: 
122. 1942 TYPE: Mexico. Oaxaca: bord del 
l'Océan Pac ‘cis Oct. 1840, Galeotti 3190 


(holotype, BR (not seen): isotypes, 6. K, LE). 


MC us socorrensis l. M. Johnston, Contr. Gray Herb. 
rau Anneslia socorrensis (1. M. Johnston) 
Hein & Rose, N. Amer. Fl. 23: 64. 1928. TYPE: 


Mexico. Colima: Isla Socorro, Mar.-June 


(0: 4 924. 


V. Anthony s.n. (holotype, UC no. 83534). 
later note, Johnston (Proc. Calif. Acad. Sei. 20: 62 


1931.) pointed out that Anthony made two unnum- 
bered collections on Socorro Island. Since he implied 
that they were made from ne localities, the 

at UC . 89257), should 

not be considered an isotype. 
dnne sstia de aay Britton & Rose, N. Amer. Fl. 23: 67. 
: Mexico. Baja € california Sur: San José 
del C ae da Sep. 1890, Brandegee 196 
pus s here designated; isolectotypes, A, 

GH, NY, UC (2), US). 

Anne 'slia interior Brian & Rose, N. Amer. Fl. 23: 66. 
Mexico. Duran mountains of south- 
ern Du 15 iis 1897, Rose 2317 (lectotype 
US no. 301227, here designated; isolec istae NY). 
Shrubs to 3(-4) m tall; stems usually slender, 
erect or drooping. Pinnae 1-3-jugate; petioles 1— 

4.2(-5) em long: rachis 0.3-3.4 em long: rachillae 
1.9-6.5(-11.3) em long; leaflets (3-)4-12(-17 
pairs per pinna, distal leaflets oblong-obovate 
rarely oblong, the 


— 
c 


o 
oblong-very widely obovate, 
proximal ones sometimes becoming elliptic, the dis- 
tal leaflets oblique at base sometimes becoming 
acute proximally, tounde d to acute at apex, usually 
2] mm, usually becoming 


w 


mucronate, 8-31 
smaller proximally, glabrous to villous; stipules lan- 
ceolate to ovate-lanceolate, to 9 mm long, glabrous 
to villous. Capitula axillary, sometimes in simple 
pseudopanicles to 13.5 em long: peduncles 1-4.7 
(-5.8) em long at anthesis. Calyx 1.5-2 mm long, 
glabrous, the teeth triangular to oblong-lanceolate: 
corolla 3-5 mm long. glabrous: filaments red-pur- 
ple, or white in the basal half and red-purple or 
pink in the distal half. Pods to 9 x 0.9 cm at 
maturity, glabrous. Seeds rhomboid-ovoid to widely 
4-5 X 4 mm, brown or gray, the 


areola lighter usually: 


rhomboid-ovoid, 
pleurogram irregular. So- 
2n = 26 


matic chromosome numbe r 


Distribution (Fig. 41). This subspecies occurs 
along the Pacific slope of Mexico, from northern 
Sonora south and southeast to the Isthmus of Te- 
huantepec. It has also been collected in southern 
Baja California and on the Revillagigedo Islands 
(Isla Socorro and Isla Clarión). A number of col- 
lections have been made from the districts of Yau- 
tepec and Tlacolula. Oaxaca. and from a number 
of localities in the Balsas Basin. It is relatively 


common in disturbed or undisturbed habitats of 


tropical thorn woodlands and tropical deciduous or 
tropical semievergreen forests. It is found on steep 
calcareous slopes, on volcanic rock, or along the 
margins of seasonal streams at 01,430 m. 
Common names Huaje de caballo (Oaxaca). 
tosapolo (Sonora). 


Additional specimens e bg id MEXICO: BAJA CAL- 


IFORNIA SU 10 mi. S Todos Santos-Cabo San 
Lucas DM on es je to Du ho La Burrera, Breed- 
love & Axelrod - 3 (CAS, MEXU). cHlaPas: Mpio. 


of tes e ie hal 195 near Monte Bonito, Jon 
3405 (CAS , US). CHIHUAHUA: Mpio. de Batopilas, 
E of La Bufa, near Arroyo La Buta, Bye 7720 (ME M 
SW Chihuahua, Palmer 297 (BM, G H, K, MEXU, NY, 
, US); La Junta, 13 km NE de Batopilas, Tenori 
Torre »s 4445 (MEXU). COLIMA: Socorro Teles 1d, 
s.n. (UC); Revillagige o = m Clarión Island, 
Sulpl ur Bay, Howell 8371 (CAS, GH) Revillagigedo 
Isl ecu Clarión Island, Mason 1566 (CAS, GH); Revillagi- 
Mason 1661 (CAS); above Rio Salado; * 5 
Mel augh & Koelz 1090 (MICH): 
, Isla Socorro, Miranda 8732 (ME XU 
IEXU); Manzanillo, Palmer 
NY, UC, US); 19 km al SW de 
Colima, carr. a Manzanillo, Soto & Cortés 257 1 (MEX 
along Colima side of Rio 2 in below hwy. 
& Fairhurst 1834 (CAS, GH, MICH, } 
ulco, Ls ve 
19 


~ 
~ 


above 


gedo Islands, 
mi. 5 of Colima, 
cumbre ie “rman 


Stevens ¢ 
GUERRERO: Pie de la Cuesta, near 
(F, US); 14.7 mi. N of € cna Ingo, ijo et al. 205 

(ENCB, MO); Distr. of Coyuca, Pungarabato, Hinton et 
al. 6400 (A, BM GH, LL, MO. ne US) Distr. 
Galeana, Tecpan, pe 10894 (BM, I i 
MEXU, MO, NY, ; Distr. Gale ana, 


388 (G. G H. K, MEXU. P. US); ip C anyon, Pringle 
10057 (CAS, F, GH, LL, MICH, UC, WIS); Achotla, 
Reko 4946 (NY, US); Mpio. de E del Río, Casa 
Verde, cerca de ree Rzedowski 10087 (CAS, 
2): 10 km al NW dee por el camino al Ca- 
IS): Bahia de Zihuatanejo, 
Vs NY); 45 mi. 5 of Iguala, 
ME XU, NY, TEX, US). 


Tomatlán, cerca de Puerto 


Sousa 6159 (CAS, MENT, 
f . 17M721 (BM, 
a de 


3 km de la Boe 


JALISCO: 
Vallarta, Delgado et al. 139 (CAS, MEXU, MO, TEX); 
Mpio. La Huerta, Estación C hame la, H. Hernández & 
Lott 230 (MEXU, MO); hills between Barra de Navidad 
and La Manzanilla on Bahía Tenacatita, Wel augh 20988 
(MICH); cerros al N de San Patricio, Pérez & Pérez 
1256 (MEXU): carretera al C ;oastecomate, Puga 2 
(ENCB); Corcovado Canyon, 11 n ¡E of Autlán me 
d Wilbur & Il ibur 23: 85 (MENU, MIC 
MEXICO: Distr. of Temascaltepec, Tenayac, Pintan 1 ri 


(GH. k, py Distr. of dum altepec, Ac xl Peta 
6545 (F, ( , K, LL, MICH, NY, UC, US); Distr. of 
Temase uc Pistes: Hinton kie E MIC H, MO, 

, US). MICHOACÁN: Aqui km al W de la Cruz de 
Ca San. Guerrero et al. 1012 Y fes AL); Mpio. Jungapeo, 
entre Agua Blanca y Lázaro Cárdenas, H. Hernández « 
Chacón 185 (MEXU, MO): du Coalo omán, Villa V ic- 


toria, Hinton 12558 (BM, NY, 
UC, US); Distr. Zitácuaro, San ke Purüa, Bintan l. ud 3 
(BM, BR, CAS, F, G, GH, K, MEXU, MICH, NY, TEX, 


850 


Annals of the 
Missouri Botanical Garden 


Me drano et al. 
ar els eo e los Coyotes 

ayabo, Soto 197 (C AS, MEXU y 
Mpio. de Arteaga, en cerros aledaños a pui. Soto 
& Silva 1036 (MO); carr. Cuatro Caminos- Playa Azul, 
> Arteaga, & Silva 1699 (MEXU, TEX, 
WIS); 7 km al 5 de Arteaga, Sousa & Soto 8033 (CAS, 
MEXU); Mpio. de e Puerto de los Cirianes, 6 
km al h a Huetamo, carr. Huetamo- Zitácuaro, Tenorio 
et al. 8 (MO). MORELOS: 10 km N de Teuixtla, Gómez- 
rd pos (MEXU); s rro Acatlipa, Pázquez 2046 
(MEXU). Oaxaca: Distr. de Jamiltepec, de Rio Verde a 
Jamiltepec, Conzatti je (US S); Ixtepec, Fisher . 35491 
(F, MO, NY, US); Mpio. Totolapan, 9 km E Totolapan 
rr. a Tehuantepec, H. Hernández & Tartes 207 
, MO); M San Matéo de Mar, 13.3 km NW 
d Hernández & Torres 209 (MEXU, 
, 17 km NW La Reforma por carr. 

El Camarón, 7 n 'rnández & Torres 210 (MEXU, 

MO) Mpio. Nejapa, 3 i 5 Nejapa de Madero por camino 
que desemboca en carr. Oaxaca- Tehuantepec, H. Her- 
nández & Torres 512 (MEXU, MO); Distr. Tlacolula, 
entre San José de Gracia y Totolapan, H. Hernández & 
Torres 215 (MEXU, MO); Distr. Tlacolula, enfrente de 
la Virgen Juovila a 12 km E de Tatolapan, H. Hernández 
& Torres 217 (MEXU, MO); Distr. = hitan, 3.3 km E 


de La Ver 


ntosa por carr. Tapanatepec, H. Hernández & 
Torres 337 (MEXU, MO); Distr. Tehuantepec al pie del 
Cerro Guingola, vertiente E rada a las Ruinas 


Arque ológie as, H. Hernández & Ramírez 918 (MEXU); 
Distr. Yautepec, 2.6 a desviación a San Bartolo 
Yautepec (Portillo pa San Bartolo Yautepec) de la carr. 

. Hernández & Ramírez 930 
San José de Gracia, 
e San José de Gracia y San 
Ius, Guegoyache, "IL onde. & Ramírez 932 
Tlacolula, NW de Totolapan por 
carr. a nata: H. Hernández & Ramírez 943 (MEXU); 


sobre carr. 


6 146 (MO); Distr. Juquila, 7 km al NW de Rio Grande, 
Sousa et al. 707 79 (BM, CAS, MEXU, MO); Distr. Juquila, 
> Pto. Escondido, Sousa et al. 8408 
(CAS, MEXU, MICH); Distr. Juchitán, Mpio. de San 
Miguel Chimalapa, Vista Hermosa, 3 km al NE de El 
Ae Sousa 8 8087 (CAS, MEXU, NY); 2 km al SW 
» San José de Garcia, cerca de Los Cantiles, carr. Oa- 
xaca huertas Téllez £ Magallanes 113 (CAS 
MENU); 1 km al SW de El Chacal, 
El Camarón, Télle yi & Magallanes 253 (ME 
e El Limón, 
al SW de svg D carr. Tehuantepec -Oaxaca- Bue- 
2 (MEXU, MO). siNALOA: Bahia 
A in hes Sun Navachiste, Gentry 11460 
, MICH); cerros Fuerte r 24 mi. N of 
Los Moc rin Ge ntry 1442 L, MEXU, 
ui 7305 (CAS, G, K. va uh 
mi. 5 of Elota, Skorepa 82 (MO); 
camino de Herradura El Alamo- Cerro auc '0, 
NE del Ejido de los de Ponce, Tenorio et al. 2816 (MO). 
NORa: Cañón de Nacapules, ca. 6 km NE Bahía San 
Carlos, Felger et Sa 84-124 (MEXU); Ense idea Grande 
(San Pe Fia Bay), 0.5 mi. E from S end of Bay, Felger 
et al. 115079 (ME x ); San Bernardo, xi Mave Pus 
1630 (A, F, K, LL, MEXU, MO, NY, ,U: 


= 


Tres Cruces, al S 


nos Aires, Torres 


S); Cerro 


formosa by the 


del Viejo SW of Caborca, Gentry 14459 ^ o a 
Bojihuacame SE of Cd. Obregón, Gentry 8 (LL, 
US); San Bernardo and vicinity, Arroyo bcd ies 
et al. 19286 (US); Hacienda de San dard E a- 
topilas, Palmer 16 (BM, GH, K, MEXU, 

This polymorphic, highly variable subspecies, 
along with Z. formosa subsp. formosa, represents 
the phylogenetic core within the species. Leaflet 
size is highly variable within and between popula- 
tions, and a clinal variation pattern is evident, 
especially in the northern portion of its range, 
possibly a reflection of decreasing precipitation from 
south to north. There is some variation in filament 
color within Z. formosa subsp. rosei, although this 
character is consistent at the populational level. 
The distribution of the different color morphs is 
not known with precision. The following generali- 
zations can, however, be drawn: the most wide- 
spread pattern of filament color is the white/red- 
purple pattern, which occurs from southern Oaxaca 
to Sonora and Chihuahua. The populations from 
the Isthmus of Tehuantepec, at the eastern distri- 
butional limit (e.g.. Hernández & Ramírez 
918, Sousa & Téllez 8687), and a few populations 
from Michoacán (e.g., H. Hernández & Chacón 
185, Soto 1636) 


ments. Populations from the districts of Tlacolula 


have entirely red-purple fila- 


and Yautepec, Oaxaca, have white pos filaments 
(e.g.. H. Hernández & Ramírez 930, 943). 
Zapoteca formosa subsp. rosei is closely related 
to subsp. formosa and to subsp. mollicula and 
subsp. schottii. It can be distinguished from subsp. 
color of its filaments and by its 
usually smaller leaflets. However, plants of these 
two subspecies sometimes resemble each other 
closely. In some localities of Guerrero, the two 
subspecies become sympatric and apparently in- 
tergrade morphologically. These collections are not 
assigned to a particular subspecies; they have been 
listed separately under subsp. formosa (and see 
discussion under this subspecies). 
will be discussed below under the respective 
subspecies, patterns of morphological variation, es- 
pecially in leaflet size, are essentially continuous 
between Z. formosa subsp. rosei on the one hand 
and subsp. mollicula and subsp. schottii on the 
other. Although each of these is relatively easily 
identifiable, their integrity breaks down because of 
the existence of morphological intermediates. Za- 
poteca formosa subsp. rosei can be distinguished 
from subsp. mollicula by having more-developed 
leaf parts (e.g., longer petioles and rachillae, usu- 
ally more numerous and larger leaflets, and usually 
larger stipules (Table 
can be distinguished from subsp. schottii primarily 


14)). Similarly, subsp. rosei 


Volume 76, Number 3 
89 


Hernández 
Zapoteca 


by having more numerous and larger leaflets (Table 
14). Nevertheless, the differences between these 
two subspecies are less marked than those sepa- 
rating subsp. rosei and subsp. mollicula. Geo- 
graphically, subsp. mollicula and subsp. schottii 
are peripheral to but contiguous with subsp. roset. 
This fact, along with their close morphological af- 
finities, suggests that subsp. mollicula and subsp. 
schottii were derived from subsp. roset. Further 
discussion of the presumed relationships of these 
three taxa is provided under the respective sub- 
species. 

12e. Zapoteca formosa subsp. mollicula (M. 


Martens & Galeotti) H. Hern., stat. et comb. 
Acacia mollicula M. Martens & Galeotti, 


nov 
Bull. Acad. Roy. Sci. Bruxelles 10: 313. 1843. 
Anneslia mollicula (M. Martens p Psion) 
Britton & Rose, N. Amer. Fl. 23: . 1928. 


Calliandra mollicula (M. Martens A Galeotti) 
du “ield Columbian Mus., Bot. 
Ser. 1929. rype: Mexico. Puebla: 
Plaine de Sec 5.500 ft., H. Galeotti 
3216 (lectotype here designated, BR). 


Standley, 


cce E re Rose, Contr. U.S. Natl. Herb. 5 
S unijuga (Rose) Britton & Rose, 
. Amer. oe . 1928. exico. Oaxaca: 
Cuisailán, l, a y 24 Sep. 1894. L. C. Smith 
203 (lectotype here designate d; GH: isolectotypes, 
2 fragments). 
Calliandré unijuga var. pueblensts : F. Macbr., Contr. 
Herb. 59: 4. 1919. TYPE: Mexico. Puebla: 
vicinity of San Luis Tali a Oaxaca, Bar- 
ranca de Tlacuilosto, June 1908, C. 4. Purpus 3185 
e x F, pos n e pudo BM, GH, 
(UG. U»(2 


Tay 


a 


Erect shrubs to 2(-2.5) m tall; stems slender, 
sometimes with exfoliating bark; branchlets pilose 
in the younger parts, glabrate, sometimes glabrous 
throughout. Pinnae 1(-2)-jugate: petioles 3-11 mm 
long; rachis 4-8 mm long; rachillae 6-18 mm long; 
leaflets (2-)3-5(-6) pairs per pinna, oblong-ob- 
ovate to widely obovate, rarely elliptic, the distal 
ones oblique at base, 4-14 x 2-9 mm, the prox- 
imal ones smaller, becoming rounded or acute, 
rounded or rarely retuse at apex, usually mucro- 
nate, all membranous, finely pilose or villous to 
totally glabrous; stipules lanceolate to ovate-lan- 
ceolate, sometimes narrowly triangular, to 2 mm 
long, finely pilose to glabrous, usually ciliate. Ca- 
pitula axillary: peduncles (1.2-)2.3- 4.5 cm long 
at anthesis. Calyx ca. 2 mm long, the teeth tri- 
angular, glabrous; corolla ca. 5 mm long; filaments 
white in the basal half, red-purple in the distal half. 
Pods to 7.5 x 0.8 em at maturity, glabrous. Seeds 


ovoid, ca. 5 X 4 mm, brown; pleurogram irregular. 


Distribution (Fig. 44). Endemic to the Te- 
huacán and Cuicatlán valleys in the states of Puebla 
and Oaxaca, where it primarily occurs on dry, 
calcareous slopes, in arid scrub and tropical dry 
forests at 550- 1.450 m. 


Huajillo (Puebla). 


Common name. 


ida specimens e xamined. Mr EXICO. PUEBLA: 
lado W de Axus ia i al. 
san Pur ien 545 EXU e 
al. 5 (NY. US); B. al SE de 
las s (18°20'N, 97?26'W), Zavala 
OANACA: arias a, Conzatti 404.1 (MEXU, U S) Coix 
tlahuaca, La Huert NE de Tepelmeme de 


al 
NCB. MEXU): 


Morelos, Cruz 2563 (El 


CO, 


A 


Tomellín, H. Hernández & Torres 21€ ¿XU, MO); 
Distr. Cuicatlán, o por carr. a Santa Maria, H. 
M ne & Torr 1 (MEXU, m 7 km N Cur 


.a inr del Camine 
Distr. Teotitlán. 


catlá or car 
& Torte s 226 ( IEXU 
'Tecomavaca ES carr. 

227 (MEX, MO); Distr. bre 5 km Sde T 
por carr. a i Clés dán. H. Hernández & Ramirez 8/4 
(MEXU): Distr. oe l km Tomellin por el camino 
a la carr. Cuicatlán e 1, H. Hernández & Ramirez 
870 (MEXU); Dis Teotitlán. en el desemboque del 
camino viejo a Tome sin, sobre la carr. Cuicatlan nach 
H. Hernández & Ramtrez i 7 (MEXU); laderas al SE 
de Cuicatlan, EXU); Cuicatlán, Velson 
1648 (US): Tomellin Canyon, pe & Hough 467 1 (US). 
Distr. Cuicatlán, 15 S de Doria: Rzedowski 
34957 (ENCB); Cañon p» Tomellin, Sousa et al. 6905 

(BM, MEXU, MO). 

Zapoteca formosa subsp. mollicula represents 
an extreme of morphological variation within the 
species in petiole length, number of pairs of pinnae, 
and number and size of leaflets. It is distinguished 
from the other subspecies by its | 
pinnae with fewer and smaller leaflets and by its 
aflet size, however 


-, rarely 2-jugate 


somewhat shorter internodes. Lea 
is extremely variable in this sna o both at an 
intra- and interpopulational level (e.g., Cruz Cis- 
neros 2563, H. Hernández & Torres 216, Rose 
& Hough 4671, Sousa et al. 6901, 0905). Sim- 
ilarly, vestiture is highly variable, and does not 
display any coherent pattern. 

Zapoteca formosa subsp. mollicula clearly is 
closely related to the highly variable subsp. rosei. 
Perhaps the most taxonomically meaningful char- 
acters separating these taxa are the usually more 
reduced leaves of subsp. mollicula. However, vari- 
ation in leaf characters (e.g., petiole length, number 
of pinnae, number of leaflets, size of leaflets) is 
essentially continuous, especially when plants of 
from the districts of Tlacolula and 

, Delgado et al. 724, H. 
919 915, 2 


dig = 


subsp. rosei 
Yautepec, Oaxaca (e.g 
Hernández & Torres 207 , 210, 
Hernández & Ramírez 930, 932, 913, Magalla- 
nes et al. 199, Sousa et al. 0519) are compared 


Annals of the 
Missouri Botanical Garden 


ND. 
Cisneros 2563, Hernández & Torres 218, Her- 
nández & Ramirez 674, 876, 877, Rose & Hough 
107 1, Sousa et al. 6904, 6905). 


with some plants of subsp. mollicula (e.g 


12f. Zapoteca formosa subsp. schottii (Tor- 


rey ex S. Watson) H. Hern., comb. nov. Cal- 
liandra schottii Torrey ex S. Watson, Proc. 
Amer. Acad. Arts 20: 364. 1885. “Callian- 


dra portoricensis Benth. var." Torrey, Emo- 
1859; Isely, Madroño 21: 280. 
1972. This species was doubtfully referred to 
in Torrey's Botany of the United States and 
Mexican Boundary Survey (loc. cit. )as “Cal 
Subse- 
Calliandra 


ry Rep. 2: 61. 


liandra (portoricensis Benth. var.)." 
S. Watson 
schottiton the description provided by Torrey, 


quently, based his 
citing a Pringle collection (272). Anneslia 
schottii (Torrey ex S. Watson) Britton & Rose, 
N. Amer. Fl. 23: 67. 1928. Calliandra schot- 
tii subsp. schottii (Torrey ex S. Watson) Fel- 
ger & Lowe, J. Acad. Sci. 6: 83. 1970. 
TYPE: Mexico. Sonora: Pimeria Alta, Sierra 
Verde, 20 Aug. 1855, 
Schott s.n. (lectotype designated by Isely (Ma- 
280. 1972), GH; isolectotypes, NY 
S (fragment)). 


Ariz. 
Arroyo de los Samotas, 


drono 21: 

(2), U 
Erect shrubs to 1.5(-2) mm tall; stems slender, 
densely branched; branchlets glabrous to finely pi- 
lose. Pinnae 1 -2-jugate: petioles 1.2-3.5 cm long; 
rachis 0.4- 
Leaflets (3—)4-6(-7) pairs per pinna, oblong-ob- 
4-12(-14) x 2-6( 


rounded to acute at apex, usually mucronate, mem- 


-7 em long; rachillae 1.9-4.5 em long. 


ovate, 7) mm, oblique at base, 
branous, glabrous to finely pubescent; stipules nar- 
rowly triangular to lanceolate, sometimes slightly 
curved, to 5 mm long, glabrous to finely pubescent. 
Capitula axillary; peduncles 1.8-4.5(-6) em long 
at anthesis. Calyx ca. 1.5-2 mm long, glabrous, 
the teeth triangular-lanceolate; corolla ca. 3 mm 
long; filaments white in basal half, pink in distal 
half. Pods to 7 
rarely very gos and minutely pilose. Seeds 


rhomboid-ovoid, x 3.5-5 the 


X 0.6 cm at maturity, glabrous, 
mm, brown, 
areola lighter: HMM irregular. 
Distribution (Fig. 44). 


slopes in a few localities of the northern portion of 


Endemic to rocky, arid 


the Sonoran Desert in Pima County, Arizona (e.g., 
Santa Catalina Mts.) and north- 

1,100 

occupies the northernmost part of the distribution 


Baboquivari Mts., 


ern Sonora at This subspecies 


of the species. 


A specimens examined. U.S.A. pape 
Pima € icson, Robles Pass, Bartram 409 (US); 
Coyote Monts. Goodding 24-57 (CAS); iiir Se 
yon, Baboquivari Mountains, Goodding 247-45 (NY); 
Baboquivari Canyon, n Mountains, Gould & 
Haskell 3253b (CAS, F, GH, MO, NY); Catalina Moun- 
tains, Sabino Canyon, Jones s.n. (C AS, F, , MO, NY, 
UC, US); Toro Canyon, Baboqu uivari Monte Kearney 
& Peebles 10397 (CAS, MICH); South Canyon 
& Peebles 14960 (CAS, GH, NY. US); 5 ay 
Federal Prison C a Headquarte E Santa Catalina Mts., 
Parker et al. 5861 (GH, NY, MEXICO: A: 
Sierra Jojoba, ee ntry & W D 25 (A, US 
western Mts., Pringle 272 (US); 17.7 mi. SE of 
dalena, Starr & Birg p cM Canón de Agua 
Amarga, If hite 3609 (MICH?, GH). 


o 
= 


Zapoteca formosa subsp. schottii is closely re- 
lated to and was probably derived from subsp. rosei. 
The former has generally smaller stature, less nu- 
merous, smaller leaflets, and smaller pods (Table 

These two taxa show geographically contig- 
uous distributions (Fig. 44), and there is clinal 
variation in leaflet size, with subsp. schottii oc- 
Although the pat- 
terns of morphological variation are apparently 


cupying the extreme position. 


continuous between the two taxa, subsp. schottii 
is relatively easily recognizable as a separate taxo- 
nomic unit with a somewhat arbitrary boundary. 
In the Baboquivari Mountains and in northern So- 
nora and Chihuahua, a number of difficultly placed 
intermediate specimens have been collected. 
Zapoteca formosa subsp. schottii could be con- 
fused with subsp. mollicula, since they have leaflets 
of comparable size. However, subsp. schottii can 
be readily distinguished by the longer internodes 
and more elongate petioles and rachillae (Table 
4). Further, although vestiture is highly variable 
in subsp. mollicula, most of its individuals have 
denser vestiture than do plants of subsp. schottii. 


13. Zapoteca scutellifera (Benth.) H. Hern., 
comb. nov. Calliandra scutellifera Benth., J. 
Bot. (Hooker) 2: 139. 1840; Benth., 
J. Bot. 3:98. 1844; es Trans. Linn. Soc. 
London 30: 542. 1875; Benth. in €. Martius, 
Fl. Bras. 15(2) nee 1876. Pug scu- 
tellifera (Benth.) Kuntze, Revis. Gen. Pl. 
189]. TYPE: Territory of tpe t 
s Ribeirao, in siccis arenosis, May 
28, Riedel 1334 Ma K, here des- 
A, K, LE (3), NY 


London 


Brazil. 


Mina isolectotypes, 


N 


fragm., P, US). 


Scandent shrubs; branchlets terete or suban- 


gulate, the surface costate under magnification, 
glabrous to finely puberulent, usually villous on the 


younger parts. Leaves glandular, the pinnae 2-3- 


Volume 76, Number 3 


Hernández 853 


Zapoteca 


jugate; petioles terete with a conspicuous adaxial 
channel, 3.5-8 cm long, glabrous to sparsely vil- 
lous, villous in the area of the channel; rachis 1— 
5 em long: rachillae 5-7 cm long; glands close to 
the base of the petioles, at the points of insertion 
of the pinnae, and sometimes between the distal 
pairs of leaflets, scutelliform, the larger ones on 
the petioles to 2 mm diam., glabrous to puberulent 
abaxially; leaflets 6-11 pairs per pinna, oblong- 
obovate, falcate or subfalcate, oblique at the base, 
rounded to subacute at the apex, mucronate, the 
median ones 15-20 x 5-7 mm, the proximal ones 
smaller. membranous, glabrous, sometimes ciliate; 
primary and secondary leaflet veins conspicuous 
abaxially; stipules leafy, linear-triangular, to 4 mm 
long, glabrous to villous. Capitula axillary; pedun- 
cles solitary or fasciculate at distant nodes, glabrous 
to sparsely villous, ca. 6.5 cm long at anthesis. 
Flowers pentamerous; calyx cup- shaped, 1-1.5 mm 
long, finely ciliate, the teeth triangular, acute; co- 
rolla campanulate, 4-5 mm long, glabrous, the 
lobes lanceolate; filaments ca. 2.5 cm long, white 
in the basal half, pink at the distal half, the staminal 
tube ca. 1 mm long; ovary ca. 1 mm long, glabrous. 
Pods acute, 9 x 0.7 thickly 
membranous, finely puberulent, especially at the 


rostrate, ca. cm, 
margins. 

Distribution (Fig. 44). Endemic to the Am- 
azon Basin where it has been collected at scattered 
localities. In Brazil known from the states of Ama- 
zonas and Pará, and from the territory of Rondónia, 
also in Bolivia from the departments of El Beni 
and Santa Cruz. It usually occurs in open sandy 
sites. 

Additional Qi examined. BRAZIL: AMAZONAS: 
Amazonum ad o ium flum. Sra Spruce 1586 (BM, 
G, : Mun. de Itaituba, estrada 
Benterómi- Cuab. BR pes Pon 1011 com uma pun 


, GH, NY, 


tracao de 3 km dentro da mata, margem direita do. Rio 
Jamaxim sul do Pará, 70940'8, 55215 W, Amaral et al. 
1231 (MO, NY). e EL BENE prov. Vaca Diez, 


Guayaramerin, 10?40'5, 65%22'W, Krapov ickas & 
5c hinini 35129 uo. SANTA CRUZ: prov. Sará, vd 
ta, 450 m, Steinbach 7047 (A, BM, F, G, GH, K, 

MO) 


This morphologically uniform species is closely 
related to Z. formosa but differs by having scu- 
telliform extrafloral glands, scandent habit, and 
usually smaller, linear-triangular stipules. More- 
over, it can be distinguished from Z. formosa subsp. 
formosa by its white/pink, rather than white, fil- 
aments. The scutelliform glands, which are con- 
stant throughout the known populations of Z. scu- 
tellifera, are an excellent diagnostic feature, as 
they occur only in this species of Zapoteca. They 


are fully developed at the earliest stages of leaf 
development, which probably reflects their defen- 
sive function. 

14. Zapoteca tehuana H. Hern., sp. nov. TYPE: 
cultivated at MO, Jan. 1, 1986, H. Hernán- 
dez 950 (holotype, MEXL; MO). 


(Original source: Mexico. Oaxaca: Distr. Te- 


isotype, 


km SW de la entrada a Buenos Aires, 

m, R. Torres 1167 (MEXU)). Additional iso- 

types to be distributed. Figure 45. 

Frutices, on gracilibus. Pinnae (1—-)2(-3)-jugae, 
foliolis 10-21-]ugis in utraque pons oblongo-lanceolatis 
vel ovato- Duos membranaceis sub-chartaceis, 
ordinatione venarum evidenter biochido droma; stipulae 
foliosae ca. 2 mm longae. Numerus chromosomatum so- 


maticus 2n 


Shrubs ca. ] 
10 mm diam. at base, the bark smooth except for 
small exfoliations; branchlets terete, fissured under 
magnification, glabrous. Pinnae (1-)2(-3)-jugate. 
terete, with a conspicuous 


1.8 m high; stems slender, ca. 5- 


Petioles eglandular, 
).5-3 em long, glabrous, some- 
1.8 em long in 


adaxial channel, 
times sparsely villous; rachis 0.7 
2-jugate pinnae, to 3.8 em long in 3-jugate pinnae; 
rachillae to 7 cm long; leaflets 10-21 pairs per 
pinna, oblong-lanceolate to ovate-lanceolate, the 
distal pairs sometimes obovate, at base oblique, at 
apex acute to subacute, sometimes mucronulate, 
the median ones 9-15 2-4 mm, the proximal 
and distal pairs smaller, the anterior proximal leaf- 
let reduced in size, rarely lacking, all membranous 
to thinly chartaceous, usually drying light green, 
usually glabrous throughout, occasionally pilose and 
ciliate, with conspicuous brochidodromous vena- 
tion; stipules leafy, triangular, ca. 2 mm long. 
glabrous, rarely pilose. Capitula axillary; peduncles 
fasciculate at distant nodes, filiform, (2-)7- 10 em 
long at anthesis. Flowers pentame rous, or rarely 
tetramerous; calyx ca. 2-2.5 mm long, glabrous 
or pilose, usually ciliate, each tooth with a central 
vein, triangular, ca. 1 mm long; corolla campan- 
ulate, ca. 5-5.5 mm long, glabrous, the lobes ovate- 
lanceolate, acute; filaments (20-)24-25 mm long, 
white in the basal half and pink in the distal half 
or white throughout, the staminal tube ca. 2 mm 
long; ovary ca. 1 5 mm long, very shortly (< 0.3 
mm) stipitate. Pods linear, at apex subacute, ros- 
tellate, to 12 cm long when mature, thickly mem- 
branous, glabrous, reticulate-veined. Seeds widely 
ovoid to widely rhomboid, flattened, 5-6 X 4 mm, 
gray, the areola light gray: pleurogram irregular, 
demarcated by a dark line. Somatic chromosome 
number 2n = 26. 


Annals of the 


Missouri Botanical Garden 


854 
N A | 177 
VAN CS 
Nee s 
MT IKK 
ESE. l MN 
S y^ = 
FIGURE 45. Zapoteca tehuana. 
pods. —C. Leaflet.—D. Detail of venation in cleared le: 
Polyad. — I. Seeds. (Drawn from living plant, H. Hernánc 


Distribution (Fig. 34). This species is endemic 
to a restricted area southwest of the Isthmus of 
Tehuantepec in Oaxaca, Mexico, where it grows 
in secondary vegetation derived from tropical de- 
ciduous forest mixed with probably relictual ele- 


ments of temperate deciduous forests. 


A. Branchlet with inflorescences at anthesis. — B. Branchlet with dehisc 


aflet. — E. 
ez 950 (MEX 


a i / Iii 
/ A 

LC 
MESS 
MS SU D 


ed 


lower.—F. Anther.—G. Dissected flower. — H. 
KU, MO); pods from R. Torres 4167 (MEXU).) 


Additional specimens examined. MEXICO. OAXACA: 
Distr. Tehuantepec, canada en el Arroyo de las Minas 2 
km al SW de El Limon y 14 km al NW de Tehuantepec, 
Garcia & Torres 1679 (MEXU, MO); Distr. Tehuante- 
pec, El Limon, arriba de El Chicozapote, SW de Te- 
huantepec, entrando por camino a Buenos A H. Her 


nández & Torres 303 (MEXU, MO); Distr. Tehuantepec, 


Volume 76, Number 3 
1989 


Hernández 855 


Zapoteca 


El Manguito, al SW de El Limón el cual está a 11.1 km 
al SW de la entrada a Buenos Aires, Torres 4167 (MEXU, 
MO); Distr. Tehuantepec, 15 km SW de Buenos = 
Hierba Santa, Torres & 
Martinez 7375 (MEXU); subida al Cerro da: por 
ladera S, L. Torres et al. 708 (MEXU). 


Zapoteca tehuana is a narrowly endemic species 
probably not closely related to species of subg. 
Zapoteca, and further knowledge on this species 
could provide evidence for its placement in a sep- 
arate subgenus. It differs from the remaining species 
of subg. Zapoteca primarily by having conspicuous 
brochidodromous leaflet venation and relatively 


thick leaflets. It might be confused with some of 


the species with oblong or ovate-lanceolate leaflets 
of comparable size (e.g.. Z. portoricensis and Z. 
lambertiana) but differs by having leaves usually 
having fewer pairs of pinnae, smaller stipules, white 
or white/pink filaments, and shorter stature. The 
relationship between Z. tehuana and Z. aculeata 
are discussed under the latter. 


IH. Zapoteca subg. Aculeata H. Hern., subg. 


nov. TYPE: Zapoteca aculeata (Spruce ex 
nth.) H. Hern 
l-, raro 2-jugae, folia sine nectariis, glabra v 


lacels, 
spicua; stipulae spinescentes. Capitula selbe. Pollinia 
tumoribus lenticularibus praedita. Legumina coriacea gla- 


Leaves glabrous or nearly so, the pinnae l-, 
rarely 2-jugate, the petiole without nectaries, leaf- 
lets 3-7 pairs per pinna, moderately large, char- 
taceous to coriaceous, with a very conspicuous 
venation; stipules spinescent. Capitula axillary. Po- 
lyads with lens-shaped thickenings. Pods coria- 
ceous, glabrous. 


This subgenus comprises a single species, Z. 


aculeata, characterized by medium size, charta- 


ceous to coriaceous leaflets, and spinescent stipules. 


15. Zapoteca aculeata (Spruce ex Benth.) H. 
Hern., comb. nov. Calliandra aculeata Spruce 
ex Benth., London 30: 541. 
1875. TYPE: Ecuador, ad pedes m. Tungura- 
hua, Sep. 1857, Spruce 5054 (lectotype, K, 
here designated; isolectotypes, BM, F, G, (2), 
GH, K GL LE NY, P, S, OE The ue 
totypes are dated as follows: BM, 1857-9; F, 
1857-9; G, 1857-9, 1861; GH, 1857-9; 
K, Aug. 1857; LE, 1857-9; NY, 1657-9; 
P, 1857-9, 1861; S, 1857-9. Figure 46. 


, Trans. Linn. Soc. 


Trees to ca. 10 m tall; branchlets terete, costate, 
glabrous. Pinnae | -, rarely 2-jugate; petioles eglan- 
dular, terete, with a conspicuous adaxial channel, 
4-25 mm long, villous, rarely glabrate; rachis 3 
5 cm long when pinnae 2-jugate; rachillae 112.5 
cm long; leaflets (3)4—-6(- 7) pairs per pinna, the 
proximal anterior one usually reduced or lacking, 
all ovate-lanceolate, at base ru at apex acute 
0.4-2.3 


cm, the proximal ones smaller, all Pb to 


to acuminate, the distal ones 1.5-0.: 


coriaceous, glabrous, the base sometimes villous: 
leaflet venation conspicuous; stipules spinescent, 
rigid, adpressed to the branchlets, sometimes be- 
linear-lanceolate, sometimes 


coming divaricate, 


conical, beaked, 2-10 mm long, glabrous. Capitula 
axillary; peduncles usually fasciculate at distant 
nodes, rarely solitary, (2.5-)5-9 cm long at an- 
hesis. Calyx shortly. sometimes irregularly den- 
tate, ca. 2.5 mm long, glabrous, the teeth some- 
times more than 5, shallowly triangular to deltate; 


corolla campanulate, 5 


6 mm long, glabrous, the 
lobes ovate-lanceolate, acute; filaments ca. 2.5 em 
long, red-purple, the staminal tube ca. 2 mm long; 
ovary ca. 1-1.5 mm long, glabrous. Pods about 
10-12.5 x 0.9-1. 


apex, sometimes rostellate, coriaceous, glabrous, 


| em, subacute to rounded at 
the surface reticulate-veined. 


Distribution (Fig. 41). 
been collected from natural populations at the foot 


Zapoteca aculeata has 


of the Mount Tungurahua in the Ecuadorian Andes 
(Province of Ambato) and from cultivated individ- 
uals at public gardens in Ambato, Banos, and Quito. 


Collections come from elevations of 1,900- 
2,850 m. 

M d specimens examined. ECUADOR. 
TUNGURAHUA: mountain ae to S of Banos, Penland & 


IS). CULTIVATED. Ect 
4costa Solís 13599 (F). TUNGURAHUA: 
9488, 9523 (F); Rose & Rose 


RM, 104 (F, 
CHINCHA: Quito, . 
Ambato, Acosta Solís 
22365 (GH, NY, US). 


Zapoteca aculeata represents by itself a sub- 
genus of Zaboteca. It is a poorly known, unmis- 
takable species characterized by spinescent sti- 
pules, medium size, and coriaceous, ovate-lanceolate 
leaflets, which are usually larger than the essen- 
tially membranous leaflets of subg. Zapoteca. It is 
not obviously closely related to any other species 
of the genus; however, as shown in the chapter on 
morphology, an analysis of its leaflet architecture 
revealed outstanding similarities to that of Z. 
huana. Zapoteca aculeata is known from only a 
few collections made in a restricted area of the 
Ecuadorian Andes and from cultivated individuals 


856 


Annals of the 
Missouri Botanical Garden 


FIGURE 46. — Zapoteca a 
C. Detail of venation in cleared lea 


— D. Spinescent sti 


- A. Branchlet with infloresc ences at anthesis, and dehisced pods. — 
ipule 


-B. Lea 
. Flower. (From Rose & Rose 22365 (US 


Sy leaflet 


s. 


from Acosta 13599 (MO); pods d Penland & Summers 104 (U S).) 


at Ambato, Banos, and Quito; it has not been col- 


lected for more than 40 years 


IV. Zapoteca subg. Amazonica H. Hern., subg. 


TYEE: 


ern. 


nov. 


Zapoteca amazonica (Benth.) H. 


lia sine nectariis, glabra vel fere glabra, ise uni- 


vel compositis disposita. Pollinia tumoribus lenticula ribus 
praedita. Legumina coriacea glabra. Semina sine pleu 


rogramiate. 


Volume 76, Number 3 
1989 


Hernández 
Zapoteca 


80 
10 ———— 
te y 
i 
as 
o 
o 
o 
. 
"mr, 
E 
Y 
ol a --—> 
` 
3 
K 
v 
A 
` 
X 
Ñ 
C 
E 
, 
- í 
Pd NL Y 
‘ 
. , = 
ey jue 
t v z 
e ! ' 
[d 1 
N 
o Cd 
e 
- 7e 
I. ER 
o i Nm E 
N 
E v^ A 
l e. ee E Teg 
^ edi a mI 
D UA 
T ys 
> 
t le 
` 
lo) X 
í 
j 
i 
p 
& 
o 
600 Km i 
ale s. o 9 


FIGURE 47. 


Distribution of Zapoteca aculeata ( 
(dots). 


Leaves without nectaries, 
the pinnae | -jugate: leaflets | 


11% pairs per pinna, 
chartaceous to coriaceous; 


leaflet. venation con- 
spicuous; stipules leafy. Capitula in long, simple or 
compound pseudopanicles. Polyads with lens-shaped 
thickenings. Pods coriaceous, glabrous. Seeds with- 
out pleurogram. 


This subgenus comprises two relatively special- 
ized species, Z. microcephala and Z. amazonica, 
which are e los terized by a set of characteristic: 
that are highly advanced in Zapoteca: leaves re- 
duced to one pair of pinnae, with 


=— 


1/2 pairs of 
large, chartaceous to coriaceous leaflets; capitula 
organized in long. simple or compound pseudopan- 
icles. 


glabrous or nearly so, 


triangles), Z. microcephala (open square), and Z. amazonica 


16. Zapoteca microcephala (Britton & Killip) 
. Hern., comb. nov. Calliandra microceph- 
ala Britton & Killip, Ann. New York Acad. 
Sei. 35: 134. 1936. TYPE: Colombia: 1760 
1808, Mutis 107 (holotype, US no. 1561096; 
isotypes, MA ( hotos at MENU, 
MO). 
Shrubs: 
glabrous. 


1) not seen; | 


branches terete, the surface smooth, 


Pinnae l-jugate; petioles eglandular, 
slender, with a pair of channels running longitu- 
dinally (30-)40-70 mm 
long, glabrous abaxially, usually scabrous between 
the channels; rachillae 13-19 mm long: leaflets 
1% pairs per pinna, the proximal anterior leaflet 
acking, all falcately lanceolate, at base strongly 


in the adaxial surface, 


857 


858 Annals of the 
Missouri Botanical Garden 
TABLE 15. Comparison of Zapoteca microcephala with Z. amazonica. 
Z. amazonica 


Z. microcephala 


Amazonian Peru 


Distribution Mense] River Valley, Colombia 
Habit erect shrubs scandent shrubs 
Pairs of leaflets per pinna 143 l 
Petiole length (30-)40-70 mm 20-42 mm 
Leaflet pubescence ciliate glabrous 
Pseudopanicles 
‘orm simple compound 
Length 20-30 cm up to 40 em 
Stipule length 1-2 mm /-22 mn 
Filament color white white /red-purple 
oblique, sharply long-acuminate at apex, the distal J. Bot. 5: 101. 1846; Benth., Trans. Linn. 
leaflets 7.5-10.5 x 2.5-4.5 cm, the proximal one Soc. London 30: 538. 1875; J. F. Macbr., 
smaller, all chartaceous, glabrous, usually ciliate; Fieldiana, Bot. 13: 67. 1943. Inga bauranik 
leaflet venation conspicuous, with 2 primary and folia “bauhiniaefolia” Poeppig, Nov. Gen. 
at least | secondary veins prominent; stipules leafy, Sp. Pl. 3: 80, t. 290. 1845. (The epithet is 
triangular to narrowly triangular, ca. 1-2 mm long, presumably also based on Poeppig 2070 used 
glabrous. Capitula in terminal, elongate, simple by Bentham in the description of Calliandra 
pseudopanicles, ca. 20-30 cm long; peduncles fas- amazonica.) TYPE of Z. amazonica: Peru. 
ciculate, rarely solitary, filiform, ca. 4.5 cm long Loreto: in sylvis provinciae Maynas circum 
at anthesis. Calyx ca. 2 mm long, the teeth tri- Yurimaguas, 1834, Poeppig 2070 (lectotype, 
angular, < 1 mm long; corolla campanulate, ca. G, here designated; isolectotypes, F, G, LE 
3-4 mm long, the lobes ovate-lanceolate; stamens pro parte, NY, P). The specimens at G and 
white, the staminal tube ca. 2 mm long. Pods to P are dated 1834, whereas those at F and 
15 x 1.1 em when mature, at apex acute or retuse, NY are dated 1831. Figure 48. 
coriaceous, glabrous, the valves reticulate-veined. 
Distribution (Fig. 47). Endemic to a restricted Scandent shrubs 3~5(~8) m tall; stems slender, 
ca. 15 mm diam., the bark smooth; branchlets 


area of the Median Valley of the Magdalena River 
in Colombia. 


Additional specimens examined. COLOMBIA. 
SANTANDER: vicinity of Puerto Berrio, between Cararé and 
aught 1/11 (F, K, US), Haught 

Mutis 3800 (MA, US) 


Bises rivers 


1 (US); m loc ality, 


T 


This narrowly endemic species is clearly allied 
to Z. amazonica, from which it is geographically 
isolated (Fig. 47). Zapoteca microcephala can be 
readily distinguished from that species by 1!2 pairs 
of smaller leaflets per pinna, simple (nonramified), 
shorter pseudopanicles, smaller stipules, and white 
filaments. Table 15 provides an account of the 
main comparative characters distinguishing these 
closely related species. Zapoteca microcephala is 
known from only four collections, 2 has not been 
collected since Haught (1711, 1734) found it in 
1935 in the floristically rich area of the Magdalena 
River Valley, Colombia. 


17. Zapoteca amazonica (Benth.) H. Hern., 
Calliandra amazonica Benth., 


3: 94. 1844; Benth., London 


comb. nov. 


London J. Bot. 


spreading, terete, glabrous. Pinnae l-jugate; pet- 
ioles eglandular, 20-42 mm long, usually glabrous; 

hilla + corrugate, with a conspicuous adaxia 
channel, 10-21 mm long (excluding the pulvini), 
usually glabrous, but occasionally sparsely villous 
in the extremes; leaflets | pair per pinna, falcately 
lanceolate-ovate, 7-22 x 2.8-9.2 cm, strongly 
oblique basally, acuminate to caudate, chartaceous 
to coriaceous, usually drying lighter green abaxi- 
+ lustrous adaxially, glabrous or rarely gla- 
primary 


ally, 
brate; aia venation conspicuous, with 2 
and at least | secondary vein prominent; stipules 
leafy, ovate-lanceolate, acuminate to acute at apex, 
7-22 x 3-8 mm, glabrous. Capitula arranged in 
terminal, elongate, compound pseudopanicles to 40 
cm long or more, the peduncles fasciculate, rarely 
solitary, 0.8-7 cm long at anthesis. Calyx dentate, 
ca. 2.2 mm long, the teeth shallowly triangular or 
deltate, ca. 0.7 mm long, glabrous, rarely ciliate; 
corolla campanulate, ca. 4-5 mm long, glabrous, 
the lobes lanceolate; filaments white in the basal 
half and purple-red in the distal half, the tube ca. 
2.5 mm long: ovary about 2 mm long, glabrous. 


Volume 76, Number 3 
989 


Hernández 859 


Zapoteca 


48. Zapote ca amazonica. 
1 ) 


URF 
ps bar Leaflet. — D. 


Pods to 24 x | cm when mature, acute at apex, 
coriaceous, glabrous, the valves reticulate-veined. 
Seeds narrowly rhomboid to narrowly ellipsoid, flat- 
tened, ca. 14 x 5 mm, dark brown, without pleu- 
rogram. 


. Branchlet with compound pseudo] panic le. B. 


E : Detail v. venation in a 
Plow man & = hunke 11445 (E) and Schunke 4497 (G), stem from Schunke 3371 (US); seed from Se habe 4737 
(US).) 


Primary pon showing 


cleared leaflet. — E. Flower. F. Pod.—G. Seed. (From 


Distribution (Fig. 47). This species has been 
collected primarily in shady sites of undisturbed 
tropical lowland forests of Amazonian Peru, from 
Rio Alto Madre de Dios to Coronel Portillo (Rio 
Pastaza) in the Departamento of Loreto, especially 


860 


Annals of the 
Missouri Botanical Garden 


Collections come from 350 


along Rio Huallaga. 
900 m 


Additional specimens examined. PERU. JUNÍN: Sani- 
beni ad Satipo, Woytkowski 5943 (F, GH, LL, MO, NY, 
US). HUÁNUCO: above Río on at Tingo Maria, Croat 
21102 (MO); Tingo Maria, km 138, trocha lote del Agui- 
la, Ferreyra 932 (l d achitea, Puert to Inca, just N of 
isnt 6832 2 (MO); jen) 

‘odo de Pozuzo (9°40'S, "75°28 , Foster 9209 (M 

uallaga, near shee € hontalagua, Mexia 

8295 (BM, F, G GH, MO, NY, S, UC, US); 
Poa Maria (9°18'S, 75°59'W), 

9 (F, US); Tingo Maria, VÉ & 
UC i LORETO: Coronel Portillo, 4-5 
horas arriba dela Abujao, sobre al Rio HR 
Díaz 861(MO, NY). 4 MADRE DE DIOs: Pantiacolla, serranía 
across Rio Alto Madre de Dios from Shintuya, yet 
al. 27293 (MO). SAN MARTÍN: Ma al Cáceres, Tocache 
Nuevo, Cerro Sinsin, 15 km Tocache Moan along 
dies i Puerto Pizana (8°09' E p 34'W), du man & 
= hunke 11445 (F); Mariscal Caceres, Depto. 
2 km de Toc “ac le Pew Fundo la in ea se hunke 
3371 (F, G, NY, US); iscal Cacere Jepto. Tocache 
Nuevo, € Ante. margen izquierdo Rio Huallaga, Se hunke 
1497 (F, G, NY, US); áceres, Dep 'ocache 
Nuevo, Malpaso de Balsa, Probanas l e is desem- 
bocadura Rio Mishollo, Schunke 4737 (F, GH, MO, US); 
Mariscal Cáceres, Depto. Tocache Nuevo, Quebrada de 
Santiago, "i dud Pizana, Schunke 0483 (MO, NY); 
Mariscal € s, Puerto Pizana (margen derecho del Rio 
oon Schunke 6913 (MO); Mariscal Cáceres, Depto 
Tocache Nuevo, Quebrada de Huaquisha (margen Rio 
eR ee pe 13 (MO, NY); Tarapoto, Spruce 
449 1 (BM, BR, GH, K, P, NY); Alto Rio Huallaga, 
W mx 6. 518 (F, US). 


= 


Tarapoto, 

Zapoteca amazonica and Z. microcephala rep- 
resent a distinct evolutionary line within the genus; 
these closely related species probably differentiated 
from a common ancestor after becoming separated 
from the rest of the genus. Zapoteca amazonica, 
which is remarkably uniform morphologically, can 
be readily distinguished from Z. microcephala by 
its scandent habit, one pair of larger leaflets per 
pinna, larger stipules, elongate pseudopanicles, and 
The 


tween the two species were discussed further under 


white/purple-red filaments. differences be- 
Z. microcephala (see also Table 15). 

Zapoteca amazonica probably is the most spe- 
cialized species in the genus, presenting a combi- 
nation of derived characters: scandent habit; ex- 
treme reduction of leaflet number concomitant with 
an increase in size; relatively complex leaflet ve- 
nation patterns; capitula organized in complex, ter- 
minal, compound pseudopanicles. The specialized 
nature of this species was discussed in the section 
on morphology. 


LITERATURE CITED 


AB'SABER, A. N. 1982. The paleoclimate and paleo- 
ecology of Brazilian Amazonia. Pp. 41-59 in G 


Prance (editor), oe al Diversification in the Trop- 
bia A. ress, New York 

Quaternary palynologic 'al studies in 
. Prance (editor), 


ics. Columbi 
1982 


the um Bur . 61-1 


ad dori in the Tropics. Columbia 
ress, New i 

ALEXANDER, M. P. 1 969. Diff f aborted 

A4: 117-122. 


and nonaborted n Stain Technol. : 
Arroyo, M. T. K. 81. Breeding systems and polli- 
nation biology in ep Pp. 723-769 in R. 
M. Polhill « aven (editors), (Fio in 
L egume Systematics, Part 2. Royal Botanic Gardens, 


Ke aplan 
Banker, H. G. & I. BAKER. 1983. Floral nectar ta 
PENS in relation to pollinator type. 
E. Jones & R. J. Little (editors) Handbo ok 
a ae Biology. Van Nostrand 
Reinhold, New Yor 
B HARRIS. 1957. 
a by bats and its attendant evolutionary prob- 
lems. E 11: 449-460. 
Bawa, K. S 974. Bre 


a » Tue tropical community. Evolution 28: 85 


The pollination of 


ding systems of tree species 


pai 


be G: 840. Contributions towards a flora of 
South A enumeration of plants collected by 
Mr. , Schomburek in British Guiana. Hook. J. Bot. 2: 
12 

me Notes in Mimoseae, vith a synopsis of 
he Species. London J. Bot. 3: 82 2 

187 evision of the suborder Mimoseae. 
dans. Linn: Soc. London 30: 335-004 

BrvrLEY, B. 1977. Extrafloral nectaries and uq 
by Loa aida, Ann. Rev. Ecol. 
40 i. 

BERNHARDT, | p. 1983. Insect pollination of Australian 
4cacia. Pp. 85-97 in E. Williams, R. B. Knox, J 
H. Gilbert & P. Bernhardt (editors), Pollination *82. 
University Press, Melbourne, Australia. 

Kenrick & R. B. Kxox. 1984. Pollination 
biology and the breeding system of Acacia retinoides 

Leguminosae: Mimosoideae). Ann. Missouri Bot. 

Gard. 1-29. 

BERTIN, R. 4 -1982. The a p maintenance of 
andromonoecy. yo y 6: 32 

BIGARELLA & puces SR 1982. Paleoen- 
rece i anges in Brazil. Pp. 27-40 in G. 
Prance (editor), Biological Diversification in the Trop- 
ics. Columbia Univ. Press, New York. 

BRANTJES, N. 1978. Sensory M to flowers in 
night-flying moths. Pp. 13- in A Richards 
(editor), The Pollination of NM by Insects. Linn. 
Soc. Symp. Ser. 6. 

BREEDLOVE, D. E. 1981. Introduction to the Flora of 
Chiapas. eg Academy of Sciences, San Fran- 


~ 


cisco, Galilor 
Brrrrox, N. L. " . Rose. 1928. Anneslia. N. 
Amer. H. 23: Pe í 
BUNTING, G. S. 1967. 


n Conner type of Callian- 

dra. Taxon 16: 469 472. 

Bi uper, H. M. 1979. sia ad botanicorum gra- 
ise m. Conservatoire et Jardin Botani oua Ger 

CANDOLLE, DE. 1825. Pp. 61-122 ; De p 

germination des Le ene Mémoires sur rla fam- 

ille des Légumineus 


CARDENAS, L. 1974 bs géneros venezolanos de las 


Volume 76, Number 3 


Hernández 
Zapoteca 


Mimosoideae (Leguminosae). Revista Fac. Agron. 
(Maracay) 7: 109-194. 
COMPTON, R. 1912 
structure in vilis Leguminosae. J. Linn. Soc., 
1-122 
CnEPET, W. LAD L. DILCHER. 1977. Investigations 
of angiosperms from the Eocene of North America: 
714- 


Amer. J. Bot. 64: 


c 


An investigation of the see ane 


Bot. 


a oe inflorescence. 


CRUDEN "n W. 1977. à conser- 


vative indicator RÀ } 
E volution 31: 
& S. M. 


Pollen-ovule ratios: 
j; : q 


ring plants. 
46. 
> RMANN. MU 
Some observation: on the Pp. | B. 
: T editors), "The clo el ME 
ies. Columbia Univ. ress, New Yor 
& S. MiILER-Wakb. 1981. Plus ovule ratio, 
pollen size, and the ratio of stigm ! 
pollen-bearing area i = pollinator: an hypothesis. 
Evolution 35: 964 
————, SH. KINSMAN STOCKHOUSE, II & Y. 
LIN T. 1976. y, and a a 
wA al moth-flowered plants. Biotropica 8: 
210. 
D'Arcy, W. G. 1971 
their typification. 7 axon 19: 554-500. 
DAGHLIAN, P, W Cum & T. DELEVORYAS. 
l€ ry angiosperms: a new 


Studying nec ctar? 
l 


atic area to the 


E E 
Pollination, fecundity 


eal names, some notes on 
1-560 


^- 
T 


Investigations 
flora a La a plumosa from the Oli- 
Amer. J. 67: 309 


e of eastern Texas. Amer. Bot. 
820. 
Ducuer, D. L. 1974. Approaches to the identification 
of anya porn leaf remains. Bot. Rev. 40: 97 
DUKE, eR. M. Porm. 1981. Sue of the 


Leguminosae. Pp. 941-949 in R. M. Polhill & P. 
H. ra en (editors), Advances in Legume Systematics, 


Part 2. Royal Botanic Gardens, Kew, Richmond, 
Su a 

iue R., A. Levy & D. Parevrireó. 1981. An- 
dromonoecy in Solanum marginatum. Bot. Gaz. 


€ 


Eom 


C rawfordsy ille) 142: 25€ 
s 1.5 Extrafloral nectaries: their structure 
and b Pp. 174-203 in B. Bentley € T. 
lias (editors), The Biology of Nectaries. Columbia 
Univ. Press, New York 
Farkcri, K. & L. VAN DER Pur. 197 
of Pollination Biology, 3rd ub. "Pe rgamon Press, 
Toronto. 
Forp, H. A. € N. Fonpr. 1976 
pollinators of Acacia aia Austral. J. Bot. 
i 193 195. 
FORERO. E. 84. Revision of Calliandra: 
apris approach. Bull. Groupe Int. Étude Mimos. 
12: 14-15 
GENTRY, A. 1974 
in M al eo eae. Bi iotropica 
stribution patterns of neotropical Big- 
nonlaceae: some phytogeopraphic implications. Pp. 
339 n K. Lar & L. Holm-Nielsen (editors), 
ropes gom ne se mic Press, London. 

1982. Phytogeographic patterns as evidence 
for a Chocó refuge. Pp. 112-136 in G. Prance 
(editor), Du Div Sie ME in the Tropics. Co- 
lumbia Univ. Press, New Yo 

GERTSCH, W. 1979. pum erican ides 2nd edition Van 
Nostrand Reinhold, New 
GOLDBLATT, P. 19 C ee UN the phylogeny of 


The Principles 
Birds as possible 
Bot. 24: 

a multidis- 


Flowering phenology and diversity 
58 


= 
wal 
> 


Leguminosae. Pp. 427-463 in R. M. Polhill & P. 


aven rela Advances in Legume Systematics, 


Pan 2. Roval Botanic Gardens, Kew, Richmond, 
Surrey, England. 
& A. GENTRY. 197 9. Cytology of Bignoni- 
aceae. Bot. N 132: 475-482 
Guiner, PH. 1965 tude das 's caractéres du pollen ids 


. Pollen & Spore 


— 


le genre Calliandra (Mimosaceae 
7 $ : 157 í ie n 

"les Mimosacées. Etude de palynologie 
ene il c orrelations, & ue ~ Sect. 

.I ondichéry 9: 

1981. Mid D kie acters of the 
pollen ies . Pp. 83 ¿ R. M. Polhill & P. 
n Ra AA Adv ances in f egume Systematics, 
Fart 2 oyal Gardens, Kew, Richmond, 
Surrey, Engla id. 

& ( babe 1969. 


(Mimosa ) rvée en mic 


otanic 


L'exine des Calliandra 


‘roscopie phorobiqus et 
en microscopie E 'ctronique. Pollen & Spores 9: 211 
221 

SAL, C. Evans & B. Mastin. 1980. 
'omposition and affinities of 


J. Linn. Soc., Bot. 


Acacia (Mimosoideae): 
the series Pulchellae Ronin. 
. s 


¿AE and seeds of E nera in the 
. Department 


y (1 abaceae “a 
of tret Tech. Bull. 
HABER, & G. FRANKIE. C Tbe acteristics and organi- 
zation i a -— al hawkmoth community. Biotropica 
(in press). 
Hare : R, J. 1982. 
Pp. 6 24 in 
fication in the Tropies. Columbia Univ. 


General aspects of the refuge theory. 
G. Prance (editor), Biological Diversi- 
y 


ORs; New 


Cork. 
Hammen, T. 


VAN DER. 1982. Paleoecology of tropical 

South America. Pp. 60-66 in G. Prance (editor), 

ata nir had in the Tropies. Columbia 
. Press, New York 


Un 
Herrmaus, E., P. OPLER & H. BAKE R. 1974. Batactivity 
and pollination of Bauhinia uu pa polli- 
nator coevolution. Ecology 55: 412-419 
HERNÁNDEZ, 1984. ( eie Soria to m system- 
atics of Calanda with particular reference to its 
infrageneric relationships. Bull. Groupe Int. Etude 
Mimos. 12: 16-18. 
1986a. Zapoteca: a new genus of neotropi vical 
Mimosoideae. Ann. Missouri Bat. Gard. 73: 5 
(63. 
86b. Proposal to amend the conservation 
of Calliandra (Leguminosae: Mimosoideae). Taxon 
35: 747-74 
Hickry, T 
dicotyle donous leaves. Amer. J. Bot. 

1977. Stratigr "s and pleobetan of the 
ertiary) of Western 
Amer. p 0. 

bases of angiosperm 


973. Classification of the architecture of 
00: 17-33. 


Galde n V alley Formation (E [SR 
an Pal on ag Geol. 
& J. 1975. 
ee fonum morphology. Ann. Missouri Bot. 
Gard. 62: 538-589. 
Hopkins, H. p 1984 
ogy of the neotropical species of Pa . J. Ecol. 
y 1 f 
gar ARD, R- 1973, The 
1-38 in A. Graham (editor), 


al History of Northern Latin America. Elsevier, 
dam 


Floral biology and Wes ecol- 
f Parkt E 


> vegetation of the Antilles. Pp. 
Vegetation and. Vege- 


mster 


862 Annals of the 
Missouri Botanical Garden 
Janzen, D. 1967. Synchronization of sexual reproduc- Pottitt, R. M., P. H. Raven & C. STIRTON. 1981. 


tion of trees within es dry season in Central America. 
eg e e 620-637 
KENRICK, J. B. “Knox, 1982. 
polyad in pad of Acacia. Ann. Bot. 50: 
727 


Function of the 


MARGINSON, G. BERESFORD & R. B. KNOX. 


"^ Birds pa: pollination of Acacia te rminalis, 
Pp. 102-109 i Wiliams, R. B. Knox, J. H. 
Gilbert P. Bernhardt (editors), Tina '82. 


University Press, Melbourne, Aust 
Kevan, P. 1972. Floral colors in the Hig e Arcti 
reference to ue flower Tera and pollination. 


ic with 


Canad. J. 2289-231 
Knox, R. B. € Kenic K. mu Polyad function in 


relation to the breeding of Acacia. Pp. 411-418 in 
D. Mulcahy & E. Ottaviano (editors), Pollen Biology. 
Elsevier, North Holland Press, Amsterdam. 
Koprur, S. 1983. Flowering phenology and floral bi- 
ology of nem (Fabaceae: Mimosoideae). Syst. Bot. 
354-368. 
1984. Outerossing and pollinator limitation 
of fruit set: breeding systems of neotropica nga 
lor ui Mimosoide ^ae). Evolution 38: LI: 


114 

KUPICHA, F. K. 1977. The 
Vicieae (Leguminosae) and the r elat 

L. J. Linn. Soc., . Bot. 74: 131-16 

Lrrrsv run P. & N. Ramayya. 1982. Trichomes in 
relation to taxonomy: Mimosoideae. Geophytology 
12: 6-21. 

LEONARD, J. 1957. 
Amherstieae africaine 


de a of the 2 
ishi 


ups of Cice 


a des Cynometreae et des 
. Essai de blastogénie appli- 
da Acad. Roy. Sci. Bel- 


Gener 


J. F. TUN Caliandra. 

s. her Hist., Bot. l 

McVaucu, R. 197 Galeotti's d e in Mex 

ico: the number of its collections ce a brief itiner- 

. Michigan Herb. 11: 291-297. 

n Erias. 1970. n ud un pol- 

linia, and syste matic implications. Amer. J. Bot., Suppl. 
50: 


in d of Peru. 
3: 66-73. 


&- a 971. Calliandra haematoceph- 
ala: history: morphology, and taxonomy. J. Arnold 
Arbor. 52: 09- 


& Cu. Niet ODA. 1978. On the genus 
Sc hleinitz ia (Leguminosae: Mimosoideae). Adanso- 
nia, Ser. 2, 18: 345-363. 
NiEZGODA, CH. J., S. M. Fever & L. I NEVLING. 198 


Pollen ultrastructure e ia tribe Ingeae ES 
deae: Leguminosae). Amer. J. Bot. 70: 650-667. 
OPLER, P. A. 1983. Nectar produerion. v a tropical 
ecosystem Py 30-79 in !B. Bentley € T. Elias 
(editors) The Biology of Nectaries. Corumbia Univ. 
, New 


od and systematics of the Leguminosae. Pp. 
1-26 in R. M. Polhill & P. de ie ee (editors) 
Adva ances in ni sth Systematic l. Royal Bo- 
, Kew, Ric hmond: ud. England. 
Forest refuges: evidence from woody 
angiosperms. Pp. 137-157 in G. Prance (editor), 
Biological Diversific ation in the Tropics. Columbia 
Univ. ws New York 
& D. G. LLOYD. 1980. 
in the New Zealand montane shrub manuka, 
tospermum scoparium (Myrtaceae). Amer. J. Bot. 
67: 361-368 
Raven, P. H. € R. M. Porn. vie Biogeography 
of the Leguminosae. Pp. 27-34 in R. M. Polhill & 
P. H. Raven (editors), Advances in Legume System- 
Part 1. Royal Botanic Gardens, Kew, Rich- 
ar England. 
30. The Dispersal of Plants Throughout 
W or id. L. Reeve & Company, Kent. 
i. P. J. & H. P. VAN DER ScHIFF. 1971. The 
genus Acacia Miller in South Africa —5 (with special 
reference to the seedling structure as a taxonomic 
pedes EDU Mitt. Bot. Staatssamml. München 10: 


Andromonoecy 
ep- 


L . 

um P T. 1986. Distribution of nonprotein imino 

Par ae amino acids in Zapoteca. Ann. Missouri 
73: 764-767. 

Ruiz, b e M. T. K. Arroyo. 1978. Plant reproductive 
ecology of a secondary deciduous tropical forest in 
th n Biotropica 10: 221-230. 

Rzrpowskrn R. 1978. Vegetación de México. Limusa, 
México. 

Simpson, B. B., J. L. Nerr € A. R. MOLDENKE. 1977. 
Prosopis flowers as a resource. Pp. 84-107 in B. 
B. ja e (editor), Prosopis: lts Biology in Two 
m Ser BP Bund Series 
t, Nr os Pennsylvania 

SORSA, P. 1969. Pollen morphological studies in the 
Mimosaceae. Ann. Bot. Fenn. 1-34. 

STANDLEY, P. 109: Cacia in: Flora of Yuc atan. 
Publ. Field € olumbian Mus., Bot. . 

STEPHENSON, 19 Flower dnd al abortion: 
pn causes and tale functions. Ann. 

253-279. 


Ecosystems. US 


. A 
: 21 i 


L- 


Ecol. Syst. 12: 
VASSAL, I. "1969. LR à l'étude de la mor- 
phologie des peres d' Acacia. Bull. Soc. Hist. Nat. 

l 


Toulouse 108: 15. 
WickLER, W. 1968. Mimicry in Plants and Animals. 
World Univ. Library, New York. 
Woopson, R. . SCHERY. 1950. po 
Ann. Missouri Bot. Gard. 37: 


i Mie of Panama. 
266. 


POLLEN MORPHOLOGY OF 
THE MELANTHIACEAE 
AND ITS SYSTEMATIC 
IMPLICATIONS! 


Masamichi Takahashi? and 
Shoichi Kawano? 


ABSTRACT 


Pollen grains of several selected genera of Melanthiaceae were observed with scanning and transmission electron 


microscopes. Helonias and Ypsilandra 
and spinules, which agrees w 
pollen grains with clavate exine 
arly different from Tofieldia, ainc th is characte 


. Japonolirion, b us 


range of infrageneric variation in exine eub. patera -foveolate, reticulate, 


Pollen of Metanarthecium has distinctive reticulate e 


lumina 


ave the distinctive feature 
vell ii He aes Chionographis and Chamaelirium ees have distinctive four- -porate 


zed by disuleate grains with retic (s exines. 


. The palynological data indicate that the Nee consist of a 


e composed of an amorphous layer 
I orp 
e grains with gemmate exines, 
4letris shows a v 
ui bri mi 


Japan, has monosulca 
and dada t 
nee two genera are distinguished by t 
culpture characterized by verrucate pror 
t least three heterogenous 


ro wide among 


which Helonias, Heloniopsis, and Ypsilandra represent one group, and ina and Chancetaphis another 


distinct group 


The Melanthiaceae comprise ca. 24 genera dis- 
tributed in North America, Japan, and China (Dahl- 
gren & Clifford, 1982). These genera are Aletris, 
Amianthium, Chamaelirium, Chionographis, He- 
Metanarthecium, Narthe- 
Pleea, 
Ste- 


Toxicoscor- 


lonias. Helontopsts. 


Nietneria, Oceanoros, Petrosavia, 
Schoenocaulon (= Sabadilla), 


Tofieldia, 
) psilandra, Ae- 


cium, 
Protolirion, 
nanthella, Stenanthium, 
dion, Tracvanthus, Veratrum, 
rophyllum, and Zigadenus (Dahlgren & Clifford, 
1982). Krause (1930) recognized the tribe To- 
fieldieae (Liliaceae- Melanthioideae), in which To- 
fieldia (Temperate-Arctic Northern Hemisphere), 


Pleea (eastern North America), Varthecium 


— 


and 


— 
n 


(Northern Hemisphere Vietneria (Guinea 
were included. However. in his treatment, Aletris 
was placed in the different subfamily, Alectroideae. 
Hutchinson (1934, 1959) grouped these genera 
into two tribes, Heloniadeae and Narthecieae. The 
Heloniadeae, characterized by the absence of bracts, 
consist of Helonias. Chamaelirium, Y psilandra, 
and Chionographis. and the tribe Narthecieae, 
characterized by the presence of bracts, comprise 
Pleea, Tofieldia, Xerophyllum, He- 


loniopsis. Clara, Metanarthecium, 


Hewardia, 
Varthecium, 


Aletris, and Vietneria. Hutchinson (1934, 


1959) 
regarded the Heloniadeae as the most primitive 
tribe in the Liliaceae, and that the tribe is closely 
related to the Narthecieae. Dahlgren et al. (1985) 
placed these genera in the family Melanthiaceae 
Melan- 


Chionographi- 


and recognized six tribes, Petrosavieae, 


thieae, Narthecieae, Tofieldieae, 
deae, and Xerophylleae. 
in the tribe Narthecieae. 
Although many brief notes on pollen morphology 

of these genera using light microscopy have been 
given by Beug (1961). Erdtman (1952), Faegri et 
al. (1964), Huang (1970, 1972), Huynh (1976), 
(1956), Kuprianova (1948), Radurescu 
1973), Schultze (1978), Shimakura (1973), 
Thanikaimoni (1970), Ting-Su (1949), Utech 
979), Wunderlich (1936), and Van der Hammen 
(1956), a comprehensive pollen morphological study 
has not been made for these genera with electron 


They included Lophiola 


Ikuse 


Ss — 
~ 


microscopy 

Although Schultze (1978) classified Chiono- 
graphis, ) psilan- 
dra, and Helonias in the the 
different pollen types and exine structure suggest 
that the treatment is inappropriate. Recently, Tak- 


Chamaelirium, Heloniopsis, 


tribe Heloniadeae, 


express our gratitude to the directors and curators of the following herbaria for providing materials used in 


this study: U.S. 
are also grateful to George Roge 


H. Utech, Carnegie Museum of Natural 


rs, Missouri Botanica 


' Department of Botany, Faculty « 
Tohoku University, Sendai, 980 Japan. 


National Arboretum (NA), Kunming qu of ew (KUN 
arden, 
History, for ees iu comments on e study. 
Mid of Biologv, Faculty of Education, Kagawa University, Takamatsu, 760 Japa 

of Science, Kyoto University, Kyoto, 606 Japan, he p dus of Genetic Ecology, 


) and Tohoku University (TUS). We 
. Ambrose, University of Guelph, and Frederick 
1 


ANN. Missouni Bor. Ganp. 76: 863-876. 1989. 


864 


Annals of the 


Missouri Botanical Garden 


TaBLE l. List of voucher specimens and pollen sizes. 


Species 


Collection Data 


Pollen Sizes (um) 


Helonias bullata 

H. bullata 

Ypsilandra yunnanensis 
Chamaelirium luteum 
C. luteum 

C. luteum 

C. luteum 
Chionographis japonica 
C. japonica 

C. koidzumiana 

C. koidzumiana 


Tofieldia calyculata 
T. calvculata 


T. coccinea 
T. coccinea 
T. coccinea 
T. coccinea 
T. coccinea 
T. glabra 

T. glutinosa 
T. glutinosa 
T. intermedia 
T. japonica 


T. Japonica 
T. japonica 


T. nuda 
T. nuda 
T. occidentalis 


T. occidentalis 


U.S.A. Maryland: Anne Arundel Co., 27 Apr. 
1941, M. B. Was 


O. M. & G. G. Free 
China. es 19 Tune don. T. T. Yu 19036 
(KUN) 


U.S.A. New bo Bargin Co., 3 May 1928, 
. (NA 


U.S.A. South Carolina: etus Co., 1 May 
1981 Koyama et al. € A 
S.A. West Virginia: Mineral PA 19 June 
1938, O. M. Freeman s.n. (NA) 
U.S.A. Florida: Duval Co., Apr., 4. H. Curtiss 
2903 (NA 
U.S.A. Pennsylvania: Vernango Co., 19 June 


—~ 


1930, O. M. Freeman s.n. (NA) 
Japan. Oomi: 13 May 1954, M. Togashi s.n. 
CYO) 
Japan. Hyougo: Awaji-shima, 1 June 1978, N. 
Fukuoka & N. Kurosawa 1586 (KYO) 


Japan. Kii: Mt. Nachi, 11 Nov. 1974, G. Murata 
Hn 


5715 (KYc 


Poland: July 19 
Italy. Pralongia: 18 July 1879, Larsen 35105 


(NA) 
Japan. Shikotan Ins.: Matakotan, 19 July 1931, 
0) 


Japan. Hokkaido: Kushiro, 13 June 1982, K. 
Takita 999 (KYO) 

Japan. Kyusyu: Hiuga, 28 July T 2, M. Ogata 
s.n. (KYO) 

Japan. Hoki: Mt. Daisen, 15 Sep. 1914, N. Ki- 
nashi s.n. (KYO) 

U.S.A. Alaska: 22 July 1948, J. A. Calder s.n. 
(KYO 


U.S.A. Washington: Garfield Co., 18 Oct. 1930, 


O. M. Freeman s.n. ( 


A) 
U.S.A. Minnesota: Polk Co., 12 July 1939, O. A. 


Canada. aa Columbia: 26 June 1971, J. Po- 


a is 110 (K 


S.A. Washington: Yakima Co., 15 July 1940, 


k W. Thompson 15079 (NA) 

Japan. Akita Pref.: 1 Aug. 1978, Fujita 807 

(KYO) 

Japan. Niigata: 5 Aug. 1978, Terao 972 

Japan. Shiga Pref.: 29 Sep. 1974, G. Murata 
22326 (KYO) 


~ 


Japan. Bizen: Kurokami-yama, 6 July 1919, Z. 
Tashiro s.n. (KYO) 


Japan. Shiga Pref.: 30 May 1962, Fukuoka 
754 (KYO) 


U.S.A. California: Del Norte Co., 10 Aug. 1937, 


C. B. Wold s.n. (NA) 


U.S.A. California: Trinity Co., 7 July 1949, E. K. 


Balls 13885 (NA) 


52, Wisniewska et al. s.n. (NA) 


KYO) 


28.5 


19.3 


+ 1.6 x 19.3 


I+ 


I+ 


mm 


— 


ho 


to 


x 


x 


x 


x 


20.0 


21.0 


16.0 


16.0 


ae 


— 
— 


— 
Ww 


co 
OOo 


pd 
— 


Volume 76, Number 3 
1989 


Takahashi & Kawano 
Melanthiaceae Pollen 


865 


TABLE 1. Continued. 


Species 


Collection Data 


Pollen Sizes (um) 


T. okuboi 

T. palustris 
T. palustris 
T. palustris 
T. pusilla 

T. pusilla 

T. racemosa 
T. racemosa 
T. racemosa 
Pleea tenuifolia 
P. tenuifolia 
P. tenuifolia 


Japonolirion ozense 


Xerophyllum asphodeloides 


X. asphodeloides 
X. asphodeloides 
X. asphodeloides 
X. asphodeloides 
X. tenax 

X. tenax 

X. tenax 
Narthecium americanum 
N. americanum 
N. americanum 
N. americanum 
N. californicum 
N. californicum 


N. californicum 


Japan. Aomori Pref.: 13 July 1962, H. Ohashi 

68772 (KY 

Canada: s kenzie, 16 July 1940, J. J. Lynch & 
C. »illham 1597 (NA) 

TE ris Lapmark, 21 July 1935, C. G. 
Alm s.n. (KYO) 

U.S.A. Alaska: Carlisle Island, 1 July 1940, 7. N. 
Gabrielson s.n. (NA) 

Greenland. Blomsterdalen, 13 July 1966, F. J. 4. 
Janiels et al. 103 ( ) 

Sweden: Jämtland, 18 June 1963, Folkeson s.n. 


U.S.A. Louisiana: 15 d 1978, 5. Parwin « E. 
ce 592 (KY 
E MIA ae Co., 25 June 1938, 
p S. & H. B. Correll 9107 (N 
U.S.A. North Carolina: Onslow Co., 30 June 
1950, H. B. Fox & S. G. Boyce 3721 (NA) 


U.S.A. "ad Carolina: Aug. 1884, C. M. Carthy 


4 (N 

U.S.A. Ned carolina: Onslow Co., 26 Aug. 
1949, W. B. Fox 3228 (NA) 

USA: M Walton Co., 4. H. Curtiss 2904 
N: 


(NA) 
Japan, Tochigi Pref.: M. Shibutu & M. Taka- 
pde s.n. (TUS) 


U.S.A. New Jersey: Newfield Aug. 1877, C. Mar- 


d s.n. (NA) 

U.S.A. New Jersey: Burlington Co., 23 May 
1963, C. F. Parker s.n. (NA) 

U.S.A. North Carolina: 23 May 1963, Freeman, 
sn. (NA) 


U.S.A. New Jersey: Pomona, 12 June 1892, F. 
L. Lewton s.n. (NA) 
J.S.A. North € de Burke Co., 2 July 1891, 
m K. Small & . . Heller s.n. (NA) 
U.S.A. Oregon: 22 Pw 1975, IF. Hesse 3633 
(NA) 


U.S.A. Oregon: € - TF Co., 20 June 1939, C. 
) 


L. Hitchcock d Martin 4786 (NA 
U.S.A. Idaho: Vi yee 5 Aug. 1950, Q. Jones 
299 (N 
U.S.A. New Jersey: Bridgeton City, 31 Aug. 
1879, Unknown collector s.n. (NA) 


~ 


U.S.A. New Jersey: Bassto, July 1877, I. C. Mar- 


tindale s.n. (N/ 
U.S.A. s Jersey: eg m: Co., 18 June 
1933, F. J. Hermann 4460 (NA) 
U.S.A. E. Jersey: 23 June 1932, Hermann 
3315 (NA) 
U.S.A. California: Mount Dyer, 3 Aug. 1880, R. 
(A 


M. Austin s.n. 


california: Trinity Co., 7 July 1948, E. K. 


SA 

Balls 13880 
U.S.A. Oregon: 19 2 June 1946, P. L. Ricker 

6180 (NA) 


19.1 


— 

^ 
No) 
Qo 


20.8 


20.0 + 


T 


+ 


0.9 x 


1.4 x 


1.6 x 


0.8 x 


0.9 x 


15.0 


i 


ps 


— 


— 


866 Annals of the 
Missouri Botanical Garden 


TABLE l. Continued. 


Species Collection Data Pollen Sizes (um) 

V. californicum U.S.A. California: Del Norte Co., 16 June 1977, 18.3 + 1.4 x 13.2 
L. Debuhr 2625 (KYO) 

N. ossifragum Sweden: Smaland, 16 July 1957, J. Christoffers- 19.8 + 1.2 x 13.4 
son s.n. (KYO) 

V. ossifragum Denmark. Raabjierg Mile, 31 July 1938, Sehjottz 19.0 + 1.6 x 13.0 + 1. 
s.n. (KYO) 

Aletris aurea U.S.A. Florida: Duval Co., May 1897, 4. H. Cur- 25.9 + 1.6 x 17.1 + 
tiss s.n. (NA) 

A. aurea U.S.A. Texas: Tyler Co., 4 June 1968, D. S. 26.0 + 1.7 x 16.8 + 


Correll 35835 (NA) 


4. aurea U.S.A. North Puce New Hanover Co., 19 23.9 + 1.2 x 17.3 
May 1973, D. E. Boufford 8877 (KYO) 

A. bracteata U.S.A. Flo rida: Monroe Co., 25 Mar. 1980, W. 24.8 + 1.9 x 18.3 
C. Brumbach 9 

A. bracteata U.S.A. Florida: 4 t Pa 1904, J. K. Small & 25.2 + l.l x 17.3 
P. Wilson s.n. (N 

4. bracteata U.S.A. Florida: Dade Co., 10 Feb. 1948, O. M. 24.3 + 1.7 x 17.3 
Freeman s.t 1. (NA) 

A. farinosa U.S.A. Florida: May June 1875, C. Reynolds 26.9 + 1.6 x 16.3 
s.n. (NA) 

4. farinosa U.S.A. Virginia: Norfolk, June 1840, F. Rugel 23.4 + 1.8 x 16.9 
s.n. (NA) 

4. farinosa U.S.A. New Jersey: Burlington Co., 1 July 1932, 25.7 + 2.1 x 17.8 
F. J. Hermann 3379 (NA 

1. farinosa U.S.A. North Carolina: Transylvania Co., 13 July 21.8 + 1.5 x 15.8 
1950, W. B. Fox 3986 (NA) 

4. foliata Japan. Kai: 16 July 1962, G. Murata 27002 27.5 + 1.6 x 20.6 + 
(KYO) 

A. foliata Japan. Niigate Pref.: Aug. 1960, Koyama 0581 21,0 £257 X-18,2 £ 
(KYO) 

A. lanuginosa China. Szechuan: Sep. 1938, Maclaren 23 (KYO) 22.8 + 1.5 x 15.1 

4. lutea U.S.A. A piss pel iis Garden, 2 May 24.0 + 1.7 x 15.4 
1962, J. E. Peele 827 (NA 

A. lutea U.S.A. Nese De kal Co., 9 June 1936, C. 22.9 + 1.9 x 16.8 
Q. Erlauson 783 (NA) 

1. lutea U.S.A. Florida: Lake S 26 May 19606, C. R. 21.8 + 2.1 x 16.0 + 
Gunn 3399 (NA) 

4. lutea U.S.A. Florida: Gulf Co., 4 May 1979, D. E. 23.8 + 1.4 x 16.8 + 
Boufiord 20070 (KYO) 

A. obovata U.S.A. Florida: EO Co., 17 May 1940, R. 22.7 t 1.8 x 16.4 + 
E hi buds 1441 (NA) 

4. obovata U.S.A. Ge qon 18 May 1930, F. S. 25.5 + 2.0 x 18.7 
ed poo 

A. obovata U.S.A. Georgia: Liberty Co., 25 May 1943, WF. 22.7 + 1.6 x 14.6 + 
C. Jrimson s.n. ( ) 

A. pauciflora Nepal: between Pis B we top la, alt. 32.4 + 3.8 x 20.5 
inus m, 23 July , K. Nishida 681 
(KYO) 

4. sitimensis Bhutan: Sela la, alt. 3,200 m, 27 July 1958, S. 26.0 + 1.7 x 14.0 + 
Vakano 135 (KYO) 

4. spicata Japan. Kochi Pref.: May 1889, T. Makino s.n. 24.2 + 1.7 x 18.5 + 
(KYO) 

A. spicata Japan. Kagoshima Pref.: 1 May 1965, F. Konta 22.4 + 2.0 x 17.2 


5015 (KYO) 


— 


Volume 76, Number 3 
1989 


Takahashi & Kawano 867 


Melanthiaceae Pollen 


TABLE l. Continued. 


Species 


Collection Data 


Pollen Sizes (um) 


Metanarthecium luteo-viride 
hashi 1617 (KYO) 
M. luteo-viride 


M. luteo-viride Japan. Ehime Pref.: 


Japan. Nagano Pref.: 21 July 1973, H. Taka- 
Japan. Bitchu: 31 July 1962, € 
(KYO) 


27 July 1983, G. Murata 


18.0 + 1.2 x 13.1 + 1.4 


5. Murata 27319 


4755 (KYO) 
Lophiola americana U.S.A. Alabama: 20 May 1979, H. H. lltis s.n. 18.5 + 1.60 x 12.7 + 1.9 
(WIS) 
Amianthium sp. U.S.A. — Carolina: Macon Co., 28 Jur 23.2 + 1.4 x 15.4 + 1.2 


1982. F. H. Utech & M. Ohara 82-27 " ( CM) 


Stenanthium occidentale 
1978, F. H. 
Schoenocaulon pringlei 


collector s.n. (WIS) 


U.S.A. p Whatcom Co., 
Utech 78-287 (CM) 
Mexico. Pederal District: Aug. 1960, Unknown 


11 July 


ahashi (19822) showed that Heloniopsis orientalis 
Tanaka is characterized by a peculiar exine struc- 
ture without columellae. He also pointed out that 
palynological evidence on the related genera would 
be useful to reveal their systematic relationships. 

In the present study, pollen grains of 92 samples 
of 40 species of 15 genera in the Melanthiaceae 
were studied in detail with transmission and scan- 
ning electron microscopes. The probable system- 
atic relationships of these genera are proposed based 
on the palynological evidence thus obtained. 


MATERIALS AND METHODS 


Pollen grains were taken from specimens in the 
herbaria of Carnegie Museum of Natural History 
(CM), U.S. National Arboretum (NA), Kunming 
Institute of Botany (KUN), Kyoto University 
(KYO), and Tohoku University (TUS). Voucher 
specimens are listed in Table 1. Pollen grains from 
each specimen were observed using scanning elec- 
tron and transmission electron microscopy. Prep- 
aration methods followed Takahashi (1982b). An 
image analyzing system IBAS-1 was used for mea- 
surements and calculations of pollen grains. 


OBSERVATIONS 
Helonias and Ypsilandra (Figs. 1-4) 

27.7 + 1.3-28.5 
+ 1.6 um in major axis of ellipse and 19.3 + 
2.2-20.0 + 2.0 um in polar axis of Helonias 


bullata, and 30.0 + 
and 21.0 + 2.3 in polar axis of ) psilandra yun- 


Pollen grains monosulcate, 


1.8 in major axis of ellipse 


nanensis, provided with a sulcus. Except for sulcus, 


exine ornamentation finely granulated with spinules 


(Figs. 2, 3), these ovate, 1 um high, with acute 
tips. Exine (spinules excluded) 0.4-0.0 um thick, 
consisting of an amorphous layer and spinules. 
Outer surface of exine granular and the inner layer 


lamellated (Fig. 4). 
There is no difference in pollen type and exine 
structure between //elonias and Y psilandra. 


Chamaelirium and Chionographis (Figs. 5-9) 


Pollen grains p" al, 4-porate (Figs. 5, 8), 
16.2 + 1.2-19.0 + 1.5 um in major axis of ellipse 
and 16.0 + 1.0-18.1 + 1.3 um in polar axis of 

and 13.2 + 0.9-10. 1.2 in 


major axis of ellipse and 16.2 + 1.8 um in polar 


Chamaelirium. 


axis of Chionographis. Porus with exinous gem- 
mae. Except for the porus, exine sculpture clavate. 
Clavae with small projections on the head (Fig. 7). 
Exine structure intectate, consisting of 0.4-0.5- 
um-high clavae and 0.4-0.4-um-thick foot layer. 
Inner part of foot layer lamellated. 


There is no difference in pollen type and exine 


structure between Chamaelirium and Chiono- 


graphis. 


Tofieldia (Figs. 10-17) 


Pollen grains disulcate, 16.7 + 0.9-25.2 + 2,1 
( 


10-103 


+ 1.4 um in polar axis. Pollen size variable among 


2 
um in major axis of ellipse and 10.9 + 


species. Furrows extending near the ends of pollen 
grains. Except for the furrows, exine ornamenta- 
tion reticulate, consisting of 0.5-um-thick muri and 


868 Annals of the 
Missouri Botanical Garden 


7 


FicURES 1-7.— 1. Pollen grain of Helonias bullata, SEM x2,600.— 2. Exine sculpture of Helonias bullata, 
SEM x6,000.— 3. Exine sculpture of Ypsilandra yunnanensis, SEM x6,000.— 4. Exine structure of Ypsilandra 
yunnanensis, TEM x34,000.— 5. Pollen grain of Chamaelirium luteum, SEM x6,000.—6. Gemmate exine 
sculpture of Chamaelirium luteum, SEM x74,000.— 7. Exine structure of Chionographis koidzumiana, TEM 
x 49,400. 


Volume 76, Number 3 Takahashi & Kawano 869 
Melanthiaceae Pollen 


FIGURES 8-13.— 
Chionographis koidumiana, SEM x 18,000. — 10. Reticulate exine sculpture of Tofieldia glutinosa, SEM x 10,000. 
11. Pollen grain of T. glutinosa, TEM x 4,100. — 12, 13. Disulcate pollen grain of 7. occidentalis, SEM x 3,200. 


870 Annals of the 
Missouri Botanical Garden 


20 


FIGURES 14-20. — 14. Reticulate exine sculpture of Tofieldia occidentalis, SEM x10,000.— 15. Exine structure 
of T. glutinosa, TEM x 43,600. — 16. Pollen grain of T. pusilla, SEM x 3,600.— 17. Discontinuous reticulate exine 
sculpture of T. pusilla, SEM x 7,600. —18. Pollen grain of Japonolirion ozense, SEM x3,000.— 19. Gemmate 
exine sculpture of J. ozense, SEM x 7,400. — 20. Exine structure of J. ozense, TEM x 2.400. 


Volume 76, Number 3 
1989 


Takahashi & Kawano 
Melanthiaceae Pollen 


871 


lumina 0.2-0.7 


gins. Exine structure semitectate, consisting of 0.3- 


7 um diam. with nonangulate mar- 


um-thick tectum, 0.1-um high columellae, and 0.2 
0.3 um-thick foot layer. 


The reticuloid exine sculpture of 7. pusilla, 


which has discontinuous muri, is unique in the 
genus. 
Pleea 

Pollen grains disulcate, 20.0 + 1.5-22.7 + 1.3 


um in 1 major axis of ellipse and 15. 8 + 1.3-17.9 


+ 1.2 um in polar axis. Furrows extending near 
the ends of pollen grains. Except for the furrow, 
exine ornamentation reticulate, consisting of muri 
and lumina 0.2-0.3 um diam. with. nonangulate 
margins. Exine 0.5 um high, consisting of 0.2-um- 
thick tectum, 0.1-um-high columellae, and 0.2- 
um-thick foot laver. 


Japonolirion (Figs. 18 20) 


Pollen grains monosulcate, 19.2 + 2.0 um in 


major axis of ellipse and 15.6 + 1.8 um in polar 
axis, provided with a furrow. This with no marginal 
thickening, exinous islet absent. Except for the 
sulcus, exine ornamentation gemmate. Gemmae 
0.2-0.3 um diam., 


foot layer 0.2- 


spheroidal. Exine composed of 
0.3 um thick and gemmate protru- 
sion 0.3-0.4 um high. 


Yerophyllum (Figs. 21-24) 


44,0 E 154290 
+ 3.2 um in major axis of ellipse and 14.9 + 
1.4-19.9 


a furrow, this lacking exinous islets. Except for the 


Pollen grains monosulcate, 
1.3 um in polar axis, provided with 


furrow, exine ornamentation reticulate and. char- 
acterized by narrow muri, these 0.1 -0.2 um thick, 
the angulate lumina 0.5-1.0 um diam. Exine 1.0 
um thick, semitectate, consisting of 0.5-0.6-um- 
thick tectum, 1.0-um-high columellae, and 0.5- 
0.6-um-thick foot layer. 


There is no conspicuous difference in pollen 


morphology between X. asphodeloides and 
tenax. 
Varthecium (Figs. 25-27) 

Pollen grains monosulcate, 17.7 + 1.3-20.6 


+ 1.5 


um in major axis of ellipse and 11.9 + 
1.2-13.5 + 1.2 um in polar axis, provided with 
a furrow. Exine 0.5 um thick, semitectate, con- 
sisting of 0.2-um-thick tectum, 0.2-4m-high col- 


umellae, and 0.1-um-thick foot layer, reticulate, 


the reticulum consisting of 0.3-um-thick muri and 
lumina 0.1-0.5 um diam. 

No conspicuous interspecific variation was rec- 
ognized in pollen size, pollen type, and exine sculp- 
ture within Varthecium. 


4letris (Figs. 28-32) 


21.8 + 1.5-32.4 
+ 3.8 um in a axis of ellipse and 14.0 + 
3.4—-20.5 t 


a sulcus with exinous gemmae. 


Pollen grains monosulcate, 


.2 um in polar axis, provided with 
This extended to 
the ends of grains. 


The genus shows wide variation in exine sculp- 
ture, e.g., perforate-foveolate, reticulate, and gem- 
mate. The perforate-foveolate type is shown in 4. 


A. bracteata, A. ee and 


aurea, A. farinosa, 
A. lutea. The perforatum are 0.1-0.3 um in di- 
ameter. 


The reticulate type is represented by 4. folia, 
A. spicata, and 4. ossifragum. Here the lumina, 
0.5-1.0 um in diameter, are roudish rather than 
angulate. The foveolate or reticulate exine consists 
of tectum, columellae, and foot laver. 

The gemmate type is observed in 4. pauciflora 


and 4. lanuginosa. The gemmate exine is com- 


posed of 0.6-0.8-um-gemmae and 0.2-0.3-um- 
foot layer. 
Metanarthecium (Fig. 33) 

Pollen grains monosulcate, 17.2 + 1.3-18.3 


+ 1.0 um in the longest axis of ellipse and 11.6 
16-14 € 1.5 


a furrow having a membranous surface and no 


um in polar axis, provided with 


marginal thickening. Except for the furrow, exine 
sculpture reticulate, the reticulum. characterized 
by small projection on lumina (Fig. 33). Lumina 
1.0-1.5 um diam., muri 0.2-0.3 um thick. Exine 
composed of tectum, columellae, and foot layer. 
Lophiola (Figs. 34-35) 

19.9 e 
the longest axis of ellipse and 12.7 


Pollen grains monosulcate, 1.6 um in 
+ 1.8 um in 
polar axis, provided with a furrow. Exine sculpture 
reticulate, the reticulum composed of 0.1 -0.3-um- 
wide lumina and O.1-um-thick muri. 


Stenanthium, and 


30-39) 


Amianthium. 
Schoenocaulon (Figs. 

Pollen grains monosulcate, 23.2 + 1.4-25.2 
+ 1.6 um in the longest axis of ellipse and 15.4 


+-1,2-17,2 


1.4 um in polar axis, provided with 


872 Annals of the 
Missouri Botanical Garden 


FIGURES 21-27. — 21. Pollen grain of Xerophyllum asphodeloides, SEM x 3,000. — 22. Reticulate exine sculpture 
of X. asphodeloides, SEM x 3,000. — 23. Pollen grain of X. setifolium, SEM. x 2,600.— 24. Exine structure of X. 
TEM x42,000.— 25. Pollen grain of Narthecium americanum, SEM x3,000.— 26. Reticulate 
exine sculpture of N. californicum, SEM x12,000. — 27. Exine structure of Narthecium, TEM x44,000. 


asphodeloides, 'TE 


Volume 76, Number 3 Takahashi & Kawano 873 
1989 Melanthiaceae Pollen 


ete 


- 
Sea. 
pe 


FIGURES 28-33. — 28. Perforate exine sculpture of Aletris aurea, SEM x 7,400.— 29. Exine structure of A. 
aurea, TEM x40,000.— 30. Reticulate exine sculpture of 4. farinosa, SEM x 7,400.— 31. Pollen grain of 4. 
lanuginosa, SEM x 7,600. — 32. Gemmate exine sculpture of 4. lanuginosa, SEM x 7,600.— 33. Reticulate exine 
sculpture of Methanarthecium luteo-viride, SEM x 7,200. 


874 Annals of the 


Missouri Botanical Garden 


` 
A 


at LI 
Pare m ^ 
Lo. 

Far 


ex 


" 
‘ Rie Bnet 


i a 
dete — 


FIGURES 34-39. — 34. Pollen grain of Lophiola americana, SEM x3,800.—35 
Lophiola americana, SEM x 7,600. —36. Pollen grain of Amianthium sp., SEM x3,000.— 37. Pollen grain of 
Stenanthium occidentale, SEM x3,000.— 38. Pollen grain of Schoenocaulon pringlei, SEM x3,000.— 39. Re- 
ticulate exine sculpture of Schoenocaulon pringlei, SEM x 7,600. 


. Reticulate exine sculpture of 
x LI 


a furrow, the furrow with an operculum. Exine 
sculpture reticulate, the reticulum consisting. of 
0.3-um-wide lumina and 0.3-0.6-um-thick muri. 
The lumina angular. 


DISCUSSION 


Helonias and Ypsilandra are characterized by 


a distinctive exine structure that consists of an 


Volume 76, Number 3 
1989 


Takahashi & Kawano 
Melanthiaceae Pollen 


— 


amorphous layer and spinules, and the same exine 
structures were observed in Heloniopsis (Vaka- 
hashi, 1982a). Utech (1978, 1980) and Utech & 
Kawano (1981) suggested that //eloniopsis and 
Helonias should be maintained in the same tribe 
based on floral vasculature and karyology despite 
Hutchinson's (1934, 1959) placement of /7elonias 
in Heloniadeae apart from Heloniopsis in the Nar- 
thecieae. The present pollen morphological evi- 
dence supports the suggestion by Utech & Kawano 
(1981) and further indicates that ) psilandra should 
be included in the same tribe with /7elonias and 
Heloniopsis. The three genera are closely related 
to each other and are distinctive because of the 
amorphous exine structure with spinules. Dahlgren 
et al. (1985) included /lelonias and Heloniopsis 
in the tribe Narthecieae, with Methanarthecium, 
Vietneria, Narthecium, and Lophiola. We now 
regard Helonias, Heloniopsis, and Y psilandra as 
congeneric, and a more thorough taxonomic ac- 
count of this controversial group is needed. 

Chionographis from Japan has clavate exine 
sculpture and four-porate pollen grains. Chamae- 
lirium from the eastern United States has the same 
pollen. This type of grain and exine structure are 
unique in the Melanthiaceae, and suggest that Cha- 
maelirium and Chionographis are closely related 
to each other as an isolated group in the Melan- 
thiaceae. The pollen morphology supports the 
treatment of Dahlgren et al. (1985), who distin- 
guished the tribe Chionographideae as made up of 
Chionographis and Chamaelirium. 

Japonolirion, which consists of only the Jap- 
anese endemic J. ozense, is sometimes included in 
Tofieldia (Willis, 1966; Hutchinson, 1934, 1959). 
The pollen grains of Japonolirion are monosulcate 
with gemmate exines, as opposed to those of 7o- 
fieldia, which are disulcate with reticulate exines. 
The pollen morphology of Japonolirion implies 
that the genus should not be e in Tofieldia 
but should be treated as a se e genus. The 
report of gemmate exine of Tofieldia puo (Ikuse, 
1956) appears to be incorrect, as we have found 
reticulate exine in this species, conforming to other 


para 


species of Tofieldia. 

There is a wide range of variation in exine sculp- 
ture, perforate-foveolate, reticulate, and gemmate 
types in the genus dletris. The species with gem- 
mate exine are 4. lanuginosa and A. pauciflora, 
which are distributed in China and Nepal. The other 
species of Aletris have reticulate exines. Similar 
variation in exine sculpture occurs in Clintonia 
(Takahashi & Sohma, 1982) and Paris (Taka- 
hashi, 1984). A parallel derivation from tectate- 
perforate to intectate exine appears to have taken 


place independently within these genera (laka- 
hashi, 1984) 

Ambrose (1985) transferred the genus Lophiola 
into the Narthecieae. The exine sculpture of the 
genus has a reticulum similar to that of genera in 
the Narthecieae; thus the palynological evidence 
supports Ambrose's taxonomic position of Lophiola. 

| the evidence available at present suggests that 
Heloniadeae and Narthecieae sensu Hutchinson 
(1959) consist of at least three or four heterogenous 
groups, and therefore the revision of the classifi- 
catory system of these groups is now in order. 


LITERATURE CITED 
AMBROSE, J. D. 1985. A re-evaluation of the Melan- 
thioideae (Liliaceae) using numerical analyses. /n: € 

; M. Gregory editor 
Linn. Soc. Symp. 


Brickell, D. F. Cutler 
e Monocotyledons. 


Buc, T ds Leitfaden der Pollenbestimmung fur 
Mitteleuropa und angrenzende Gebiete. Leiferung 1: 
1982. The 


Academic 


H. T. CLIFFORD. 
A Comparative Study. 


DAHLGREN, R.M 
Monocotyledons: 
Press, London 

Families 

an 


The 


Evolution 


Yro. 1985. 
Struc ture, 


= 


Q,——— QU Pt. 
of the Monocotyledons. 
Taxonomy. Ha Verla 
19 Pollen Morphol and Plant Tax- 

Wik 


onomy. b is Almqv sell, Stock- 
ge 

FAEGR , J. Iversen & H. T. »d VEERBOLK. 1964. 
iP s of Pollen Analysis, 2nd rev editio 


Books, Mode eo 


Scandinavian Univ. 
1970. lants 
Taiwania 15: 73-1 
ae Pollen ed of Taiwan. 
v Un , Botany Dept. Press. 
HUTCHINSON, T 1934. The Families of Flowering E 
Volume II. Monocotyledons. MacMillan & Co. 
don 


— 


gen. 
Huanc, T. C. 
(6). 


Pollen grains of Formosan 
79, 


National Tai- 


(oa. 


The Families of Flowering Plants, 
London. 


1959. 
Monocotyle dons, 2nd edition. Oxford, 
197 The Families of Flowering Plants, 3rd 
el London. 
Huyxn, K. L. 1976. Arrangement of some monosul- 
cate, disulcate, trisulcate, dicolpate and tricolpate 
pollen types in the tetrads and some aspects of evo- 
lution in n angiosperms. Linn. Soc. London Symp. 
: 101-124. 


edino. 


Ser. 

Ikuse, M. Tm 
2 s 

KRAI sE, K. A. Engl e K. eae Die Na- 
turlichen aa LN tion. 2: 227 6. 

Kuprianova, L. A. 1948 decenas i pollen et 
phylogónie d E. b x e zomm. Komarow 
Inst. Acad. 1(7): 163-262. [In Russian. | 

RADURESCU, 197 7:3. Recherches morpho-palynolo- 
giques sur la famille P Acta Bot. Horti Bu- 

. 1972-1973: 248 
Ww. 1978. foe zur Taxonomie der Li- 


Hirokawa 


dum Grains of Japan. 


T n 


LL 


"T€ LEZE; 


Annals o 
Missouri Botanical Garden 


liifloren. IV, Melanthiaceae. Wiss. Z. ddp Schil- 
ler-Univ. Jena, Math.-Naturwiss. Reiche 27: 87-95. 
SHIMAKURA, M. 1973. eg inar of n plants. 


a Mus. Nat. Hist., . Publ. 1-60. [In 
usse 
TAKAHASHI, M. 1982a. 


Pollen morphology in the genus 
p ae). Grana 21: 175-177. 
——— Pollen morphology in TUN deno 
species ol Trillium. Amer. J. Bot. 69: 1185 
1984. Pollen inert Bl in Paris P its 
related genera. Bot. Mag. (Tokyo) 97: 233 
& K. SonmMa. 1982 Pollen mophr d 
e a (Liliaceae). Sci. Rep. Tohoku Imp. 
Un , Biol. 38: 157-164. 

ERU a 1970. Les elos palynologie et 
Sg rur oF Pondichery Trav. Sect. 
Sci. Tech. 

Tinc-Su. 1949. a ne grains of some 
Chinese pu. Bot. Not. 282 

Urrcu, F. H. 118. Vasc Pu foral Reston of He- 
lonias us (Liliaceae- Helonieae), with a com- 

Asian A loniopsis nia Ann. 

egie Mus. 47: 169-191. 

1979 die analysis, palynology and 


E 
3 

P 
e; 


^D. 


external seed ip ied of Tofieldia tenuifolia 
(Michx.) Utech (Liliaceae- Tofieldieae). Ann. 
negie Mus. 48: 171- 84. 
. 1980. Somatic karyotype analysis of Helonias 
bullata L. (Liliaceae), with a c MA er to the Asian 
Heloniopsis orientalis (Thunb.) C. Tanaka. Ann. 
ae Mus. 49: 153-160. 
1984. Floral vascular anatomy of Japonoli- 
rion osense Nakai pou and its tribal relation- 
ship. Ann. ee Mus : 447-46 
—— &€ VANO. 1981. E ascular floral anatomy 
of the East oe Heloniopsis orientalis (Thunb.) € 
e M Helonieae). Bot. Mag. (Tokyo) 94: 
295-3 

VAN DER n T. 1956 
nomenclature. Bol. Geol. (Bogotá) 4: 63- 
I-XII. 


Palynological systematic 


101, pls. 


Wirus, J. C. 1966. A Dictionary of the Flowering 
Plants c Ferns, 7th edition. Cambridge Univ. Press., 
Cambri 

RUE x 1936. Vergieiheude Untersuchung- 

enkórnern einiger Liliac 


en von een und Amar- 
yllidaceen. ire terr. Bot. Z. Sonderabdr. ., Heft 1, Bd. 
>: 30-35. 


A NEW SUBGENUS AND Linda K. Escobar? 
FIVE NEW SPECIES 

IN PASSIFLOR A 

(PASSIFLORACEAE) 

FROM SOUTH AMERICA! 


ABSTRACT 

Five new species of Passiflora (Passifloraceae) are described: Passiflora solomonii from Bolivia; P. caatingae 
trom Bahia, Brazil; P. sierrae from the Sierra Nevada ih: Sant pi of northeastern Colombia, and P. amazonica 
and P. pascoensis from Peru. Passiflora sierrae is pla in a new subgenus Porphyropathanthus. All are lianas 


or herbaceous climbers with axillary tendrils and simple jam es, found on roadsides or in recent Res ngs. 


Passiflora amazonica L. Escobar, sp. nov. TYPE: — 1.4 cm wide, acuminate at apex, densely pubescent 
Peru. Amazonas: Bongara Prov., km 385- with tightly curled trichomes on inner surface, 
365 Moyobamba Bagua road (77?45'W, | moderately pubescent with slightly curved tri- 
5°45'S), 2,000-1,800 m, 17 Feb. 1984, D. chomes on outer surface; flower stipe ca. 4 mm 
N. Smith 6012 (holotype, MO). Figure 1. long. Flowers salverform, 9.5-10.5 cm long, « 

6.5 cm diam. at apex, probably pendent; AN 

n subg. Tacsonia juxta P. mixtam Linn. f. collocanda, 

ab hac foliis rugosis rigide coriaceis margine grosse den- 
tatis diversa. 


thium cylindrical, ca. 6.5 cm long, 1.0-1.2 cm 
wide (pressed), densely pubescent on outer surface 
with straight to slightly curved trichomes up to | 
Liana. Plants pubescent with straight to curling mm long; sepals ovate-oblong, 4.0-4.3 cm long, 
yellowish, transparent trichomes 0.2-1.0 mm long. 1.2-1.8 em wide, acute at apex with subapical 
Stems subangular, striate. Leaf blades 3-lobed to arista ca. 1 mm long, coriaceous, pink; petals sub- 
ca. 12 their length, widely depressed ovate in out- equal to sepals, up to 1.5 cm wide, rounded 
line, (8.5-)11-11.7 em long, (9.0-)1 4-15.2 em apex, membranaceous, pink; corona reduced to an 
wide, acute at lobe apices, deeply cordate at base, undulate, irregularly toothed band; operculum de- 


coarsely dentate at margins, stiff-coriaceous, ru- pendent, nonplicate: ovary pubescent with straight 
gose, scarcely pubescent (mostly on veins) on adax- trichomes ca. 0.5 mm long. Fruit unknown. 


ial surface, densely pubescent on abaxial surface UE e "e 
i Distribution. Passiflora amazonica is known 


with curved trichomes ca. 0.5 mm long; leaf lobes l 
` ong E only from montane forest at 1,800-2,000 m el- 


ovate, lateral ones (6.0-)7.2-8.2 cm long, (3.6-) 


. : : us evation in Amazonas Department in northern Peru. 
4.2-5.6 cm wide, divergent from mid-lobe ca. 75°, P 
this (7.3-)9.3-9.7 cm long, (4.2-)5.4-5.8 cm wide; esq c ollec tions examined. PERU. AMAZONAS: 
petioles 2.3-2.6 cm long, with up to 10 elongated dnd e 0-43 km E of Pomacoc i near — Cumbre, 
1 376 of ple etera Marginal, 5°45'S, SW, l: 


jectaries ca. | long scattered along adaxial 
nectaries ca mm long scattered along adaxia le 1986. S. Ruane d = gode 25 Lo 


surface; stipules auricular, ca. 1.5 em long, ca. 
0.7 em wide, acuminate with arista 4.6 mm long Passiflora amazonica is placed in subg. Tac- 
at apex, lacerate at margins. Peduncles solitary, sonia near P. mixta L. f., a widespread Andean 
5-6 cm long: bracts 3, connate ca. 14 their length, species (Escobar, 1980; Killip, 1938) from which 
forming a closely fitting tube around base of hy- it differs by the stiff-coriaceous, rugose leaves with 


panthium, each bract lanceolate, 3-4 em long, ca. coarsely dentate margins. 


! The present study was initiated in November 1987 while on a leave of absence, and completed in 1988 while 
on a sabbatical leave supported by the University of Antioquia. I am grateful to the curators of the herbaria of the 
Missouri Botanical Garden, The New York Botanical Garden, and the Smithsonian Institution for help and use of the 
facilities during my stay at their institutions. Special thanks are due Dr. Rupert arneby, who reviewed the manuscript 
and provided the Latin diagnoses, and to Dr. John MacDougal for criticism of the manus 

* Departamento de Biología, Universidad de Antioquia, Apartado Aéreo 1226, Medellin, ‘Colombia, South America. 


ANN. MISSOURI Bor. GARD. 76: 877-885. 1989. 


878 Annals of the 
Missouri Botanical Garden 


MISSOURI 
BOTANICAL GARDEN 
HERBARIUM 


N2 3433072 


GURE 1. Holotype of Passiflora amazonica L. Escobar, deposited in the herbarium of the Missouri Botanical 


F 
Garden (MO). 


Volume 76, Number 3 879 
1989 


scobar 
New Passiflora 


MISSOURI 
BOTANICAL GARDEN 
ARIUM 


N? 3110137 


cO 2 
i tu 
may 

ae 
4 
boa 

2A 
> 
H “u 
^ 
2 
A 
e Ss 
o 
= 

N Fa 

MN 

- - 

x 


FicuRnE 2. Holotype of Passiflora caatingae L. Escobar, deposited in the herbarium of the Missouri Botanical 
Garden (MO). 


880 


Annals of the 
Missouri Botanical Garden 


Passiflora caatingae L. Escobar, sp. nov. TYPE: 
razil. Bahia: 27 km NE of Cándido Sales, 
along Highway BR-116, caatinga, 890 m, 30 
Mar. 

& D. M. Vital 116354 (holotype, MO 


ure 2. 


— 


Fig- 


-- 


Subgeneris Tacsonioidi referenda, P. mendoncae 
Harms proxime accedens, sed ab hac stipulis join rad 
ceolatis bracteisque minoribus diversa. 


Liana. Plants glabrous with terete, striate stems. 
Leaf blades 3-lobed to ca. ?4 their length, depressed 
3.6-6.9 cm long, 6.4-11.2 cm 
wide, acuminate at lobe apices, cordate at base, 


ovate in outline, 


shallowly serrate at margins (glandular in sinuses 
of serrations on very young leaves), coriaceous, 
lustrous, yellow-green; leaf lobes elliptic to oblong, 


lateral lobes 3.8-5.5 cm long, 1.5-2.2 em wide, 
divergent from mid-lobe ca. 85°, mid-lobes 4.0- 
6.8 cm long, 1.6-2.8 cm wide; petioles 0.9-2.4 


€ 


cm long, with 2-4, paired, scarlike nectaries at or 
below mid-point on adaxial surface; stipules linear- 
lanceolate, soon deciduous, ca. 1 mm long, ca. 0.2 
mm wide at base. Peduncles solitary, 2.0-2.5 cm 
long; bracts 3, free to base, verticillate, ovate, 1.0— 
1.5 em long, 0.8-1.0 cm wide, borne 0.5- 1.1 cm 
below base of hypanthium, acuminate at apex, with 
arista ca. 0.5 mm long, narrowed at base, minutely 
serrate at margins: flower stipe 0.5- 1.1 cm long. 


Flowers salverform 3.2-5.0 cm long, ca. 4 cm 
diam. at apex; hypanthium slightly funnel-shaped, 
1.3-1.5 em long, ca. 9 mm wide at apex, ca. 6 
mm wide at base then inflated, with inflated portion 
5 mm long. | cm wide; sepals oblong, 2.6-3.3 cm 

1 with keel 2 


long, ca. 7 mm wide, 
abaxial surface, subcoriaceous, red-purple; petals 


3 mm wide on 


oblong, ca. 3.1 em long, ca. 0.7 em wide, rounded 
at apex, cone denis poten corona in 2 
series ca. 2 mm apart, filamentous, vithi inner series 
ca. 5 


€ 


mm long, outer series ca. 2-3 mm long; 


operculum erect, nonplicate, ca. 4 mm long, lac- 
erate at margin, transparent. Immature fruit ellip- 


soidal, 3.9—4.4 cm long, 2.0- 2.9 cm wide (pressed). 


Distribution. Passiflora caatingae is known 
only from two collections near Cándido Sales in 
the state of Bahia in eastern Brazil. It was collected 
along the highway in caatinga vegetation at 890 
m elevation. 

BRAZIL. BAHIA: 
30 Mar. 1976, 
M. Vital 11625 


Additional collections examined. 
along ne BR-116, caatinga, 890 m, 
G. Davidse, T. P. Cue d & D. 
(MO). 

Passiflora caatingae is placed in subg. Tac- 


sonioides, where it most closely rese uos. Passi- 


1976, G. Davidse, T. P. Ramamoorthy 


flora mendoncae Harms (Killip, 1938) but from 
which it differs by the linear-lanceolate stipules and 
The bracts resemble those of P. 
racemosa Brot. of the monotypic subg. Calopa- 


smaller bracts. 


thanthus, but this species possesses flowers with a 
folded operculum and strongly keeled sepals. 


Passiflora pascoensis L. Escobar, sp. nov. TYPE: 
Peru. Pasco: Oxapampa, Cordillera Yanacha- 
ga, 12 km E of main Oxapampa- Villa Rica 
road, remnant cloud forest, LON- 2,200 m, 
10935'S, 75°15'W, 2 Mar. A. Gentry 
& S. Smith 35939 (holotype, MO). Figure 3. 


Subgeneris Plectostemmatis sect. Decalobae referen- 
da, foliorum ambitu et coronae filamentorium exteriorum 
orma P. alnifoliam et arcte affines simulans, sed floribus 
et Aa magnis necnon foliis inibi crasse reticulatis 


diver 


> 


Herbaceous vine. Plants pubescent, with irreg- 
ular, curved to wavy, white to yellowish transparent 
trichomes 0.1-0.4 mm long. Stems 3-angular, 
striate. Leaf blades entire, ovate to elliptic, (5.3-) 
9.5-14.8 cm long, (3.8-)6.5-8.6 cm wide, 2-3- 
lobed at apex, shallowly cordate to rounded at base, 
entire at margins; leaf lobes subdeltate, lateral lobes 
0.8-1.5 em long, 0.7-1.3 
(lacking on some leaves) 0.2-0.4 cm long, 1.7- 


cm wide, mid-lobes 
3.0 cm wide, mostly glabrous adaxially, pubescent 
on abaxial surface, with expanded fine veins form- 
ing an incrassate reticulum, slightly coriaceous; 
aminar nectaries variable in number, 5-ca. 40, 


mostly concentrated at base and between major 
lateral veins in upper 14 of abaxial surface; petioles 
1.0-2.5 cm “p eglandular; stipules falcate, 0.5 

1.0 cm long, « 
20-2 
upper 4 of ean le, 0.4-0.6 em long, 0.5 mm 


. 0.5 mm wide. Peduncles paired, 


.0 em long: bracts setaceous, scattered on 


wide; flower stipe ca. 3 mm long. Flowers widely 
campanulate, 7-8 cm diam., pendent; hypanthium 
1.8-2.( 
-3.0 cm long, 1.8-2.0 cm wide, 


patelliform, ca. 0.4 cm long, cm wide; 
sepals ovate, 2.7 
obtuse at apex, coriaceous, greenish white; petals 


subequal to sepals, membranaceous, pinkish or white 


edged with pink; corona in 2 series, filamentous, 
cream-white banded with red-purple, the inner fil- 
aments 0.3-0.7 mm long, 0.5-0.8 mm wide and 
outer ones ca. 1.5 cm long, ca. 2 mm wide, dilated 
at apex; operculum closely plicate, erect, ca. 6 mm 
long. minutely crenulate at margin; ovary glabrous. 


Fruit ellipsoid, 2.5-3.0 em long, 2.3-2.6 cm wide 


(pressed), vellow: seeds obovoid, ca. 3.5 mm long, 


ca. 2.0 mm wide, with ca. 7 rugulose transverse 


ridges, dark brown at maturity. 


Volume 76, Number 3 881 
1989 


scobar 
New Passiflora 


MISSOURI 
BOTANICAL GARDEN 
HERBARIUM 


No 2983759 - 


PERU 


FicuRE 3. Holotype of Passiflora pascoénsis L. Escobar, deposited in the herbarium of the Missouri Botanical 
Garden (MO). 


882 Annals of the 
Missouri Botanical Garden 


Plantae, colombianae 
CUATRECASAS 


NS 24615 


FIGURE 4. Holotype of Passiflora sierrae L. Escobar, deposited in the herbarium of the Smithsonian Institution 
(US). 


Volume 76, Number 3 


Escobar 883 
New Passiflora 


Qt 


FiGURE 5. Holotype of Passiflora solomoni L. Escobar, 


Garden (MO). 


BOTANICAL GARDEN 
HERBARIUM 


N? 3479310 


a d 
$ 
ALO manel 4 mt en MN 


Xenda K uto E 


BOLIVIA 


MISSOURI BOTANICAL GARDEN HERBARIUM (MO) 


eidwork supported by the Nationa! Science Founda! 


deposited in the herbarium of the Missouri Botanical 


884 Annals o 
Missouri Botanical Garden 
Distribution. Passiflora pascoensis is found cles paired, 1.5-2.0 cm long; bracts 3, scattered 


on the borders of disturbed cloud forest at eleva- 
tions of 1,850-2,500 m in the Department of 
Pasco, Peru. 


= 


Additional collections examined. PASCO: 
Oxapampa, Cordillera Yanachaga, new rr in construc- 
tion from Oxapampa to Villa-Rica, 7-9 km E of main 
00-2,500 m, montane cloud forest, 10°45'S 
l Mar. 1982, 4. Gentry & D. Smith 35857 
(MO): Valle de San Alberto, E of Oxapampa, above hy 
droelectric plant, at base of C ig a Yanac x dads 
side forest remnants, 1,900 m, 1 n 1984, R. Foster, 
Alban 7714 (HU | MOX pastures in 
y E of ^ village, secondary 
growth, 1,850 m, 10?35'S, 75°35'W, 10 Mar. 1984, S 
Knapp, J. Mallet & W. Brack 6319 (MO, NY, US). 
This beautiful, large-flowered species is placed 
in subg. Plectostemma sect. Decaloba (Killip, 
1938), where it resembles members of the Pas- 
siflora alnifolia complex in leaf shape and form 
of the outer coronal filaments. It is easily distin- 
guished from those species by the large flowers and 
fruits and by the incrassate reticulum formed by 
expansion of the fine veins of the lower leaf surface. 


Passiflora subg. Porphyropathanthus L. Es- 


cobar, subgen. nov. TYPE. Passiflora sierrae 


L. Escobar. 


ectariis petiolaribus, operculo haud plicato necnon 

Passio lane ifoliae A L N aen similis, 

s 1.3-1.8 em longis, seminibus 

ransverse tubae et nectariis in foliorum lamina sitis 
incongruum. 


Passiflora sierrae L. nov. TYPE: 


Colombia. Magdalena: Sierra Nevada de Santa 


Escobar, sp. 


arta, southeastern slopes, hoya del Río Do- 
nachui, near Col, 3,100-3,070 m, 9 Oct. 
1959, J. Cuatrecasas & R. Romero-Casta- 
reda 24675 (holotype, US; isotype, COL). Fig- 
ure 4. 


lisdem notulis jam sub subg. Porphyropathantho in- 
dicatis unica. 

Herbaceous vine. Plants pubescent, with straight 
to wavy, reddish trichomes ca. 0.5 mm long. Stems 
iia, striate. Leaf lobes entire, ovate to lan- 
ceolate, (3.8-)4.3-9.0 cm long, (1.4-)2.4-4.5 cm 
wide, acuminate and mucronate at apex, rounded 
at base, entire at margins, stiff-coriaceous, lustrous 
on adaxial surface, yellow-green, pubescent on veins 
on both surfaces; laminar nectaries 0-3, scarcely 
visible between major veins of abaxial surface; pet- 
ioles 3-6 mm long, with 1 pair of stipitate, cup- 
shaped nectaries ca. 1 mm diam. at apex; stipules 


linear-lanceolate, ca. 5 mm long, reddish. Pedun- 


1.5 
2.0 mm long, ca. 0.3 mm wide; flower stipe 1.6- 


on upper 2 of peduncle, linear-lanceolate, 


2.2 cm long. Flowers campanulate, 1.7-2.0 cm 
long, 3.5-4.0 cm diam., pendent, red-purple; hy- 
panthium patelliform, 3-4 mm long, 0.5-1.0 em 
wide; sepals oblong, 1.7-2.0 cm long, 2-6 mm 
wide, obtuse at apex, subcoriaceous, pubescent on 
abaxial surface; petals oblong, 1.3-1.8 cm long, 
1.3-3.5 mm wide, rounded at apex, delicately 
membranaceous, pink to light purple; corona in | 
series, filamentous, ca. 1.5 mm long, but irregular, 
each filament ca. 0.1 mm wide; operculum erect, 
ca. 2 mm long, nonplicate, crenate at margin; 
ovary ellipsoid, glabrous, light green. Fruits glo- 
bose, ca. 1.5 em long, 1.3 cm wide (pressed); seeds 
with 


obovoid, ca. 6.0 mm long, ca. 3.1 mm wide, 


ca. 13 rows of transverse rugulose ridges. 


Distribution. This species is known only from 
subpáramo vegetation in the Sierra Nevada de San- 
ta Marta in northeastern Colombia, where it is 
found in dense shrubby growth above 3,000 m 
elevation. 


Acciona cid examined. COLOMBIA. MAG- 


DALEN rra Nevada de Santa dri entre San pes 
y cabeceras d al Río Sevilla, 3,195 m, 31 Jan. 195 

H. Bare lay & P. Juajibioy € 67 783 (C OL. Moy; Transect 
d Alto Buriticá, cuchilla y cerros a 3,300 m, lev. 33, 


ug. 1977, Jaramillo M., T. van Pf pande 


M A. Cleef & 0. Rangel 5472 (COL). 


Jo 


The presence of petiolar nectaries and. purple 
flowers with a nonplicate operculum, suggests a 
relationship to P. lancifolia Desv., of subg. Chlo- 
ropathanthus (Harms, 1925; Killip, 1938), which 
is known only from Jamaica. Unlike that species 
and others of subg. Chloropathanthus, P. sierrae 
has well-developed petals, poorly developed lami- 
nar foliar nectaries, and seeds with transverse ru- 
gulose ridges. This unique combination of char- 
acters requires the creation of a new subgenus. 
Data from pollen studies (Prėsting, 1905; Spirlet, 
1965; Escobar, unpubl.) suggest that several pur- 
ple- or red-flowered species (usually with a tubular 
hypanthium and therefore presumably humming- 
bird-pollinated) have arisen independently from the 
large and variable subg. Plectostemma. They do 
not suggest that all species with the hummingbird- 
pollinated syndrome are monophyletic. 


Passiflora solomonii L. Escobar, sp. nov. TYPE: 
Borivia. La Paz: Prov. Nor Yungas, 9.1 km 
NE (below) Chuspipata on road to Yolosa, 
2,400 m, 16?16'S, 67°47'W, 29 Oct. 1984, 


Volume 76, Number 3 
1989 


scobar 
New Passiflora 


J. C. Solomon & M. Nee 12592 (holotype, 
MO; isotype, HUA). Figure 5. 


Foliorum forma, nisi ambitu transverse lineari (nec 
subg d exoperculatam Mast. 

ra pedunculis longioribus, folio- 
ectariis petiolar Pus eae 
necnon coronae Pens capitellatis differ 


Herbaceous vine. Plants sparsely pubescent with 
wavy, transparent trichomes ca. 0.1 mm long. 
Stems angulate, striate. Leaf blades transversely 
linear, coriaceous, sparsely pubescent on abaxial 
surface, dull yellowish green, 2-lobed, the lobes 
divergent by ca. 90? from mid-vein, each 1.1-1.5 
7.1-10.0 cm 


mucronate at apex, shallowly cordate at base, 


cm long, wide, acuminate and 
tire at margins; petioles 4-9 mm long, eglandular; 
stipules linear-lanceolate, ca. 2.5 mm long, ca. 0.2 
mm wide. Peduncles paired, 6-12 mm long, very 
slender; bracts 3, setaceous, ca. 2 mm long, drying 
black, scattered along peduncle; flower stipe ca. | 
mm long. Flowers campanulate, 2-3 cm diam., 
pendent, greenish white; hypanthium patelliform, 
2.5-3.0 mm long, ca. 3.5 mm wide; sepals tri- 
angular, 8-12 mm long, ca. 10 mm wide at base, 
acute at apex; petals lanceolate, 11-12 mm long, 
3-6 mm wide, delicately membranaceous, greenish 
white; corona in 2 series, the outer series linear, 
7-8 mm long, ca. 1 mm wide, the inner filaments 


ca. 3 mm long, 0.2 mm wide, operculum dependent 


then recurved, loosely plicatez ovary globose, pu- 


bescent. Fruits unknown. 


Distribution. Passiflora solomonii is known 
only from the type locality in La Paz Department 


of Bolivia at 2,400 m elevation. 


The unusual leaf shape of Passiflora solomonii 
is reminiscent of P. exoperculata of subg. Plec- 
tostemma but is transversely linear rather than 
transversely ovate; P. exoperculata also differs 
rom P. solomonii in possessing prominent petiolar 
nectaries and capitellate inner coronal filaments as 
well as longer peduncles and shorter leaf lobes. 


LITERATURE CITED 

Escobar, L. K. 1980. 
of d 5 Centering A 
sima (H.B Bailey, as Tacsonia. 
Dissertation. x re sity of Texas, Austin, Texas. 

HanMs, H. 1925. Passifloraceae. a Natürlic :hen Pflan- 
zenfamilien, 2nd edition 21: —5( 

Kini, E. P. 38. The He an species of Passi- 
zm Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 

-6 


Interrelations of the Edible Species 
round Passiflora mollis- 
h.D 


Sue D. 
der Pas M RAM n. 
SPIRLET, M.- 1965. 
x de pollen de Passifloracees I. Poller 


Pollen 
Utilisation taxonomique a omi 


1 & Spores 7(2): 


Zur Mg ace der cue r 
& S 7(2): 1 


A REVISION OF 
MESOAMERICAN 
PSYCHOTRIA SUBGENUS 
PSYCHOTRIA (RUBIACEAE), 
PART III: SPECIES 48-61 
AND APPENDICES’ 


Clement W. Hamilton? 


GROUP 7. THE TEVUIFOLLA GROUP 


Shrub or small tree; young stems glabrous or 
puberulent; stipules often sheathing, ovate to tri- 
angular with apex long-biacuminate in P. tenuifolia 
(Fig. Ze), uniform in color, glabrous or puberulent, 
caducous. Leaf blades elliptic (to sometimes slightly 
ovate in P. limonensis), drying green-brown or 
red-brown; secondary veins 8-21(-24) pairs, di- 
verging (60°) 70?-90?(—95?), 
mous with collector vein near margin, the axils 


brochidodro- 


lacking domatia or hairs; tertiary veins orthogonal 
reticulate (to sometimes percurrent in P. limonen- 
sis). Inflorescences umbelliform panicles of cymes 
(Fig. 7f), the peduncle lacking or no more than 2 
em long; secondary axes in one pair per rank; 
bracts not conspicuously enlarged. Corolla tubes 
1.5-2 mm long, the lobes without apical exten- 
sions. Fruit when dry ellipsoid; persistent calyx 
inconspicuous or a beak; seed dorsal surface with 

-5 longitudinal furrows (often irregular in P. li- 
monensis), the ventral surface with 2 longitudinal 
furrows (often incompletely divided in P. limonen- 
sis). 

The two species in the group, Psychotria li- 
monensis and P. tenuifolia, are both widespread 
32, 33) and often 
occur together in lowland forest. 


throughout Mesoamerica (Figs. 


oth species appear to be normally distylous 
with no between-morph asymmetry in floral part 
lengths. 


48. Psychotria limonensis K. Krause, Bot. 
Jahrb. Syst. 54, Beibl. 119: 43. 1916. TYPE: 
Costa Rica. Limón: Puerto Limón, July 1898 
(fr), Pittier 12681 (holotype, B, destroyed, 
fragment, F, photo, F ne no. 518; isotypes, 

, US). Figures 7f, 3 


hae prio s K. Krause var. laxinervia Loese- 
r, Repert. Spec. Nov. Regni Veg. 18: 361. 1922. 

TYPE: Mn: :0. Chiapas: Ruinas del Palenque, 10 
Mar. 1911 (fr), C. Seler & E. Seler 5463 (holotype, 

B, destroyed, fragment, F, photo, US; isotype, GH). 


Shrub to small tree 0.5-3(-5) m tall; young 
stems glabrous or sparsely puberulent, the bark 
smooth; stipules sheathing, rounded-ovate to tri- 
angular, 6-9 -10 mm, glabrous, caducous, 
leaving a pale or red-brown ridge with red-brown 
fringe. Leaves petiolate; petioles 2.5-6 cm long, 
glabrous, flat or channeled above; blades membra- 
nous to chartaceous, elliptic to slightly ovate, the 
apex acuminate, the base narrowly cuneate to at- 
tenuate, (12-)1 7-29 x (6-)7.5-15 em, glabrous 
above and below, drying red-brown to green-brown; 
secondary veins (10—)15 
(609—)709—90*(—959?), 


lector vein undulating near margin, straight or 


-21(-24) pairs, diverging 
brochidodromous with col- 


slightly arcuate, prominulous below, glabrous, the 
axils lacking domatia or hairs; tertiary veins in- 
conspicuous to evident, orthogonal reticulate to 
percurrent. Inflorescences terminal or pseudoax- 
illary, umbelliform panicles of cymes (Fig. 7f); pui: 
icle branched to 4-5 degrees; main axis 1.5-5.5 
cm long, the peduncle usually lacking or to 2 cm 
long; sec ondary axes in (4-)5-6 ranks, the first- 
rank axes 2, 1.5-5.5 cm long, the second-rank 
axes 2, 1-3 cm long, the third-rank axes 2, 0.6- 
2 cm long, the fourth-rank axes 2, 0.3-1 cm long, 
the fifth-rank axes 2, 0.3-0.5 cm long, the sixth- 
rank axes 2, 0.1-0.2 cm long; cymes branched to 
l-2 degrees; bracts and bracteoles acute-trian- 
gular, 0.5-1.2 mm long, minutely ciliate. Flowers 
pedicellate, the pedicels 0.5-1 mm long; calyx cup- 
shaped, the tube 0.2 mm long, the lobes (4-)5, 
triangular, to 0.5 mm long, ipie or puberulent; 
corolla white, the tube cylindrical, 1.5-2 x 1-1.5 


' Part I: Introduction and Species 1-16, and Part II: Species p id Appeared: in the preceding two issues of the 


e of the M 


issouri Botanical Garden, Volume 76: 67-111, 


Center for Urban Horticulture, GF-15, University of AEN AA Washington 98195, U.S 


ANN. Missouni Bor. GARD. 76: 886-916. 1989. 


Volume 76, Number 3 Hamilton 887 
1989 Mesoamerican Psychotria, Part III 


e 

e 
- 
o. 


e © 
cS 
e 

o 

L 10 4 
300 km 
b 
Q 

105 100 95 90 85 80 
A ia | de E 4 E 


FIGURE 32. Distribution of Psychotria limonensis in Mesoamerica. 


mm, white pubescent in throat, the lobes (4)5, on rd. to Ocosingo, 300 m, 27 July 1972 (f), Breedlove 
Dr 22 (MEAT MO); Mpio. Faixa, 6-8 km NE of 


lanceolate, 1.5-2 x 0.8 mm; stamens (4)5, the 
9 y < 

filaments 1.5-2 mm mee in pins, 373.9 mm long Breedlove 28501 (ME XU, Mi )). Mpio. La Independen- 
in thrums, the anthers 0.7-0.8 mm long: style : 9 cia, valley of Sta. Elena along rd. to Ixcan, 30 km E of 
3.5 mm long in pins, A mm long in thrums, the paved rd. at Montebello, 800 m, 29 Nov. 1976 (fr) 
Breedlove 4197 1 (MEXU); Mpio. Ocosingo, ie ont. to 
a \ : runa Oc oe 800 m, 27 Nov. 1972 (fr), Breed- 
4(-4.5 long, (3-)3.5(-4 liam., matur- guna ad 

A Dc cum Mu and ler 29855 (MENU, MO, NY); Mpio. Oco- 
ing red, drying dark red-brown; persistent calyx zocoautla de Espinosa, 18-20 km N of Ocozocoautla along 
inconspicuous or a beak to 0.2 mm long; seed dorsal to Mal Paso, 800 m, 20 e 1971 (fr), Breedlove 
& Thorne 20976 (MEN ; NE of € haps nango, 
410 m, 2 Aug. 1965 (sry PN "Chavelas 194 

um ES : MEXU); along gravel rd. ana Palenque and Bo- 
pletely divided longitudinal furrows. nampak, 88.90 mi. SW of Palenque, 350 370 m, 5 
July 1977 (fl), Croat 10211 (MO); Río Lacantum 2 km 
. : por arriba de la unión con el Rio Chajul, 16?05'N, 
through Panama, especially southern. Mexico to 90°57’W, 19 Apr. 1976 (f : 


branches linear. Fruit when dry ellipsoid, (3.5-) 


= 


surface with 4-5 deep often irregular longitudinal 


furrows, the ventral surface with 2 often incom- 


Distribution (Fig. 32). Common from Mexico 


Sr 
a 
E 
a 
~ 
= 
S 
o 
Pe 
UH 
~ 
~ 
= 
I 
A 
= 


western Honduras. The species occurs at e levations dg ). OANACA: Dto. Pochutla, Cafetal Concordia, 700 
of 0-1,700 m, most frequently at 700 m and , 14 Apr. 1917 (fl), Conzatti et al. 3001 (GH, MEX U); 
T camino al Naranjal, 12 Sep. 1947 (fr), Mi- 
randa 4208 (MEXU — 2 sheets). PUEBLA: Mpio. Hueyta- 

ialco, Rancho Ss ripa 17 Aug. 1975 (fr), López 
(usually equatorial-tropical) climate. Psychotria li fomes id: FUMEXUX km 247 de la carret. México Te 


below, and in tropic al moist to wet to sometimes 
premontane wet forest with equatorial to tropical 
monensis has been collected in flower brane ee pan, 27 Feb. vu (fr), Sarukhán et v 1. (MEXU). 
the year—it appears to flower throughout the year SAN LUIS Porosí: Huic ‘ps ayan, km 402 p la carret. 
in Panama but just primarily May- August in the México- Nuevo Laredo, 250 m, 30 Oe. JN (fr), R. 
jut sf tte i Fruns fur e ý occa MES ve Cruz C. 1448 (ENCB). Tabasco: Cerro las Campanas 3 
res o i s range. ruits orm throughout the year, |, p, f Teapa, 50-100 m, 12 Aug. 197 4 (fr d. JJ amd 
primarily August April. et al. 2890 (MO); Paca Retiro, 19-25 June 1939 
(fl) Matuda 3437 (A, F, K, MO, NY). VERACRUZ: Mpio. 

Selected. specimens examined. MEXICO. CHIAPAS: San Andres Tuxtla, Laguia Tisatal 3 km al NE Ta T apan, 
Mpio. Rayón, Selva Negra 10 km above Rayón Mezcalapa 400 m, 19 June 197 202 dr aman 0205 (F, MEXU) 
alins rd. to Jitotol, 1,700 m, 13 July 1972 (f), Breedlove Mpio. Playa Vicente, 3 Sep. ¿1 (fr), LM he et al. 
20 139 (MEXU); Mpio. Pale nque, 6-12 km S of Palenque E£S-4125(MEXU); camino Ve pio a Catemaco, 12 km 


888 


Annals of the 
Missouri Botanical Garden 


de Catemaco, 350 m, 6 July 197 7 (fi), Fay & C. Her- 
nández 826 (F, NY, US S); km 156- 7 carret. Veracruz 
Coatzacoalcos, 27 Feb. 1960 (st), González L. & Garza 
604 (MEXU); Mpio. Hidalgotitlán, brecha Hermanos Ce- 
dillo- La Escuadra, 150 m, 26 Aug. 1977 (fl), Márquez 
346 (ENCB); Estación de Biologia Los Tuxtlas, Sep. 1978 
(fr), Torquebiau 1012 MERNE Mpio. Minatitlán, 6.9 
km al N de la terraceria La Laguna- Rio Grande, 17°20'N, 
94°22'W, 130 m, 13 July 1980 (A ), Wendt & Villalobos 
2500 (ENCB, MEXU). GUATEMALA. ALTA VERAPAZ: Semo- 
coch, 16 km from Sebol on Cobán rd., 14 May 1964 
(fl), Contreras 4679 (MEXU); 8 km W of Tucurü, 600 
7 (f). Croat 41510 (MO); 6 km NE of 
20 July 1977 (f), Croat 41594 (MO); 
Cline, 350, m, July 1903 (A), Tue peng 8404 (US — 
2 sheets) ESCUINTLA: Torolá, 300 m, Mar. 1890 (fl), 
Donnell-Smith 2042 (F, GH, US» sheets); between 
Rio Jute and Río Pantaleón, on rd. between Escuintla and 
Sta. Lucia Cotz, 540-720 m, 24 Jan. 1939 (fl), Standley 
63017 (A, NY); wooded barranco of Rio Gavilan, NE of 
Ese 'untla, 720 m, 16 Mar. 1941 (fl), Standley 89562 
( 2 sheet IZABAL: El Estar bordering Río Sarco, 24 
May 1975 (fl), Lundell & Contreras 19352a (LL); be- 
tween Escobas and Montana Escobas, across bay from 
Puerto Barrios, 1 - 100 m, 13 Apr. 1940 (fl), Steyermark 
39302 (F); along Río Frío and Herpes 75-150 m, 
18 Dec. 1941 (fr), Uds 41570 (NY, US). PETÉN: 
Uaxactün, 19 Mar. | (fr), Hi eee 7 NY); 
Lacandón, 3 km E on Tenosique trail, E 
(fr), Contreras 3385 (F, NY); La Libertad E. "e. 
June 1933 (6, e ll 4226 (F); Yaltutú entre Us 
>y ) 11 Nov. 1965 (fr), 4. Molina R. 
15566 (F—2 s | NY); $ San Pedro, en bui el 
camino para € bn enas a km 165, lado N salier 15 Dec. 
1970 (fr), Ortíz 1509 (ENCB, F, MO, NY. en S allende 
e p camino ak Santo Toribio, km 70, lado NE de Do si 
7 Feb. 1971 (fr), Ortíz 1595 (F, US); forest betweer 
Finca pane sind Rio San Diego and San Diego on 
Río Cancuen, 50-150 m, 25 Mar. 1942 (fr), Steyermark 
45393 (US). RETALHULEU: sin loc., May 1877 (fl), Ber 
noulli & Cario 1717 (K). SUCHITEPÉQUEZ: vic. Tiquisate, 
100 m, 17 June 1942 (fl), Steyermark 47054 (A, F). 
BELIZE. BELIZE: between Belize River bridge and Belize 
sawmill on Northem Hwy., 20 June 1973 (f), Dwyer 
11313 (F, bo 2 sheets). CaYo: Hummingbird Hwy. 
[ Belmonan 22 Jan. 1974 (fr), Dwyer & 
Liesner 12085 (F, GH, MO, NY); Valentin, e: 
valley, June-July 1936 (fl), Lundell 6250 (F, Y 
US). STANN CREEK: Mullins River-Stann Pu ia 14 
Aug. 1940 (fr), Gentle 3374 (A); Middlesex, 60 m, 14 
July 1929 (fl), Schipp 242 (A, F, GH, K, MO ds E 
TOLEDO: San José, 11 km N of Columbia Forest Statio 
13 June 1973 (A), Dwyer 11 186 (DUKE, MO). HONDU RAS. 
VTLANTIDA: Lancetilla, S of Tela, 20-50 m, 25 June 1970 
(fl), Davidse & Pohl 2183 (F, MO); base of hills S of 
San Alejo nr. Rio San Alejo, Rd m, 2 -27 A T. 


~ 
= 
a] 


— 


4 (fl), ARRA 4591 - MO: 
b d of Mt. Crd 16 July 
1938 (early f) Yuncker et al. 8467 (F, GH, K, MO, 
NY, US). CORTÉS: — de Rio Lindo, 1 km Fi 
Canaverales, 600 m, 25-26 Aug. 1955 (early fr), 4. 
Molina R. 5. a? (F): San Pedro Sula, 300 m, June ae 
(fl), Thieme 5274 (F, US); NE shore of Lake Yojoa, 630 
m, 30 Dec. 1952 (fr), L. 0. Williams & R. P. Williams 
18750 (F, DE US). sANTA BARBARA: alrededores de El 
Mochito, 22-25 July 1976 (fl), Vargas 695 (TEFH). 


YORO: rn 930 m, Oct. 1937 (early fr), Hagen & 
Hagen 110 Y). NICARAGU CHONT: ALES al 
NW de Sto. Domingo, 12°17'N, 85?06'W, 280 m, 12- 
13 May 1984 (fl), Grijalva 3822 don MANAGUA: Sierras 
de Managua, Hda. Encanto, 920 m, 1981 (fl), A. 
ones 5 (HNMN). MATAGALPA: summit of Matagalpa 
Tuma rd., 9 km E o 3 intersection, 12°57'N, 
85051" W, 1,000 m, 24 May 1981 (fl), D. Stevens 
& Henrich 20335 (MO). ZELAYA: Guamil o Bonas sobre 
areas pantanosas a lo largo del "e Grande, 0- c 
Apr. 1949 (fl), .4. Molina R. 7 (F, GH); 1 
N de Monkey Point, 11°36'N, du 5- 10 m, 27 
Oct. 1981 (fr), P. P. Moreno & Sandino 12609 (MO); 
10 km NE of Siuna gallery forest along Cano Madriguera, 
_ Ne ill 3726 (MO); nr. Rio 
5 June 1978 (fl), Neill 
Rica. LIMÓN: Port Limón, O m, 1 May 
1903 (fl), Cook & Doyle 440 (US); Isla Uvita, 13 Oct. 
1941 (fl Echeverría 46 (F); 1 mi. toward Zent from 
Moin Junction, 30 m, 1 Sep. 1946 (fr), Morley 819 (F); 
entre Barra Parismina y Rio Chiquero, 9 Oct. 1951 (fr), 
Shank & Molina 4354 (F, GH, US). PANAMA. BOCAS DEL 
TORO: Changuinola to 5 mi. S at junion of vs e 
guinola and Terebe, 30-60 m, a 19 Dec. 1966 (fl, fr), 
Lewis et al. 822 (GH, JS); Old Bank hs 21 
Feb. 1941 (fl), Wedel 2137 WF. GH, MO). CANAL AREA: 
Coco Solo, Tropic ii Center study site, 27 Mar. 
(A), Gentry & Tyson 4848 (MO, NY); Barro Colorado 
Isla ir 2. í Won 197 0 (fr), Luteyn 760 (DUKE, NY); 20 
Nov. A), Shattuck 416 (F, MO); nr. old Fort San 
Laren e Mar. 1923 (fl, 2 iud 5980 (US). c 
"ortobelo, 7 May 1971 (f y 
` MOX a to Rio Indios, : 
Pina, 50 m, 7 July 1976 Ai fr), G. Sullivan 133 
DARIÉN: ridge N of Ensenada El Guayabo, 
Cee Rio Jaque ng Es Pacific Ocean, 7?26'N, 


4465 (MO). Costa 


78°05'W, 300-500 m, 26 Jan. 1982 (fl), Knapp «€ 
Mallet 3 174 (MO). SAN BLAS: 3-4 hours up Rio Mulutupu, 
17 Aug. 1967 (fr), Kirkbride 226 (MO, NY). 


Psychotria limonensis may be recognized by 
its broad-elliptic, thin-membranous leaves, brochi- 
dodromous secondary veins with collector vein near 
margin, broad umbelliform inflorescences, and small 


— 


usually 4 mm long) ellipsoid to subspherical fruit. 
Leaves on Mexican and Guatemalan specimens 
have fewer secondary veins than those from else- 
where; the secondaries diverge characteristically 
at acute (vs. right) angles in Nicaragua. 


49. Psychotria tenuifolia Swartz, Prodr., 43. 
1788. Uragoga tenuifolia (Sw.) Kuntze, Re- 
vis. Gen. Pl. 2: 963. 1891. TYPE: Hispaniola: 
Swartz s.n. (holotype, S, n.v.). Cf. also Swartz, 
Fl. Ind. Occid., 402. 1797. Figures 2e, 33. 

Psychotria sessilifolia M. aber i rae Bull. Acad. 

oy. Sci. Bruxelles 11: 228 . Uragoga ses- 
E jun Martens & Cale eie Kuntze, Revis. 
Gen. Pl. 2: 962. 1891. s exico. Veracruz: 
Mirador b Zacuapan, 750-1,050 m, June 1840 
(fl), Galeotti 707 qe G, n.v., photo, F neg. 
258206; isoty 


Psychotria granade nsis "Bentham in Oersted, Vidensk. 


Volume 76, Number 3 
1989 


Hamilton 889 


Mesoamerican Psychotria, Part Ill 


Medi del "gos Naturhist. Foren. Kjebenhavn 1852: 
3. Uragoga granadensis (Benth.) Kuntze, 
960. 


Reis, pm Pl. 2: 1891. TYPE: oe 
Granada, Oersted s.n. (holotype, C, ., photo, F 
neg. 22833). 

Psychotria nert Small, Fl. Miami, 176. 1913. No 
type ci ite 


Shrub 0.5-2(-3) m tall; young stems glabrous 
the bark smooth; stipules ovate, 
2e), 8-13 x 


caducous, leaving pale ridge 


to puberulent, 
long-biacuminate (Fig. 3-6 mm, gla- 
brous to puberulent, 
with red-brown fringe. Leaves petiolate; petioles 
(0.3-)0.5-4 cm long, glabrous to puberulent, 
narrow- to 


grooved above; blades membranous, 


medium-elliptic, the apex acuminate or sometimes 
acute, the base attenuate, (4-)7.5-23 x (1-)2.5 

— 1.1) em, glabrous above, glabrous to puberulent 
below, the midvein puberulent below, drying green- 
brown to sometimes red-brown; secondary veins 
8-18 pairs, diverging 70°- 80°, brochidodromous 
with collector vein fairly near margin, slightly ar- 
cuate, prominulent and puberulent below, the axils 


E 


lacking domatia or hairs; tertiary veins inconspic- 
uous to evident, orthogonal reticulate, the tertiary 
loops between secondary loops and margin often 
evident. Inflorescences terminal or pseudoaxillary, 


— 


panicles of cymes; panicle branched to 3-4(-5 


degrees; main axis 1.4-4.5 cm long, the peduncle 


— 


to 2 em long or lacking; secondary axes in (2)3(4 
ranks, the first-rank axes 2, (0.9-)1.8-3 cm long, 
the second-rank axes 2, (0.4-)0.8-1.2 
the third-rank axes (1)2, 0.1-0.5 em long, the 
1-0.3 cm long; cymes 


cm long, 


axes 2, 0, 
3 degrees; bracts and bracteoles 


fourth-rank 
branched to 2- 
lanceolate to linear, 0.5- 1 mm long, puberulent. 
Flowers pedicellate, the pedicels 0.5-1.5 mm long; 
calyx cup-shaped, the tube 0.2-0.3 mm long, the 
minutely 


lobes 5, triangular, 0.3-0.5 x 0.5 mm, 


ciliate; corolla white, the tube cylindrical, 1.5-2 


x 0.8-1 mm, white pubescent in throat, the lobes 
5, triangular, 0.7-1 x 0.5-0.7 mm; stamens 5, 
Ad filaments 1-1.5 mm lona in pins, 2.5 


0.7 mm long; style 


3 mm 
long in thrums, the anthers 0.5- 
2-3 mm long in pins, 1.5 mm long in thrums, the 


branches linear. Fruit ellipsoid when dry, 4-5 mm 
long, 3-4.5(-5) mm diam., maturing red, drying 


eil eater persistent calyx inconspicuous or a tu- 
bular beak to 0.5 mm long; seed dorsal surface 
with 4-5 shallow longitudinal furrows, the ventral 


surface with 2 longitudinal furrows. 

Distribution (Fig. 33). 
Mexico through Panama at 0-2,000 m elevation, 
mostly below 1.000 m, in tropical dry to premon- 


Widespread from 


tane moist forest with equatorial to tropical climate. 
Psychotria tenuifolia occurs also in Florida, Cuba, 


and Hispaniola. It has been collected in flower 
throughout the year, primarily May July. and in 
fruit throughout the year, primarily August Feb- 


ruary. 


Selected specimens examined. MEXICO. CAMPECHE: 
Tuxpena, 16 Feb. 1932 (fr), Lundell 1339 (F); Cham- 
P dn. July 1932 (fl), Steere 1586 (F, MEXU); km 
de la carret. Escárcega € id eum M la terraceria 
iam Carranza, 5 Feb. Tellez et al. 
62 203 (MEXU). CHIAPAS: Mpio. slopes nr. 
Habenal, paraje of Mahben E 1,050 m, 26 Nov 
1964 (f P Eraeatooe £626 (ENCB, NY): Mpio. Ocozo- 
Selva del D 32 km NW of 
m ront, 600 m, 27 1972 (fr), ee 
MEXU, MO): Mpio. eee inta, 11 i 
of E alo ui e to La sri and Chicoasén, 1,000 
m, 23 Feb. 73 (A), Breedlove 33806 (MEXU, 
ee aa de od nr. Fr ; 
km N of Cintalapa, 1,250 m 5 Aug. 1974 (fr), Breed- 
ds 3003 o ) Mpio. C sara Playas de Catazaja, 
m, 20 ay 1978 (fl), Sanches 1029 (ENCB). 
re : : a RO: fed de Oca, Va 7 
(fl), Hinton et al. 10610 ^K. NY, 
meld. 8 Sep. 1939 (fl), Hinton et al. la E 9 3 (F, MO 
Acapulco, 13 Nov. 1956 (fr), Paray 2310 (ENCB). ni- 
DALGO: at San Luis Potosí border, km 343 
below Chapulhuacán, rocky slopes, 6020 m, 30 June 19: i 


Ta 6s Ps , 


~ " 


(A), Moore & Wood 3660 : A, MEXU). JaLisco: 10- 
mi. S of Talpa de Allende, headwaters of Rio de Talp ya, 
3 mi. above jp Sauces, steep mountains, 26 Nov. 19 60 


(fr), MeVaugh 21463 (LL). MICHOACÁN: Mpio. Apatzin- 
gan, Tancitaro region, Hda. California, 1,200 m, 12 Aug. 
1941 (fl), Leavenworth & i iar 1400( F). OAXACA: 
18.4 mi. 5 of Val Hu T3 p Sierra 
de Juárez 197 ^ (A), Croat 48077 
(MO); Dto. iude predio particular Cafetal Natividad 
y Anexas, San José de las Flores, 810 m, 14 Nov. 1979 


,9 re 
(MEXL a Temas 'al, 20 May 1962 
(M EAS. PUEBLA: km 247.3 de la carret. ! 
, 1 May I 962 (fl). Sarukhán el rar 
Di! 
Lunde ll & Lundell ; 7677 í S) 
11 Jan. 1980 (fr), Tellez 1 Calrera l 205 (MEXU). san 
LUIS POTOSÍ: Tamazunchale, 12 July 
wards 533 (F, MO). 13 mi. S 
VERS 2 Sep. 1948 (fr), old & Crum 3954 (A) 
Mpio. de Ciudad del Maiz, 0.5 km NE de Las Abritas, 


800. m, 30 June 1959 (fl), Es edowski 11126 (ENCD). 
TABASCO: a 12 an de Villahe rmosa, por la carret. a 
Escár 1D aw 5 (fr), L onzülez L. & Pérez 


4004 VENC B, ME U); Balancán, San Isidro, 7-11 J 
1939 (fl), Matuda . E A.F, SXU, NY). TAMAULIPAS: 
nr. Nuevo Morelos, 1 Sep. 1948 (fr), Kenoyer & Crum 
1002 (: , nacimiento del 
Rio Sabinas, , Valiente et al. 249 (MEXU). 
VERACRUZ: ber 20 carret. Alemán ( sosamaloapan, 13 Nov. 
1964 (fr), Chavelas i Pérez 223 (ENCB); San Lacen nzo 
Tenochtitlán, 30 Nov. 1967 (st), Chavelas et al. 
2469 (MEX U); Mpio. de ie wma, a 8 km de Ozulauma 
hacia ee | Dec. 70 (fr), Chiang 250 (MEXU); 
20 mi. W of Tuxpan on a from Alazán, 5 Jan. 1904 
(fr), Cor n & Correll 28710 (LL); Mpio. de Coatzintla, 
Palmar de Zapata, 110 m, 11 Oct. 1982 (fr), M. Cortés 


> 
= 
> 
pr 
p 
d 
8 
e 
f 
= 
2m 
£s 
= 
E 
E 
= 
> 
a 
< 
& 


890 


Annals 
Mesa nad Garden 


——— -T T T T 
PN \-+- T.C.4 
| A Ü 
ice 
E o, 20 4 
. de y + 
a ve o 
e 
e 
. & ? ° 
1 + g ^ = 5 s 
ES + A] "m 
e? % 
- 
s] 
300 km + p 
ove 
105 100 95 90 85 s 80 


FIGURE 33. 


195 (MEXU); Mpio. Papantla, Cerro del Carbón, 240 m, 
11 Dec. 1981 (fr), M. Cortés 60 (MEXU, MO); Mpio. 
Espinal, brecha Entablader ro ro l km antes de Ojo 
de Agua, 140 m, 11 Nov. 1981 (fr), Cortés-Vázquez 48 
(MEXU) a 11.6 km de B de las Hayas, 700 m, 24 
, Lot et al. 2022 (F, MEXU ; Mpio. 
Totutla, 3 km SE of El Mirador, 19°1 2'N, 9655 LW, 900 
, Nee 22995 (MEXU, MO); Mpio: 
P 5 km NE of Tempoal, 50 m, 22 
> 1980 (fA), Nee & Danse (n 18372 (FE); Zac uapan, 
. 1906 (fr), Purpus 2268 (GH, 
, US); Mpio. Minatitlán, orilla del Río Uxpanapa, 
un poco ahaha del poblado de Uxpanapa, 1 a m, 26 May 
1981 (fl), Wendt et al. 3322a ( MEXU ). YUCATÁN: 
rida, Cance rillo, 6 July 1805. (fl), . 7 

) ALTA VERAPAZ: Seno! 


ic. Laguna Sapalá (Chajvovuch), 
: 280 m, 11 Mar. 1942 2 (st), id 
Cubilgüitz, Aug. 1903 (A), 
heim 11-796 (F, US). BAJA VeRAPAZ: Unión a 4 
972 (fr), Contreras 11 101 (MO). CHIMALTEN 
nr. Sibaja, 1,050 m, 6 Jan. 
(F). CHIQUIMULA: 4 km N of "d n 
1,100 m, 27 bd 1971 (fr lina R. & 4. 
CUINTLA: 6 mi. zi of Escuintla on de to 
Alotenango, 650: n, 27 July 1977 (early fr), Croat 42052 
(MO); below Las Lajas, 900-1,200 m, 9 Feb. 1939 (fr), 
Standley 64775 (F). HUEHUETENANGO: along Río pos 
below Jacaltenango, Sierra de los Cuc humatanes, 1,40€ 
m, 4 Sep. 1942 (fr), Steyermark 51862 (F —2 she A 
IZABAL: Puerto Méndez/ Cadenas, dii Rio Cadenas, 


— 


Qda. Resimie nto, 
Molina 


Distribution of Psychotria tenuifolia in Mesoamerica. 


3 June 1970 (fl), Contreras 9932 (MO); Modesto Méndez, 
30 m, 13 Mar. 1970 (fl, fr), Harmon & Fuentes 2090 
(F, GH, MO); Finca Murcielago ca. 16 E a of El Estor, 
8 July Vd, oe Oed E 18 c. Quiriguá, 
15-225 m, 15-31 2 db. "dun Y 9. 4045 (US). 
JUTIAPA: nr. P? Molino. Dept. Santa Rosa), Ae m, 26 
Nov. 1940 (fr), Standley 78446 (F). PETÉN: Tikal Na- 
Sanal Park, in secondary growth bue airfield, 23 
. Contreras 115 2 (LL) Dolores, on old 
16 Sep. 196 1 (fr) Contreras 2914 
(LL); Macanche, dn 34-35 of Melchor de Mencos Rd., 
24 May 1966 u^ uu 5655 (GH); dos Rd., 
m 82 367 (fr), Contreras 7242 (MO); Say- 
xché, Cabal, 17 MS 1970 (fl), Contreras 9773 (MO); 
Paso Caballos, 80 m, 14 July 1956 (fl, 
Guzman 2892 (ECON); La Cumbre, 
1976 (fr), Lundell & Contreras 202. 
mino a La Libertad, a km 7 lado E del camino, 1 3 
Sep. 1971 (fl, early fr), Ortíz 1929 (F, US); en el camino 
a la aguada Paxcaman, a 3 km lado N de Uaxactün, 17 
May 1973 (fl), Ortíz 2572 (ENCB, F, MO, NY). SANTA 
ROSA: nr. e 900 m, 20-27 Nov. 1940 (fr), Stand. 
ley 27958 (F); S of Guazacapán, along the Avellana rd., 
150 m, 6 Dec. 1940 (fl), Standley 79472 (F). BELIZE. 
on Western Hwy., 5 July 1973 (fr), 


~ 
pæ 


N 
2 
— 
ZO 
c 
= 


BELIZE: mi. 34.5 
Dwver ves 7O(F, GH, MO, NY); Bomba, Northern River, 
Dec. 1933 (A), Gentle 1034 (A—2 sheets, F, GH 
MO, NY); Aud nr. Manatee Lagoon (Southern Lagoon), 
5 Sep. 1905 (fl), Peck 959 (GH, K, NY). cayo: El C 
Mar. 1933 (fr), Lundell 3984 (F); 
of Western and Purus) dn hwys. 
18 July 1970 (f), Sp an & Newey 175 

COROZAL: 1931-1932 ^ "d Gentle 226 (F). ORANGE 
WALK: N of Orange Walk, vic. Nev 23 June 1973 


T 


ayo, 

4 km 5 of junction 

; quarry hill, 75 m, 
3 (MO 


v River, 


(fl), Gentry 8552 (MO); Freshwater Creek Reserve, co- 


Volume 76, Number 3 
1989 


Hamilton 891 


Mesoamerican Psychotria, Part III 


hune ridge, Feb. 1933 (fr), Pelly 9 (F). TOLEDO: N n 
of Río Sarstun, opposite Modesto Méndez, 10 m, 29 Jur 
1970 (early fr), Harmon & Dwyer 2844 (MO). HONDURAS. 
COMAYAGUA: S shore of Lake Yojoa, 2] Aug. 1973 (fr). 
Hazlett 684 (MO —2 sheets); Farallón El Pizote, Cerros 
El Cedral, 6 km NE de Taulabé, 1,940 m, 28-29 
1975 (fr), C. Nelson & Vargas 2859 (MO, TEFH); Agua 
Caliente, vaguada de rios beo y Humuya, 35 km E 
f 220 m, 22 Nov.-31 Dec. 1980 (Ir), G: 
Velson et al. 6. 500 (TEFH). coPÁN: Copán River 2 km 
` of à s, La Vegona, 500 m, 23 Nov. 1969 
na R. & Molina 24794 (F). CORTÉS: 
Montana La Cumbre, Cordillera de Omoa al NE de San 
Pedro Sula, 200 m, 19 Apr. 1956 (fl), 4. Molina R. 
t ? (F, US); orilla del Rio Humuya, 40 km N Santa 
ruz de Yojoa, 100 m, Nov. 1980 (fr), C. Nelson et al. 
En 38 (TEFH). FRANC e O MORAZÁN: El Zamorano, 800 
m, 13 June 1945 (fl), . ad cinis 3054 (F); Sabana 
Grande, 1,100 m, 26 e 1945 (st), J. |}. Rodríguez 
3270 (F). ISLAS DE La BAHIA: E end of Utila Island, 5 m, 
15 Aug. 1954 (fr), Johannessen 499 (TEFH). SANTA 
BARBARA: nr. Santa Barbara, thickets along Sesecapa ‘Riv 
er, 500 m, 22 Aug Molina R. 22021 
(NY —2 sheets). YORO: Aguan River valley, vic. Coyoles, 
ur. Medina, 200 m, 26 July 1938 (fl), donis et 
8628 (F, GH). El SALVADOR. AHUACHAPÁN: vic. Ahua 
chapán, 800- l, E m, 9-27 Jan. 1922 (f), under 
2 US). SAN SALVADOR: VIC. 
. 1922 (fl), Standley 22739 


— 


(G H, 
, Standley & Padilla 3776 
(E). SONSONATE: vie 1922 (fl), Stand- 
ley 21849 (US). NICARAGUA, CHINANDEGA: cauce pluvial 
al SO del Volcán Cosigúina, 29 Sep. 1981 (fr), Fonseca 
131 (MO). GRANADA: Vales an Mombacho, 2 km al SE de 
la hda. Cutirre, "El Chipote," 48'N, 85?554'W, 
m, 11 June 1982 2 (fl, fr), P. P. Moreno 16450a, 10450b 
(MO). LEÓN: Volcán Santa Clara, 18 Aug. 1981 (fr), 
Quezada 52 (MOL i \NAGUA: 7 km W of airport on Hwy. 
l to Tipitapa, nr es 150 m, 17 Mar. 1977 
nd fr), A "oe (MO). Masaya: Sierra de Man- 
gua, region of Las Nubes, 650-950 m, 14 May 1947 
$, Standley & Garnier 8403 (F). MATAGALPA: Paiwita, 
15 km al E de Matiguas, yir a 12252 
"500 m, 5 May LP. MG no 24 Y 19 
(MO): 20 km E of eae Rio Yasica, nr. rd. to E 
Tuma, 25 May 1977 (fi), Veil 1980 (MO): along rd. 
between Muy Muy an Pa e eee ca. 9.5 km W of 
bridge at Muy Muy, at e, 12245'N, 85°42'W, 
310 m, 30 July 1978 (fl), D. Cod 0545 (MO); 
Qda. Sta. Cruz, có D. Viejas and ka 
12°35'N, 85°50'W, 420 m, 27 July 1983 (f), W. 
D. Stevens 223. 38 (MO). NUEVA SEGOVIA: El ic “Fila 
de Monterric o," 13?44'N, 86°04" 
1981 (fr), P. P. Moreno 13759 (MO). ZELAYA: region of 
p ah s Bluff (Puerto Cabezas), 1928 (fl), Englesing 
228 (F); descent of Cerro Saslaya, down Cerro El Inocente 
Et o El Horas ro, 300 600 m, 6 May 1977 (fl), n 
1905 (MO); Cerro Waylawas, 13°38 N, 84°48'W, 
150 m, 9 Mar. 1979 (bud), Pipoly 4171 (MO). Rin 
Rica. ALAJUELA: San Miguel de San Ramón, May 1934 
(fl), ien 19218 (CR, F, NY); Guanacaste: 28 km N 
` Cañas. OTS area, 100 m, 2 Oct. 1969 (fr), Frankie 
296a (MO); Comelco area N of Bagaces, nr. Potrero 
ae O June 1971 (fl), Ge ittry BOTE, MO, US); Comelco 
of Bagaces, 23 May 1971 (fl), Hetthaus 113 (C R, 
SN Santa Rosa National Park, 10%50'N, 85?37'W, 0- 


rT] 


p= 
me 
o 


— 


~ 


7 km 
5 Feb. 


320 m, 20 Aug. 1981 (fr), Janzen 12122 (MO): 
SW of Curime, 10?04'N, ie 31'W, 100-300 m, 


197 8 (fr), A dv 1994 (MO). PUNTARENAS: Los Bar- 
rancas, along Pan-Am. Hwy. nr. Miramar turnoff, 2 June 
171 (fl), Gentry T78 MON Monteverde, La Pa Pas. 5 


km N of Canas, 
PANAMA. CANAL AREA: 
(fr), Croat 7316 (MO); 8 June 


June 1979 (fl), Haber 380 (MO). 
Barro Colorado rue 8 Jan. 1969 
White 123 
lache Bridge, 
: (fr), a 
& Palmet 2051 (MO): Balboa, i ae Jan. 1924 
(fr), Standley 25566 (US); Las Cascadas Plantation, nr. 
Summit, 2 Dec. 1923 (fr), Standley 25087 (US); 
France Field and Cativá, 9 Jan. 1924 (fr), Standley 
30312 (US). DARIÉN: Río Chico, 1 hr. by iis from 
Yaviza, 19 Dec. 1966 (fr), Burch et al. 1118 (MO). 
PANAMA: dam site nr. Rio Bayano beyond Carita, a Sep. 
1971 (early fr), Gentry 
of Panam. Hwy.. 90D0N, 
1982 (early i je ud >: 5 (CR, ur 
of Panam. Hwy., 9903'N, 78?25'W, ca. 100 m, 
1982 (fr), Halón & D’ e 13280 MO) Chimán, 12 
Dec. 1967 (fr), Lewis et al. 3310 (MO); Taboga X 
Dec. 1923 (fr), Standley 27049 (US); between 
banas and Matías Hernández, 21 Jan. 1924 (fl), Standley 
). 


1 (US 


between 


As 
ur 


This variable species includes P. sessilifolia 
(Mexico; 
ragua; broad leaves). and P. sulzneri (Florida; small, 


narrow leaves). P. granadensis (Nica- 


narrow leaves). 


Psychotria tenuifolia may be recognized by its 


narrow-elliptic, thin-membranous leaves; brochi- 
dodromous secondary veins with collector vein near 
the margin; small umbelliform panicles of cymes: 
and small (4-5 mm long) ellipsoid fruits. Specimens 
from Honduras and further north are more pu- 
berulent, especially on the leaf underside, 
inflores- 


from Nicaragua and south. Honduran 


cences have especially conspicuous linear bracts. 


GROUP 8. THE C4ALOPIDLLA GROUP 


Shrub or subshrub or herb (sometimes P. ham- 
melii and P. rosulatifolia) or small tree (P. cal- 
ophylla); young stems glabrous or red-brown pu- 
bescent; stipules usually sheathing, ovate or 
truncate with apex usually biaristate or bifurcate 
(Fig. 2g), uniform in color, glabrous or ferrugin- 
eous-pubescent (or fringed in P. insueta), caducous 
or often persistent at terminal 3-5 nodes in P. 
hammelii and P. a cos Leaf blades el- 
liptic or obovate or oblanceolate (or lanceolate in 
P. rosulatifolia). drying red-brown to green-brown, 
sometimes paler below; secondary veins (7—)10- 
25(-42) pairs, 5°-)50°-85°(-1 10°), 


usually brochidodromous with collector vein (Fig. 


diverging (45 
3c) or sometimes without collector vein or eu- 
camptodromous, the axils lacking domatia; tertiary 
veins orthogonal reticulate (or sometimes percur- 


892 Annals of the 

Missouri Botanical Garden 
F -4- a } | TOA 
@ 204 


300 km 


b 
105 100 95 90 85 * izo 
L sk l L 
FIGURE 34. Distribution of Psychotria calophylla. 
rent in P. calophylla, P. insignis, and P. rosu-  alfaroana—shows any between-morph asymme- 


latifolia). 
glomerules, pedunculate except for P. insignis, P. 
psychotriifolia, and often P. calophylla; second- 
ary axes in 1—-2(-3) sometimes size-differentiated 
pairs per rank and/or in clusters (P. alfaroana, 
P. chitariana, P. dressleri; Fig. 7d), delicate only 
in P. rosulatifolia; bracts often conspicuous. Co- 
rolla tubes (1.5-)2-5 mm long (10-12 mm in P. 
dressleri), the lobes usually with apical exten- 
sions, protuberances, or thickenings. Fruit when 
dry ellipsoid; persistent calyx inconspicuous or 
a beak or a tube to 3.5 mm long; seed dorsal 
surface with (3—)4—5 usually deep longitudinal fur- 


rows, the ventral surface with 2 shallow, often 


incompletely divided longitudinal furrows. 
34) and P. 


psychotriifolia (Fig. 35) are not narrow endemics; 


Only Psychotria calophylla (Fig. 


only the former has been collected north of Nica- 
ragua. Psychotria rosulatifolia and P. dressleri 
occur in eastern Panama (Figs. 35, 36) and share 
inflorescence secondary axes often in size-differ- 
entiated pairs, and therefore may be closely related; 
but phylogenetic relationships among these mostly 
narrowly endemic species are difficult to hypoth- 
esize. 

Of the twelve species in the group, four are 


normally distylous; only one of those — Psychotria 


Inflorescences panicles of cymes or of 


try in floral part length. Psychotria rosulatifolia 
appears pin-monomorphic, based on five flowering 
collections examined. Psychotria monteverdensis 
appears long-homostylous. The remaining six species 
are represented by few or no flowering collections 
and cannot be assessed. 


50. Psychotria alfaroana Standley, J. Wash. 
Acad. Sci. 18: 273. 1928. TYPE: Costa Rica. 
Guanacaste: El Arenal, 500 m, 18 Jan. 1926 
(fr), Standley & Valerio 15179 (holotype, 
US). Figure 36. 


Subshrub 0.3-0.0 m tall; young stems glabrous, 
the bark irregularly furrowed longitudinally; stip- 


ules sheathing with the sheath 5-7 mm long, biar- 


istate with the extensions 7-12 mm long, glabrous, 
caducous, leaving a pale ridge with red-brown fringe. 
30) mm long, gla- 


membranous, ob- 


Leaves petiolate; petioles 7-25( 
brous, grooved above; blades 
ovate, the apex acuminate to cuspidate, the base 
attenuate, 11-29 x 4.5-10 cm, glabrous above, 
glabrous below with minute organic detritus ad- 
hering to midvein, drying dull green; secondary 
veins (7-)1 1-14(-16) pairs, diverging (60?—)70?— 
80%—90°), eucamptodromous, 
vein—then straight—then arcuate near margin, 
or constantly arcuate, elevated below, glabrous, 


arcuate at mid- 


Volume 76, Number 3 
198 


Hamilton 893 
Mesoamerican Psychotria, Part III 


12 4 
o 
e. e 
1 4 
- , | 10 4 
Q y d 
v LA 
o oQ 7 B 4 
100 km M (y 
& 7 de 
85 84 83 82 81 80 79 78 | 
L L n L 4. L L L L 7 


FIGURE 35 
(open circle), 


the axils lacking domatia or hairs; tertiary veins 


evident, orthogonal reticulate. /nflorescences ter- 
; panicle 
)2-4 cm 


secondary axes 


minal, compact- EN d panicles of cymes 


)ranche« o 2 degrees; main axis 
I hed t leg ( 


long, the MR 1-3 cm long; 


in | cluster, several, 5-7 mm long: cymes branched 
to 1-2 degrees: bracts not evident. Flowers ped- 
) 


icellate, the pedicels 1-2 mm long: calyx drying 


pale brown, cup-shaped, the tube | mm long, the 
lobes 5, | mm, glabrous to sparsely 
2.5-4 
X 2 mm, white pubescent in throat, the lobes 5, 
2-3 x 


linear, 1-2 x 


ciliate; corolla white, the tube cylindrical, 


linear, thickened at apex, mm; stamens 
5, the filaments 2.5 mm long in pins, 6 mm long 
in thrums, the anthers 0.8-1.2 mm long; style 4-5 
thrums, the 


8-10 


maturing red, drying black: 


mm long in pins, 3.5 mm long in 
branches linear. Fruit when dry ellipsoid, 
mm long. 4 mm diam., 
persistent calyx 1-3 mm long, drying pale brown; 


seed dorsal surface with 4-5 deep irregular lon- 
gitudinal furrows, the ventral surface with 2 irreg- 


ular, incompletely divided longitudinal furrows. 


Distribution (Fig. 36). 
Costa Rica, from upland Guanacaste to southeast- 


Fairly widespread in 


ern Limón, collected in tropical moist forest with 
equatorial to tropical-equatorial climate at eleva- 
tions of 100 to 900 m. Psychotria alfaroana has 


Distributions of Psychotria insignis (solid squa 
P. psychotritfolia ( (solid circles), and P. rosulatifolia (open triangles) in Mesoamerica 


re), P. insueta (solid triangles), P. monteverdensis 


and 


March, 


August, 


been collected in flower January, June. 


July, and in fruit July (immature). January, 


and February. 
Selected specimens examined. Edw TA RICA. ALAJI E- 
v2 km N of Bijagua on trail to Si 
of Rio Naranjo, 400-500 m, 8 Jul; | 
Utley € A. Utley 5310 (DUKE). CARTA 


900 m, 7-8 Feb. 192 26 (fr), Standle y «| 5 
JS). GUANACASTE: Silencio, nr. Tilarán, 750 m, 13 
an 26 (fr), Standley & Valerio 44003 (US), El 


l alerio 62 (US). 


Arenal, . 
, OTS field station, Rio Puerto 


(DUKE); 17 Mar. 19€ 80 (fl), CE MOV (DUKE); 4 
Aug. 1980 (fr), Viu 944. 8 Aug. 
(fr), Hammel 9489 (DUKE): s ise ca. 10 km 
from Cariblanco a from bridge on way to La 


Virgen del Socorro, 10 June 1982 (fl), di dn et al. 
irs (DUKE). Limón: Los Diamantes, Rio Sta. Clara, 


6 km E of Guapiles, ca. 200 m, 9 July 1949 | " Holm 
5 due 354 (MO); 2 km N of Bribri, along Rio Sand Box, 
18 July 1976 (early fr), J. Uie; & K. Utley 
IKE). TORO: Rio Teribe nr 
Qda. Lukulon, 1968 (fl), Kirkbride 
& Duke 511 (M 


Psychotria alfaroana may be recognized by its 


suffruticose habit: leaves drying. green; compact 
globose, pedunculate inflorescences with clustered 


secondary axes only 5-7 mm long: and large (8 


X 4 mm) ellipsoid "s usually drving black. 


894 


Annals of the 
Missouri Botanical Garden 


12 4 
o 
114 
- e 10 4 
e 
6 . 
o m 94 
al 
Jo 84 
100 km " ^ 
—— 
oe 
85 84 83 82 8i 80 79 78 7 
Fic ;36. Distributions of Psychotria alfaroana (solid circles), P. chitariana (open square), P. dressleri (open 
Tun ) P. hammelii (open circles), P. pacorensis (solid square), and P. sixaolensis (solid triangles). 


The corolla tube of the pin morph appears con- 
spicuously shorter (2.5 


thrum (3-4 mm). 


3 mm) than that of the 


Əl. Psychotria calophylla Standley, Contr. 
U.S. Natl. Herb. 18: 129. 1916. TYPE: 
ama. Panamá: Chagres, Isthmus of Panama, 
15 Mar. 1850 (early fr), Fendler 60 (holo- 
type, US; isotypes, GH, MO). Figures 3c, 34. 


Pan- 


Psychotria cerroaz zule nsis Dwyer, Ann. joe Bot. 
Gard. 5; . 1966. TYPE: Panama. Panamá: Ce- 
rro Azul, Goofy Lake area, 1 Sep. 1963 (fl), je 
2875 (holotype, 

Psychotria pd Dwye sr & Hayden, Ann. Missouri 
Bot. Ga à 5 1967. TYPE: Panama. Cana 
rem are db ón ee ge un Locks and Gatun 
Lake, 5 ix 195 5 (ea rly fr), Johnston 1554 (ho- 
lotype, MOS isotypes, LS, d ts). 

Psychotria grandicanpe Dwyer & Hayden, Ann. 
souri Bot. Gar« 


Mis- 


2 Mar. 1967 (fr), 
4 (lectotype, ar 
i. 'totypes, MO 


Dwyer et al. 
2002447, here designated; 
2 sheets). 


Tree 3-8(-15) m tall; young stems brown-pu- 
berulent, the bark sme stipules ovate with ir- 
regular lobes and 2 aristate to narrow-triangular 
extensions 3-6 mm long, 8-15 x 5-15 mm, 
fringed, densely red-brown pubescent, caducous, 


leaving a pale ridge with red-brown fringe. Leaves 


petiolate; petioles 1-3 cm long, red-brown pubes- 
cent, grooved above; blades membranous to sub- 
coriaceous, elliptic to obovate, the apex ac uminate, 
the base attenuate, (10-)13-26(-31) x 


glabrous above, red-brown tomentose to sometimes 


x 5-12 cm, 


glabrous below, drying green-brown above and red- 
veins (8-)10-15(-18) 


brochidodromous with 


brown below; secondary 


pairs, diverging 50°—-75°, 
collector vein undulate (Fig, 3c), constantly ar- 


cuate, elevated below, red-brown tomentose, the 
axils lacking domatia or hairs; tertiary veins in- 
conspicuous to evident, orthogonal reticulate to 


percurrent. /nflorescences terminal, panicles of 
glomerules: panicle branched to 3(4) degrees; main 
axis 2.5-15 cm long, the peduncle lacking or 1.5- 
axes in (2)3(4) ranks, the 
first-rank axes 2, 1.2-8 cm long, the second-rank 
axes 2, 0.6-4 cm long, the third-rank axes 2, 0.2 

2.5 cm long. the fourth-rank axes 2, 0.7 


long: bracts inconspicuous, triangular, | mm long, 


4.5 cm long; secondary 


] cm 


brown pubescent. Flowers sessile; calyx cup-shaped, 
the tube 1-2 mm long, the lobes 5, broad, barely 
evident, to 0.5 mm long, red-brown pubescent; 
corolla green-white, the tube cylindrical, 3.5-4 x 
2-3 mm, very densely white pubescent in throat, 
5(6). 


2.5 X 0.7 mm; 


puberulent without, the lobes ovate to lan- 


ceolate, thickened apically, 2 sta- 
mens 5(6), the filaments 2.5-3.5 mm long in pins, 


Volume 76, Number 3 
1989 


Hamilton 895 
Mesoamerican Psychotria, Part III 


4.5 1.5 mm 
long; style 4-6 mm long in pins, 
thrums, the branches linear or club-shaped. Fruit 
when dry ellipsoid, 10-14 mm long, 6-9 mm diam., 


maturing red, drying red-brown, sometimes red- 


mm long in thrums, the anthers | 
2 mm long in 


brown puberulent: persistent calyx 2-3.5 mm long; 
seed dorsal surface with 4-5 longitudinal furrows, 


the ventral surface with 2 longitudinal furrows. 


Distribution (Fig. 34). Known from southern 


Mexico and Izabal, Guatemala, and disjunct to 
Colón and Coclé to Panamá Province, 
This species is found at elevations of 30 to 1,000 
m, mostly below 400 m, in tropical moist to pre- 
montane wet forest with equatorial to tropical cli- 
mate. It has been collected in flower in March and 
May in Mexico and July September 
cember in Panama. Fruiting collections date from 
December in Guatemala and August and Novem- 


Panama. 


and in De- 


ber- March in Panama. 


Selected. specimens examined. MEXICO. OAXACA: 
Mpio. Matias Romero, 3.2 km al 550 del Ase pias La 
Floresta, 16 km al 5 de NIMMT 17?03'N, : 
300 m, 28 Ses 1981 (A), Fendt et al. 3332 y ). 
VERACRUZ: M - Hidalgotitlan, Rio Solosuchil 5-6 km al 
ESE de Heron Cedi m, 25 Mar. 1982 (fl) 
Lorence et al. 3935 (MENU). GUATEMALA. IZABAL: along 
Rio Bonita, d i da m, 21 e 1941 iu Steyermark 


~ 
= 
= 


1040 (A, F); along Rio Tameja, 50 1 1 Dec. 1941 
(fr), Stevermark 41785 (^, F); Cerro San e 300-900 
m, 25 Dec. 1941 (st), Stevermark 41924 (F). PANAMA. 


CANAL AREA: e Road 5-10 mi. N of Gamboa, 21 

V1 (fr), Gentry 26. 59 (GH, MO, Dh 11 Feb. 
Vee 9600 (F, MO); nr. Pina, 100 

. 1973 (fr), Gentry & Nee 8721 (MO); Pi veline Road, 

9e] SN. 79°45'W, 50 m, 10 Mar. 1983 (st), Hamilton 


et al. 3255 (MO). cocrfé: hills above El Valle, ca. 1,000 
m, 24 Dec. 1972 , Gentry 6898 (F, GH, MO, NY) 
coLÓN: Miguel de la Borda, hill nr. beach, 24 Apr. 1970 


(early fr), Croat 10036 (F, MO — 2 sheets); Rio Guanche, 
l-4 km upstream from Portobelo Road, 0-100 m, 10 
Dec. 1973 (fr), Gentry 8791 (MO —2 sheets): headwaters 
of Rio Boqueron nr. fork with Rio Nombre de Diosito, 
150 m, 21 ds 978 (Il), Hammel 3959 (MO); joa 
. 1971 (fr), Lao & rris 207 
2 we us Rita Ridge rd., 10.2 in from em 


=$ 


Roosevelt Hwy., 350 m, 15 Mar. 1975 (fr) Mori & 
Kallunki 5065 (CH. MO): Rio Guanche, 2.5 km up- 
stream 24 bridge on Portobelo Road, x m, 27 Aug. 
1975 Mori & Il ithe de ¿973 (N IS). PANAMÁ 
Cerro Her 23 Jan. 1967 (fr), Duke 930. (DU KE, MO): 
erro Jefe, to 850 m, | ek 1972 (early fr), Ge MPs B 


al. 3513 (ENCB, MO, NY); El Llano Cari rd., 
km from Inter-American Hwy., 3 de e ) m, 4 Dec. 
(fr), Mori & Kallunki 3528 (GH, 


ae 4 


The three species named from central Pana- 


gatunensis, 


ma— Psychotria cerroazulensis, P. 
and P. have morphologies 
within the range of P. calophylla and are therefore 
included. One of the three specimens at MO of 


grandicarpa well 


Dwyer et al. 7244, cited as the type of P. gran- 

dicarpa, is hereby chosen as the lectotype. 
Psychotria calophylla may be rec ib by 

its arborescent habit, large (usually 1: x 

12 em), elliptic to obovate leaves 

red-brown tomentose below), undulate collector vein, 


iss uso 


large panicles of glomerules with secondary axes 
usually in 3 ranks, and large (10-14 x 6-9 mm) 
ellipsoid fruits. Collections from Guatemala are 
noteworthy in that the leaf blades are obovate, as 


opposed to the more comon elliptic shape. and have 


ewer secondary veins than elsewhere in the range 

of the species. 

32. Psychotria chitariana Dwyer & C. Ham- 
ilton in C. Hamilton, Phytologia 64: 221. 


13 km E 


1988. TYPE: Costa Rica. Cartago: 
of Turrialba, hwy. to Limón, canyon of Rio 
Chitaría. 99555'N, 83%36'W. 750-800 m, 10 


May 1983 (fl). Liesner et al. 15100 (holotype, 
MO; isotype, CR). Figure 36. 


Shrub | m tall: voung stems glabrous, the bark 
irregularly furrowed, with organic material accu- 
mulated in leaf axils; stipules not seen. Leaves 
petiolate; petioles ca. 1.5 em long, glabrous, winged, 
flattened blades charta- 
ceous, obovate, the Pow rounded to obtuse, the 

28- x glabrous 


ned pale brown to greenish 


above: membranous to 


base attenuate. 15 cm, 
above and below. 
red-brown; secondary veins 16-19 pairs, diverging 
(50° )60°- 65% 70°), brochidodromous, straight to 
arcuate near margin, elevated below, glabrous, the 
axils lacking domatia or hairs; tertiary veins evi- 
the 
terminal, dense glo- 


orthogonal reticulate, intersecondaries 
Inflorescences 
panicle branched to 3 


dent, 
conspicuous. 
bose panicles of cymes; 
degrees; main axis 7.5 cm long, the peduncle 4.5 
cm long; secondary axes in | rank plus terminal 


cluster, the first-rank axes 2, | em long, the clus- 


tered axes ca. 8, equal, 0.71 cm long: cymes 
branched to 1-2 degrees; bracts linear to trian- 
gular, 5 x 2.5 mm, glabrous. Flowers pedicellate, 
the pedicels 3-5 mm long; calyx cup-shaped, the 
tube 1 mm long, the lobes 5, 
brous; corolla greenish white, 

1.5 mm, white pubescent in throat, the 
lobes 5, triangular, 2.3 x 1.5 the 
filaments 4.5 mm long in pins, not seen in thrums, 


barely evident, gla- 
the tube cylindrical, 


mm; stamens D5 


the anthers 1.5 mm long; style 7 mm long in pins, 
not seen in as the branches minute. Fruit not 


seen. 


Distribution (Fig. 296). Known only from the 
type collection from Cartago, Costa Rica, at 750 


800 m in an area of premontane to lower montane 


896 


Annals of the 
Missouri Botanical Garden 


rainforest. with equatorial-mountainous climate. 
Psychotria chitariana was collected in flower on 


May 10. 


Psychotria chitariana may be recognized by 
its large (28-33 x 13-15 
with rounded to obtuse 


cm), obovate leaves 
apex, and inflorescence 
secondary axes in one rank plus a terminal cluster. 


he only flowering collection seen was a pin. 


53. Psychotria dressleri (Dwyer) C. Hamilton, 

Phytologia 64: 223. 1988. Cephaelis dress- 

leri Dwyer, Ann. Missouri Bot. Gard. 67: 68, 

fig. 15. 1980. TYPE: Panama. San Blas: SE 

of Puerto Obaldia, 18 Aug. 1971 (fl), Croat 
16801 (holotype, MO). Figures 7d, 36. 


i ali ice oensis Dwyer, Ann. Met tn P 
67: 78. ryPE: Panama. Darién: S slop 
C pi e 'una e Rio Pucro e camp, Ob 
1 25 Jan. 1974, Gentry & Mori 13895 
pee MO, n.v. v). 


Shrub or subshrub 0.2-1.5 m tall; young stems 
glabrous, the bark irregularly fissured and fluted, 
with organic material accumulated in leaf axils; 
stipules sheathing, slightly cleft at apex, 17-26 x 
8-10 mm, glabrous, caducous, leaving a pale ridge 
with. interrupted. red-brown fringe. Leaves sessile 
to subsessile; petioles to 1 em long; blades mem- 
branous, obovate, the apex acuminate to cuspidate, 
the base attenuate, (13-)20-47 x (8-)10-24 cm, 
glabrous above and below, drying red-brown to 
green-brown, paler below; secondary veins 17-25 
pairs, diverging 50?-70?, eucamptodromous to 
brochidodromous, straight to slightly arcuate to- 
ward margin, elevated below, red-brown puberulent 
below, the axils lacking domatia or hairs; tertiary 
veins very conspicuous, orthogonal reticulate, the 
the 
looping tertiaries between the secondary loops and 


percurrent tertiaries especially prominent, 


the margin also prominent. /nflorescences termi- 


nal, panicles of glomerules; panicle branched to 2- 
3 degrees; main axis 2-9 cm long, the peduncle 
2 ranks 
7d), the first-rank axes 2 or 4, 
1.5 em long, the shorter pair 


(0.5-)1.5-5 em long: secondary axes in | 
or clustered (Fig. 
the longer pair 0.5 
when present 0.5-0.7 em long, the second-rank 
axes 2 or 4, the longer pair 0.5 cm long, the 
shorter pair when present 0.2 cm long; bracts and 
bracteoles spathulate, 10-15 x 6-8 mm, gla- 
brous. Flowers sessile; 


2 mm long, the lobes 5 


calyx cup-shaped, the tube 
. linear, 1.5-2 x 0.8 mm, 
fringed; corolla white, the tube cylindrical, 10-12 

1-2 mm, white pubescent in tube in zone ca. 
8 mm from base, the lobes 5, linear with apical 
appendage | mm long, 2-3 x | mm; stamens 5, 


the filaments 10 mm long in pins, 13-14 mm long 
in thrums, the anthers 2.5 mm long; style 14 mm 
long in pins, 8-10 mm long in thrums, the branches 
linear, puberulent. Fruit when dry ellipsoid, 8- 
mm long, 3.5-4.5 mm diam., maturing color un- 
known, when dry deep red-brown; calyx persistent, 
1-3 mm long; seed dorsal surface with 3-4 deep 
longitudinal furrows, the ventral surface with. 2 
incompletely divided longitudinal furrows. 


Distribution (Fig. 36). 
ern Panama near the ridge running along the Ca- 


Known only from east- 


ribbean coast south to Cerro Pirre at elevations of 
20- 1,000 m and across the border into Colombia 
in tropical wet to premontane wet forest with equa- 
torial-mountainous to equatorial-tropical climate. 
Psychotria dressleri has been collected in flower 
January-August, and in fruit August- December 


and March. 


Additional pii A PANAMA. COLÓN: 
Santa Rita lumber road, Aqua Clara rain gauge, 20 
July 1974 (fl), Dressler - 1677( MO- 

idge, ca. 12 km from Transisthmian Hwy., 28 
1978 (early fr), Hammel 3665 (MO); Río Guanche, 
km pure am from bridge on rd. to Portobelo, 20 m, a 
Aug. 1975 (f r), Mori & W ri 7954 (MO); along 
ru of Río € EUN above bridge, 50 m, 8 July 1976 
(fl), € Se. 183 (M DARIÉN: La Boca de Pir 
13 Oct. 1967 (saris fr), Bristan 1277 (MO); trail hen 
Tacarcuna Village on Rio Tacarcuna to Cerro Mali, 800 
1,300 m, 16 Jan. 1974 (fl), Gentry 13598(MO); Serranía 
del Darién, trail from Cerro Malí to Río Pucro camp, 20 
July bae x Gentry et al. 16826 (MO). PANA a El 
., 4 mi. beyond hwy., 500 m, 5 Dec. 1979 

(fr), en 49145 (MO). 


Material annotated. Cephaelis panchocoensis 
by Dr. Dwyer does not differ substantially from P. 
dressleri and is therefore included. 

Psychotria dressleri may be recognized by its 
large (usually 20-47 x 10-24 cm), obovate leaves 
with organic material accumulated in the leaf axils 
and by its contracted (2-9 cm long) panicles of 
glomerules with clustered secondary axes and en- 


arged (10-15 x 6-8 mm) spathulate bracts. 


Ann. Mis- 
383, fig. 72. 1980. TYPE: 
Panama. Veraguas: mountains W of Alto de 
die W of Santa Fe, 960—1,080 m, 8 Sep. 
Hammel 1634 (holotype, 
MO— 2806155) 


54. Psychotria hammelii Dwyer, 
souri Bot. Gard. 67: 


8 (early fr), 
Ha — 2080014; 
Figure 36. 


Isotype, 
Herb or subshrub 0.1-0.3 m tall, stoloniferous; 
voung stems glabrous, angled, the bark smooth; 
stipules sheathing, bifurcate, often split down cen- 


ter, 15 


2 x 5-10 mm, glabrous, caducous, per- 


Volume 76, Number 3 
1989 


Hamilton 
Mesoamerican Psychotria, Part III 


sistent at top 3—5 nodes, leaving a red-brown ridge. 
Leaves petiolate; petioles 7-18 mm long, glabrous, 
grooved above: blades membranous, bullate, ellip- 
ic, the apex cuspidate, the base narrowly cuneate, 
(5-)8-12 x (2-)3-4.5 cm, glabrous above, pu- 
berulent on veins below, drying deep red-brown: 
pairs, diverging (45?—)60?— 
ele- 


secondary veins 11-15 
70°, eucamptodromous, constantly arcuate, 
vated below, red-brown puberulent below, the axils 
lacking domatia or hairs; tertiary veins conspicu- 
ous, orthogonal reticulate. /nflorescences terminal, 
panicles of cymes; panicle branched to 3 degrees; 
main axis 4-4.5 cm long, the peduncle 1.5-3 cm 
long; secondary axes in 2 ranks, the first-rank axes 
2, 1.2-1.3 cm long, the second-rank axes 2, 0.4 

0.5 em long: cymes branched to | degree; bracts 
linear, 8-9 x 3 mm, glabrous. Flowers pedicellate, 
the pedicels ca. 1 mm long; calyx cylindrical, the 
tube 1-2 mm long, the lobes 5, linear, 1-2 x 0.7 
mm, glabrous, fringed: corolla white, the tube cy- 
lindrical, mm, white pubescent in throat, 
the lobes 5, ovate with apical extension 0.5 mm 
long, 1 x 0.7 


long in pins, not seen in thrums, the anthers | mm 


mm; stamens 5, the filaments 3 mm 
long; style 5 mm long in pins, not seen in thrums, 
the branches linear. Fruit when dry ellipsoid, 5 

7 mm long, 3.5-4 mm diam., maturing red, drying 


deep red-brown; persistent calyx. 1-2 mm long; 


seed dorsal surface with 4 deep longitudinal fur- 
rows, the ventral surface with 2 shallow longitudinal 


furrows. 


known only from the 
1.000 m el- 


evation in low montane wet forest to rainforest with 


Distribution (Fig. 36). 


western cordillera in Panama at ca. 


atorial limate. This species has 
ben collected in ces: in March and June- July, 
in fruit in April, and with immature fruit in June 


and September. 


Sin ue an rues examined. PANAMA. BOCAS DEL 
: 1 Zorra, 19 Apr. 1968 (tr), Kirkbride 
& Duke 812 (MO—2 s )eets). CHIRIQUÍ: vic. Gualac 
. from Planes de Hornit T 

980 


e 


. La Fortuna, 1,30( 
{ntonio 5060 (MO); behind ibd 
1 N of Los Planes de i 
IRHE F ges Hy a a Project, 8°43'N, 
17 June ak ? (fl, early fr), Knapp & 
adis ka 55. E (MOX ds de Arena, N of Carretera de 
E e ode to, IRHE Fortuna drole Bic Project, 8*46'N, 
W, 1,100 m, 12 Mar. 19 (fl), Knapp et al. 


— 


pre (MO) 


Psychotria hammelii may be recognized im- 
mediately by its bullate leaves, small (4-4.5 em 
long) panicles of cymes with enlarged bracts, and 


ellipsoid fruits with conspicuous persistent calyx. 


The only two flowering collections seen are of 


the pin morph. 


Contr. U.S. 


l ragoga 


59. Riana insignis Standley, 

. Herb. 18: 130. 1916. Not 
io (Muell. Arg.) Kuntze (based on Rud- 
gea insignis Muell. Arg. in Martius). TYPE of 
P. insignis: Panama. San Blas: hills of Sperdi, 

Puerto Obaldia, 20-200 m, Sep. 1911 
(A), Pittier 1110 (holotype, US: isotype, F). 
Figure 35. 


Shrub; young stems densely ferrugineous-pu- 
bescent; stipules ovate, 12 X 4 mm, ferrugineous- 
pubescent, caducous, leaving a pale ridge with red- 
brown fringe. Leaves petiolate; petioles 
cm long, ferrugineous-pubescent, pe rfectly terete 


2.5-5.5 


or often grooved above; blades membranous, ovate, 
the apex acuminate to cuspidate, the base cordate, 
14-22 x 7 
adaxially puberulent midvein, ferrugineous-pubes- 
cent below, drying deep red-brown; secondary veins 
17-20 pairs, diverging 100*-105?, brochidodro- 
mous with collector vein undulating near margin, 
constantly arcuate, elevated and densely pubescent 


10 em, glabrous above except for the 


below, the axils lacking domatia or hairs; tertiary 
veins evident, orthogonal reticulate to percurrent. 
Inflorescences terminal, panicles of glomerules; 
panicle branched to 3-4 degrees; main axis 2.5 
cm long, the peduncle lacking; secondary axes in 
l rank, the first-rank axes 1-2, 1.5 em long; bracts 
triangular, 2-2.5 mm long. 5 mm across, ferru- 
gineous-pubescent and fringed. Flowers sessile; ca- 
lyx cylindrical. the tube 1 mm long, the lobes 5. 
linear, 1-1.2 0.8 mm, red-brown puberulent: 
mature corolla, stamens, and style not seen. Fruit 
not seen. 


Distribution (Fig. 35 Known only from the 
type collection from near Puerto Obaldia, San Blas. 
Panama, at ca. 100 m elevation in tropical moist 
forest with tropical-subtropical climate. It was col- 


lected in flower in September. 


Psychotria insignis may be recognized imme- 
diately by its large (14-22 x 7-10 cm) ovate 
leaves with bases cordate and drying deep red- 
brown, numerous (17-20) secondary veins diverg- 
ing 100?-105?, and by small (2.5 em long) fer- 
rugineous-pubescent panicles of glomerules. 


56. Psychotria 
Phytologia 64: 
Ann. 


insueta (Dwyer) C. Hamilton, 
226. 1988. Cephaclis insueta 
Missouri Bot. Gard. 07: 73. 


Panama. Veraguas: 11 km from 


Dwyer, 
1980. TYPE: 


898 


Annals of the 
Missouri Botanical Garden 


Escuela Agricola Alto de Piedra, along Rio 
Dos Bocas, Atlantic slope, 15 Nov. 1974 (ear- 
ly fr), Mori & Kallunki 3108 (holotype, MO). 
Figure 35. 


Shrub 0.4—1.7 m tall; young stems glabrous to 
minutely puberulent, the bark furrowed longitu- 
dinally, with organic material accumulated in leaf 
axils; stipules with a sheath 9 mm long and wit 
2 linear extensions 2 mm long, minutely fringed, 
caducous, leaving a red-brown ridge without fringe. 
Leaves sessile; blades membranous, obovate to ob- 
lanceolate, the apex cuspidate, the base decurrent, 
30-42 
drying dull green or brown-green above, paler dull 


8-19 cm, glabrous above and below, 


-— 


green below; secondary veins 34-42 pairs, di- 
verging 75?-110? with higher angles toward base, 
brochidodromous, the collector vein straight along 
margin, constantly arcuate, elevated below, gla- 
brous, axils lacking domatia or hairs; tertiary veins 
evident, orthogonal reticulate to percurrent. /nflo- 
rescences terminal, rounded-umbellate panicles of 
cymes; panicle branched to 4 degrees; main axis 


17-18 cm long, 


the peduncle 3.5-7 cm long; 


secondary axes in 4 ranks, the first-rank axes 2, 
9 cm long, the second-rank axes i 3), 3-4.5 cm 
long, the third-rank axes 2, 2-2.5 em long, the 


0.7 


to 3 degrees; bracts subtending branches, broac 


fourth-rank axes 2, | em long; cymes branched 
irregular, 1 mm long, irregularly fringed. Flowers 
pedicellate, the pedicels 0.5-1.5 mm long; calyx 
cup-shaped, the tube 1-1.5 mm, the lobes 5, barely 
evident to none, with pubescent fringe; corolla white, 
the tube cylindrical, 2 1.5 mm, white pubescent 
in throat, the lobes 5 
4, the filaments 2 mm long in pins, not seen in 
4 mm 


. linear, 2 X | mm; stamens 


thrums, the anthers | mm long; style 3.5 


long in pins, not seen in thrums, the branches 
Fruit ellipsoid when dry, 
long, 4-5 mm diam., maturing blackish (?), drying 
black; to 0.5 


seed dorsal surface with 5 longitudinal furrows, the 


spathulate. 7-10 mm 


persistent calyx a beak mm long; 
ventral surface with 2 incompletely divided lon- 


gitudinal furrows appearing as | furrow. 


Weird (Fig. 55). Known only from the 
continenta ide north of El Cope, Coclé, Panama, 
at ca. 750 m and north down to ca. 200 m ele- 
vation, and from the Atlantic slope in Veraguas; 
it is found in tropical wet to premontane rainforest 


with equatorial-mountainous climate. This species 
has been collected in flower on June 21 and in 


fruit October- November. 


Additional specimens examined. PANAMA. COCLÉ: 
El Copé, on Pacific side, half-hour walk from sawmill, 


780 m, 16 Oct. 1979 (fr), Antonio iud a 

Aserradero El Copé, N of El s PE eam E of 
sawmill, 700-750 m, 2 Nov. oon 5964 
(MO); E of El Copé sawmill, cn Siu. ca. 660 m, 


Hammel 3595 (MO). VERAGUAS: trail 
Rio Barrera junction with Rio 
17 Oct. 1978 (early fr), Ham- 


2] June 1978 (fl), 

from ( Coquyito mine to 

Burr ion, 300 m, 
7 (MO 


mel 52 


Psychotria insueta may be recognized by 
large (30-42 x 8-19 cm), glabrous, obovate to 
oblanceolate leaves with many (34-42) secondary 


°-110° and joining in a collector 


veins diverging 7: 


vein; large (17-18 cm long) panicles of cymes; 
and ellipsoid fruits drying black. 
The only flowering collection seen is a pin morph. 


57. Psychotria monteverdensis Dwyer & C. 
Hamilton in C. Hamilton, Phytologia 64: 230. 
1988. TYPE: Costa Rica. 
renas: al oeste de Reserva de Monteverde, 
1,520-1,590 m, 24 June 1977 (fl), Dryer 
1531 (holotype, MO; isotype, CR). Figure 35 


Guanacaste-Punta- 


Shrub 2.5 m tall: 


bark irregularly furrowed longitudinally; stipules 


young stems glabrous, the 


sheathing, truncate with 2 aristate appendages 2 
mm long from apical corners, ; mm, gla- 


brous, caducous, leaving a pale ridge with long red- 
brown fringe. Leaves sessile to petiolate; petioles 
to 0.5 cm long, glabrous, grooved above; blades 
subcoriaceous, elliptic, the apex acuminate, the 
base cuneate, (9-)13-15 x (2-) 
brous above and below, drying pale green-brown; 
secondary veins 9-11 pairs, diverging 45°-60°, 
brochidodromous with collector vein undulating near 


3-4.5 cm, gla- 


margin, constantly arcuate, prominent below, gla- 
brous, the axils lacking domatia or hairs; tertiary 
veins inconspicuous, orthogonal reticulate. /n/lo- 
rescences terminal, few-branched panicles of glom- 
erules; panicle branched to 2 degrees; main axis 
3.6 cm long, peduncle 2.4 cm long; secondary 
axes in 2 ranks, the first-rank axes 2, 0.0-0.7 cm 
long, the second-rank axes 2, reduced; bracts lin- 
ear, 8 mm long, ciliate; bracteoles ovate, 3-4 mm 
long, ciliate. Flowers sessile or with pedicels to 1.5 
mm long; calyx cup-shaped, the tube ca. 1 mm 
long, the lobes 5, triangular to lanceolate, to 1 mm 
long, ciliate; corolla green-yellow, leathery, the tube 
cylindrical, 3 X 2 mm, pubescent in throat, the 
lobes 5, triangular with robust. keel-like append- 
ages, 
long, 


2 X ] mm; stamens 5, the filaments 2.5 mm 
the anthers 1.2 mm long; style 4 mm long, 


the branches clublike. Fruit not seen. 


Distribution (Fig. 35). Known only from the 
type collection from near the Monteverde Reserve 


Volume 76, Number 3 
1989 


Hamilton 


il 899 
Mesoamerican Psychotria, Part III 


near the Guanacaste- Puntarenas border, Costa 


Rica, at ca. 1,550 m elevation in low montane wet 
forest with. equatorial-mountainous climate. Psy- 
chotria monteverdensis was collected in flower on 
June 24 


Psychotria monteverdensis may be recognized 
by its subcoriaceous medium-sized (usually 13-15 
x 3-4.5 em) leaves with collector vein, small (3.6 
cm long) inflorescences with secondary axes 2 per 
rank in 2 ranks, those of the second rank sessile, 
and leathery greenish corollas with keel-like abaxial 
appendages on the lobes. 

Inly one flower was dissected, but the minute 
separation (0.3 mm) between stigma and anthers 
suggests that the species is long-homostylous. 


58. Psychotria pacorensis C. 
tologia 64: 232. 1988. TYPE: 
má: vic. Cerro Jefe, Altos de Pacora, 720 m, 
18 Dec. 1980 (early fr), Antonio 3241 (ho- 
lotype. MO: isotype, ENCB). Figure 36. 


Hamilton, Phy- 


Panama. Pana- 


Shrub 2-3 m tall; young stems minutely red- 
brown puberulent, the bark longitudinally grooved 
and fluted: stipules ovate, X 4 mm, glabrous, 
caducous, leaving a red-brown ridge with or without 
fringe. Leaves petiolate: petioles 5-13 mm long, 
glabrous, grooved above; blades membranous, el- 
liptic to obovate, the apex acute to caudate, the 
25 x (4-)5-10 cm, gla- 
brous above, glabrous below except on veins, drying 
12-14 


eucamptodromous to 


base cuneate, (13-)16 


red-brown, paler below: secondary veins 


70-80", 


brochidodromous, constantly arcuate, elevated and 


pairs, diverging 


minutely puberulent below, the axils lacking do- 
matia or hairs; tertiary veins conspicuous, orthog- 
onal reticulate, the percurrent tertiaries especially 
conspicuous, the tertiary arches near margin nu- 
merous. Inflorescences terminal, panicles of cymes; 
panicle branched to 3 degrees; main axis 4.5 cm 
long, the peduncle 3.5 em long; secondary axes in 
3 ranks, the first-rank axes 4, the longer pair 0.5- 
0.6 cm long, the shorter pair 0.3 cm long, the 
second-rank axes 4, the longer pair 0.2-0.3 cm 
ipf: the shorter pair reduced, the third-rank axes 
. 0.2 em long: cymes branched to 1-2 degrees; 
bus and bracteoles broad, to | mm long, fringed. 
Flowers sessile to pedicellate, the pedicels to 2 mm 
mature flowers not seen. 


long: Fruit when dry 


ellipsoid, 5.5-7 mm long, 3.5-5 mm diam., ma- 
turing purple, drying red-brown; persistent calyx 
cup ca. 0.5 mm long: seed dorsal surface with 4- 
5 deep longitudinal furrows, the ventral surface 
with 2 shallow longitudinal furrows. 


Distribution (Vig. 30). 
Altos de Pacora region near Cerro Azul and Cerro 
Jefe. eastern Panamá Province, 


Known only from the 


Panama, at ca. 
750 m elevation in premontane wet forest with 
tropical-equatorial climate. Psychotria pacorensts 
has been collected with immature fruit in December 
and in fruit in February. 


Additional specime ns examined. due PANAMÁ: 
Cerro Jefe, 6 mi. past Cerro Azul « rd. jm be 
Pacora, 780 m, 19 Feb. 1981 (fr). Sytsma e D 


3689 (MO —2 sheets) 


Psychotria pacorensis may be recognized by 
its large (usually 16-25 x 5-10 cm) leaves drying 
red-brown and with brochidodromous veins with 
secondary loops (not marginal collector vein) and 
conspicuous 


tertiary veins including numerous 


arches between the secondaries and the margin. 


59. Psychotria psychotriifolia (Seemann) 
Standley. U.S. Nath Herb. 18: 133. 
1916. Cephaelis ponents Seem., Bot. 
Voy. Herald: 138. 1854. vlde. ra psycho- 
triifolia (Seem.) ea Revis. 
962. 1891. 
auctt. TYPE: 
type, BM. n 


Contr. 


Gen. Pl. 2: 
Psychotria psychotriaefolia 
182 (holo- 
v., photo, MO). Figures 2g, 35. 


Panama: Seemann 


Psychotria Publ. 
Mus., Bot. Ser. 7: 114. 1930. 
Bolivar: Boca Verde, on Rio Sinu, 
Pennell 4572 (holotype, NY 


Field Columbian 
lYPE: 
4 Mar. | 


sinuensis Standle y, 
Colombiz 


1918 (fl) 


Shrub 0.5 
pubescent, the bark smooth: stipules with a sheath 


e13-X* 4-5 


-2 m tall; voung stems ferrugineous- 


mm. truncate, biaristate with the 


extensions 1.5-4 mm long from apical corners (Fig. 
2g). bent to ferrugineous- E scent, « 'aducous, 
often persistent at terminal : 
ridge. Leaves petiolate; ne 2-15 
ferrugineous-pubescent, grooved above; 

the 
acute. the base attenuate or subcordate, (5.5-)7.5 
17(-20) x (1.5-)3-6.5( 


glabrous to red-brown puberulent below, the mid- 


membranous. oblanceolate to elliptic, apex 


8) em, glabrous above, 
vein puberulent, drying green-brown to red-brown, 


veins (10-)14-20(-23) 


brochidodromous, 


paler below; secondary 


70°=859,; 


collector vein almost parallel near margin, con- 


pairs, diverging with 


stantly arcuate, prominulous below, puberulent be- 


low, the axils lacking domatia or hairs; tertiary 


veins inconspicuous, orthogonal reticulate. /nflo- 
rescences terminal or pseudoaxillary, panicles of 
glomerules; panicle branched to 2(3) degrees; main 
axis 8-17 mm long, the peduncle absent; second- 
ary the first-rank axes 2 


axes in 1-2(-3) ranks, 


900 


Annals of the 
Missouri Botanical Garden 


or 4(6), equal, 0.8-2 cm long, the second-rank 
axes 2 or 4, equal, 0.5-1 cm long; bracts sheath- 
ing, biaristate, 3-5 mm long, glabrous to puber- 
ulent; bracteoles subtending glomerules enlarged, 
cuplike, 3-4 mm long, glabrous to puberulent. 
Flowers sessile; calyx cup-shaped, the tube 0 
x 0.7 
puberulent, Br corolla green-white, the tube 


mm long, the lobes 5, lanceolate, mm, 


cylindrical, -1.5 mm, white pubescent in 
throat, the e e lanceolate with minute apical 
appendage, | X 0.6 mm; stamens 5, the filaments 
1.5 mm long in pins, 2-2.5 mm long in thrums, 


the anthers 0.5-0.7 mm long; style 2.5-3 mm 
long in pins, 1.3-2 mm long in thrums, the branch- 
es short. Fruit narrow-ellipsoid when dry, 5-6 mm 
long, 3-4 mm diam., maturing red, drying red- 
brown; persistent calyx conspicuous, tubular, 0.5 
2 mm long: seed dorsal surface with (4)5 longi- 
tudinal furrows, the ventral surface with 2 longi- 
tudinal furrows. 
Distribution (Fig. 35). Known from southern 
Nicaragua through Panama, with very discontin- 
uous distribution, at 0-800 m elevation in tropical 
moist to premontane wet forest with equatorial to 
tropical climate. This species occurs also in Colom- 
bia, Venezuela, and Ecuador. It has been collected 
in flower and in fruit throughout the year. 
Selected specimens examined. NICARAGUA. RIVAS: N 
de Volcán Concepción, Isla de Sade 11?32'N, 
: j )JO-1,000 m, Aug. 1982 (fr), eae 
3469 (MO). ZELAYA: along Rio y ey W of Nue 
Guinea, 11?42'N, 84°26’W, 130 m, 21 Aug. 1983 (fr) 
J. Miller & Sandino 1254 (MO); Mpio. de des Santa 
Rosa, 6 Sep. 1982 (fr), Ortíz 115 (MO TA Rica. 
CARTAGO: 12 km S of bore, by air, 4 km E of Pejibaye 
along Rio Gato, 9°48'N, 2'W, 700 m, 16-17 Apr. 
1983 (fr), Liesner etas MO). HEREDIA: Finca La 
Selva, OTS field station on the Rio Puerto Viejo, 100 m, 
24 May 1980 (f), Hammel 8790 (DUKE); 1 Aug. 1980 
(fr), W ilbur . 30166 (DUKE). LIMÓN: camino a La Estrella, 
0 5 28 Feb. 1958 (fl), vi 470 (MO); Golden 
28 July 1936 (fl, fr), Dodge & Goerger 
9468 (CR, F, 'G H, MO, US); Las : valora: -Guapiles, md 
l (fr), León 653 (E); l mi. toward Zer 
30m, I Sep. 1946 (fr), Moe 
9) eu de los rios Parismina y Reventazón, 
1951 (fr), Shank & Molina 4195 (F, GH 
US) s SAN JOSÉ: vic. El General, 61 ) m, Jan. 1939 (fl, fr), 
3683 (GH, K, MO, ANAMA. BOCAS 
DEL TORO: Changuinola to 5 mi. S at junction of ríos 
C an Es Terebe, 30-60 m, 17-19 Dec. 1966 (fr), 
is et ¢ 63 (GH, MO, NY, US); Chiriquicito to 5 
mi. S Sin AA Guarumo, 5-7 June 1967 (fl, fr), Lewis 
et al. 2103 (G H, MO, US). CANAL AREA: Frijoles, 17 June 
, Ebinger 90 (F, MO, US); Fort Sherman and 


nearby, 9e YN, 80%00'W, 0-100 m, 16 June Wr s 
ow : a 3692 (MO); Pipeline Rd., ay 
1973 fr), Kennedy & Wilder 3103 (F, oo y 


(ez 
Pedro a nr. East Paraiso, 7 Jan. 1924 (tr), Standley 


29941 (US). CocLÉ: Cerro Gaital Caracoral, 800-1,000 
m, 25 July 1969 aes Dwyer & Lallathin 8950 (MO), 
COLÓN: alon 4 mi. SW of de Y 7 Apr. 
1971 (fl), Don 14114 (MO); Rio ig 
above bridge on Portobelo Rd., 50-200 m, 8 july 
(early fr), Croat 36982 (MO, NY); trail from headwaters 
scandaloso, route 
oned railroad, 75- , 21 July 1981 (fr), 
ond 3995 (MO); between France Field and Catival, 
9 Jan. 1924 (fl), Standley 30370 (US); Palmas Bellas, 
30 May 1971 (fl), Thoms 31 (ENCB, us ARIÉN: from 
Punta Guayabo to Río Jaqué, 50-200 m, 24 Apr. 1980 
(8), Antonio 4447 (ENCB, MO); de Aruza, 24 Oct. 1967 
(fr), Bristan is (GH, MO); Cativo Swamp ca. 6 mi. 
NW of El Real, 15 June 1962 (8). Duke 4824 (F, GH, 
MO); Rio Me 250 m, 18 Sep. 1967 (fr), Duke 14185 


— 


— . 

= 
FD 
O 


, fr), dy. et al. 4165 (MO); 
1.5 mi. E of Manené, 21 Dec. 980 (st), se Edd 12099 
(ENCB, MO); Rio Tuquesa, AH Venado, d 4 July 
1975 (A), Mori 6933 (MO). PANAMA: Rio € nere. l mi. 
above Madden Lake, 7 Oct. 1961 (fr), Duke py 4 (GE 
2 sheets, MO); 25 km NE of Cerro Azul on Rio Piedras 
;orgas Memorial Labs, 550 m, 25 Nov. 1974 (fr), . 
MO — 2 sheets); 
924 (fr), Standley 2613 3 (US); Arraiján, 15 
m, 22 2 June 1 1938 (fl), W oodson et al. 7; 76 (F, MO). SAN 
BLas: Ri i, ca. 6 air mi. SW of Mulatupu, ca. 100 
m, 20 Sep. 1967 (early fr), Duke 14210 (F, MO, NY). 
VERAGUAS: Coiba Island, 31 Aug. 1924 (fl), Collenette 
466 (US) 


pon 


The original spelling “psychotriaefolia” has been 


corrected to “psychotriifolia” in accordance with 
the [International Code of Botanical Nomenclature 
Article 73.8) 


Psychotria psychotriifolia may be recognized 


— 


by its oblanceolate to elliptic leaves drying green- 
brown to red-brown with collector vein near mar- 
gin, epedunculate inflorescences with enlarged cup- 
like bracts subtending the ultimate glomerules, and 
narrowly ellipsoid (5-6 3-4 mm) fruits with 
conspicuous persistent calyx tube. 


60. Psychotria rosulatifolia Dwyer, Ann. 
Missouri Bot. Gard. 67: 423, fig. 77. TYPE: 
Panama. Darién: trail from Rio Pucro base 
camp up ji 2 of Cerro Mali, 640—1,000 
m, 6 Feb. 5 (fr), Gentry & Mori 14174 


(holotype, do Figure 35. 


Subshrub or herb 0.5 m tall; young stems gla- 
brous or puberulent, the bark irregularly furrowed; 
stipules sheathing, ovate, often split down center, 
15-20 x 6-7 mm, glabrous, caducous, leaving a 
red-brown ridge. Leaves sessile; blades membra- 
nous, lanceolate, the apex caudate, the base sag- 
ittate, 16-28 x 4.5-8.5 cm, glabrous above and 
below, drying deep red-brown or sometimes green- 
brown; secondary veins 27-32 


pairs, diverging 


Volume 76, Number 3 
8 


milton 
Mesoamerican Psychotria, Part III 


80°—95°, brochidodromous with collector vein near 
elevated below, gla- 
the axils 


margin, constantly arcuate, 
brous or red-brown puberulent below, 
lacking domatia or hairs; tertiary veins evident, 
orthogonal reticulate to percurrent. /nflorescences 
terminal, spreading panicles of panicle 
branched to 4 degrees, the axes delicate; main axis 
8-13 cm long, the peduncle 2.5-6.5 cm long; 
secondary axes in 3 ranks, the first-rank axes 2, 
2.5-4 cm long, the second-rank axes 2 or 4, the 
longer pair 1-2 cm long, the shorter pair when 
present 1-1.2 cm long, the third-rank axes 2, 0.5 

0.9 em long; cymes branched to 1 -2(-3) degrees; 
bracts linear, 4 mm long, glabrous: bracteoles tri- 
angular, to 1.2 mm long, glabrous. Flowers ped- 
the pedicels 1-2 mm long: calyx cup- 
mm long, the lobes 5, 


cymes; 


icellate, 
shaped, the tube 0.5-0.7 
often barely evident, triangular, to 0.2 mm long, 
glabrous; corolla white, the tube narrow-cylindrical, 
2.5-4 x 
lobes 5. linear with 1-mm-long extension from apex, 

not spreading. 2.5-4 x | the 
filaments 2 mm long in pins, not seen in thrums, 
the anthers narrow, 2 mm long; style 6 mm long 
in pins, not seen in thrums, the branches clublike. 
Fruit ellipsoid when dry, 5-6 mm long, 4-4.5 mm 
diam., maturing red, drying red-brown; persistent 
calyx a beak ca. 0.5 mm long: seed dorsal surface 
with 5 deep longitudinal furrows, the ventral sur- 
face with 2 barely evident longitudinal furrows. 


1.2 mm, white pubescent in throat, the 


mm; stamens 95, 


Distribution (Fig. 35). Known only from the 
regions of Cerro Pirre and Cerro Mali, eastern 
Darién, Panama, at ca. 700- 1,300 m elevation in 
premontane to low montane rainforest with equa- 
torial-mountainous climate. Psychotria rosulati- 
folia has been collected in flower in April, July, 
and August, and in fruit in August, November, and 


February. 


Additional ud examined. ANAMA. DARIÉN: 
967 (f), Bristan 515 (MO); ascent 
Rio Pirre, 5 of El Real, 750- 1,030 m, 
; Cerro 


^ 
Ne) 
~ 
- 
2 
— 
—_~ 
=> 
= 
= Y 
— = 
— 
— 
c 
= 
=~ 
e 
ma 


Fot 
13788 (MO); Cerro Pirre, =E nr. 

, 1,200 m, uly 1977 (fl), un 4295 
(MOX Catia US valley between Pirre and next most 
southerly peak, 10-20 July 1977 (f), Folsom 1428 (M0); 
top of ridge Tei to Cerro Pirre, nr. Rancho d 


1,200-1,400 m, 13 Nov. 1977 (fr), Folsom et al. 62 
(MO) 


= 


Psychotria rosulatifolia may be recognized by 
its large (16-28 4.5-8.5 cm), oblan- 
ceolate leaves with 27-32 secondary veins plus a 


collector vein; delicate panicles of cymes; flower 


sessile, 


buds appearing narrow and acuminate due to nar- 


row |-mm-long apical extensions from corolla lobes; 
and medium-sized (5-6 x 4-4.5 mm), ellipsoid 
fruits. 

Five pin and no thrum collections suggest strong- 
ly that Psychotria rosulatifolia is pin-monomor- 


phic. 


Hamilton, Phy- 
Costa Rica. 


61. uibus de sixaolensis C. 
1988. TYPE: 
Limón and Shiroles, 
Sixaola, 0.9 mi. SW of Bambu, 0.5 mi. SW 
of Bribri, 50 m, 12 Aug. 1977 (fr), Croat 
13303 (holotype, MO). Figure 30. 


tologia 64: 234. 


món: rd. be tween 


~ 


Shrub or subshrub to 1.5 m tall; young stems 
ferrugineous-pubescent; stipules sheathing, ovate, 
10 x 6 


leaving a pale ridge with red-brown fringe. Leaves 


mm, ferrugineous-pubescent, caducous, 


petiolate; petioles 1 3.5 cm long, red-brown pu- 
berulent, grooved above; blades membranous, el- 


the apex acuminate, the base 
attenuate, (14-)16-30 x (5.5-)8-13 cm, gla- 
brous above, sparsely red-brown tomentose below, 


liptic to obovate, 


the midvein more densely tomentose, drying red- 
brown; secondary veins (20-)25- 25 pairs, diverg- 
ing 75°-80°, brochidodromous with collector vein 
near margin, constantly arcuate, elevated below, 
red-brown tomentose below, the axils lacking doma- 
tia or hairs; tertiary veins inconspicuous, orthog- 
onal reticulate. /nflorescences terminal or pseu- 
doaxillary, panicles of glomerules; panicle branched 
o 2-3 degrees; main axis in fruit 9.515 em long, 
the salian le in fruit 7 13.5 em long; secondary 
the fest-rank axes 
long, the second-rank axes 2, 0.5 em long; bracts 
x 0 


axes in 2 ranks, 2, 1.5-2 cm 


and bracteoles ovate, í mm, ferrugineous- 


pubescent. Flowers sessile; calyx cup-shaped, the 
tube 1-2 mm long, the lobes 5, linear, 2-3 x 0.8 
mm, ferrugineous-pubescent; corolla white, the tube 
cylindrical, 4 x 2.5 mm, white pubescent in throat, 
the lobes 5, ovate with blunt robust protuberance 
from near apex, 3 X 1.5 
filaments 3 mm long in pins, not seen in thrums, 
the anthers 1.3 mm long; style ca. 6 mm long in 
pins, not seen in thrums, the branches linear. Fruit 


mm; stamens 5, the 


ellipsoid when dry, ; 
drying red-brown: persistent calyx 2-3 mm long: 
| deep longitudinal fur- 


9 mm long, 5 mm diam., 


seed dorsal surface with 
rows, the ventral surface with 2 shallow longitudinal 


furrows. 


Distribution (Fig. 36). Known from south- 


eastern Limón. Costa Rica, and Bocas del Toro, 
Panama, at 0-50 m elevation in tropical moist 


forest with equatorial-tropical climate. This species 


902 Annals of the 
Missouri Botanical Garden 


€ 


has been collected in flower in April, in mature Group 2 
fruit in August. 8. P. balancanensis C. Hamilton 
9. P. costivenia Griseb. 


> > COSTA “A, LIMÓN: ` : 
pom diea UE ns examined. Costa Rica LIMÓN e altorum (Standl desea dE Hamilton 


nr. Río Sixaola, ca mi. SW of Bambu, ca. 3 mi. NE me : 
|! Bratsi, 15 m, 12 1977 (early fr), Croat 43250 | MAP CONUS 
dl Aug. y b pu 10. P. fendleri Star nal 
(CR, MO). PANAMA. BOCAS DEL TORO: Rio Teribe between 3 
11. P. flava Oerst. ex Standl. 
Qda. Treglo and Puerto Palenque, 2 Apr. 1968 (fl, early 12. P. grandis S 
fr), Kirkbride & Duke 536 (MO — 2 sheets). 13 P. dina setas sd 


E P. papantlensis Ps erst.) Hemsl. 


; : . . P. pleuropoda Donnell-Smith 
its ferrugineous aspect, large n 16-30 x B- a Pp Mn ee Standl. 


F 


Psychotria sixaolensis may be recognized by 


13 cm) leaves with usually 25-35 secondary veins 


and a collector vein, and eel pedunculate panicles — ,, , 
Group 3 


17. P. bakeri Dw 
18. P. graciliflora Benth. in Oerst. 
: 7 19. P. jimenezii $ 2 adl. 
EXCLUDED TAXA 20. P. laselvensis C. Hamilton 
. , 21. P. liesneri Dwyer 

Psychotria microdon (A. P. de Candolle) Urban, 55 p marginata Sw. 

Symb. Antill. 9: 539. 1928. Rondeletia mi- 23. P. orosiana Standl. 

crodon DC., Prodr. 4: 408. 1830. Mapouria 24. P. parvifolia Benth. in Oerst. 

microdon (DC.) Bremek., Recueil Trav. Bot. P. philacra Dwyer 

Néerl. 31: 286. 1934. 


of glomerules with large (7 x 6 mm) ovate bracts. 
The lone flowering collection seen is a pin morph. 


. . T" n . . . Group 4 
Psychotria pinularis Sessé & Mociño, Fl. Mexic. ed. 2 
51. 1894 26. P. remota Benth. 


Group 5 


Steyermark (Mem. New York Bot. Gard. 23: 
47 


446-447. 1972) included this species in subgenus ; 

‘had : 27. P. aguilarti Stan ¿ Steye 
Pereira, from which it differs, however, in hav- S /, guilarii Standl. € Steyerm. 
p f 28. P. boquetensis Dwy 
ing leaves drying pale green, a seed cross-section 59. p chagrensis Stand 
with only a central keel on the dorsal surface (vs. — 30. P. erythrocarpa Schldl. 


several longitudinal furrows), marcescent stipule — 31. P. fosteri C. 
32. P. fruticetorum Standl. 
33. P. jinotegensis ie Molina & Standl. 
a. Var. jinote rens 
b. var. moraz sane ic Hamilton 
related to any neotropical members of P. subg. — 34. P. idas E Hamilton 
1 Sw. 


remnants persisting at the nodes, and lateral 
branches in opposite pairs (vs. diverging singly from 
a main axis). This species does not appear closely 


: x . " : 
Heteropsychotria Steyerm., either. Instead, the — 35. P. nervosa 
above character states plus the leaf secondary veins 
arching broadly toward the margin suggest affinity Group 6 
with African P. subg. Tetramerae (Hiern) E. Petit, — 36, p. cascajalensis C. Hamilton 
many species of which have bacterial "nodules" — 37. P. chiriquina 24 idl. 
on the leaf undersides (Petit, Bull. Jard. Bot. Etat 38. P. cocosensis C. Hamilton 
30. -B: Ansara 
+. ]- 29 36: 65-190, 1966). Psyc ho- 3 J. di dwyeri C. pus . 
: erod IP S beti | 40. P. hornitensis Dwyer & C. Hamilton 
tria miero on ane , DES Ow. may be the sole 41. P. lundellii Stan 
neotropical species — aside from P. punctata Vatke, 42, P. mexiae Standl. 
introduced in Florida —of that subgenus. 43. P. olgae Dwyer & Hayden 
44. P. panamensis Standl. 
a. var. compressicaulis (K. Krause) C. Hamilton 
b. var. ixtlanensis C. Hamilton 
APPENDIX I. Numerical list of taxa. c. var. magna (Standl.) C. Hamilton 


G l 
roup 45. P. sar neto Stan dl. 


l. Psychotria carthagenensis Jacq. 46. P. stockwellii C. Hamilton 

2. P. clivorum Standl. & Steyerm. 47. P. trichotoma i Martens & Galeotti 
3. P. lamarinensis C. Hamilton 

4. P. micrantha Kunth G E 

5. P. neillii C. Hamilton & Dwyer bano E 

6. P. quinqueradiata Polak. 48. P. limonensis K. Krause 


7. P. viridis Ruiz & Pavón 49. P. tenuifolia Sw 


Volume 76, Number 3 Hamilton 903 
1989 Mesoamerican Psychotria, Part III 


Group 8 Beach, J., 1339 (29); 1361 (22); 1365, 1366 (29); 1418, 
50. P. alfaroana Standl. B dx ie di 1495 (22); = 511 (185 1515 (44a) 
l. P. calophylla Standl. each, d. x fera yunt, : ae) Sane =: 
2 a e d TC Beaman, J., 5162 (11) 5190 (14); 5307 (43): 5310 
re ela bags E m (18); 5410 (42); 5564 (44d); 5573, 5594 (42); 6007 
23. P. dressier: (Dwyer) C. Hamilton (44d); 6081 (11); 6094 (34) 6183 (6); 6187 (42): 
ed Y Ale Dwyer 6205 (48); 6391 (44d); 6448 (48) 
99. P. e Standl. , Beaman, J. & €. Alvárez, 5364, 5772, 5976 (42) 
56. P. insueta (Dwyer) C. Hamilton Bequaert J. 101 (25) 
57. P. monteverdensis Dwyer € C. Hamilton ls ; 
58. P. pacorensis C. Hamilton NR P Aye) 
BO. Po pevehntritblns (S ) Standl Perroi, C. & lo Cario, 1714 (13); 1717 (48) 
e E Paes or Gor 17321 00) 
61. p. sixaolensis C. Hamilton a 5. & G. Heath, 1956 EN 
XO da CE Blake, S. F., 7619 (9b); 7755 (4); 6 (9b) 
4 ES 99 5 29 9 
APPENDIX II. List of exsiccatae. A nid 1221 ass i eal (22) [599 (4. LADO 
Acosta, M. A 29, 31 (19) Blum, K. & J. PY. 395 (59); 398 (12); 2107 (22); 
Aguilar: I., 21 (13); 126, 215 (27) 2108 (48); 2148, 2152 (22); 2459 (35); 2494 (1); 
Aguilar, J. E 750, 1140, 1143, 1371, 1729 (30) 2652, 2657 (444) 
Aguilar H., M., 260 (13); 292 (35) 499 (18); 509 (22) Blum, K. et al., 1699 (18); 1708 (22) 
Aguilera R., V., 2 (12) Boege, W., 1239 (47) 


Allen, P., 11 (22); 25, 27 (18); 308 (44d); 936 (1); 951 Boster, J., s.n. (15) 
UR 1187 (22); 1387 (44d); 2035 (12); 2186 (29); Botteri, M., 542 (49); 920, 1056 (44d) 
2563 (13); 3507 (44d); 4060 (1); 4144 (22); 6498  Bourgeau, E., s.n. (49); 2321 (9b), 2454 (49) 


(32) ourgeau, M., s.n. (35) 
Alonso, D 26 (30) Bravo H., H., 116, 208 (49) 
Alston, 1853, 8716 (6), 8881 (35) Breedlove, D., 6319 (49); 6334 (6); 0485 pee 6489 
a N., s.n. (13) (32); 7544 (9a); 7626 (49); 9537, 9744, 9792 (9a); 
Antonio, T., 1586 (37); 1771 (29); 1804 (44d); 2090 9993 (9b); 10593, 11392 (30); 12034 M 13969 
(56) 2305 (22) 2339 : 2h 3241 (58); 3338 (22); (9a); 14126 (30); 15397 (9b); 20102, 20672 (30); 
3461 (44d); 4447 (59); 4545 (13); 4796 (22); 4908 21140 (44d); 23196 (9b); 23269 (42); 23390 (9a); 
(6); 5060 ES 23901 (32); 23974, 24698 (30); 24913 (18) 24948 
s T. € W. Hahn, 4317 (29); 4340 (22); 4407 (9b); 25337 (9a); 25348 (9b); 25671 (9a); 25966 (6): 
26) 25975 (13); 26006 (30); pi (48); 26311 (44d): 
E. M., 3168 (44a); 3247 (22); 3248, 3403 26314 (9b); 26461 (47); 26522 (48); 26989 (13); 
(29); 3645 (37); 3818 (1) 27173 (30); 27286 (13); n (30); 27427 (32); 
x rs ain, M. & P. P. Moreno, 833 (6), 876 (1); 901a 27437 (49), 27455 (48); 27577 (9a); 27821, 27877 
3 (44d); 2490 (18) (9b), 28107 (48); 28265, 28360, 28450 (30); 28501 
Arnason, i s.n. (49) (48) 28958 (30) 28971 (42); 31270 (30) 32880 
Arnason, T. & a Lambert, 17081 (13); 17293, 17320 (9a); 32945 (48) 33059 (18); 33093 (47); 33806 
35); 17615 (49) (495; 34380 (18); 34530 (9b) 34538 (42): 34559 
Atencio, G., 1 ney (9b); 35102 (9a); 35141 (44d); 35316 (34); 36637 
Atwood, J., 57, 212b (44d); 224, 275, 306 (18); 4235 (49); 37190 (30); 38394 (13); 38672 (11); 38753 
22 1078. (29); 4819 Es 5299 (2 29) (9b); 39816 (13); 39944 (30); 40061 (13); 41341 
Atwood, J. & A. D. Moore, 364 (29) (9a); 41971 (48); 42043 (30); 42076 (9a); 42189 
Atwood, J. & F. Seymour, 2990 (35) (30), 42841 (13); 44372 (30); 53213 (9a) 
vendano, S. & J. Calzada, 516 (47) daa D. & R. Dressler, 29565 (44d); 29846 (32); 
Avendaño D., M., 39 (2); 174 (18) 29855 (48) 
Aviles, S., 9 (13); 932 (22); 973 (48) Breedlove. D. & E. McClintock, 34065 (22) 
Bailey, L. H. € E. Z. Bailey, 63 (4); 161 (22); 393 (13) Breedlove, D. € P. Raven, 8205 (9a); 8336, 8347, 
Baker, : F., 609 (6) 13161, 13205, 13331 (30); 13568 (9b) 
Bakes , 17 n > 20 (35) Breedlove, D. & A. R. Smith, 21542 (30); 21713, 22174 
aan, W., 235 (22) (9b); 31291 AE 31292 (9b); 32414 (44d) 
.«& E L Ta M (6) a D. & R. Thorne, 20640 (47); 20960 (42); 
die E D., 3/5A (6); 10/5, 10/5B (14); 14/4 (6); 20976 (48); p 9 (9a); 21140 (9b); 21140a (44d); 
30; 14B, 30, 14D (4 A 21156 (9a); 21203 (42); 21313 (30); 30180 (18); 
arrera, A. et al., 228 (9b); 298 (47) 30660 (47); eer 2 (44d); 30828 (9b) 
Bartlett, H., 11435 (35); 11499 (6); 11622, 11869 (32); ai J. et al., 14484 (30) 
12038 (7; 12043 (22); 12043a (13); 12205 (48); Brenes, A. M., s.n. 0% 1902 (18) 3902, 3908, 4233. 
12244 (35); 12397 (32); 12533 (41); 12534 (42); 4547. 5377 (6); 5534 (23); (24) 6249 (6); 
12633 (9b; 12686 (35); 12722 (6); 12809 (9b) 12603 (13); 1 2624 " Do 1266 (18); 14281, 
12819, 12824 (32); 12834 (9b); 12835 (32); 12884 14310, 14531, 15528 (6); 19204 (13); 19218 (49); 


(35), 12907 (9b); 13002 (6); 13122 (32); 13143 (6) 21427, e (6); 22006 (19), 22484 (1) 
Bartlett, H. € T. Lasser, 16343 (22); 16444, 16957 — Bristan, N., 83 (18); 515 (60); 1018 (18); 1037 (59); 
(35) 1051 n 1170 ( A 2 1277 (53); 1368 (59); 1386 
Basurto-Rafael P., F., 185 (9b); 246 (11) (22) 1457 (12); 1494 (18) 
va, K. S., 139 (1) Broome, C., 704 Pd 


904 


Annals of the 
Missouri Botanical Garden 


Brown, B. & M. Artavia, 725: 
Brown, M., 143, 161, 163 (22 


434 (32); 521 (6); 855 (12); 


Burger, W. & R. Stolze, 5032 (29 
41 (22) 

Burger, W. et al., 10654 (22) 

Busey, P., 522 (12); 627 (18); 770 (13) 


Busey, P. & T. Croat, 239 (49) 

Bustillo, S., 187 (44d) 

Cabrera, E. & L. Cortés, 145, 226 (35); 308 (32); 326 
(9b); 343 (32) 

Cabrera, E. « G. Ibarra, 1144 (35) 

Cabrera, E. & S. Tarate, 1529 (35 

Cabrera, E. et al., 1875 (35); 1934 (48); 2037 (35); 
2700 (22 


A 


` (1) 
; 68 (14d) 116 (11) 157 (6); : Mg 
1); 1564 (30); 


(11); 1037 (14); 1273 (29); 1429 (I 
4965 ved 7740 a )) 
Calzada, J. & W. ae 4527 (30) 
Campbell, E. J. F., s.n., 11 (3 2k 9] (35) 
29a 12 


, M. C. 1009, 1100 (x 1745 (9a); 2449 (44d) 
Carrasquilla, L. " al., 
Carvajal, A., (18% 260 ü 3) 449 (18); 474 (44d) 


() 
oe 501 (31 


J) 
228, : 2263 (6) 2264 (35) 2536b (37); 


) 

) 

Castro M., M., 57 (1); 83 (33a) 

Castro T., N., 170 (32) 

Cedeno, J. deD., 24 (1) 

Cedillo T., R., 2 (42); 155 (14); 197 (11); 420 (44d) 
Cedillo T., R. & J. Calzada, 45 (29); 162 (18) 

Cedillo T., R. & D. Lorence, 2345 (42); 2397 (44b) 
Cedillo T., R. & R. Torres C., 1143 (11); 1481 (18); 


= > 
L 


‘ 


) 
Chacon, I., 76 D i (44d) 
6 (23) 


B 5) 
. Hernández X., ES-794 (9b) 
C. Madrigal, ES- 125 (3 5) 
Chavelas P., J. & L. A. Pérez J., 4 (22); 223 (49); 410 


al., 1245, 1256 (11); 1947 (48); ES- 
152 (35); ES-1050 (9b); ES-1247 (35); ES-1464 (9b); 
ES-2442 (1); ES-2469 (49); ES-2832, ES-2843 (39); 
ES- 4125 (48); ES-4245 (47) 
Chávez, E., s.n. (30) 

Chávez L; O. & he E biau, 910 (14) 

Chazaro B., M., 7 (45 

Chazaro B., M. & i Dorantes, 21 (30); 110 (44d); 255 

0) 


Chazaro B., M. & Oliva, 2582 (47) 
250 (49); 


Chiang C., F., 224 (30); 251 (6) 

Chickering, A., 103 (48); 131 (18) 

Churchill, H., 5688 (13) 

Churchill, H. & G. de Nevers, 4348 (22); 4400 (18); 
4 (22) 

lin A., 4554 


(35) 
, A. & G. Cruz, 4082 (22) 
.& M. 3039 (44d) 
3346 (44d) 


¿mández 
. & J. Martínez, 
Cle . & E. Tyson, : 
Ex bra Cs 466 (59); 469 (22) 
Conrad, J. & R. Conrad, 3006 (13); 3246 
, 2890 (48 
5 (32); 54 (9b); 19, 188, 217 (49); 269 
(32); Pu (49). 337 (32); 351, 500 (41); 659, 660, 
717 (49); 936, 940 (11); 1069, 1077 (35) 1082 (6); 
1097 (32); 1098 (9b); 1152 i 1192 (32); 1225 
(9b); 1305 (35); 1308 (32); 1377 (49); 1431 (35); 
1443, 1450 (32); 1454 (35); 1462 2 (41); 1586 (9b); 
1590 (49); 1762 (6); 2046, 2058, 2070, 2089 (35); 
2090 (9b); 2117 (22); 2140 (7 » 2175 (6); 2190 (49); 
2250 (42); 2266, 2269 (49); 2304 (11); 2305, 2335 
(9b); 2362 (49); 2368 (6) 2424, gu 25; 245 n Zr 
(49); 2474 (9b); 2476, 2477 (49); 2 
2539, 2540 (49); 2554 (42); 2558 pA 2603 2 
2607, 2622 (49); 2650 (35); ni ( 
2913 (9b); 2914, 2915, 2916, 2 f 
2968 (42); 3025 (22); 3038, 3095 (49), 3170, 3181 
(9b); 3287, 3343 (13); 3383 (49); 3385 (48); : 
(49); 3403, 3466 (13); 3522, 3540 (9b); 3578 (32) : 
3610 (49); 3678 (42); 3703 (9b); 3709 (41); 
(9b); 3809 (41); 3815 (9b); 3896 (49); 3938 (9b); 
3966, 3998 (13); 4074 (41); 4080 (49); 4105 (13); 
4118 (41); 4122 (13); 4130, 4132, 4133, 4135 (15); 
4172 (41); 4319 (9b); 4345 (42); 4407 (49); 4485, 
4638, 4652 (18); 4679 (48); 4708a (35); 4712, 4774 
(9b); 5281 (4); ans (12); 5291, 5360 (9b); 5416 
592 (9b); 5609 (41); 5682 (35); 
5850, 5851, 5855 (49); 5907 (12); 
; 5990 (18); 6055 (13); _6065 (32); 6097 
(9b); 6129 (49); 6145 (18); 6157 (6); 6159 (35); 
6166 (15); 6198 (32); 6274 (9b): 6275 (6); 6282 
(42); 6312 (22); 6330, 6334 (9b); 6340 (49); 6342 
(18); 6357 (41); 6388 (9b); 6389 (42); 6400 (49); 


a 
| 
ELLE: 

>>; 


242 "^ 


(30) 


Ww 
[es 
© 
o 


(6); 7397, 7486 (49) 1524 (35). 7674 4 Aly 7685, 
7690 (6); 7704 (32); 7720 (6); 7721 (9b); 7724 (6); 
7725 (9b); 77 ps 49 (14); 7793 


(32); 7958 (14); 7976, 7997, 824 1 (9b: 8289, 8363 
(32); 8364 (9b); 8547 (32); 8550 (35); 8627 (49); 
8692 (32); 8744 (49); a (48); 8819 (32); 8839 
(41); 8876 (9b); 8882 (32); 8899 (9b); 8929 (49); 
8988 (14); 9004 (41); 9099, 9117 (32); 9159 (35); 
9374 (49); 9462 (9b); 9485 (9b); 9708 (35) 726 
(9b); 9740 (13); 9741 en 9773 : 9777 (6); 
9798 (9b); 9809 (9b); ME (35) 9851 (9b); 
(49); 9888 (12); 9901 (48); 9932 (49); ha (12); 
9943 (48); 10002, 10055 (32); 10073 (9b); 10227 
10266 (32); 10396 (13); TUE 
10430, 10432 (41); 10447 (48); 


(48); 10424 (9b); 


Volume 76, Number 3 Hamilton 905 
Mesoamerican Psychotria, Part Ill 
10468 (42); 10485, 10557 (9b); 10602 (41); 10604 34579 (35); 34591 (1); 34697 (13); 34713 (22); 
(7); 10605 (48); 10610 (7); 10684 (41); 10706 (9b); 34982 (37); 35200 (12); 35700 (29); 36338 (12); 
10795 (48); 10854, 10856, 10858 (9b); 10859 (13); 36449 (18); O (12), 36785 (18); 30867 (6); 
10860, 10864 (6); 11062 (33a); 11101 (49); 11268 36947 (53); 36982 (59); 37602 (22); 38092 (12); 
(41); 11345 (9b); 11521, 11523, 11524 (12) 38228 (4); xod es 39115, 39118 (6); 39126 
Conzatti, C., 856 in part (49); 3807 (47) (44d); 39229 (30); 39266 (49); 39569, 39598 (30); 


(48); 3265 (30 
Cook, O. & C. Doyle, 440 (48); 672 (1) 
Cook, O. & R. Griggs, 369 (1 e 540 us Jo 
Cook, O. & R. Martin, 149 (35 

Cooper, G. P., 242 (22); 453 (2 JS 


Conzatti, C. et al., 3061 ( 


— 


558 (0b) 


7 (35) 
Slater, 33, a 148 (35); 


C dl G. P. € G. 186 
(12) 

Córdoba, J. E 470 (59) 

Correa A., M., 111, 125 (13); 205 (4); 228 (13 

Correa A ; R. Dressler, 600 (44d); 836 (29); 1015 


Correll, D. S., 12 
Correll, D. S. € ci B. 
s M. E., 60, 281, 
Cortés-Vázquez, 48 (49) 
Cosentino, K., 133 (18) 
2069 (15); 2284 (32); 
T 7 (32); Sia (9b) 
B C. A. Magaña, 3267 (47) 
Cowan, C. & E Niño, 3385 (13) 
Cox, D. & M. ES 2892 (49) 


6 (35); 12272 (13) 
Correll, 28710 (49) 
495 (49) 


2890 (11); 2996 (35); 


Crankshaw, W., 
Crisman, G. & W Wil 290 s : 
roat, T., 607 (1 3): € 2 (12); € Au 401 Aa (48); 


4033, 4099, 4105 (22): - 
(48); 4138 (13); 4303 (22); 4383 (13); 4552 (48) 
48); 4922 (48); 4985 
,5518(13) 5678, 5681, 


a 
ho 
bo 

Lo b 
e 
ec 


(2 2); 6465 (48); 6481 (22) 6519 (13); 6548 (22); 


(48): 9226 (13): 10023 (10): E 10315 (2: 
10337 (13); 10347 (1); 10582 tel 10758 (1: 
10821 (12); 11150 (29); 11972 (48); 12045 ( 
12150 (44d); 12357 (29); 12389 (32); 12395, 12515 
(35); 12757 (12); 12883 (59); 13594 (18); 13737 
(37); 14114 (59) 14150a (35); 14474, 14553, 14643, 
14703 (29); 2); 14930 (29); 
14997, 15023 (49); 15090 (12); 15110 (1); 15125 
(4); 15139 (35); 15361 (4); 15402 (35); 15710 (44d); 
15906 (18); 15959 (12); 15990, 15993 (44d); 16206, 
16510 (49); 16583 (4); dde (53) 16874 (12); 
16888 (35); 17049 (49); 2 2326 a 2 (45); 
23345, 2s 23465a 


24922 (32) 
: oc ie 25303 (18); 
26028, 26048 (44d) 26103 


2 9); 27289 (2: 2); 27548, 27641 (44d); 
32755 (1) 33538 (22) 34212 (44d); 34403 (1); 


— 


39771 (44d); 40005 (11); 40157, 40166 (48); 40180 
(14); 40211, 40284 (48); 40288 (9b); 40294 (41); 
40308 (11); 40371 (35); 40606 (48); mE (9b); 

(30); 41138 (33a); 41510 (48); « 
41594 (48); s 42052 (49); 

(6); 42128 (49) 42608 (22); 
42883 (1); 43250, 43303 (61); 43779 (13); 43826 
(6); 43860 (13); 46651 (9a); 48055 (44d); 48077 
(49); 48079 (44d); 49053 (18) 49145 (53) 


Croat ¿ J. Folsom, 34052 (26); 34264 (18) 

Croat, T D. Porter, 15532 (4); 16116 (37); 16266 
(12); 16272 (4); 16403 (48) 

Cruz C, R., 1 

PIERDE, G., 5 (59): 4 6 (12); 516 (59) 
Arey E (13) 2j i) mr (22); 4002, 
4014 (35); 4020 (10); 4036b (35); 4222 T 9316 
(13); 9364 (35); 9448 » 9454 (22); 9498, 9524 (35); 
9681 (29); 9686, 10605 (22); 10674 (37); 10876 
(44d) 11263, 13615 (29); 13676 (10); 14246 (12) 
14248 (35); 14613 (22); 14987 (18); 15115, 15123 
(7); 151: 54 (13) 

D'Arcy, W. & J. D'Arcy, 6393 (24) 

D'Arcy, W. & C. Hamilton, 14977 (22) 

D'Arcy, W. & B. Hammel, 12301 (13), 12315 (22) 

aridi R., 295, 498, 597 (1) 

vidse, G., 23825 (37); 24230, 24362 (44d); 24415 

Eos 24526, 24602 (44d) 

Davidse, G. & W. D'Arey, 10062 (2 

Davidse, G. & J. Davidse, 9406, 94: 28 (6); 9428a (49) 

Davidse, G. & C. Harlan: 23483 


Davidse, C. € G. Herrera Ch., 26175 es 19 (37) 

Davidse, G. & R. Pohl, 1473 (1); 1668 (44d); 2070 
(13); 2183 (48); 2204 (6); 2245 (33a 

Davidse, G. et al., 20033 Ga 20178 (9b); 20183 (32); 
20385 (35); 23342 (6); 23455, 23772 (18) 

Davidson, M. E., 172, 363 (46); 709, 849 (28); 1120 
12 

51 C. C., 6036 (18); 6047a, 6075 (30) 

E. Siegerist, 118 (48) 

Dillon. , 1834 (14) 

Dillon, va et al., 1840 (11) 

Dodge, C. & V. Goerger, 9468 (59); 10078 (22 

Dodge, C. et al., 16636 (13); 16637 (35); 16673 (22) 
16674 (13) 

Donald, G., 2 (42) 

Donnell- Smith., J., 1821 (1); 2042 (48); 2074 (1); 2754 

4832 (22); 6594 (12); 6598, 6601 (22); 6602 

Dorantes, B., 2537 (22); 2731 (2); 2880 (11); 2897 

(18); 2915 2933 (2). 3022 (6): 3028, 3066, 3197 


3583 ( D 
e J. & W. Márquez, 609 (30) 


Dorantes, J. et al., 743 (49): 869 (30); 913 (6); 968, 
785, 1822 (30); 2275 (48); 3779 (6); 5190 (30) 
Dressler, R., 1530 (48); 4077 (53); 5964 (56) 
ressler, R. & Q. Jones, 93 (34) 
Dressler, R. & W. Lewis, 3732 (26) 
Dryer, V., 120, 653 (23); 790 (19); 1105 (24); 1243 
(19); 1531 (57); 1539 (45) 
Duke, J., 3852 (7); 3886 (22); 3988, 3997 (13); 4002 
(1); 4197 (13); 4264 (48); 4474 (59); 4701 (13); 


906 Annals of the 
Missouri Botanical Garden 
4728 (35); 4759 (12); 4764, 4769, 4779 (13); 4794 Dwyer, J. € M. V. Hayden, 6 (22); 7 (59); 22 (35); 23 
(18); 4802 ns 3) 4803 (2: 2); 4824 (59); 4953, 4954 (13); 7693 (44d); 8042 (51 
(13); 4993 (18); A. fos 2); 5114 (4); 5172 (59; Dwyer, J. € J. Kirkbride, 7440 (13); 7783 (10); 7869 
5217 (4); 5223 (22); 63 (59); 5339 (60); 5414 29) 
(59); 5517, 5517a (1) pn (12); 6130 (595 8190 Dwyer, J. & B. Lallathin, 8629 (44d); 8664 (18); 877 
(18); 8397 (12) 8443 (13), 8466 (4); 8597 (12); (44d); 8813 (35); 8816 (13); 8822 (4); 8950 (59) 
8745 (26); 8773 (12); 8801 (59); 8952 (35); 8989 Dwyer, J. & R. Liesner, 12085 (^ 
(32); ees E 9239 (12); 9259 (13); 9273 (59); Dwyer, J. = a Pippin, 10098 m 10193 (42) 
i: (13); 9363 (51); 10027 (29); 10042, 10048 Dwyer, J. . Robyns, 8, 90, 93 (1); 135 (22); 136 
2) 10176 (35); 10230 (7); 10568 (12); 10708 (7); (48); oe 160 (48) 
10% 66, 11701, 11743 (13); 11744 (7); 11897 (12) Dwyer, J. et al., 58, 84 (9b); 97 (22); 167, 281 (13); 
540 (35); 12549 (13); 13081 (59); 13138a (13); 375 (42); 436 (32); 558 (22); 4553 (2); 7244 (51); 
ee (18); 13151 (2); 13246 (6); 13701 (44d); 7288 (43); 7558 (13); 8193 (43) 
14178 (13); 14184 (18); 14185, 14210 (59); 14214 i caste: c 30 526 108 (22); 129, 168 (48); 251 (13); 
(18); 14238 (13); 14248 (12); 14271 (22); 14313 í 556 (13); 615 (22); 878, 1058, 1071 (13); 
(13); 14342 (22); 14393 (18); 14470 (13); 14560 1144 
(7), 14571, 14572, 14575 (13); 14782 (44c); 14869 pd J., 46 (48); 773, 783 (22) 
(22) 15228 (51); 15237 (21); 15538, 15618, 15690 ane J.. 469 (18); P- 291. P-613 (6); P-618 (13); 
(22) 26 (9b) 
Duke, J. € N. Bristan, 288, 342 (13); 362, 396 (22); S eh M., 502 (9b); 533 (49) 
8255 (18); 8296 (22) Egler, F., 42-264 (9b); 42-302 (32) 
Duke, J. & G. Child, 10680 (29) Elias, 1 En 1721 (35); 1729 (4) 
Duke, J. & M. Correa A., 14672 (44d) Englesing, F., 185 (22); 228 (49); 252 (13) 
Duke, J. & T. Elias, 13788 (60) Enriquez, 679 7 ) 
Duke, J. & J. Kirkbride, "i 36a, 14040 (22) Erlanson, C., 61 (35); 91 (3 2); 2 213, 236 (13); 279 (32); 
Duke, J. & H. Mussell, 6640 (35) 407 (35); 447 (13); 475 (35); 516 (32); 517 (35); 
Duke, J. et al., 3616 (59); Ro (18); 3644 (22); 3052 556 
(12); 3655 (59) Ervendberg, L., 246 (30) 
Dunlap, V., 229, 229b (59); 463 (12); 470 (7); 473 Fairchild, D., s.n. (22) 
(22); 538 (3: Fay, J. & J. Calzada, 927 (30); 929 (6) 
Duran, R. & à oer 126 (35) y, J. & C. Hernández, 826 (48); 832 (30 
Durkee, L., 7 3 (22) Fendler, A., 58 (59); 59 (10); 60 (51); 64 (22); 110 (29) 
Dwyer, J., s.n. pos 100a (1); adn o 1319 (4); 1411 Fernández N., R. € Guadarrama-Zamudio, 1379 (11) 
(13) 1540 (12% 1725 (35) 1959 (22) 2007 (35) — Ferreyra, R., 15953 (35) 
2047 (1); 2105, 2189 (22); 2 en (26); 2239 (125 Ferris, R., 5417 (13) 
2314, 2348, 2384a, 2408 ES 2445 (6); 2489 (1); Flores, G., 36 (1) 
2620 (13); 2622 (1); 2858 (13); 2875 (51); 2898, Flores, J. S., 50 (45) 
4264 (35); 4286 (13); 4374a (59); 4387 (12); 4393 Folsom, J., 1202 (44d); 1304 (29); 1388 (7); 1603 (2); 
(22); 4398 (4); 4399 (59); 4400 (22); 6540, 6560, 2280 (1); 2980 (18); 3089 (1); 3513 (13); 3534 (7); 
6585, 6588 (1); 6626 (48); 6630, 6638, 6706 (35); 3817 (43); 3858 (22); cd (13); 4025 (18); 4295, 
6762 (13); 6815 (35); 6820, 6822 (13); 6838 (4); 4428 (60); 4662 (13); 5859 (22); 8756 (18); 8774, 
6907 (35) pad (13); 6911 (12); 7083, 7098 (21); 8864 (44a); 8927 (22); ds 29 (44a); 9204, 9370 (22); 
7128 (35); 7179 (12); 7315 (35); 8326 (2); 8394 9412 (29); 9644 (59); 9880 (22); 9936 (59); 9957 
(35); 8526 e 8572 (51); 8591 (22); 9123 (9b); (18); 9958, 10004 (20); 10041 (18); 10093 (29) 
9162 (35); 9170 (4); 9837 (48); 9842 (12); 9843 . Folsom, J. & L. Collins, 1539 (29); 1615, 1621 (44d); 
(35); 9861 (48); 9902 (22); 9903 (18); 9917 (12); oe 1664 (7); 1771 (44d) 
10000 (35); 10164 (32); 10744 (6); 10763 (35) Folsom, J. arp, 1368 (43) 
10769 (42); 10784, 10828, 10845 (9b): Folsom, J. et al., 1968 (44d); 2079 (37); 2250 (46); 
10846 (42); 10847, 10858 (32); 10905 (42); 10912 3327 (13); 3723 (22); 4185 (59); 4190, 5020 (22); 
(6); 10917 (42); 105500 3); s (3 2); en MADERA 5103 (29); 5454 (23); 5505, 5569 (45); 5790 (29); 
11060 (6) 11084 (32% 110 5 (22); 6285 (60); 6589 (29); 6599, 6791 (22); 6821 (49); 
der 1115 2 Ur NT 2); e 8); 6823 (18) 
11 a(42); 1 263 (9b); 11302, 11313 (48); 11347 Fonseca Z., A., 118 (1); 131 (49) 
(3 2; n (9b); 11373a (32); 11373b (9b); 11374 Fosberg, F. R. 5525 ^ (22) 55256 (29); 55264 (48); 
55624, 55626 (22 


(32); 11387, 11391 (32); 11391a (9b); 11470 (49); 
11472 (13) 11474 (9b); 11491 ( 

11529 (9b); 11575a (32) 11: : 
12424 (35); 12436, 12451 (6); 12459, 12462 (3 
1246 2a (9b); 12536 (9b); 12537 (35); 12552, 1258 


2 


`: 


(6); 12588 (9b); 12589 (32); 12594 (49); 12595 (9b); 
12676 (49); 12700 (6); 12706 (35); 12746, 12787, 
12805, 14705 (32); 14760 (35); 14779 (32); 14793 
(12); 14933 (32); 14943 (22); 15115 (32); 15141 
(18); 15166 (35); 15370 (44d 
Dwyer, J. & M. Correa A., 7983 (18); 8423 (44d) 
Dwyer, J. € L. Dieckman, 10441 (9b) 


Dwyer, J. & A. Gentry, 9426 (44d) 


Feet, F. R. & Hatheway, 47789 (23); 47806 
(2 


4) 
Foster, R., 600 (48); 641 (4); 789 (13); 791 (22); 793 
854 (13); 899 (29); 1060 (12); 1365 (13); 1433 


(48); 
(29); 1575 (13); 1605 Pu 1607 (13); 1614 (22); 
1622 (13); 1687 (29); 6 (22); 1770 (49), 2040 
(48); 2322 (49); 2842 d 

Foster, R. Kennedy, 1274 (13); 1851 (21); 1974 
(13); 2010 (18); 2013 (13); 2017 (35) 

Frankie, G., 224a (44a); 296a (49) 

Frost, S., 100 (22); 105 (48); 222, 248 (1) 

Fuentes, S., s.n. (11) 


Volume 76, Number 3 
1989 


Hamilton 907 


Mesoamerican Psychotria, Part III 


Gabriel H., I., s.n. (18) 
tateari, m 2653 (42) 2684 (34); 2686a (47); 7078 
3 (42) 95 (30) 


128 (49): 249 (13) 


112 (13) 
Gaumer ut 27, 147, 974, 24019 
Gentle, P 4 (32) 150 (9b); a (49); 440 (32); 
532 (Ob): pes ys 928 (9b); 946 (3: 2); m (49) 
11 (32), 1215 (2b); 1255 (6); 1687 (13); 1952 
(32), 2307 (9b); 2595 (6); 2685, 2818 (22); 2851 
2871 (48); 2946 (22); 3002 (32); 3336 (22); 337 
(48); 3546, 3600, 3881, 4164 (32); 4539 (9b); 4581 
(18); 4688 (12); 4699 (32); 4862 (35); 4913, 4914 
(32); 5241 (22); 5 32), 6085 (12); 6689 (18); 
94. 29 


2); 9040, 9116 (13); 9126 


( 22); 9169 (48) 
Gentry 778, 820 (49); 895 (1); 2292 (35); 2294 
(22); 2296 (13); 2297 (35); 2304 (13); 2314 (18); 
3281, 3310 (51); 3318 (12); 3648 (45) 


4 5); 5736 (13); 
(45); 6414 (12) 6417 (12); 6475 (18) 6600 (29); 
; 1814 (6); D (32); 7849 
(15); 8212 (32) ae 8406 
8790, 8791 


a 


- 13490 (1); 13598 (53) 


Gentry, A. & A - Clewell 6938 (59); 7135 (22) 

Gentry, A. & J. Dwyer, 3361 (51); 3383 (12); 5514 
2) 

Gentry, A. & E. Lott, 32499 (29) 

Gentry , 13895 (53) 13902 (7); 13916 


A.&M. Nee, 8702 (13); 8721 (51); 8741 (22) 


Gentry, 
Gentry, A. & E. Tyson, 1669 (49); 1693 (13); 4848 
18) 
Gentry, A. et al., 3513, 8867 (51); 16826 (53); 16958, 
28589 a 28593 (59); 43930 (44d); 44001 (6) 
oe J. 7101 (30) 
Gilly, E E. i e X., 339 (47); 378 (9b) 
heh L., J., 658 (30) 
Glassman, S., 1830 
Godfrey, R., 66170 (2: an 67041 (1) 
Goldman, E. A., 141, 840 (30) 
7 (18) 18, 19 (35); 22 (13) 


Gomez, J. A.. 


Gómez, L. D. et al., 20357 (5); 20381 (26); 20510, 
20521 (35); 21509 (37); 21970 (44d) 
Gómez-Pompa, A., 2019 (18); 4468 (14) 
Gómez-Pompa. A. & L. Nevling, 1158 (6) 
Gómez-Pompa, A. & M. Rosas, 3970 (14) 
Gómez-Pompa, A. et al., 3065 (44d); 3436 (21) 
González L., L. A. & V. Garza, 604 (48); 8567 (9b) 
González L., 4 A « He jura P., GH134 (9b) 
iow bi A. & L. A. Pérez J., 3896 (9b); 4004 
(49); de (47) 
Pad L. L. A. et al. 9422 (44d) 


e Ortega, J., 75, 103, 614, 4013 (13); 5875 (30) 
Conde O Lo 139 (30): 1485 5 (48) 1789 (0b) 
uer Medrano, F. et al., 2 (18) 

Gouin, s.n. (30 

Graham, S. A., 197 (22 

Grant, V., 849 (6) 


M., 2114 (59); 2745 (12); 2874 (50 
947 (444 jy 999 (13); 1961 (6); 237 
3793 (12); 3822 (48); 3 

(47); 61, 


Grayum, 
Grijalva, Á., 
1); 2872 (47); 
Grijalva, A. & M. Araquistain, 41 
37% 212 P Os 656 (44d) 
Grijalva, A. ". Burgos, 1522, 1579, 1588 (13); 1601 
(32) IA 1651 (22)2 1T 12 (6s I712a (13): 
1721 (6); A Y 
Grijalva, A. & M de Grijalva, 1282 (13) 
Grijalva, A. & P. P. Moreno. 1063 (6); 1102 (E 
(13) 1198 (32 
Grijalva, A. & D. Rios, 3489 (2: 
Grijalva, A. & D. Rr. 3736 
ria. A. et al., 99 (6): 
917 (6); 2998 Pip 3376 (18); 


Guevara, A., 3 (18) 


1 (6) 175 


22); 1159 


14 (44d); 2893 (49); 
3081 (6) 


1 
, 39 (6); 273 (11 
, 234 (11) 
55 (49 


Guzman, M. et al., 345 (1); 1212b (13) 

Haber, W., 380 (49); 447 (19): 470 (49) 531 (45) 

Haenke, T., 2495 (59) 

Hagen, C. von & W. von Hagen, 1103 (48); 1114 (18): 
1194 (35); 2018, 2024 (37) 

Hahn, L., s.n. 

Hahn, W., 101 aay 123 (7); 163 (18); 296 (12); 373 
(44d); 380 (6) 473, 482 (18); 484, 492, 510 (44d): 
633 (22) 

Hall, J. & 5. o 7542 (44d); 7572 (6) 

Hamilton, ~ 52 25 551 (49); 567 (13); 3677, 3678 
(1); 36 oa 3081 (13) 

Hamilton, fs '& W. D`: Arcy, 686 (29); 1328 (49): 1335 
(18); 1336 (1); 1411 ( 

Hamilton, C. & I Dese i 2986 (44 ud) 

Hamilton, C. & A. Gentry, 3272 (3! 

Hamilton, C. & K. Krager, 3194 (22); 3197 (26); 3773 
(44d) 3784, 3949 (18); 4167 (29) 

Hamilton, C. & Y. Palmer, 2951 (49); 2963 (22) 

coro, C. & H. Stockwell, 1481, 1504, 3121 (22): 

4 (18); 3082 (29): 3692, 3693 (59); 3695 (22); 
ed 3697 (35); 3706 (13) 

Hamilton, €. et al., 513 (13); 7 7760 (44d) 1196 
(29); 1290 (18); 3255 (51); 3296 (1); 4043 (29) 
Hammel, B., 908 (29); 1124 (22) 1130 (7); 1195 (22 
1256 (18); 1311 (22); 2221 (23); 2558 (44d); 272 
(22) 3595 (56); 3665 (53); 3765 (44d); 3821 (4): 
3905 (1); 3959 (51); 3995 (59) 4178, 4332 (13); 
4634 (54); 5227 (56); 5321 (12); 5371 (13); 5413 
22) 5428 (18); 5583 (29); 6061 (44d); 6072 (37): 
6189 (40); 7227 (36); 7320 (12); 7323 (13); 7778 
(20); 7790, 7800 (59); 7992 (44a): 8005 (22): 8107 


(50); 8294 (22); 8706 (20); 8730 (22); 8790 (59); 
9032 (29); 9220, 9420 (44a); 9443 (50); 9447 (12): 


9461 (44a); 9486 (22); 9489 (: de 9927 (22); 9977 
(44a); 10091 (22); 10938 (20); 179 (22); 1149] 
(20); ea 12400 (22); er T 

Hammel, B. € J. Trainer, 12769 (59); 13085, 13142 
(44a) 13154 (22) 

Hammel, B. et al., 6863 (18); 12808 Pu 

Hansen, B Nee, 2 d to 2 (3( 

Marmot W., 40 B 1 (27): RU (49), 2513 
(9b); 2 538 " Au ae un 5811 ( 

ion M . & J. Dwyer, 2768 (35); 2786, 2793 (9b); 
2815 (35) 2837 (12); 2844 (49); 2867 (9b); 2960 


Annals of the 


908 
Missouri Botanical Garden 
(35); 2961 (6); 2971 (49); 3105 (30); 3711 (13); 1071 (32); 1218 (22); 1551 (29); 1554 (51); 1636 
3953, 3955 (6); 4055 (13); 


3887 (35); 3945 (13); 
4332 (9b) 
Harmon, W. & J. A. Fuentes, 2090 (49); 2375, 6432 


(13 

Harris, S., 7, 12 (35) 

Hartman, R., 12028 (59); 12088 (22); 12099 (59); 
12211, 12490 (18) 


Hartshorn, G., 955 (29); 1107, 1261 (44a); 1390 (22); 
2 


6) 
., 9039 (22 
Hatheway, W., 1460 t 
Hayden, M., 8 ui , 14 (22 PA 5 (13); 26 (48); 28 
(22); 37 (48); 3 9 (13); 66, 76 T 100 (29); 
104 e v 2); Ie. 3); 174 (49); 175 a 
e , 164, 200 (22); 396 (1); 477 (59); 562 (22); 
70 v 1458 (59) 
Halen D., s.n. (44d); 684 (49) 
Heithaus, E., 96 (13); 109, 113 (49); 114 (13) 


22 (6) 


Heller, A., 
y J. & W. D. Stevens, 167, 174 (13); 307a (18); 


325 (42 
Heriberto: De 74 (1); 164 (35); 207 (13) 
Hernández, A., 2 (18) 
Hernández M., R., 1250 (11) 


Hernández M., R. & B. Rosales He 1559 (32) 


Press An R. xr — Kirkbride, J. & N. Bristan, 1449 (59); 1460 (12) 
Hernandez P., & M. Hernández M., 5160 (12) ao * 12 64 4): APT ru em te 
pl Mas F. et al., ES-367 (35) 1357, 16 48 (22) ; 
i yde, s 0&8 E om Te 10610 (49) Kirkbride, J. & M. Hayden, 298 (29) 

Eur 7 eta 6 > Knapp, S., 952 (12); 955 (22 2 3); 1037 (22); 

11 , 13912 (13); 14166 (42); 14352 (13); 14598 MM (13); eue 35); eerie. (59 — oe TES us 

( 49) (22) 1300 (35); 1339 (22); 1344 (59); 1537, 1580 
Hoffmann, C., s.n. (6); s.n. (19); 55 (24) (18); 1646 (46); 2182 (13); 2277, 3877 (22); 3882 
Holdridge, L., 2331 (9a) (13); 5963 (1) 

Holm, R. € H. Iltis, 22 (12); 231 (1); 342 (22); 354 Knapp, S. & W. J. Kress, 4379 (44d) 

(50); 355 (12); 861 (18) iron S. & J. Mallet, n os 2948 (32) 2959 
Hoover, W., 130 (48); 198 (9b); 237 (35) (13); 2988, 2995 (22): 22); 3054, 3063 (26) 
Horvitz, C., 206 (14) 3102 (29); 3174 (48); Pen Wt 2) 4649 (355 57 
Huft, M., 1776 (1) 22) 

Huft, M. & K. Barringer, 2030 (12) Knapp, S. € R. Sensus 3595 (22); 3631 (59); 4841 
Huft, M. et al., 1950 (7); 1974 (13); 2077 (35); 2108 (1); 4857 (2: 2); 4 se 

(1) Knapp, S. & Sytsma, 2313 (29); 2342 (7); 2368, 
Hunter, A. & P. iad 240 (32); 461 (22); 699 (35) 2444 (18); 2513 (44d) 2517 (18) 

Huston, J., 620 (32) Knapp, S. & M. Vodicka, 5538 (54); 5595 (23) 
Ibarra, J j 10 (49) Knapp, s. et al., 2029 (44d); s (54); 4153 (45); 
Ibarra M., 22 (14) 4178 (18); 4196 (37); 4198 (4 
Jaén, E., ` 5) Knight, D., 69-4 (22) 
Janzen, D., 10426 (13); 10443 (35); 10611 (13); 10684 Koch, S. et al., 79288 (9b); 79293 (18); 79294 (13) 

(35); 10766 (1); 10979 (13); 10980, 10112 (35) Koptur, S., 122 (19); 146 (6); 171 (24) 

11534, 11545 (22); 11711 (13); 11750, 11786 (35); cess, W. 77-784 (29) 

11814, 11898, 12006, 12007, 12008 (13); 12021, Kress, W. J. et al., 9594 (19) 

12022, 12023, 12024 (35); 12025, 12034, 12074 ruse, H., 277 (13) 

(13) 12091 (3: na 12121 (13) 12122 (49; 12123 Kuntze, O., 1857 (22 

(13) 12151, T-107 (1) Laguna, A., 23 (12); 127 (29); 131, 143 (44a) 
Jaramillo, S., 4 (1) Lamas, R., 368 (9a) 
Jiménez M., A., 5 er e 503 ir 661 (22); 1059 Lamb, F., 54, 62 (15) 

(29), 1150 (6); 1937 (18); | (19), 2366 (22); pun t 3769 (30) 

3103 E 3269 m 3308 e cae 5 (24); 3448 (59); Lao, E., 9 (45) 

3831, 3 (22) T E. 3 " Holdridge, 207 (51) 

Johannessen, C., 499 s ao, E. 392 

Johnson, E. P., 61 (9b); 79 (49) Miren Ms 695 (9b); 771, 1329, 2134 (30) 

Johnson, H., 90, 512 (49 ) Lauvert ` Barkley, 39552 (33a) 

2, 297, 314 (35); 349, Lazor, R., 2276 (1) 


18, 18a (13); 142, 2 


Johnston, I. M., 


9) 
Johnston, M. C., 4945 (30) 


Jones, G. et al., 3108 (9b) 

Juan, A., 135 (35 

Juan, A. & M. VEI 12 (11) 

Judziewiez, E., 445 3) 

Karwinsky, W. von, 300, 300b (6); 1262 (49); 1263 
30 

Kellerman, W., 7580 (22) 

kelly, I., 183 (9b) 

Kennedy, H., 1215 (18); 1216 (13); 1857 (12); 2334 
(22); 2671, 2775 (13); 3241 E 3259 (29) 


2) 
Kennedy, H. & 


2) 
Kennedy, H. & R. Foster, 2151 (51); 2215 


903 (2 
2135 (59); 
Kennedy, H. & 5. Gra, 2232 (22) 
Kennedy, H. & C. Wilder, a (59) 
Kennedy, H. et al., 2047 
Kenoyer, L., 562 (22); Eon (13); 566 (48); A265 (30) 
Kenoyer L. & E. Crum, 3715 (30); 3903 (49); 3917 
49); 4013 (9b); 4062 (49) 

Killip, E., 3118 (13); 3407 (1); 39960 (22) 
King, R., 1955, 2742 (13), 3918 (30) 
Kinlock, J., (32 ) 


Kirkbride, J., 39 (13); 192, 202 (35); 226 (48 


Volume 76, Number 3 
1989 


Hamilton 


909 
Mesoamerican Psychotria, Part Ill 


5364 (22) 5396 (59) 
H. Hongsrand, 1460 (49) 
18) 


Lazor, R. & E Pim, 


pus rl 


LeClezio, 19 de 2 (59); 2 

Lent, R., 1307 (24): 17 pus n 2246 pores 2471 (12); 
2589 (18); 2727 (44d); 2924 (19); 3282 (22); 3863 
(19) 

Lent, R. et al., 3412 

León C., J. M., 114 (11); 179 (22); 653 (59); 673, 715, 
949 (22); 1556 (59); 1690 (1: 2), 1856 (22); 1872 


1883 


(4); 1880 (12); (35) 
568 (30); 633 (6) 


LeSueur, H., 352 (6); 
Lévy, P., 510 (18) 
Lewis, W. & R. Sharp, 14 (29) 
Le 


ewis, W. et al., 156 (4); 253 (22); 822 (48); 963 (59); 
1512, 1540 (1); 1830 (22); 2103 (G9) RA 
2249 (17); 2915 (13); 3248 (22); 3310 (49); 35 
5205 (1); 5320, 5340 (22) 


Lewton, F., 25 

Leyva, C., s.n. (18) 

Liebmann, F., s.n. (6); s.n. (47) s.n. (495 11576, 11577 
(9b); 11578 (1); 11580, 11582 (6); 11595, 11598 
(3 011602. 11603, 11604, 11605 (11); 11614 (9b); 

3), 11630 (11); 11645 (18); 11645a, 11646, 

30); 11658 (6); 11662 (9b); 


26); 767 (18); 
; 1718, 2011 (22); 
5, 2991 (13); 3263 (44a); 
4342 (35); 4401 (13); 4863 
218 (44d); 5222a (44d); 


MK 


; 4250 (Ly 
; 5060 (6): 5 


i 14383 (59) 
Liesner, R. € J. Dwyer, 1464, 1598 (13) 
Liesner, R. & E. Judziewiez, 14537, 14914 


(18) 
Liesner, R. & R. Lockwood, 2396 (13); 2409 (35); 2452 


(1); 2529 (49); 2637 (35 
Liesner, R. et al., 2779, 3470, 15125 (1); 15222 (12); 
15230 (18) 15298 (22); E 7); 15315 (12); 
15377 (23) 15400 (52); 15563 (44d); 15585 (24) 
Linden, J., s.n. (22); 787 (49) 
oera, F. et al., €-3 (13) 
Long, L., 202 (29 
López, A., 5 (48): 19 (13) 
López, R., 17 (13) 
López-Forment C., W., 7, 224 iss 
Lorence, D. & G. Castillo C. i 4283 (30) 
Lorence, D. & A Cedillo T., 12) 
Lorence, D. & T. P. animan , 3289 (14); 3290 


(11); 3863, Mo (3€ 
Lorence, D. & R. Tone 4018 (44b) 
et al., L (42); 3934 (18): 3935 (51): 


41 38 a 


3) 
310 (32); 


: 3984 (49) 


2. pu 6105 (35) 6193 (42) 
6234, 6249 (41); 6250 (: B) 6260 ery 6t 
6702 (32); o [5232(42) 1 4 (41); 
(12); 15404 (41); 15705, 15738 (9b: 157 


"LES tu 
2 ho 


15891 (9b); 16152 (6); 16158 (41); 16214 (32); 
16253 (35); 16270 (22); 16283 (48); 16378 (32); 
16380 (42); 16394 (9b); 16429, 16510, 16628 (49); 
$ : 17025, 17041, 17633 (41); 
1) 17681, 17735 (13); 17801 (1); 
21 (9b); 17856 (13); 17987 (48); 
18247 (7); 18339 (12) 
js : ntreras, 18861 (49); 18920 (Qa); 
18946, 18953 (44d) 19012 (32); 19025 (9b); 
(12); 19063 (15); 19180 (9a); n e dis 
(9b); 19352 (49); 19352a (48); 
19440, 19443, 19449 (44d); 


ce 
— 


— 
OREA! 
ec 
uw 


5) 19964 ; (ob) 
19987 (48x 19997 
(42); 20099. (ny 


20207 (9b); : 
(32); 20443 (44d); 
20540 (42); 20545 (49), 20555 
20637 (9b); 20678 (15); 20808 (9b); 

7 (41); 21068 (9a) 
Lundell, C. ^ A. Tandel 7144 (30); 7677 (49) 


ia 
20483, mu dd ane 
(35) 20570 (13); 
20811 (42); 


Luque, J., 45 (1 

Luteyn, J., EE (48); 1083 (13); 1406, 1416 (22); 3988 
(29); 4022 (18) 

Luteyn, J. & R. Foster, 1382 (13); 1387 (1) 

Luteyn, J. & H. Kennedy, 4157 (12) 

Luteyn, J. & R. Wilbur, 4655 (43) 

Lyonnet E., 2173 (30) 

McDade, L., 569 (13); 570 (22); 578 (12): 580 (59) 

McDaniel, S., 5012 (22) 5051 (4); 8169 (22) 

McDaniel & R. TE 14859 (1) 


McDiarmid, M- a 73-32 (1): 73-58 (13); 74-31 
(1) 


MacDougall, J., El 4a) 

MacDougall, T., s.n. (2); H28 (30) HI: e He 02 o 

McDowell, T., 306 us 341, 360 (59); 735 (44a); 77 
(22); 1013 (18); 1066 (29) 


McPherson, G., 737 (48) 
31 463 (49) 


Mac Vani: C. 43 (44d) 

Magallane s, J. A. S., 708, 2456a (13) 
Magaña A., M. A., 49 (6) 

Magana A., M. A. & R. E 189 (14); 201 (18) 
Mancias, J. & J. Hernández, 1138 (35) 
Marcks, B. & C. Marcks, E. e 
Marin, S. & R. Cisneros, 164 (47) 
ae R., W 16 (48 

Márquez R., W. et al., 62W (30) 
Martin, P. & B. Harrell, 100 (18) 
Martinez, J. & C. Bejarano, 236 (33a) 
Martinez, M., 48 (9b) 


34 (4 | 47 
enn : 
209] (49) 


Martinez Calderon, G., 13 (1); 30 (30); 
(9b) 166a (49); 166b (30): p : x 296 ( 


(49): 1762 (48); 1785 (29); 1790 (14): 

Matuda, E., 528 (13): eats m 30); 2277 (6); 
2926 (9a); 3025 (1); 3146 (6); 3264 (9b); 3266 (32) 
3286 (8) 3366 (35) 3373 (9b): 3390 (49) 3437 
(48); 3710 (11); 5573 (9b); 5815, 15813, A 
(30); 16387 (13); 16438 (22); 16461, 16727 (47) 
17144 (1) 17762 (9a) 17985 (0): 18002 (1); S-4 


Msi, P pL die = (48) 
8 (2 


Maxon, W. 


910 


Annals of the 
Missouri Botanical Garden 


Maxon, W. & A. 3 
6634 (13); 6635 (35); 6676 (13); 6694 (22) 
Maxon, W. & R. Hay, 3243 (12) 
Maxon, W. a A. Valentine, d 6937 (13) 
Meave, J. et al., B-8 (22); B 35); B-120 (48); 
(9b); p.128 5295 (3 A 1); B-5 
Medina, Ls 


B-127 
94 (48) 


160, 205 (22) 

Mendon. A. & R. Bracho, 826 (30) 

Menéndez L., F., 67 (44d); 114 (6) 

Mexia, Y., 1262 (42); 1686, 1846 (47); 
9265 (39) 

Meyer, F. G., 22 00 (12) 

Meyer, W., 98 


9221 (11); 


Miller, G., 1900 (13) 2047 (22 

Miller, J. & M. Nee, 137 Du 1375 (1) 

Miller, J. & J. Sandino, 1068 (12); 1104 (18); 1165 
(44a); 1254 (59 

Miranda, F., 1935 (30); 4208 (48); 4859 (42); 4945, 


; 0395 (34); 5445 (30); 5834 (34); 5836 
7461 (9b); 7584 (14); 78: 30 (: 44d); 8518 (30); 9150 
(36 ee 

Mitchell, E., 
Molina, D., 


87 (6) 
109 5 
- 2 (13) 920 (44d); 967 is 
(44d); 1842, i (32); EA (22). 2 
(4); 2317 (48); 2 37 0 (4); 2 


a 3653 (6 " í 
: 9) 5046 (33a); 5404 (13); 
6464 (44d); 6710 (6); 6722 (49); 6 
(9b) 7013, 7048 (6); 7068 (35); 219 (18) 
(44d); 8337 (1); 10086 (13); 10309 (22); 10062, 
10823 (44d); 10946 (18); 11760 (47); 11910 (13); 
12991 (9b); 13685 (44d); 14150 (9b); 14185 (35); 
14245 (9b); 14251 (33a); 14378 (13); 14623 (9b); 
14630, 14653 (33a); 14749 (32); 14925 uot 15041 
(32); 15060 (22); 15083 (12); l 5088 (4); 
15498 (9b); 15508 (6); 35 (35) 
549 (13); 15566 (48); 15578 (42); 15644 (13); 
15645 [33h 15653 ties 15669, 15775 (32); 15786 


(42); 15827 (32); 15829 (42); 15851 (7); 15867 (22); 
15877 (48); 18035 (22); 18648 (33a); 18652 (9b); 
18705 (32); 20462 (6); Be (35); 20516 (37); 
0580 (18); 20717 (13); 2 En (22); 21983 (13); 

2021 (49); 22447 (13); 22557 (9b); 22615 (44d); 
22920 (37); 22970 (47); 23013 (13); 23303 (32); 
25412 (9b); 25424 (33a); 27968 (48); 27969 (22); 


30464 (35); 30578 (44d); 30609 (13) 

Molina R., Molina, 12183 (9a); 13882, 
13983, 1. 1095 ( Hd 24350 (3: 2); 24394 (44d); 24627 

; 24794 (49); 24804, 26735 (13); 26778 (49); 

784 (30); 27876 (32); 30521 (37); 

307 46 (49); 30835, 31059 (9b); 31091 (32); 31105 

(47) 

Molina R., A. & E. Montalvo, 21607, 

Molina R., A. & L. O. Williams, 20025 (44d) 

Molina R., A. et al., 17281 (29); 17325 (44d); 
(29); 17649 (6); 18047 (22 
31280 (32) 

Moloney, C. A., s.n. ( 

Monsalvo G., F., 

Montalvo, E., 4695 (33a) 


21831 (41) 


17609 
; 18146 (4); 18192 (22); 


ey, 6582 (1); 6606 (35); 6614, 
D29 


a E. & D. Vargas, 3229 (1) 

34 (1) 

+ Wood, 3660 (49) 

Morales dde J., 3 (30) 

Moreno, P., 209 ves 

Moreno, P. P., m (495 1369, 4121a, 4175 (44d); 
5449 (1); 6299 (49); 6420 (6); 6462 s 6664 


(47); 7459 (44d); 7713 (22); 7752, 7759, 8036 (37); 
9468, 9514 (6); 10232 (47); 10242 (6); 11123 (42); 
11692 (13); 12371, 12415, 12448, 12524 (12); 


13261, 13270 (22); 13759 (49), 14805, 15023 (18); 
15040, 15072, 15148 (22); 15162 (44a); 15167 
(18); 15476, 15487 (42); 15513 (6); 15556 (44d); 
15628 (6); 15666 (47); 15939 (6); 15956 (49); 16009 
(44d); 16126 (6); 16146, 16188 (44d); 10224 (35); 
16450a, ee (49); 16451 (6); 16953 (13); 17167 
22) E iiu. O 18264 (47 
(18% 18822 (6); 824 (: 1882 
35) 18854 (37); 
22): 19636 (6); 19687 (22 
ere 19696 (18): 19769a opas ie 88a (22); 19806 


~ 
7T = 


(35); 19941 (1); 20073 (37); 20193 (47); 20217, 
20232 (18); 20240, 21132 (44d); 21678 (37); 22110 
(65 22151, 22981 (22); 23060 (5); 23117, 23783 
22); 23840 (49); 23901 (29); 23904, 24101 (22); 
24119 (49); 24236 (13 

Moreno, P. P. & J. Henrich, epe 8395 (6); 8469 
(49); 8479 (6); 9116 (18); 9 a ») 

Moreno, P. P. & A. López, a 

Moreno, P. obleto, 2 (44d); 20651, 


20695, 207 43 (22); 20758 (29); 20839 (22); 21016 

Morena: P. P. & L. Rot 

Moreno, P. P. & 
(6); (1: 
(35); 12017, 12049, 12064 (48); 12195 (2: 
(48); 12215 (22); 12259 (44a); 12494 (12%. 
12609 dins 12617 (12); 12850 be. S 


, 4972 (12) 
A 7583 (37); 7599 TE 
11905b (1 2) 11913a (48); 11913b 


1848 


13042, 
Moreno G 
M 3, 7003 (59); 7032, 
18); 7122, 7133 (43); 7734 (44d) 

1190 (44d); 7240 (46); 7328 (46) 
, 7798 (44d); 7809 (37 
1963 (59); 2010 (29) 


22); 7080 ( 
¿ A. Bolten, 
R 


3002 (59) | 


37 19 (51); 4745, 41 
(51), 5213 (59); 
0507 (43 

Mori, 5. & J. T Poen 


5324 (18); 5325 (45); 5660 (37); 


1951 (51); 7952 (59); 7954 


pom 9 (5 39); 
T. s n 2y 
Moron. IL 

Morton, C. 4 o 
Mueller, F., 424 (39) 


7537 (18) 
819 (48); 868 (59) 


(48) 


Muenscher, W., 12370, 12381 (1) 
Narvaez Montes, M. & A. Salazar, 614 (13 
Nee, M., 7082 (22); 7104 (4); 7207 (13); 8036 (1); 
637 ee. 8946 (51); 8993 (35); 9091 (22); 9419 
(59); 9523 (13); 9598 (22); 9600 (51); 11682, 11688 
(10); 2 2375. 22995 (49); 23880 (44d) 
Nee, M. pS J. Dwyer, 9235 (12) 


Volume 76, Number 3 
1989 


Hamilton 911 
Mesoamerican Psychotria, Part Ill 


Nee, M. & A. Gentry, 8688 (51); 8725 (59) 

Nee, M. & B. Hansen, 18372 (49); 18483 (30) 

Nee, M. & G. Schatz, 19873, 19927, 19977 (44d) 
Nee, M. & D. Smith, 11105 (22) 

Nee, M. & E. Tyson, 10937, 10939 (26) 

Nee, M. et al., 8763 FA 

Neill, M 129 (37); 2 m. c is 2n 652 


(37); 2789 (49); 3167 (37) : 

3658, 3720 (49); 3726 (48); 3791 (1 2 3876 (32) 
4149 (12); 4188 (13); 4227 (35); 4249 (6); 4267 
(13) 5); 3d 4465 (48) 


ee (18); 3264 (44d); 
Neko, Gs 
Nelsoi, m 
e E. nera: 4104 ma 4366, 4488 (13); 
> & E. Vargas, 2359 (49); 2632 (35); 2738 
(22) 2740 (12) 3456 (35) 3473 (13) 4906 (32) 
Nelson, C. et al., 91 (32); 3415 (35); 3542 (9b); 5574 
(13); 5843 (1); 2T 3); 5938 (49); 6022 (6); 6093, 
6125 (1); 6566 (49) 
Nelson, E. B., 


, . Gómez-Pompa, 557 (30); 903 (47); 

25 10 24 (6); 1163 (30); 1667 (9b); 2359 (30); 
2483 (44d); 2564 (18); 2603 (48) 

Nichols, C., 407 (29) 
velo, A. et b 51, 129 (9b); 204 (13) 

Oersted, A., (13); s.n. (49) 15 (45); 11575 (1); 

11611 (18); 11621, 11623, 11628 (13). 


11594 (35 5) 
11634 (22); 11637 (24) 

Oliver, R. et e 3691 (29) 

Opler, P., 139 (1); 325 Ae 3); 584 (6); 628 (22) 662 
(6); 098 (18); 834 (13); 883 (59); 980 (44a); 1683 


inui A , 3106 (30); 3161 (6); 3226 (11); 3405 (30); 
345 


Nos A. D. L., 384 (6) 
Ortega O., R., pr (30) 
Un O., R. et 1225 (11) 


Ortiz, B. & a 37 (11); 71 (2) 

Ortiz, F., 115 (59); 204 (4); 770 (49); 967 (22); 
35) 

Ortiz, R. a SUE 29 (9b); 11 (41); : 


36 (9b); 54 (12); 
E 104 (6); rey i: |: 
» 


5€ 1) 314, 


1016 ul Bre (13); 


1097, 1142 2 dre f L64, 1197 (35); 1234, po (9b); 
1344 (22); 1357 (35); 1388, 1447 (12); 1477 (6); 
1480 (12); 1482 (35); 1500, 1508 (13); 1509 (48); 
1521 (4); 1578 (41); 1583 (22); 1586 (9b); 1595 

3 1 y 35); 1926 


Palmer, E., 117 (30); 118 (49); 347, 384 (6); 418, 478, 
638 (30) 


Paray, L., 322 (30); 1946 (34); 1962 (14); 1971, 1975 
(6) 2316 ee 2661 (13) 

Paredes, E. J., 97 (33a 
arrales, N., s.n. (44d 

Peck, M., 23 (32); 385 (35); 709 (22); 803 (6); 888 
(12), 915 (18); 959 (49) 

Pelly, R., 9 (49) 

Pérez, A. & J. González, 39 (13) 

Pérez J., L. A., 279 (30) 

Pérez J., L. A. € C. s UN 1620 (35) 
érez S., G., jus 57 

Pfeifer, H., 140 2 REM (13) 

Piper, C., 5554 (32). : 9822 (22); 5986 (48 

Pipoly, J., 3572 (29); 3597, 3650, 3790, 4000 (22); 
4045 (32) 71 (49) 4295 (35) 4377 (6); 4396 

3) 4507 (35) 4586 (22% 4748 (13): 4719, 5030 


289 (9b): 377 (12) 23722. »372b (35) MUR 
(29) 2483 (22) 2805 (35) 2 


bo 
m co 
"M jmi 


A 32a (13); 3936 (35) 1092 2 ( bus ) 4181 (35) 
99); 5742 (22); 6969 (13); 7593 (22); 9657 (29); 
11090 (18) B es 12412 (44a); 12681 (48); 
13395 (22) 16279 (38) 
a H., I: 171 (€ e 132 (7) 
once C., F. & R. Cedillo T., 13 (44d) 
Poole, J., 1 ] (48 
Popenoe, H., 50 (49) 
orter, t al., 4052 (35); 4165, 4811 (29); 4993 
22) 
oveda, L., 129 (4); 1028 (12) 
Pina. R. et al., 196 


1, 9041 (18); 5292, 
ra G., 32118 (29); 32124 (18); 36591 (2: 
(35 


7682 (30); 8198 (47) 
2y 38954 


Proctor, G. et al., 26916 (22); 27134 (29); 27233, 
21305 (22 
Puga, B., 19 (13); 22 (35) 


Puga, L. M. V. de, 5 
; 669 (11); 3270 (18); 
; 4637 (30); 4939 (6); 


Purpus, i sum (Tk -3 5 (30) 2208 (49); 7023 
5 (39), LOLOL nn 10265 (30); 10705 (39); 
V QUNM 10713b (30); 10730 (9b); 10756 (30): 
10815 (49); 10828 (9b); 10889, 13024 (30); 13024a 
(49); 13086 (30); 14124 (39); 14132 (9b); 14138, 
14163 (30); 14264 (9b); 14315 (49); 14334 (30); 
14340 (47); 14398 (9b); 16464 (49); 16676 (18) 


Ramirez C., D., 9 e 154. 319 (49); 373, 411 (30); 


, D., 4623 (14 
Rendon, G. & M. Hernández M., 5379 (1) 
Riba, R. & A. Gómez-Pompa, 312 (30) 
Richards, P., 6248 (18) 


— 


Ro 
Robleto, W., 218 (22); 229, 483 (44d); 524 (12); 544 


) 
(1) 572 (22); 595 (12); 692 (1) 


912 


Annals of the 
Missouri Botanical Garden 


Robyns, A., 65-5, 65-10, 65-27 (22); 65-30 (48) 


Rodriguez, J. V., 2 (18); 431, 505, 836 (49); 1841 
(33a); 2744, 2860 (32); 3054, 3270 (49); 3271 (13) 

Roe, K. & E. Roe, 2235 (49) 

Roe, K. et al., 801 (48); 900 (11); 903 (14) 

Romero, H. O., s.n. (13) 

Rosas R., M., 1293 (4 7), 1325 (14) 

Rosas R., M. & R. Hernández, 133 (47) 

Rose, J., 3284 (13) 

Rovirosa, J. N., 156 (47); 2 ae 545 (49); 732 (6) 

Rowland, L. & 5. Snedaker, PN 2) 

owlee, W. & H Rowlee, 166 

Rzedowski, J., 4818, 7449 (30); es 29 (9b); 7772 (30); 
9741, 10144, 10479 (9b); 10577 (18); 10664 (30); 
11126 (49); 20348 (48); 30658, 32973 (44b); 33188, 
33189 (30) 

alazar, L. € 5 (1) 

Sanchez L., V., 1029 (49) 

i (6); 483 (35); 492 (44d); 


PAREM. hc C. 41 bs 47 
5( Sis 32); 954 (6); 1283 (6); 1353a 
"2180 (35); 2321, 2: 


48 

ES: Aldubin, 4379 2 

Martinez, 3886, 3906 (1) 

; Mckearin, 410 (44d) 
3074 (13) 


i. (48); 2000, 2119 (9b); 
9); 2360, 2373, 2805 (9b); 3880 (49) 

563 (35); 635 (6); 1039 (35); 
E M. Nee, 233 ( 
. 258 (30); 


2132 
1215 (22) 
45) 

261 (9b); 385 (30); 515b 


04 (32) 242 (48); 245 (22); 644 An 
Ib 03 S. 394 E ae ee 5-608 (49); 5-637 
15); S-640 (42) S-714 (32) 
Schlumberger, H., i (49) 
Schott, A., 524 Tm 
Schubert, B., 612 (22); 1052 (6); 1315 (18); 1373 (59) 
Schultes, R. & B. Reko, 732 (18); 757 (47) 


(44d); 358 (18); 520 (35) 

Seler, C., dus v 5036, 5441 (6) 

. & E. Seler, 2800 (30); 5463 (48); 5502 (14) 
(13x 1612 (49); 1617, 5471 (30); 


(7) 4520 (22) 4564 (35) 4689 (13) 4708 (48) 
4713, 4772 (22); 4846 (12); 4857 (22) 

Shapiro, G. & D, Elliott, 437 (48) — 

acc :k, O; j, 274 (22); 416, 

3 (48); ue 1115 (4); n" 
Be B. & C. L. Smith, 1893 
Skutch, jo acis (4d); 1486 (9a); e 3 (2); 2087 (9b); 

: 3137 (18); 3282, 3330 (45); 3883 (59); 


61 (48); 932 (22); 
52 (29) 


2873 


e 


3904 (7); 3955 (22); 4318 (13); 4763 (22); 4949 
(12) 
Smith, A., 3, 91 (44d); 105 (19); 165 (46); 334 (19); 
383 (44d); 485 (19); 543 (46); 619 (18); 646, 699 
(19); 847 (44d); 917 (10); 1057, 1105 (44d); 1708, 
: 2587 (29); 2612 
(18); 2643 (19); 2647 (18) 2688, 2753, 4224 (44d); 
10015, 48-228, 48-261 
Smith, C. L., 590 (6); 1132, 1369 (30 
WE Damon, 90 (18); 531 (22) 
) 


y 1844 (39) 


~ 


= 


ped 
ts ud 529 (44d) 
Solis R., F., 567 (59 
Solomon, J., 2421 (49); 2436 (13); 5375 (24) 

Sousa, M., s.n. (49); 936 (18); 1374 (49); 1455 (18); 
1532 (49); 1749 (6); 3358, 3365 (11); 3615 (44d); 
4183 (30); 4434 (9b) 

Sousa, M. & M. Pena S., ) 
ig D. & W. Newey, 1 670 (13); 1701 (6); 1714, 
3 (49); 1913, 1993 (9b); 1996 (32) 
En D. & Stoddart, 2341 (35) 
(48) 


2104 (44d 


A (48); NU. (6); 


nns 11510 1); 13377, 133778 ( 4d); 19222 
(29); VARON (ee) ae 2 yia md 20007, 


© 
D 
a 
+ bo 
c 
= 
ho 
© 
© 
wn 
— 


(49); 21043 «(x IN 21849 (19% 22330 x 


24045 (49); 24: 248 (32) 2429] (18); 24491, 24534 
22); 24589 (12); 24666 (22); 24701 (13); 24857 
24925, 24940 (18); 25084 (48); 
; 25492 (49); 25501 
, 25967 (13); 26079 
(49); 26132 (35), 26133 (59); 26140 (22); 26276 
(35); 26310 (49); 26586 (13); aped a 26859, 
27049 (49); 27168 (1); 27181 (13); 2 
(22); 27604 (49); 27772 (35); Um (22) 28129 
(59); 28378 (22); 29178 (13); 29198 (4); 29268 (1); 
29279 (49); 29287 (13); 29327 (35); 29355, 29410 
(1); 29426, 29433, 29442 (35); 29480 (59); 29662, 
29663 (49); 29725 (35); 29733 (13); 2 
29845, cia [o 30178 (18); 30189 (22 
(59); 30580 (22); 30667 (35) 30819 (13 
(35); 31036 (12) 31037 (22); 31070 (18) 
(59); 31109 (29); 31247 (22); 31370, 31373 (29); 
(22) 31831 (49); 3: 21 19, 32129 


37892 (19); 
39803 


38200 (18); 
39861 (23); 40173 fe 40768 as 35 


, 40998 (4 d 
" 146 (29): 41924, - 
raid 2050, 52 


42 ( 
56197 (32); 56547 (48) pel aap 56733 (35); 
a (18) 56876 (6); 58081 (35); 59208 (30); 
60207 (47); 60657 (13); 62278 (49); 63602 (35); 


63604, 63617 (48); 63622 (35); 64078, 64191 (1): 


Volume 76, Number 3 
1989 


Hamilton 913 
Mesoamerican Psychotria, Part III 


64770 (35); 64775, 64818 (49); 65024 (27); 68129 
(44d); 68196 (18); 68772, 68783 (2); 70305 (12); 
70328 (9b); 70418 (12); 71396 (9a); 72313 (12); 
72345, 72351 (13); 72549 (22); 72986 (48); 76317, 
76783, 77070, 77633 (30); 77925 (13); 77958 (49); 
78006 (13); 78446, 79472 (49); 80900 (27); 85583, 
86991, 87008 (9a); 87667 (1); 88882 (47); 89562 
(48); 89965, 90065, 92157 (9a 

Standley, P. & H. Garnier, 8403 (49) 

Standley, P. & H. Lindelie, 7348, 7454 (13) 

Standley, P. & E. Padilla B., 2631, 3776 (49) 

Standley, P. & R. Torres R., 47459 (18); 47566, 47571 
(24), 47573, 47600, 47856 (18); 51005, 51161, 
51173 (23) 

Standley, P. & J. Valerio, 43187, 43268 (44d); 44076 
(24); 44098 (44d); 44193, 44231 (6); 44585, 44603, 
44620 (50) 44897, 44926 (1); 44986 (6); 45153 
(1): 45179, 45200 (50); 45344 (45); 45411 (19); 
45433 (44d); 45479, 45691 (6); 45770 (42); 45845 
(50); 45852 (18); 45855 (50); 45904, 46135, 46219 
(6); 46375 (42); 46668, 16670 (1); 46985 (50); 48555 

; 48647 (59); 48781 (29); 48826 (59); 49194 

; 49989 (16); 50271, 50396, 51953, 52044, 
í y 


. Williams, 670 (44d) 
de 164 (13); 304 (48) 
NE I an 19) 
, 1303 (37); 1337 (7 
“Hamilton, a 31 (29) 

l 

; 15, 19 (13); 145, 160 (35); 
22 (4); 325 (13); 416 (22); 426 

2); 735, 921 (18); 935 (12); 


3) 
, pue 2955 (13); 3466 (47); 3635 (13); 
; 4088 (13); 4089 (6); 4327 
Ms (6); 4953 (18); 4976 (22) 5 
(44a); A (13) 5625 (37); 6054 (13) 6316 (22); 
6: 366 (44a); 6608, 6866 (22); 7043 
, (416 (35) 7597, 7742, 7761, 7799, 787 1 
= ee 5); 8145, 8208 (32); 8399 (12); 8465, 
22): 8705 (6); 8746 (35); 8867 (29) 8956 
: 9004 (12); 9277 (13); 9545 (49); 9597 (18); 
7); 10453 E 10564, ae (13); 10826 
11243, 11256 (1); 11638 (18 
i 2495, 


TH 


yo: "18423 


2338 (49); 22435 (35); 22461 (37); 
22696 a 22698 (13 

Stevens, W. D. & A. Grijalva, 147 

sis (47) 

.D. & W. Hahn, 18943 (47) 

D. & J. Henrich, 20332 (6); 20335 (48); 


3 (6); 15098, 15136 


Stevens, da 
1 (1) 
5, W. ^ & O. M. Montiel, 
Stevens, W. D. & P. P. Moreno 
Stevens, W. i et al., 14473 (13 ) 
(1); 21144a (47); 
Ste yermark, J., 29457, 29458 (1); 30684 (30); 31237 
(32); 31300 (9b); 31321 (32) 31675 (3 3a); 
h 33417 (44d); 33543 (2); : 


1/516 (6) 

(22) 
i 16530 (22); 
21145b (37 


E 
AS 


16569 


(30); 32472, 


33744 (44d); 34302, 36714 (9a); 371% 78 (44d); 37 388 


37415 (44d); 37628 (2); 37866 (1); 37948 
(44d); 38056 (35); 38096 (9b); 38103 (12) Be 6, 
38561, 38644 (9b); 38733, 3886 

39447 (9b); 39499 (18); 39830 (32) 41 
41678 (14); 41740, 41785 (51); 41882 | 
(51); 41949 (29); 41968 (44d); 42705 cor 
(33a); 43776 (9a); 44111 (49); dun 35) 
(32); 44286 (22); 44433 (35); 44444 a 44480 
(6); 44594a (15); 44833, 44837 (12); 44873 (9b); 
44912 (49); eden iio", 44991 SUE 15077 
(22); 45228 (9b); 4 
14); 45352 (9b); 


(9a); 


T 
— 
£i 
to 
B y 


ae 


45665 (32); 45800 ü2i ' 45802, 
(18); 45864 (9b): o ): 46101 01: 3) - 
- 47694 (13) 


47361 (9a); 47 1638 (13) 47 654 


47770 (1); 47995, 48002 (9b); 48844 (9a); 48850, 
O (12); 49228 (22); 49271 (18); 49405 (9b); 
225 27% 51230 (13); 51423, 51630 (30); 51862 
(49); 52052 : 52059, 52060 (9a 
Steyermark, J. I 17110 (18); 17456 (22) 
tier, F., 39 s 102, (35) 
Stimson, W., 5115 (12); ere (22) 
tork, H., 1150, 1736, 1739 (24); 2169 (23); 2364 
29) 
Sullivan, G., 30 (22); 133 (48); 141 (59); 183 (53); 589, 
600 (13); 693 (22) 
Sullivan, J., 644 (18) 
Sytsma, K., 1212 (44d); 1279 (12), 1384 (59); 1388 
(13) 1504 (43); 1643, 1713 (59); 1937, 1939 (22) 
1942 (5¢ >); 1990 (43); 2920 (29); 2941 (44d) 
Sytsma, K. V. D'Arcy, 3292 (22); 3390 wre 3425, 
3443 age 3643 (44d); 3689 (58); 3725 (4 
RAN K. al., 2769 (29); 4215 (53); TA 4286 
29) 
a D., 42 (13) 
te, R., A. 20 (35); 209-334 (48) 
Taylor, B., 4424 (32) 
Taylor, C., 2620 (9a) 
Taylor, J., 17883 (19); 18059 (1) 
Tellez, O. & E. Cabrera, 1265 (49); 1794, 1807 (35) 
1953 (49); 1954, 2040, 2287 (35) 
l'ellez, O. & J. L. bes uy d: (30) 
Pellez, O. et al., 3327 (35); 4898 (18); 4907 (9b); 4963 
(44a); 4972 (12); 5120, E rf (44d); 5156, 5200 (2); 


0293 (49); 6490 (30) 
Tenorio L., P. . Romero de T., 579 (30) 
I & R. Tery, 1 lAa (22); 1436 (26) 

"ONES UE 8 (1) 5285 (6) 
. M. de, 31 (5 9): 6 (4) 
40456 (6) 


hornos: R. & E ; ahy. 

odzia, C., 1162 (2 

Toledo, V. M., 163 E x 164 (11) 

Tomlin, S., 60 (37); 126 (1); ce 32) 157 

Yon, A. S 28, 1573, 2270 (9b); 2499 Pu 2618, 
3034 (30); 3525 (9b); 3581 (44d 

Ponduz, A., s.n. (2); 1974 (19); 2910 (1); 6670, 7050 
(22), 7053 (12); 7054, 8134, 8667 (22); 8726 (6); 
8730 (22); 8890 (13); 9318, 11468 (22); 12610 (18); 
13018 (2); 13625, 13844 (1); 17087 (6); 11-711 
44d) 

d E., 1012 (48); 1013 (11) 

Torres ¿ R. Cedillo T., 101 (11) 628 (13); 730 
(9b) 1 1397 (18); 1461 (44b) 

Torres C., R. et al., 845 (18); 1646 (13); 1672 (49) 


914 


Annals of the 
Missouri Botanical Garden 


Trochez, L., 41 x 

Trott, S. et al., 19) 

Tucker, J., 610 (ib 814, 948 (1) 

Tuerckheim, H. von, 1364 (9a); 7910, 8404 (48); 
(22); 8529 (15); 8533 (22); 11-796 (49); II-1482 
11-1828 (12); 11-1908 (15); II-2365 (9a) 


P" 


1913 (1); 2233 fa 3); 2953 (1) 

5561 (13); 6741 (1); 7022 (44d): 707 : (37) 

n uon E. & R. Ah Chu, 1699 (29); 171 D 

Tyson, E. & K. Blum, 1659 (22); 1677 (59); : 
3988 (13); 4066 

Tyson, E. & A. Clewell, 587: 


l 
Pes ELA 9), 1185 (12); 


4472 (13); 4481 (12) 4532, 4556 (51); 4575 (1) 


4562 (18); 4633 (22); 4634 (18); 4692 (22) 
Utley, J., 148 (2: 
Utley, J. & K. Utley, 1052 (22); 3134 (13); 3923 (19); 


4805 (18); 5137 (44a); 5316 (50); 5394 (35); 5511 


Utley, K., 6054 
Valerio, M., 


22) 


( 
496 (1); 708 (24); 1267 (22); 1333 (13); 


1) 
R., J., 62 (50); 111, 124 (6); 132 (1); 1366 (23) 
> " & L. Viveros, 161 (30) 
el tal. ., 249, 259 (49); 322 
, 603 (22); 037 
argas, D 30 (6); 095 (48) 
Vaughan, ]. et al., 241 (35); 244 
E 4 76 S l 


(30) 
(15) 641a (12 


— 


N 
— 
t 
w 


E EN 418 (2); 
48) 


; 779 (47); 
1 (47) 
531 (6) 575, 614 (30); 


1586 (11) 


Vázquez Yanes, C., 781, 812 
(6) 

Vega, S. & A. Grijalva, 32 (6) 

Ventur, a 267 (12) 

Ventura A., F., 103 (47); 1121 (9b); 2536, 2635 (30); 
2811 (49); 3338, 3339 (9b); 4190 (47); e (30); 
4715 (49); 4740 (9b); 4763 (42); 5288 (44d); 5927 


(111); 7279 (30); 8094 (44d); 9492 (47); 9972 (44d); 
10733 (42); 10809 (30% 11269 (42); 
12073, 12320 (49); ; 
13543 (9b); 13658 (42) 13932 (30); 13946, 14951 
(9b); 15007 (6): 15208, 16268, 16897 (9b); 16942 
(44d); 17035 (11) 172 (42); 17977, 
18139 (11); 20151 (1 4): roh (6) 

Viereck, H., 680 ) 


20548, 


30 (11); 46 (29); 77 (1: 


t) 
Vincelli P.. , 274 (1); 388 397 a 2) 533, 584 (32) 
Mcd B. & A. Sanchez, 1-2 
son, S., 162 (22); 218, T 9) 318 (12) 
aa H., 164, 253 (30); 690 (6); 707 (30) 
Weaver, R R. Foster, 1440 (22); 1442 (13); 1443 
(12); 1577 (48); 1600 (4) 


Webster, F. & M. Webster, 90 (18) 

Webster, G. et E 126 

d H., : 
Wedel, H. 160 (22 


), 364 (59); 1079 (22); 1123 
(48); 1163 (4) 1191, 


1347 (22); 1407 (48); 1722 


22): 1770 (48); 1809, 1869 (4); 2011 (22); 2137 
(48); 2160 (22); 2508 (4) 

Wende hake, C., 40 (13) 

- P (19 

& A. Villalobos C., 2560 (48) 

; t al., 2258 (44b) 2345 (9b) 2534 (35) 
2574 (18); 2630 (9b); 2751 (11); 2821 (14); 2939 
(9b); 3163 (11); 3285, 3307 (44d); 3322 (18); 3322a 


(49); PEE 3352 (18); 3476 (11); 3490 (9b); 
3555 ne 0 (11) 
West, R., "d 
Weston, A., e a 5) 
a R. «€ E. (22) 
», G., 16, 69 VEN 39 (13) 
n G. & P. White, ip 4d) 
White, P., 123 (49); 159, 187 (44d); 248 (1) 


White fanrd. C., 1119 ü Ee 
(13); 2549 (32) 

Whitmore, 28 ( 

Wilbur, R., 20099 (18); 20128 (35); 20148 (18); 28182 
(22); 30166 (59); 30202 (18) 

Wilbur, R. J. Luteyn, 19316 b ) 

Wilbur, R. & D. Stone, 9665 (18 

Wilbur, R. et al., 12951 (1); 1 n ge 13211, 13544 


2078 (35); 2368 (32); 2428 


Wiley, J., 213, 214 (59); 531 (49 
Williams, Ll., 8677 (2); 8811 (18); 8976 (11); 9011 
(2); 9012 (18); 9307 (48); 9475 (39); 9587 (48); 


Williams, L. O., 8318, 8332, 8418, 8543, 8634, 8798, 
8898 (14); 9160, 9325, 9546 (11); 15953 (13) 
Williams, L. O. & A. Molina R., 10128 (49); 10390, 

10396 (44d); 10841 (13); 11081 (44d); 11436 (22); 


12512 (9b); 14426 (48); 14487 (18); 14513 (9b); 
14515 (42); 14562 (18); 17929 (22); 18075 (44d); 
42652 (37 

Williams, L. O. € R. P. Williams, 18750 (48) 

Williams, L. O. & T. P. Williams, 24555 (6) 

Williams, L. O. & M. Wilson, 40739 (12) 

Williams, es O. et al., 15588 (44d); 23342 (37); a 
(42); 23861 (6); 23951 (18); 24196, 24220 (22 
oe 24669 (6); 24803 (47); 24823 (18) Te 
(37) 26372 (35); 26632 (6); 27594 (37); 28596 (24); 
29176 (37); 40581 (9a); Pus (30) 41958 (9a); 
42086 (18); ei (44d); 43559 (33a) 

Williams, R. Mn 238, 32 i. 1004 (22) 

Wilson, C., zd 351 (9a) 

Wilson, P., n 148 (18) 

V-1 (32 


Winzerling, H., 
Wi . & F. Witherspoon, 8297, 8301 (59); 
8630 (22); 8693 (13) 
. & R. Schery, 
Woodson, R. et al., 

1745 (35); 


Woodworth, 
l 9 


946 (12); 29) 

772 (22) 7 76 (59); 1459 (13); 1588, 
1918 (12) 

R. & P. Vestal, 310 (48); 653 (22); 7 

Wooton, E., s.n. (30) 

Wright, C., s.n. (1); s.n. (49) 

Wunderlin, R. et al., 314, 384 (49); 386 (48) 

Yuncker, T., 4591 (48); 4664 (35); 4975 (12); 
(6) 

Yuncker, T. et al., 5939, 6013, 6118, 6143 (44d); 6229 
(33a); 8019 es 8038 (6); 8040 (1); 8173 (35); 

(48); 8567 (12); 8628 (49); 8673, 


507 


5 (6) 
Zelaya, L., 191 (32) 


Volume 76, Number 3 
1989 


milton 915 


a 
Mesoamerican Psychotria, Part Ill 


APPENDIX III. Index of latin names. 


Di KU E in boldface type refer to desc riptions; numbers 

with ! refer to pae ns; names in italics refer to 

synonyms. Page numbers are all in volume 76 of the 

Annals of the Missouri gra al Garden: pp. 67-111 

86-429 in Part II (number 
3). 


in Part I (number 1), PP- 
2), and pp. 886-914 in ES III (number 


i RUN Schott 68, 75, 77 
Bactris 
Cavendishia Lindl. 75, 77 
Cephaelis Sw. 67, 68 
dressleri Dwyer 896 
insueta Dwyer 897 
panchocoensis Dwyer 896 
d dd. ean Seem. 899 
nthae G. Don 
Heliconia L. 68. 15.7 17 
Mapouria Aubl. we pn 2 
chamissoana Loe 


graciliflora (Benth.) ) P 388 
] l. 80 


7t 7^4 
(Os d 


` 


micrantha (Kunth) Wernham 90 
microdon (DC.) Bremek. 902 
miradorensis Oerst. 100 
obovata Oerst. 91 
papantlensis Oerst. 108 
parvifolia (Benth.) ed 396 
remota (Benth.) Muell. Arg. 398 
Miconia Ruiz & Pav 
Monochaetum (DC.) Naudin 75, 77 
Myrstiphyllum P. Br. 80 
horizontalis (Sw.) Millsp. 105 
marginatum (Sw.) Hitche. 393 
undatum (Jacq.) Hitche. 410 


Valetonia Bren 7 

Vonatelia l. 64 

Votopleura cow: & J. D. Hook.) Bremek. 67 
Ouragoga 

Palicourea "dd 


nigrescens M. Martens & Galeotti 418, 420 
Piper L. 68 
Psychotria L. 67, 68, 71 
subg. TEMOR Steyerm. 67, 68, 902 
subg. Psychotria 67, 68, 70-78, 80, 902 
subg. Tetramerae (Hiern) E. Petit 67, 902 


p group 79, 80, 412 
remota group 79, 80, 398 
dur group 79, 80 ) 


costivenia complex : 
graciliflora complex 71, 79, 386 
horizontalis complex 77, 79, 95 


. & Steyerm. 8o. p don 401! 
TK. Krause 398, 
alfaroana "Standl, 80, 83, 2. eon 
altorum Standl. & Steyerm. 99 


anceps Kunth 398 

asiatica L. 68, 80 

bakeri Dwyer 79, 82, 386, 388! 

meer C. Hamilton 79, 83, 95, 97! 

bimea L. Riley 105 

ca Dwyer 76, 80, 82, 400, 401! 

calophylla Standl. 71!, 78, 80, 81, 891, 892!, 894 

carthagenensis Jacq. 69!, 70!, 71!, 731, 751, 761, 77, 
, 84, 85', 88, 94, 95, 428 

mundos tent auct. 85, 87 


io nid Hamilton 76, 80, 82, 83, 412, 413, 
41 


cerroaz wale nsis Dwyer 894- 

chagrensis Standl. 80, 81, SUN wen 405, 428 

chamissoana (Loes.) Standl. 403, 40 

chimarrhoides DC. 410 

chiriquina Standl. 80, 82, 84, 412, 413, 414, 418 

chitariana Dwyer & C. Hamilton 76, 80, 83, 892, 
894!, 895 

clivorum are & Steyerm. 79, 83, . 87, 88! 

cocosensis . Hamilton 151, 80, 82, 1 9. 413! 415 


3, 95, 98, 103, 105, 110 
var. altorum (Standl. 8 oa ) €. Hamilton 97!, 
99 
costivenia 76!, 97!, 99, 100 
eles (Muell. F ) Standl. 398 
dressleri (Dwyer) C. Hamilton 73!, 80, 82, 892, 894!, 
896 


durilancifolia Dwyer 423, 42 

dwyeri C. Ben n 82, B4. 412, 413, 414! 416 

elongata Benth. 4 

eni oaa Sc la E 731, 80, 83, 84, 399, 400, 
403' 


fendleri Standl. 77, 79, 83, 95, 96!, 101 

flava Oerst. ex Send, 731, 761, 79, 82, 95, 961, 102 

fosteri C. Hamilton 80, 81, 399, 400, 4031, 405 

a dee 80, 83, 399, 109; ps 405 

gatunensis Dwyer & soli 894-89 

glaucescens Kunth 1 

glomerata Kunth 94 

grac cilif a Benth. 76!, 79, 84, 386, 387!, 388, 392, 
"306 


granadénsis Benth. 888, 

grandicarpa Dwyer & Hayden om ds 5 

grandis Sv. 73, 76!, 79, 83, € . 103 

grandistipula Merr. 423 

grandistipula 2 423, 425 

hammelii Dwyer 76, 80, 83, 891, 894!, 896 

hirta Roem. & Sc d 410 

horzontalis Sw. 71!, 751, 76!, 79, 83, 95, 96!, 102, 
105 


subsp. basicordata Dwyer 105, 108 

var. glaucescens (Kunth) pkey adm; 105 

var. msilophylla Steyerm. 
hornitensis Dwyer C. Hamilton 76, 80, 84, 412, 


"T 
í 


~ 
© 
e 


insignis Standl. 77, 80 . 892, 893!, 897 
insueta (Dwyer) C. B 76, 80, 81, 891, 8931, 
897 


jimenezii Standl. 79, 81, 386, 391, 
Ber Nelson, Molina & Standl. 
100, 407 


, 


5, '. 80, 82,84, 


var. 'jinotegensis 4031, 408 

var. morazanensis C. Hamilton 4031, 408 
a C. Hamilton 19, M 84, 88, 80! 
. Hamilton 77, <83; 386. 391, 392! 


laselvensis 


916 


Annals of the 
Missouri Botanical Garden 


393 
428, 886, 887! 


liesneri Dwyer 76, 79, 84, 386, 388, 

limonensis K. Krause 68, 735 80, 81, 
var. da Standl. 87 

Loes. 886 


105 


var. rite 
longicollis Ben 


undai Standl. a 83, 84, 412, 413, 414!, 417 
magna Standl. 422 
72!, 73!, 74!, 78, 79, 83, 84, 386, 


marginata Sw. 
387!, 39: 
mexiae Standl. 76!, 80, 82, 412, 413, ET wi 
micrantha Kunth 71, 79, 82, 84, 88!, 90, 410 
microdesmia Oerst. 
microdon (DC.) Urb. 902 
drena nsts (Oerst.) Hemsl. 
i i uci hs 82, 
bur nl a 
monteverdensis Dos r Es c. 
893!, 898 


js n: 
, 408, 409! 


Hamilton 76, 80, 81, 892, 


morae Polak. 91, 93 

neillii C. d & Dwyer 79, 83, 84, 88!, 9I 

nervosa Sv , 82 90, 399-401, 409!, 410 
subsp. s scens (Kunth) Steyerm. 410, 412 
ME Mus var. hirta (Roem. & Schultes) Stey- 

410 

nicaraguensis Benth. 393 

nigrescens De N ild. 418, 

nutans Sw. 902 


e 


Le 


420 


388, 390 


oaxacana Aeon 


oerstediana (Kuntze) Standl. 

olgae nd bad & Hayden 76!, 
414!, 420 

oli solvit ha DC. 

orosiana Standl. d 161, 79, 83, 84, 386, 
396 


P 80, 82, 412 


392!, 393, 


pacorensis C. Hamilton 77, 80, 81, 83, 894!, 899 
panamensis | ml 70!, 73!, 80, 82-84, 412, 416, 
420, 42 
var. minds (K. Krause) C. Hamilton 4.131, 
421 


413!, 421, 422 
421, 422 


var. ixtlar >. Hamilton 80, 82, 
var. magna ome ) C. Hamilton 413), 
- paname nsis 4131, 421, 423 
"eir: nsis (Oerst.) Hemsl. 75!, 79, 83, 95, 961, 
parvifolia Benth. 19, 82, 380, 388!, 396 
philacra Dwyer 77, 79, 81, , 388, 397 
pinularis Sessé & Moc. 902 
ple wi Donnell-Smith 79, 83, 95, 96!, 109 
portoricensis DC. 410 
psychotrifolia (Seem.) Standl. 
899 


108 


70!, 78, 80, 81, 891- 


punctata Vatke 902 
quiinifolia Dwyer 410, - 


eo Polak. !, £a 81, 82, 84, 89!, 91, 
remota Benth. 72!, EE , 80 . 398, 399! 


0, 83 
rosulatifolia Dwyer 77, ^ 8l. 891 892. 893! 900 
rufescens Kunth 90, 410 


rufescens Roem. & Schultes 90, 410 


sagraeana Urb. 

sarapiquensis i 80, 82, 84, e 2, m 425, 426! 
schippii Standl. & Steyerm. 418 

sessilifolia M. ce & Galeotti nae 89] 
sinuensis Standl. 

Ham d de E 81, 894!, 901 
Hamilton 71, , 76, 80, 84, 412, 413, 


sixaolensis C. 

stock wellii C. 
417! 426 

sulzneri Small 889, 89 

sylvivaga Standl. 76, 

tenuifolia Sw. 68, 70!, , 428, 886, 888, 890! 

tric p M. Martens & Gale otti 80, 82, 84, 412, 

13, 427 


22, 4261, 


4, 95, 96!, 110 


Noch Griseb. 94 

undata Jacq. 410 

vallensis Dwyer 388, 

viridis Ruiz & P 

wendlandiana Standl. 

yunckeri Standl. 423, D 5 
Psychotrophum P. Br. 80 
Rondeletia L. 


mic rodon Dc. 


, 19, 82, 84, 85!, 94. 


902 


glomerata (Kunth) Kuntze 94 
graciliflora (Benth.) Kuntze 388 
granadensis (Benth.) Kuntze 889 
brandi is (Sw.) Kuntze 103 
horizontalis S 2 105 
insignis (Muell. Arg.) Kuntze 897 
ie uanha T S Baill. 81 
vyt Kuntze 396 

longicollis (Benth. ) Kuntze 


~ 


~ 


ensis (Oerst.) Kuntze 100 
morae (Polak.) Kuntze 91 
nicaraguensis ( dioe .) Kuntze 
oerstedtiana Kuntz 


e oido (DC.) Eos 4 


393 


psychotriifolia (Seem.) Kuntze 899 
quinqueradiata (Polak.) Kuntze 91 
remota (Benth.) Kuntze A 
rufescens (Kunth) Kuntze : 
sessilifolia (M. Martens = Gak otti) Kuntze 888 
tenuifolia (Sw.) Kuntze 
trichotoma (M. at & Galeni Kuntze 427 
trispicata (Griseb.) pora 
undata (Jacq.) Baill. 
"ipi (Jacq.) Sau ze í 
iridis (Ruiz & Pavón) MR 94 


NOTES 


NOTES ON ARACEAE 


Anthurium iramirezae Dunting, sp. nov. TYPE: 
Venezuela. Territorio Federal Amazonas: Dep- 
to. Rio Negro, Serrania de Tapirapeco: aflora- 
miento de roca granítica, 1°20'N, 64%55'W, 
1.350 m, 25 Mar. 1988, 1. Ramírez & L. 
Laskowsky 310 (holotype, VEN). Figure 1A, B. 


erba perennis ca. 75 cm alta. Caulis horizontalis 
vis crassusque. Petioli subteretes sed adaxialiter can- 
19- a cm longi. Foliorum laminae cartilagineae, 
ovatae, 30 1 longae egy gs eun vel eis 
usque ad lo longiores), 19-24 cm y pani non 
sinu plus minusve arcuato vr 0-1.7 
viter acuminatae, abaxialite 
ateralibus primariis 
x costa oreuntibus. 
Spathae anguste oblongo- 
ad 11 em anes in sicco 1.8 cm latae, 
basi decurrentes per 0.8 cm, recurvatae vel reflexae, 
virides, persistentes. Spadices eS 13.3-15 
em longi, sessiles sed axe antice nud 


re 
aliculati, 


— 


profunde cordatae s 

profundo, apice bre 

obscureque fusco-punctatae, nervis 
> F e 


ao 


Perennial herb ca. 75 cm tall. Stem horizontal, 
thick, with very short internodes, some dry cata- 
phyll remains, and few leaves. Petioles subterete 
but adaxially canaliculate, the canal narrower than 
19-40 
cm long; geniculum 1.7 2.3 em long, 1.0-1.2 em 
thick. Leaf blades nearly erect on petiole, cartilagi- 
30-37 em long (sub- 


petiole width and with rounded margins, 


nous (drying stiff), ovate, 
equaling or to twice as long as petioles), 19-24 
cm wide (1.5-1.7 times longer than wide), basally 
shallowly cantar. with sinus 1.0-1.7 cm deep, 
apically obtuse and briefly acuminate ending in a 
cusp to 1.3 cm long. adaxially glossy, abaxially 
sparsely and obscurely brown-punctate; midrib and 


all veins prominently elevated; primary lateral veins 


- 


45-55? and running to margins; lesser veins form- 
ing a rather fine reticulum prominent on both faces; 
lowermost primaries not naked in sinuses. Pedun- 
cles solitary, 45-71 em long. Spathes narrowly 
oblong-ovate, to 1 1 em long (including 8-mm-long 
slender tip), drving 1.8 em wide, basally decurrent 
for 0.8 cm, recurved or reflexed, green, persistent. 
Spadices cylindric, 13.3-15.3 em long, 0.75 cm 
thick, dark purple becoming green in age, exceed- 
ing 1.5 em thick in fruit, sessile but axis naked on 
front for 0.8 cm, with 6-8 flowers visible in the 


principal spiral. Fruits unknown. 


(-8) arising from each side of midrib at angles of 


Paratype. VENEZUELA. TERRITORIO FEDER 
\MAZONAS: same locality as type collection, Ramtrez E 
EN). 


Laskowsky 318 (V 


This species is closely related to Anthurium 
guanchezii Bunting but differs by having a car- 
tilaginous leaf blade not basally stipitate, the low- 
ermost primary lateral veins not naked in the basal 
sinus, a proportionally narrower spathe, and a non- 
stipitate spadix. In 4. guanchezii Bunting, the 
or semicoriaceous, and the 


blade is pergamen 
spathe is 3.6 times longer than broad. Additionally, 
A. iramirezae grows at an altitude of 1,350 m on 
granitic outerops with thin soil and woody vege- 
tation to 10 m tall, whereas 4. guanchezu occurs 
in moist forests at 120 m altitude some 500 km 
to the north. 

rom 4. wurdackii Bunting, the new species 
differs by having leaf blades that tend to be ovate 
and proportionally broader with an arcuate basal 
sinus, while the peduncle is shorter and the base 
of the spathe is more shortly decurrent. In A. 
wurdackii, the leaf blade is often elliptic-ovate, 
about 1.9 times longer than wide with an angular 
basal sinus, and the spathe is decurrent for 3.5- 
7.5 cm. Further. the leaf blade of 4. 
apparently lacks abaxial, dark punctations such as 


wurdackii 
occur in A. iramirezae. 

The species is named in honor of a collector of 
the type material. Ivón Ramirez, a young Vene- 


zue 
into cultivation. I observed the plant in Maracay, 


an botanist who also successfully introduced it 


Venezuela, among a collection of unusual Vene- 
zuelan aroids and other plants growing at the family 
home of her husband, Germán Carnevali. 

Philodendron triplum Bunting, s TYPE: 
carretera El Dorado— 


sp. nov. 
Venezuela. Edo. Bolivar: 
Santa Elena de Uairén, en aprox. km 120 al 
sur de El Dorado, en bosque al lado de la via, 
1,200 m, 22 Feb. 1968, G. S. Bunting 3102 
(holotype, NY: isotype, MY). Figure 1C. 
erba perennis subrosulata. Caulis brevis validusque 
cataj le reliquiis fibrosis ferrugineis tectus. Pe toli 
adaxiali late e leviter 
sub- 


a s rotundati superficie 

9 cm longi. Foliorum laminae 
: cm longae, 16.5 em 
latae, basi rotundatae sinu acuto | em profundo, apice 


acutae vel obtusae et breviter acuminatae, nervis later- 


ANN. MissouRi Bor. GARD. 76: 917-919. 1989. 


918 Annals of the 
Missouri Botanical Garden 


Volume 76, Number 3 
1989 


Notes 


alibus primariis utrinque 9-11 sub angulo 55-65? ex costa 
abeuntibus. Inflorescentia ignota. 

Subrosulate, perennial herb. Stem short, stout, 
with accumulated, rust-colored, fibrous cataphyll 
remains. Petioles abaxially rounded, with adaxial 
face broadly but shallowly concave, to 19 cm long. 
Leaf blades subcoriaceous, oblong, to 58 cm long 
(3 times longer than petioles), 16.5 cm wide (ca. 
3.5 times longer than wide), at base rounded and 
notched (1 em deep), at apex acute or obtuse and 
briefly acuminate; primary lateral veins 9-11 aris- 
ing from each side of midrib at angles of 55-65*, 
with some secondary lateral veins manifest. Inflo- 


rescence unknown. 


The type collection was taken from a plant grow- 
ing as an epiphyte 2 m above the ground; it is not 
certain that it represents the fully mature adult 
phase of this species. A similar though somewhat 
less adult plant, collected in the same area as the 
type, has recently been seen in cultivation at the 
family home of Lic. Germàn Carnevali in Maracay, 
Venezuela. A juvenile specimen, Bunting 2916 
(MY), collected in the same sector at km 126 south 
of El Dorado, Bolivar, Venezuela, apparently per- 


tains to this taxon. It has leaves of similar form, 


texture, and color, but, as is typical of juvenile 
shoots, has a more slender stem with elongate in- 
ternodes and has wide-winged petioles basally en- 
circling the stem and sheathed up to the base of 
the blade. 

This species differs from Philodendron meli- 
nonii Brongn. ex Regel by the proportionally short- 
er petioles and narrower leaf blades. In P. meli- 
nonii, the blade length is 1.4 times longer than 
the petiole and 1.5- 2.4 times longer than wide. 

The specific epithet reflects the fact that the 
leaf blade is three times longer than the petiole 
and more than three times longer than wide. 


I am grateful to Lic. German Carnevali and his 
wife, Sra. Ivón Ramirez de Carnevali, for making 
available dried and living specimens of the aroids 
collected by them during recent botanical explo- 
rations. My appreciation is expressed to the Di- 
rector and personnel of the Herbario Nacional 

EN), Jardin Botánico, Caracas, for the loan of 
herbarium material. 


— George S. Bunting, 101 Linden . 4ve., Pocomoke 
City, Maryland 21651 , U.S.A. 


— 


Ficure 1. A, B. Anthurium iramirezae. — A. Holotype.—B. Paratype.— C. Philodendron triplum, habit. Type 


specimen pre vun from this pian nt. 


HOSTA JONESII 
(LILIACEAE/FUNKIACEAE), 
A NEW SPECIES 

FROM KOREA 


Hosta Tratt. is an economically important genus 
of approximately 22-25 species restricted to east- 
ern Asia. Hosta has been placed in the Liliaceae 
1981; 1964) or Funki- 
aceae (Dahlgren et al., 1985). Bailey (1930), Stearn 
(1931), and Hylander (1954) all pointed out that 
the major difficulty with the classification of the 


(Cronquist, Hutchinson, 


Hosta lies in the uncertainty regarding criteria for 
delimiting the species. The taxonomic difficulties 
have been attributed to relatively few diagnostic 
characters (Fujita, 1976), the extreme difference 
in appearance between living plants and dried her- 
barium specimens (Hylander, 1954), and possible 
widespread hybridization (Maekawa, 1940). Thus 
the treatment of species within Hosta has varied 


Maekawa 


1969) described several new species but combined 


depending on authors. For example, 
others recognized in his 1940 treatment. Fujita 
(1976) noted that Maekawa (1940, 1969) did not 
consider the variability of natural populations when 
developing his classification. Fujita recognized only 
15 species as native to Japan, while Maekawa 
(1969) had recognized 25 species there. Although 
Fujita carefully described morphological characters 
and ecological and geographical distribution, his 
studies were confined to Japan. 

In 1985, 1987, and 1988, 


trips to collect hostas from natural populations in 


| conducted field 


Korea and on Tsushima Island of Japan in order 
to understand better the variation patterns. While 
visiting several islands off the coast of southern 
Korea, | encountered a morphologically distinct 
group of Hosta DoD ie not referable to any 
described species of Hosta. The herbaria at Seoul 
National University (SNU), Tokyo University (TD, 
and Kyoto University (K YO) were visited and there 
were several undetermined specimens of this group 
collected by Young Chung (SNU) and one specimen 
collected by T. Nakai (TI). Loans were examined 
from BH, BM, E. K, L, 


specimens of the group were encountered. 


and PE, but no herbarium 
This 
series of populations is a new species as described 


below 


Hosta jonesii M. Chung, sp. nov. [subgen. Bryo- 


cles (Salisb.) F. Maekawa sect. Tarthanthae 


(F. Maekawa) F. Maekawa]. TYPE: KOREA: 
Kyeongsang Nam Do, Namhae Gun, Sangju 
Myeon, Namhae Island, Mt. Kumsan, N 13 
mi. ascent from Yangha Ri. Among rocky and 
humus soil on south-facing slope, shade, ca. 
250 m, common, 28 Aug. 1988, M. G. Chung 
& M. S. Chung 1613 (holotype, GA). Figure 1. 


Rhizoma breve repens. Folia erecto- na o 
elliptic o-lanceolatae vel anguste ovatae, 6 ongae 
3-5 em latae, obtusae vel breviter ac uminatae in apice, 
leviter angustatae in base, distincte e petiolatae, supra atro- 
virentes vel viridulae, utrinqu je 5-7 nervatae, nervis infra 
paullo laevibus; petioli 4.5-13 1 : 
Se api oblique ascendentes, 35-60 cm longi 2-4 mm lati, 


ca. cm lor f acutis, na- 
vicularibus, viridibus, clani Infra a entiam brac- 
teae ascendentes, lanceol m long 


virentes, pon "ie ania punctati. 


tia. Perianthia : mm longa, ca. 25 mm in diametro; 
pars inferior n ris d saepe alb 18; pars 


superior tuborum dilatatorum paullo campanulata, dilute 


joiga, fere aequilonga corollae; antherae c 
subflavescenti-purpureae. Pistillum 45. 52 mm longum. 


Herbaceous perennials from short, creeping rhi- 
zomes. Leaves erect-patent, spirally arranged at 
the base of stem; petioles 4.5-13 cm long, 4-7 
mm wide, dotted with purple, slightly winged; blades 
elliptic-lanceolate or narrowly ovate, 6-13 cm long, 
3-5 em wide, dark dull green, slightly rigid, obtuse 
or acuminate at the apex, gradually narrowed at 
the base into the petioles, the nerves of upper leaf 
surface inconspicuous when fresh, nearly glabrous 
on the 5-7 pairs of somewhat smooth, not-elevated 
nerves of lower leaf surface, margins thinly white- 
35-60 


cm long, 2-4 mm wide, dotted with purple dots 


lined. Scapes obliquely ascending, terete, 
basally, usually with 2 lanceolate bracts below in- 
florescence, these 15-40 mm long, 4-20 mm wide; 
bracts within inflorescence acute, navicular, green, 
8-13 mm long, 3-4 mm wide, not open at flow- 
ering, relatively persistent after flowering; pedicels 
obliquely ascending, 4-8 mm long, whitish green, 
minutely purple-dotted, usually shorter than the 
subtending bracts. Perianths 40-50 mm long, ca. 


ANN. Missouni Bor. Ganp. 76: 920-922. 1989. 


Volume 76, Number 3 
1989 


Notes 


FIGURE 1. 


Hosta jonesii (A-C, 


surface. — D. Fruits on the obliquely ascending scape. 


25 mm diam., about (1-)3-20 flowers on each 
raceme, whitish purple, greenish in bud; the basal 
narrower portion of perianth tubes whitish; the 
upper dilated portion of perianth tube somewhat 
bell-shaped; the inner nerves not intensely colored; 
lobes oblong, acute, 13-15 mm long, 7-8 mm 
wide; 6 translucent lines originating from the di- 
viding parts of the lobes extend to the middle of 
lower narrower perianth tube; stamens 39-48 mm 


holotype).— A. Habit.— B. Flower viewed from the side. — C. 


adaxial 


Leaf, 


long, nearly equal to or longer than perianth; an- 
Pistil 45- 
52 mm long. Capsule eylindric, 22-33 mm long, 
4-6 mm wide. Flowering in mid August to early 


ther ca. 3 mm long, yellowish edipi 


September. 
Korean name. | Tadohae-bibich'u. 


Pa REA: Kyeongsang Nam Do, Namhae 
Gun, Ne. : Island: Mt. Kumsan, Y. Chung s.n. (NA); 


922 


Annals of the 
Missouri Botanical Garden 


Cholla Nam Do, Yoch'on Gun, Dolsan Island, M. Chung 
957 (CA S. Lee 101 (GA), 103 (KYO), 106 (MO), 107 
(SNU), 712 (TI). Each collection from shade of pine-oak 


forest on rocky and rich humus soil near ocean. 

Hosta jonesii is distinguished from other species 
of Hosta by the navicular, green, persistent bracts; 
thick, adaxially dark dull green leaves; and white- 
lined leaf margins. This species appears to be re- 


lated to H. minor (Baker) Nakai by the shape of 


bracts and perianths and by the creeping rhizomes. 
It differs by the terete scapes, elliptic-lanceolate 
eaf blades, and flowering period from middle Au- 
gust to early September. Hosta minor flowers in 
mid July to early August and is endemic to south- 
eastern Korea, including Wando and Kojae islands. 
The new taxon also is very close to /7. tsushimensis 
1988) but differs 
by having short creeping rhizomes; longer perianth 
lobes; upper dilated portions of perianth tubes bell- 


N. Fujita (Fujita, pers. comm., 


shaped; whitish purple inner perianth nerves; scapes 
dotted basally with purple; thinly white-lined leaf 
margins; somewhat smooth, not-elevated nerves of 
lower leaf surfaces; and thick, dark dull green 
leaves. Hosta vingeri S. B. Jones is distinct from 
H. jonesii by having ovate, adaxially lustrous leaves; 
delicate raceme of flowers spread around the cen- 
tral axis of the inflorescence; decurrent, flat bracts; 
relatively longer pedicels; and distinct, exserted 
3+3 stamens. 


The new species is named in honor of Dr. Samuel 
B. Jones, who has provided encouragement with 
my studies of Hosta. I thank Dr. Richard A. LaFleur 
for correcting the Latin description and Nancy C. 
Coile for comments on this manuscript. I also thank 


Dr. N. Fujita, Dr. H. Obha, and Mr. Haynes Currie 
for discussions on the classification of Hosta. Spe- 
cial thanks go to my parents, Mun Su and Jeom 
Ryo Chung, my wife, Min Suk Chung, Mr. Jong 
Sae Son, Seok Myeon Lee, and Chin Seop Hwang 
for their company on field trips. Gratitude is due 
the directors and curators of BH, BM, E, GA, K, 


KYO, L, PE, SNU, TI, and US. 


LITERATURE CITED 
BaiLEy, L. H. 1930. 
rb. 2: 117-14 
CRONQUIST, A. An Integrated System of Classi- 
neston of Flowering Plants. Columbia Univ. Press, 
rk. 


Hosta: the plantain lilies. Genetes 
9 


New 
DAHLGREN, RA H. T. CLIFFORD & P. F. Yeo. 1985. 
he Poule: of the Monocotyledons. Springer- Ver- 
lag, Berlin 
FujrA, N. 1976. The p [OM in Japan. Acta 
Phytotax. Geobot. 27: 
Hurchinson, T. 1964. The Familie of Flowering Plants, 
se Il. Monocotyledons, Clarendon Press, Ox- 
for 


HYLANDER, N. 1954. The genus d in Swedish gar- 
dens. Acta Horti Berg. 16 -420. 
MakKAWA, F. 1940. The genus > lost E Fac. Sci. 
Univ. Tokyo, Sec. 3, Bot. 5: 317 
. 1969 Tranda Hos Tra tt. ds En- 
cycl. Hort., Volume 3 10. Seibundoshin- 
kosha, Du "i Japane 


STEARN, W. The nud or funkias, a revision 
of a plan y Gard. Chron. 2: 27, 47-49, 
88-8 10. 


—Myong Gi Chung, Department of Botany, The 


University of Georgia, Athens, Georgia 30602, 
S.A 


A NEW SPECIES OF 
ALLOMARKGR AFIA 
(APOCYNACEAE) AND 
NOTES ON THE GENUS 


Allomarkgrafia laxiflora A. Gentry, sp. nov. 
TYPE: Colombia. 
road, W of Junin, 900 m, tropical pluvial 
forest, 25 Nov. 1981, 4. Gentry, C. Barbosa, 
E. Barrera, C. Bonifaz, H. Cuadros, L. Qui- 
nones, E. Renteria & 1). 34939 
(holotype, COL; isotypes, COL, 


Narino: Tumaco Tuquerres 


Sanchez 


MO). 


Frutex scandens, glaber. Folia anguste oblonga, acu- 
minata, coriacea, prope basin supra glandulifera. Inflo- 
rescentia laxa. Flores calyce 5-lobato, corolla infundibu- 
liformi, parte superiore anguste campanulata, parte basali 
tubulosa, intra pilosa infra insertionem antherarum sag- 

Fructus linearis, 31-35 2mm 


— 


ittatarum. » em longus, ca. 2 


latus. 

Vine, stems terete, glabrous, drying dark with 
pale lenticels. Leaves very narrowly oblong, acu- 
-2.8 cm, 


coriaceous, completely glabrous, the secondary and 


minate, cuneate at base, 7-13 


tertiary venation not evident, glandular adaxially 


~ 
— 


at base of midvein, the petiole 0.5 cm long. 


Inflorescence axillary, open, dichotomously 


branched, the dichotomies subtended by minute 
triangular bracts. Calyx lobes round-tipped, 1-2 
mm long: corolla pale yellow, narrowly campanu- 


ate above a narrowly tubular base, constricted at 


intersection of upper and lower tubes at level of 
stamen insertion, glabrous outside, inside densely 
pilose in upper half of lower part of tube, the base 
plus upper tube ca. 2.5 em long, the lobes round, 
ca. 5 mm long. glabrous outside; filaments pilose, 
| mm long, the anthers sagittate, connivent, 5 mm 
long, the obtuse basal auricles ca. 1 mm long, the 
thin apical connective ca. | mm long; gynoecium 
2 mm long, glabrous, apocarpous. Follicles paired, 
long and narrow, glabrous, 31 


and ca. 2 mm wide before dehiscing, flat anc 


35 cm long, terete 
to 5 
mm wide when dehisced, the seeds narrow, acute, 
5-6 mm long. with a dense apical tuft of tan 
trichomes to 3 cm long. 


Although this distinctive new species resembles 
species of Mandevilla, it belongs to Allomarkgra- 
fia, which differs from Mandevilla primarily in the 
paniculately branched inflorescence. However, this 
species is not at all close to 4. plumeriiflora Woods., 
with which it was first identified. Among the obvious 


ANN. MISSOURI Bor. GARD. 


features differentiating A. laxiflora from A. plu- 
mertiflora are the very openly branched inflores- 
cences with small bracts, very differently shaped, 
narrower flowers, and narrower leaves. 
dllomarkgrafia laxiflora is probably more 
closely related to 4. foreroi ^. Gentry and 4. ovalis 


(Mgf.) Woodson. 


Colombia, and both of which have similar laxly 


both of which occur in. Chocó, 
branching inflorescences. The new species differs 
vegetatively from both of these species in having 
very obscure leaf venation like that of 4. plume- 
riiflora. The distinctive corolla shape of 4. laxi- 


flora likewise helps separate it from its two probable 


closest relatives. In 4. ovalis the corolla is broadly 


(vs. narrowly) infundibuliform-campanulate above 


FIGURE 1. 
34939) 


4llomarkgrafia laxiflora (Gentry et al. 


76: 923-924. 1989. 


924 


Annals of the 
Missouri Botanical Garden 


the basal tube, and in 4. foreroi it is more narrowly 
campanulate above the basal tube, which is longer 
and narrower than that of the new species. Allo- 
markgrafia foreroi differs further from A. laxi- 
flora in having slender, more tenuous inflorescence 
branches; longer, thinner calyx lobes; and differ- 
ently colored flowers (white with upper tube green- 
ish and lower tube reddish). 

There are only two other Allomarkgrafia species 
in addition to those discussed above: Peruvian 4. 
tubiflora Woodson ex Dwyer has leaves like 4. 
ovalis and a longer campanulate upper corolla tube 
much like that of many Bignoniaceae, while Costa 
Rican 4. brenesiana Woods. has white flowers with 
a gradually and narrowly infundibuliform upper 
part of the corolla tube. Another Costa Rican species 


described by Woodson is conspecific with 4. plu- 
meriiflora. 

Thus from a bitypic genus (Woodson, 1933) 
Allomarkgrafia has now grown to six spec les. As 
currently. known, 
with four out of the six species occurring in trans- 
Allomarkgrafia laxiflora and 
A. ovalis is wide- 


it is strongly Chocó-centered, 


Andean Colombia: 
A. foreroi are endemic to Choco; 
ranging from Amazonian Peru to Panama; and 4. 
plumeriifiora ranges from trans-Andean Colombia 
to Nicaragua. The two localized outliers are 4. 
brenesiana at middle elevations in Costa Rica and 
A. tubiflora in upper Amazonian Peru - 
950 m 

The species of Allomarkgrafia may be sepa- 
rated by the following key: 


la. Inflorescence closely branching with strongly ascending, crowded, conspicuously bracteate branches; corolla 
tube base with conspicuous medial or submedial swelling; flowers shorter than 3 cm, the upper tube broadly 


infundibuliform 


lumeriiflora 


Inflorescence laxly 
. tubiflora); kii tube 
campanulate or narrowly infundibuliforr 
9, 
2a 


— 
— 
A 


branching with small, inconspicuous bracts (reduced tc 
base lac xing prominent swelling; flowers E than 3 cm, the upper tube 


2-3 few-flowered branches in 


Corolla ah very gradually cade infundibuliform above the narrow basal part; Costa Rica above 


m 


. brenesiana 


2b. Corolla tube abruptly campanulate above the narrow basal part; South America to Panama, below 


; J IM. 


Leaves broadly elliptic to oblong, the mi parallel tertiary venation conspicuous; base of corolla 


tube muc h shorter than campanulate 


obconical up 


pper par 
Ipper part of corolla tube broadly infundibuliform- -campanulate, the mouth 2-3 cm across 


4b. Upper part of corolla tube campanulate, the mouth less than 1.5 cm across 
very narrowly oblong or elliptic, the tertiary venation inconspicuous or not 


Leaves narrowly to v 


Qo 
a 


Mic os base of corolla tube longer than narrowly campanulate upper part. 


or very obscure below; young 


dein acuminate, coriaceous, the secondar 
branchlets drying dark 


ary and tertiary venation not visible 
with light lenticels; inflorescences 


ca. twice hn hed, the branches drying black or dark brown; basal part of corolla tube 


pilose inside in upper half 


laxiflora 


LITERATURE CITED 
Woobson, R. E. 1933. 
IV 


Studies in the Apocynaceae. 


. The Americsn psa of o ES Ann. Mis- 
605- 


souri Bot. Gard. 


b. Leaves tapering gradually to long acumen, chartaceous, the sec ondary and tertiary veins 


imes dichotomous, the branches drying light brownish; basal part of 
corolla tube pilose inside only near ape 


A. foreroi 


Missouri Botanical Garden, 
Missouri 6316060, U.S.A. 


—Alwyn H. Gentry, 
P.O. Box 299, St. Louis, 


CHROMOSOME NUMBER 
IN WALLERIA 
(TECOPHILAEACEAE) 


The small African genus Jf alleria, comprising 
three species (Carter, 1962), is generally consid- 
ered to be allied to the Lilialean family Tecophi- 
laeaceae. The five core genera of Tecophilaeaceae, 
Tecophilaea, Zephyra, and Conanthera (all Chil- 
ean), Odontostomum (Californian), and Cyanella 
(southern African), are small seasonal perennials 
united by their tunicate cormous rootstock, basally 
clustered leaves, racemose inflorescences, and flow- 
ers with poricidal anthers, partly inferior ovary, 
and subulate style with a punctiform stigma. In 
addition, the pollen grains have an operculate sul- 
cus and tectate-columellate ektexine (Simpson, 
1985), and the smooth seeds have a multilayered 
testa with a black, phytomelan-encrusted outer lay- 
er. As treated by Hutchinson (1934), the first mod- 
ern phylogenist to recognize the family, Tecophi- 
laeaceae also included the African. Cyanastrum, 
now generally treated as the sole genus of Cy- 
anastraceae (Dahlgren et al., 1985; Simpson, 
1985). In 1959 Hutchinson transferred Walleria 
to Tecophilaeaceae from Liliaceae- Dianelleae 
(Hutchinson, 1934). Dahlgren et al. (1985: 165) 
accepted Walleria in Tecophilaeaceae with res- 
ervations; Dahlgren & van Wyk (1988) concluded 
that it did belong in the family, although they placed 
it in its own subfamily, Wallerioideae. 

Walleria is discordant in Tecophilaeaceae for 
several reasons. The rootstock is a fleshy naked 
tuber, the leaves are cauline (not basal), the inflo- 
rescence is foliose, and the ovary is completely 
superior. Also, the seeds are warty and dark brown, 
not smooth and black, and it is uncertain whether 
they contain phytomelan (Dahlgren & van Wyk, 
1988), although this is said to be absent by Huber 
(1969). Simpson (1985) demonstrated that the pol- 
len grains of Halleria resemble those of other 
Tecophilaeaceae in having a medial apertural oper- 
culate sulcus and an endexinous basal layer of the 
operculum and aperture border. Although the oper- 
culum of Walleria differs from other Tecophilae- 
aceae in being very scanty and in having a granular 
(vs. tectate-columellate) ektexine, its basic. struc- 
tural similarity with that of the core genera of the 
family supports inclusion of If alleria there (Simp- 


1985). Embryologically, W'alleria differs in 


son, 


ANN. 


having mostly campylotropous ovules (anatropous 
in other Tecophilaeaceae) and a large embryo 
(Dahlgren & van Wyk, 1988). With this back- 
ground in mind, we undertook this investigation of 
the chromosome number in the two tropical African 
members of the genus, previously unknown cyto- 


logically. 


MATERIALS AND METHODS 


Root tips were harvested at mid-morning from 
greenhouse-grown wild-collected material, tubers 
of W. mackenzii and seeds of IF. nutans. The tips 
were placed in saturated aqueous mono-bromo- 
naphthalene, and left at room temperature for four 
hours before fixation in 3: 1 absolute ethanol- gla- 
cial acetic acid for two minutes. After hydrolysis 
in 10% HCl for six minutes at 600°C, 
portions of the tips were squashed in FLP orcein 
(Jackson, 1972). 


Voucher data are as follows: 


the apical 


Walleria nutans Kirk — Namibia. Gobabis: Stein- 


ausen road, Goldblatt & Manning 8802 
W. mackenzii Kirk — Malawi, Zomba district, near 
Thondwe village, Goldbatt 7519 ( 


OBSERVATIONS AND DISCUSSION 


The diploid chromosome number in both species 
of Walleria examined is 2n = 24. The chromo- 


somes are of moderate size, 2-4 um in the method 
used. The chromosomes are mostly acrocentric but 


four of the smaller pairs are sub- ete as 


Basic chromosome number in Te 
is probably = 12 rnduff, 197 9: Cink 
0). C nets has predominantly n = 12 or 24 


ca a population of C. hyacinthoides has n 


14, and one Pp of C. lutea has n = 
(Ornduff. 1979). the New World genera ex- 
amined ae Tecophilaea has n = 12 (La 
Cour, 1955) and Odontostomum n = 10 (Cave, 
1949, 1970). The chromosome cytology of Wal- 
leria thus accords well with Tecophilaeaceae and, 
viewed in conjunction with data reviewed above, 
supports its placement here. 


Missouni Bor. GARD. 76: 925-926. 1989. 


926 


Annals of the 
Missouri Botanical Garden 


LITERATURE CITED 
CARTER, S. 19 Revision of I alleria D Cyanas- 
trum re d Kew Bull. 85-195. 
,M.S ocumented ieu num- 
ioni in plants. Madroño 10: 95. 
1970. Chromosome numbers of d California 
1-48 & 


California. Publ. Bot 


CAVE 


Liliaceae. Univ. 
plat 
NU REN, CR. & A VAN Wyk. 1988. Structure and 
ide inp af families d to or centered in 
EHE a Afr des 'a. Monogr. Syst. Bot. Missouri Bot. 
d. 
EN E CMS NM P. Yeo. 1985. The Families 
f the Monocotyledons. Springer- -V erlag, Berlin 
bum DBLATT, P. P olyploidy i in URP AR mono- 
dons. Pp. 2 W. wis (editor), 
Polyploidy: E cal Rele vance. Plenum. New York. 
Huser, H. 196 Die 
sc ichafisy ae y mee rae 
amml. München 8: 219-538. 
HUTCHINSON, J. 1934. The Families of oe ring Plants. 
, Ist edition. Macmillan, Londor 


RS 


. Mitt. Bot. 


> Samenmerkmale und Verwandt- 
Staats- 


59. The Families of n Plants. H, 

2nd od Clarendon Press, ord. 

Jackson, R. C. 1972. C abu ry evolution in no 
lopappus gracilis: acentric transposition rates. 
lution 27: 243-256. 

La Cour, L. F. 1955. In C. D. Darlington & A. 

W ylie, Chromosome Atlas " Flowering Plants. Mac- 
millan, New York. 

ORNDUFE, R. 1979. Chromosome numbers in Cyanella 
is cophilaeaceae). Ann. Missouri Bot. Gard. 66: 581 

83. 


on M. 1985. 
i te Grana 24 

— Peter Goldblatt, B. 4. Krukoff Curator of Af- 
rican Botany, Missouri Botanical Garden, P.O 
Box 299, St. Louis, Missouri 63166, U.S.A. and 
John C. Manning, National Botanic Gardens, Kir- 
stenbosch, P. Bag X7, South 
Africa. 


Pollen ultrastructure of the Te- 
77-95 


Claremont 7735, 


NEW COMBINATIONS AND 
TAXONOMIES IN 
CLUSIACEAE 


On preparing treatments of Clusiaceae for var- 
ious floras in Central America, I have found the 
need to make several new combinations. 


Dystovomita paniculata (J. D. d Hammel, 

comb. nov. Rheedia paniculata J. Smith, 

ot. Gaz. 55: 43 3. TYPE: Rica. 

San José: between Rio Union and Rio Con- 

vento, 800 m, Feb. 1898, Pittier 11957 (ho- 
lotype, US). 


Tovomita aid Engler, Bot. Jahrb. Svst. 58, Beibl. 
130: 8. 1923. Dystovomita d ri (Engler) D'Arcy, 
Ann. Mis Bon Gard. 65: 695. 1978. TYPE: 


Costa Rica o: Tuis TRUE d 1,000 m, May 
1899, Ee priv B, not seen; isotype, 


panamaensis Maguire, 

ytologia i an 1977. TYPE: Panama. Darién: 
Cerro Mali, 00 m, Jan. 1975, Gentry & Mori 
13769 fm Pn MO). 


Ch DEMOS clustifolta subsp. 
Phyto 


The following specimens probably represent most 
of the collections of this species. 


Additional specimens examined. COLOMBIA. CHOCÓ: 


Rio Truando, 40-200 m, Duke 13320 (MO); Hoya del 
Río San Juan, 5 m, Forero 4464 (MO); area of Baudó, 
8 m, Fuchs 21764 (MO); Serranía del Darién, 1,400 m 


niry a MO); eases to Tutunendo, 90 m, Gentry 
30286 (MO Rio Tagachi, 
Ô). VALLE: Bahía de Malaga, 
(MO). Costa Rica. HEREDIA 

, Folsom 9612 (DUKE), 9089 


sommet "t 1906, 11545 (DUKE); Hart- 
iei 15 520 (MO); D. Smith 17.5 (DUKE, MO). PUNTARE- 
NAs: near San Vito de Java, 1,700 m, Raven 21791 


(MO) Panama. CHIRIQUÍ: Fortuna area, 1,150 m, 
McPherson 7310, 12505 (MO). cocL£: El Valle, 1,100 
m, de Nevers 4398 (MO); 1,000 m, Duke 15102 (MOJ; 
900 m, McPherson 11 246 Apes 1,000 m, McPherson 
11959 (MO); 900 1 I i ) $ 
Mali, 1,300 m, io 3984 (MO). 1,400 m, Gentry 
13769 (MO); id o Cerro e 1,40 01 m, Ham- 
mel 16570 (MO). VERAGUAS: N of Santa Fe, Cerro Tute 

rea, 750 m, McPherson 12093 (MO), ‘Mori 2612 (MO), 


This is the only species of Dystovomita in Cen- 
tral America. Its large, ovate to nearly round leaves 
with very distant parallel lateral veins and large 
ligulate swellings at the bases of the petioles make 
The holotype 
382472, a 
twig with the remains of an inflorescence, and 
US— 1382473, leave 


even sterile material unmistakable. 
at US consists of two sheets: US— 1 


At MO we still have very little material of this 
genus from Venezuela, where a second species, D. 
clusiifolia (Maguire) D’Arcy, is endemic (one col- 
lection, Schultes 11755, is from Vaupés in south- 
eastern Colombia). Although I can see no way to 
distinguish sterile material of D. clusufolia from 
D. paniculata, seemingly minor differences in the 
staminate flowers, along the lines indicated by Ma- 
guire (1977) and D'Arcy (1978), do hold up for 
the many new collections of the latter species, 
which is restricted to Colombia, Panama, and Costa 
Rica. However, contrary to somewhat inaccurate 
and contradictory interpretations of prior authors, 
the flowers of both species have two bracteoles; 
the internode between the bracteoles and sepals is 
shorter in D. clusiifolia than in D. paniculata. 

In addition to differences in the staminate flow- 
ers, the fruits of D. clusiifolia appear to have 
somewhat longer stigmas (or at least they persist 
longer) and are slightly larger at maturity. than 
those of D. paniculata. 


Garcinia intermedia (Pittier) Hammel, comb. 
nov. Rheedia intermedia Pittier, Contr. U.S. 
Natl. Herb. 13: 454. 1912. TYPE: Guatemala. 
Alta Verapaz: Rio MN 300 m, Mar. 1902, 
von Türckheim 8180 (holotype, US, photo 

MO; isotypes. F, MO 


u oani See "mann, i Voy. Herald 89. 1853. 
R 


( Planchon & Triana, Ann. 

Sel. s Dot. . 14: is 1860. Not Garcinia 

edulis Exell, in ipn xi 1927. TYPE (of C. edule 
Seemann): Panama. Veraguas: dun Lorenzo, See- 


M, not pu ies MO). 
Rheedia tonduziana Engler, Bot. Jahrb. 58 Beibl. 
10. 1923. TYPE: Costa UM San José: ke Rio 
Virilla near San Juan, 1,100 m, May 1897, Tonduz 
11249 (holotype, B, not seen, photo MO; isotype, 
US, photo MO). 


mann s.n. (BM 


Robson (1958) discussed the logical error in 
maintaining Rheedia separate from Garcinia, es- 
sentially saying that no basis for separating them 
exists. Previously, Garcinia was not considered to 
occur naturally in the New World, only Rheedia. 
Features thought to distinguish Rheedia from Gar- 
cinia (number of perianth parts and degree of 
fusion of the stamens) are not constant in Rheedia 
and occur in both genera. Adams (1972) and Four- 


ANN. MISSOURI Bor. GARD. 76: 927-929. 1989. 


928 


Annals o 
Missouri Botanical Garden 


net (1978) treated Rheedia as a synonym of Gar- 
D'Arcy 


(1980) nevertheless kept the two genera separate 


cinia for floras within the. West. [ndies. 


on grounds that transferring the species of subg. 
Rheediopsis from Garcinia to Rheedia would be 
"more correct" than uniting Rheedia with Gar- 
cinia. Recent comparative work within the family 
supports Robson's position (P. Stevens, pers. 
comm .). 

Seemann's epithet (edule) cannot be transferred 
to Garcinia as it would create a later homonym. 
Rheedia intermedia clearly belongs in Garcinia 
and provides the oldest available epithet for the 
spec ies. 

Garcinia intermedia is the only Central Amer- 
ican member of the genus with a soft, edible, exo- 
carp. All others have a thick, tough, inedible exo- 
carp and delicious, white, acidic, sweet endocarp 
(as in the mangosteen, Garcinia mangostana L.). 
Leaf sizes and shapes are variable in the species 
and encompass the narrow-leaved material named 
as Rheedia tonduziana. 


Garcinia macrophylla C. Martius, Flora 24. 2 
Beibl. 35. 1841. Rheedia macrophylla (C. 
Martius) Planchon & Triana, Ann. Sci. Nat. 
Bot. Ser. 4. 14: 309. 1860. Garcinia mega- 
phylla Verde., Kew Bull. 31: 262. 1977, 
. (superfluous new name based on 
Martius) Planchon 
& Triana). Not Garcinia macrophyla Miq. 
(1864). Not “G. macrophylla Anderson ex 
Hooker" 
Kewensis (1895), this species published as G. 
TYPE: Mar- 


nom. illeg 


Rheedia macrophylla (C. 


(manuscript name listed in Index 
andersonii Hooker, 1872). Brazil. 
tius s.n. (not seen). 
EM macrantha Standley & ic an Field Mus. 
. Hist. Bot. 23: 65. 19 :: Guatemala. Alt: 
Pain along Rio Frio near Tact 'tic, 1,400 m, Apr. 
1941, Standley 90524 (holotype, F). 


In a footnote to a description of a new African 
species of Garcinia, Verdcourt (1977) transferred 
Rheedia macrophylla to Garcinia giving it the 
new but superfluous name, G. megaphylla. His 
citation states that Rheedia macrophylla Plan- 
chon & Triana is “non Garcinia macrophylla T. 
Mart., nec Mig.” 


& Triana gave a complete citation of G. macro- 


Anders., nec However, Planchon 
phylla C. Martius in synonymy of their R. mac- 
rophylla. Clearly, the basionym of Rheedia mac- 
rophylla (C. Martius) Planchon & Triana and oldest 
usage of the epithet 


“macrophylla” in Garcinia is 
Martius. 


rophylla Miq., from Sumatra, published more than 


G. macrophylla C. Garcinia mac- 


20 years after Martius’s Brazilian species, may 


need a new name. Ít is not certain that G. 
phylla C. Martius 
America (see discussion under Garcinia magni- 
folia Pittier). 


macro- 


occurs in southern Central 


Garcinia madruno (Kunth) Hammel, comb. nov. 
i dice madruno Kunth, Nov. Gen. & 
Sp. 5: 202. 1822. Rheedia madruno (Kunth) 
Planc lin & Tana, Ann. Sci. Nat. Bot. Ser. 
4. 14: 315. 1860. TYPE: Colombia. Antioquia: 
Magdalena and Cauca valleys, 300—1,000 m, 
Bonpland 1723 (P, not seen, photo MO). 


Verticillaria acuminata Ruiz Lopez & Pavón, Sy st. Fl. 
8 


ruv. 140. . Rheedia acu ta (Ruiz Lopez 
& Pavón) Plane a E As pees RE Nat. Bot. 
Ser. 4. 14: 3 Not Garcinia acuminata 


Planchon & n EAN TYPE: Peru: Ruíz « 
Pavón s.n. (G, not seen, photo, MO). 


Ruiz and Pavón's older epithet, acuminata, can- 
not be transferred to Garcinia as it would create 
a later homonym. Calophyllum madruno appears 
to provide the next-oldest available epithet for this 
distinctive species with tuberculate fruits. 


Garcinia magnifolia (Pittier) Hammel, comb. 
nov. Rheedia magnifolia Pittier, Contr. U.S. 
Natl. Herb. 13: 454. 1912. rYPE: Costa Rica. 
Cartago: above Rio Hondo near Matina, 400 
m, Aug. 1901, Pittier 16165 (holotype, US, 
photo MO). 


Rheedia chocoensis Cuatrec., Ann. Inst. Biol. 20: 106. 
. TYPE: Colombia. Valle: Rio Calima, La Trojita 
5-50 m, Feb. 1944, Cuatrecasas 16387 (holotype, 
F). 

| consider Panamanian material previously iden- 
tified as Rheedia lateriflora L. and R. 
phylla to be conspecific with Garcinia oo 

humilis (Vahl) €. Adams (= 


lateriflora) and G. macrophylla are lobed, har 


macro- 
Stigmas of G. 


the mature fruits are apiculate, whereas G. mag- 
nifolia has globose mature fruits with a disc-shaped 
stigma. Recent Colombian collections demonstrate 
that R. 


Garcinia magnifolia. 


chocoensis falls within the variability of 


Under the taxonomy adopted here, with fruiting 
material most important for identifying specimens, 
four native species of Garcinia occur in Central 


America. They can be distinguished by the follow- 


ing key: 

KEY TO NATIVE SPECIES OF GARCINA IN 

CENTRAL AMERICA 

la. Fruits with a soft, edible exocarp and 1 or 2 
seeds; leaves usually drying dull on both sur- 
faces "— . G. intermedia 


Volume 76, Number 3 Notes 929 
1989 
lb. Fruits with a hard, inedible exocarp and ofter D'Arcy, W. G. 1978. Dystovomita, a new genus o 
more than 2 seeds; leaves usually drying s neotropical Guttiferae. Ann. Missouri Bot. Gard. 65: 
above 694-69 
2a. bos tuberculate; mature e eae 980. Guttiferae. In: Flora of Panama. Ann. 
less than cm in diam.; lea dryi Missouri Bot. Gard. 67: ne 1043. 
DE silver-green hia. resin ihe. ms Fourxer, J. 1978. Flore Illustréi des Phanérogames 
vious on leaves G. madruno de Guadeloupe et de Marine os National de 


2b. Pu smooth; mature fruit often 6 cm 
or more in diam.; leaves drying green or 
brown below, resin lines inconspicuous. 
3a. Leaves very coriaceous, drying green- 
ish below; fruits globose, not apiculate ate 
mu s. magnifolia 
3b. Leaves + membranaceous, dryin 
brownish below; fruits ovoid, apiculate 
7. mac ella 


LITERATURE CITED 


ADAMS, C. D. 1972. Flowering Plants of Jamaica. The 
University Press, Glasgow. 


la Recherche oe a 
Hooker, 1872 e Flora of British. India. L. 


a 
Macuire, B. 1977. Notes o 
Panama. I. Phytologia 36: 
Robson, N. Guttiferae. /n: New and little g 
species from the ai Zambesiaca area. Bull. Soc 
Brot. 32: 170-17 
VERDCOURT, B. 1977 “Two new taxa of Guttiferae from 
East Africa. Kew Bull. 31: 262 


on the Clusiaceae — chiefly 
391-407 


Missouri Botanical Garden, 
Missouri 03 100-0299, 


— Barry Hammel, 
P.O. Box 299, St. 
U.S.A. 


Louis, 


A NEW SPECIES OF 
CALYCOLPUS (MYRTACEAE) 
FROM THE VENEZUELAN 
GUAYANA 


In preparation of a treatment of the subtribe 
Myrtinae (Myrtaceae) for the Flora of the Vene- 
zuelan Guayana a new species of Calycolpus has 
come to my attention. It is known to me through 
two collections, both from Estado Bolivar, Vene- 
zuela. It clearly belongs to Calycolpus as shown 
by the following combination of characters: flowers 
pentamerous and erect; stamens exserted, the an- 
thers elongate, with several glands in the connec- 
tive; seed coat lustrous with no pulpy covering, the 
outer seed coat ca. | cell thick at narrowest point, 
the superficial cells of the seed coat nearly isodi- 


ametric. 


Calycolpus bolivarensis Landrum, sp. nov. TYPE: 
Venezuela. Bolivar: entre kms 146 y 145 de 
carretera a San Juan de Manapiare, 150 m, 
5 Mar. | (fl), 4. Fernandez 3068 (ho- 
lotype, MY). Figure 1. 


C. cochleato McVaugh primo adspectu simillima, sed 
folio sessili vel subsessili, oblongo-lanceolato; petioli 
mm longi; nervi laterali prominenti subtus; pedunculis in 
brachyblastos bracheatos aggregatos in extremitatibus ra- 
SER i 
rutex circa |-metralis; pili E Barr Adi ramuli ju- 
ventute dense a folia n longa, 1.8 
3.3 cm lata, 2-3-plo longiora 2 bes apex acu- 
minatus; basis subo data; laminae coriaceae, in sicco 
pallido-brunneae vel atrorufo-brunneae; dci cam- 
panulatum, circa 4 mm longum, dense puberulum; stam- 
:a 8 mm longa, md. circ 
i -glandulis; ovarium 3 


m. 
" 


ina circa 
onga, 
lare, loculis 15-19-ovulatis; stylus 7 
ber 


e 
c 
on 
a 
S 
= 
^ 


10 mm longus, gla- 


Shrub ca. | m high, the twigs, leaves, and flowers 
sparsely to densely puberulent on most surfaces; 
hairs whitish, simple, mainly appressed, —0.1-ca. 
0.8 mm long; young twigs grayish to reddish brown, 
densely puberulent. Leaves sessile or subsessile; 
blades stiffly coriaceous, drying light brown to dark 
reddish brown, oblong-lanceolate, 4.5~ 10 cm long, 
1.8-3.3 cm wide, 2-3 times as long as wide, mod- 
erately to sparsely puberulent or densely puberu- 
lent along the midvein, the margin revolute, the 
apex acuminate, the base subcordate; midvein im- 
pressed to nearly flat above, prominent below; lat- 
eral veins ca. 20-22 pairs, nearly straight, leaving 


the midvein at nearly a 90? angle, visible or not 
visible above, prominent below; marginal veins about 
equaling laterals in prominence, running about par- 
allel to and ca. 2-3 mm from the margin; petiole 
0-1 mm long, 1-3 mm wide. Peduncles ca. 1-2 
1.5 mm wide at apex, velvety pu- 
clustered in 


cm long, ca. 
berulent, borne in the axils of bracts, 
bracteate shoots at the tips of branches; bracteoles 
clasping the hypanthium, triangular, ca. 0.6-1.5 
mm wide, 1 -2 mm long. Calyx lobes before anthesis 
about triangular, ca. 2-3 mm long, 2-2.5 mm 
wide, densely to sparsely appressed puberulent 
within, sparsely puberulent without, the hypan- 
thium tearing 1-3 mm between the lobes at an- 
thesis, the calyx lobes thus longer than before; 
petals sparsely to moderately puberulent; hypan- 
thium campanulate, ca. 4 mm long, densely velvety 
puberulent; disk ca. 5 mm across, densely puber- 
ulent on staminal ring; stamens ca. 185, ca. 8 mm 
long: anthers ca. 0.8 mm long, with ca. 10 glands 
in connective; ovary 3-locular; ovules 15-19 per 
locule, + multiseriate; style 7-10 mm long, gla- 
brous. Fruit globose, probably ca. 1 cm diam.; seeds 

ca. 12 per fruit, ca. long, the seed coat 
lustrous, ca. 1 cell thick across distal wall, the cells 


mm 


nearly isodiametric. 

Additional specimen examined. Venezuela. Bolivar 
entre Guaniamo y de 100-280 m, 9 
June 1984 (young fr), H Lopes Palacios, A. Carabot & 
C. Rosquete 4521 (NY). 


c 
H 


Calycolpus bolivarensis is most similar to C. 
cochleatus. The differences are summarized in the 


key below. 


m 
ie) 


Leaves oblong- lanceolate, sessile to subsessile, 
the petiole 0-1 mm long; lateral veins didi 
nent below: polum ii borne in the axils of brac 
clustered in bracteate shoots at the tips of UE 
branches; bendum densely pn pace 


bolivarensis 
Leaves rum to obov ate, petiolate, the peti 


1 axillar 
bract shoots along ength of young 
branches; hypanthium pee lana - 
cochleatus 


ANN. Missouni Bor. Garb. 76: 930-931. 1989. 


Volume 76, Number 3 Notes 931 
1989 


FIGURE 1. ol bolivarensis. —A. Flower bud.— B. Flower after anthesis, showing subtriangular petal 
scars, stamen scars, and tears between calyx lobes.— C. Leaf from below. —D. Lateral view of seed. A, C from 


Fernandez 3068 (MY); B, D from Lopez-Palacios et al. 4521 (NY). Drawn by Bobbi Angell. 


I am grateful to Bruce Holst for discovering and — Leslie R. Landrum, Department of Botany, Ar- 
sending me the specimen of Calycolpus bolivar- | izona State University, Tempe, Arizona 85287, 
ensis from MY that I have used as the type. U.S.A. 


CHROMOSOME NUMBER IN 


PHENAKOSPERMUM AND STRELITZIA 


AND THE BASIC CHROMOSOME 


NUMBER IN STRELITZIACEAE 
(ZINGIBERALES) 


Chromosome counts are available for most Afro- 
Madagascan members of Strelitziaceae but until 
now the neotropical Phenakospermum has been 
unknown cytologically. Chromosome cytology of 
the small southern African genus Strelitzia is com- 
paratively well known. The two acaulescent species 
5. reginae Ait. (syn. S. parvifolia Ait.) and S. 
juncea Link (possibly only a form of S. reginae) 
l4; in contrast, the 
caulescent S. alba (L.f.) Skeels (syn. S. augusta 
Thunb.) has 2n = 22 (Table 1). The closely related 
5. nicolai Regel & Koern., also caulescent, has 
been reported by Sató (1948, 1960) as having 2n 
caudata R. A. 


have diploid numbers of 25 — 


= 14. The remaining species, S. 
Dyer, is unknown cytologically. 

It has seemed puzzling that the chromosome 
number of S. nicolai corresponded with the two 
acaulescent species rather than with the allied cau- 
lescent 5. alba, particularly as the merit of main- 
taining the two species as separate (Geldenhuys, 
pers. comm.) is in some doubt. We have now es- 
tablished that at least one Transkeian population 


of S. nicolai has the same chromosome number, 


2n — 22, as S. alba. T 


[he chromosome number in 
22. 


Phenakospermum is also 2n — 


MATERIALS AND METHODS 


Seeds collected in the wild were germinated in 
the greenhouse. Actively growing root tips were 
harvested at mid morning into saturated aqueous 
mono-bromo-naphthalene and left at room tem- 
perature for four hours before fixation in 3:1 ab- 
solute ethanol-glacial acetic acid for two minutes. 
After hydrolysis in 10% HCl for six minutes at 
60°C, the apical portions of the tips were squashed 
in FLP orcein (Jackson, 
as follows: Phenakospermum guianensis Aublet 


1972). Voucher data are 


Guyana: DeFilipps s.n. (US); Strelitzia nicolai 
(L.f.) Skeels, South Africa. Transkei: coastal bush 
at Mkambati Nature Reserve, P. Goldblatt 7603a 


= 


OBSERVATIONS AND DISCUSSION 


The diploid chromosome number of P. gui- 
anensis and S. nicolai is 2n = 22. The chromo- 


TABLE l. Chromosome numbers in Strelitziaceae. 
Haploid 
Species Number n References 
Phenakospermum 
guianensis Aublet 11 this paper 


Ravenala 


madagascariensis Sonn. 11 


Strelitzia 


Cheesman & Larter (1935); Simmonds (1954); Sharma & 
Bhattacharyya (1959); Sató (1948, 1960) 


alba (L.f.) Skeels 11 Cheesman & Larter (1935); Simmonds (1954); Chakravor- 
ti (1960); Sató (1948, 1960) 

nicolai Regel & Koern. IBE this paper 

reginae Ait. f Cheesman eai rter (1935); presa in (1960); S 
(1948, 1960); Mananty (1970); Small et al. (ped 

juncea Link 1 Mahanty (19 Er s S. parv Jai var. juncea and as S. 
parv ifolia); Small et al. (1980) 

X kewensis (S. reginae x 5. alba) 17 Cheesman & Larter (1935 


orts by Satô (1948, 1960) of n = 
le. 


' Rey 
from the ta 


ANN. Missouni Bor. 


7 in this species are presumed to be incorrect and hence are excluded 


GARD. 76: 932-933. 1989. 


Volume 76, Number 3 
1989 


Notes 933 


FIGURE l. Mitotic metaphase in Strelitzia nicolai. 
Scale en = 10 um 


somes are of moderate size, ranging 1.5-3.5 um 
in the method used. The chromosomes are mostly 
acrocentric, but four of the smaller pairs are sub- 
metacentric (Fig. 1). The karyotypes are compa- 
rable to those illustrated by Small et al. (1980) for 
S. reginae and S. juncea except for the difference 
in the number of chromosomes. 

It is likely that Sató's count of 2n — 14 for 
Strelitzia nicolai is incorrect. Our count provides 
additional evidence D the caulescent species of 

Strelitzia have x . This corresponds with the 
number in the ions an Ravenala, also x — 
11 (Table 1), the least specialized member of the 
family according to Lane (1955 the arbores- 
cent members of Strelitziaceae thus have x — 

The families most closely related to Spehian: 


'ae : ex= 
and 12 respectively (Goldblatt, 1980). This makes 
it reasonable to hypothesize that x = 11 is basic 


for Strelitziaceae, as Mahanty (1970) suggested. 
If this is correct, it follows that the arborescent 
members of the family have retained the ancestral 
base number while the acaulescent species of Stre- 
This 


implies that the acaulescent condition in the family 


litzia have a derived chromosome number. 


is secondary, which is consistent with the conclu- 
sions of Lane (1955) and Tomlinson (1960). 


The alternative, that x = Y is basic, seems t 
us less likely, although Chakravorti (1960) has 
suggested that x — ll in Musaceae (including 


PES UE and Heliconiaceae) is derived from 
x — 7 by fragmentation across eight secondary 


This 


process is mechanically unlikely (Jones, 1977). In 


constrictions in the 5. reginae karyotype. 
light of the predominance of decreasing over as- 
cending aneuploidy in plants (Stebbins, 1950; Jones, 
Goldblatt, 1981), we assume 
that the chromosome number in the acaulescent 


977 & pers. comm.; 


species of Strelitzia was achieved by chromosome 
fusion leading to the reduced base number. Critical 


measurement of the quantity of DNA in Strelitzia 
may help solve this question. 

Support for this study by grants DEB 81-19292 
and BSR 85-00148 from the U.S 


Foundation is gratefully acknowledged. 


. National Science 


LITERATURE CITED 


CHAKRAVORTI, A. K. d > Megan studies with spe- 
cial reference to c nucleolus relationships 
oa its bearing on 1 the cytogenetics of Heliconia. The 
Nucleus 3: 225-250. 
CHEESMAN, E. E. & .H. LARTER. 1935. Genet 
and cytological studies of Musa: MI. Chromosome 
numbers in the Musaceae. J. Genet. 30: 31-52 
GOLDBLATT, P. 1980. Polyploidy in angiosperms: mono- 
cotyledons. Pp. 219-239 in H. Lewis (editor), 
Pli eo al Relevance. Plenum, New York. 
981. Cytology and the phylogeny of and 
ne -463 in R. M. Polhill & P. 
Raven (editors), Avanes in Legume da 
Royal Wer Gardens , Kew. 


minosae. Pp. 


Jackson, R. C bo wl evolution in Hap- 
pap gracilis acentric transposition rates, Evo- 
lu tio 24 : 243 256. 


Jones, K. 1977. The role of Robertsonian change in 
karyotype pines in higher plants. Chromosomes 
: 121-129. 


y 
Laxe, LE. 1955. ts and generic relationships in 
Musaceae. Mitt. Bot. Staatssamml. München x 3): 
114- 
MAHANTY, K. 1970. A eytological study of the 


eme with special reference to their taxonomy. 
ids ; 


ur 
- 

c 
y 
— 


Karyotype and sy (Y of Zingi- 

a Dr J. Genet. 23: 44-45. [In Japanese. ] 

960. Karyotype analysis in a ch with 

me reference to the protokaryotype and stable 

kar Pri S Ps Coll. Gen. Educ. Univ. Tokyo, 

Bio 5- 

SHARMA, A.K.& N BHATTACHARYYA. 1959. Cy- 
tology of séveral Ms of Zingiberaceae and a 
study of the Wu of their chromosome com- 
plement. Cellule 59: 299-345. 

SIMMONDS, N. W. E rap oo in 
some ya al plants Heredity in F 45. 

SMALL, J. C N DE VEN a GARNER. 
l a pons in E Sire litzia: hy- 

bridization and chromosome numbers. J. S. African 
Bot. 46: 167-171 

STEBBINS, G. L. 1950. 
Columbia Univ. Press, New York 

TOMLINSON, P. 1960. Phylogeny i the Scitamine- 
io and anatomical considerations. 
Evolution 16: 192-213 


T 


Variation and Evolution in Plants. 


— John C. Manning, 
Kirstenbosch, Private Bag X7, Claremont 7739, 
South Africa; and Peter Goldblatt, B. A. Arukoff 
Curator of African Botany, Missouri Botanical 
Garden, P.O. Box 299, St. Louis, Missouri 03100, 


National Botanic Gardens, 


ENCEPHALARTOS VOIENSIS 
(ZAMIACEAE), A NEW 

EAST CENTRAL AFRICAN 
SPECIES IN THE 

E. HILDEBRANDTII COMPLEX 


In the past 30 years, several locally endemic 
species of Encephalartos have been described for 
Central Africa. Most are from Mozambique (Dyer 
1969; Lavranos & Goode, 1985), 
Zimbabwe (Dyer & Verdoorn, 1969), and, to a 
lesser extent, Zaire (Devred, 1959; Malaisse, 1969). 


located west and southwest 


& Verdoorn, 


All of these taxa are 
of a line connecting Lake Victoria, Lake Tangan- 
1). Little work has 
taken place on Encephalartos east of this line. In 


yika, and Lake Niassa (Fig. 


fact, the distribution map in the most thorough and 


recent work on African Encephalartos (Melville, 
1957) shows no collections between the line and 
than two locally 
bubalinus Mel- 
Heenan (1977) 


repeated Melville’s (1957) descriptions and includ- 


the east coast of Africa other 
endemic species he described: £. 
ville and E. tegulaneus Melville. 
ed a discussion of three undescribed and imper- 
fectly known, presumably new taxa from this area. 
One of us (JPS) has recently collected material 
of Encephalartos in the central border region of 
Kenya and Tanzania. In these collections there is 


NAMIBIA 


q (à y 


ETHIOPIA 


,9,x,d E. sp. 
. ituriensis 

. barteri 

. bubalinus 


- 


© 

E 

E 

E 

E. chimanimaniensis 
E. concinnus 

E. gratus 

E. hildebrandtii 

E. laurentianus 

E. manikensis 

E. marunguensis 

E. munchii 

E. poggei 

E. pterogonus 

E. schmitzii 

E. septentrionalis 
E.tegulaneus 

E 
E 
E 


SsOXx(0OPOOOBRDADO + oo ^ 


FIGURE 1. Distribution map of Encephalartos species in Central Africa. Derived, in part, from Melville (1957) 
ika. 


and Devred (1959). L. Nia. = Lake Niassa, L. Vic. 


ANN. MISSOURI Bor. 


= Lake Victoria, L. Tan. 


= Lake Tanganyi 


GARD. 76: 934-938. 1989. 


Volume 76, Number 3 
1989 


Notes 935 


at least one new species, which is known to us from 
two localities near Voi, Kenya. This species appears 
in many respects, including locality, to e orrespond 


to Heenan's (1977) Encephalartos “sp. B." This 
species is described below and is followed by a 
discussion of its relationships and affinities to neigh- 


boring congeners. 


ied ce voiensis Moretti, D. Stevenson 
TYPE: Kenya: Voi, 2 Jan. 
hole: K). Figure 2 


¿ Sclavo, sp. nov 
1986 "^ Sclavo 
Truncus erectus, ad 2.5 m 
Cataphylla ac Serei pus eolat 
mentosa. Folia 2-4 m longa, oblor 
attenuata; rachis Kore duobus ls superne praedita. 
Foliola mediana alternata vel subopposita, oblanceolata, 
subfalcata, sensim acuminata, dx cun marginibus cal- 
basia cuneata in ciii 


i 35-70 cm UE 
1 to 


loso- dentatis, denti 


sus 
a iden sn 3 5-40 1 


diam.; pedunculus ebracteatus, 5-7 cm longu 
7 a mediana rhomboidea, 35 mm lon- 

40 mm lata; pedicellus 10 mm longus, tribus abaxi- 
sibus io praeditus; bulla rhomboidea, 40 mm lata, 

8 mm alta, 24 mı 

hexagona atque 
bus trapezoideis ports constituta atque tuberculata cris- 
ta tubereulateque margine praedita; lateralis bullae lobi 


7-9 mm longi, 2-4 mm lati, ad latera complanati. 


Trunk erect, to 2.5 m tall, 


straight to slightly tapering towards apex 
persistent. Leaves 2-4 m long, oblong, gradually 


35-70 em diam., 
, leaf bases 


tapered basally and becoming abruptly spinose in 
the lower 16. Petiole bulbous and tomentose. Ra- 
chis glabrous and with 2 shallow longitudinal adaxial 
grooves between leaflet insertions. Median leaflets 
alternate to subopposite, coriaceous, adaxially con- 
cave, blue-green, inserted at a right angle to the 
rachis and slightly oblique in the ab-adaxial plane, 
oblanceolate, 25-40 cm long, 4 cm 
wide, gradually acuminate apically and pungent, 
abruptly attenuate and strongly inequilateral ba- 


subfalcate, 


sally, superior margin with 1-5 strong spinose teeth 
on the basal curve and with 1-5 barely discernible 

callose teeth along the rest, inferior margin with 
0-1 small teeth basally and 1 
callose teeth along with the rest. 


-5 barely discernible 
Lower leaflets 
exhibiting an abrupt transition from those with 1 
basal spinose tooth on the superior margin to those 
with bifurcate apices to completely reduced and 
pungent lower leaflets. Wicrosporangiate strobilus 


long-cylindric, greenish yellow when young, yellow 
to tan when shedding pollen, 45-50 cm long. 10 
cm diam.; pe without bracts, glabrous, 5-7 

5 cm diam. Median microsporoph ylls, 


35-40 mm dune. 22-25 mm wide, perpendicular 


cm long. 3— 
to the strobilus axis, cuneate to deltoid, bulla de- 
flexed, terminal facet triangular to pentagonal. Me- 
gasporangiate strobilus cylindrical to ovoid-cylin- 
drical, green to yellow when young, yellow to tan 
when mature, 50-60 em long, 15-20 cm diam.; 
peduncle ebracteate and glabrous, 5-7 em long, 
3-5 cm diam. Median megasporoh y lls pedicellate 
with rhomboid bulla, 35 mm long, 40 mm wide; 
pedicel with 3 abaxial facets, the median facet 
10 mm long. Bulla 
10 mm wide, 18 
mm high, 24 mm deep; the terminal facet slightly 


inconspicuous and triangular, 
of megasporoph yll rhomboidal, 


concave and hexagonal, 15-17 mm wide, 6-8 mm 
tall; adaxial face with 2 lateral trapezoidal facets, 
a median rectangular to trapezoidal facet, and a 
short median lobe 2-4 mm wide and long. the 
margin and seminal fringe tuberculate: abaxial face 
with 2 lateral trapezoidal facets and a median rect- 
angular to trapezoidal facet with obtuse lateral sag- 
ittal ridges, the inferior ridge and sagittal crest 


tuberculate; lateral lobes 7-9 mm long, 2-4 mm 


wide, flattened laterally. 


Additional uy examined YA: i ar Voi, 


Mlilonyi Hill, 4 Jan. 1986 (3), Sello 2 2 (EA , MO, 
AP). 


Encephalartos voiensis lives in isolated moun- 
tains surrounded by dry savanna in which species 
of Encephalartos do not occur. The new species 

ccurs on steep rocky slopes cloud forest at 
s -1,800 m elevation as an understory plant. 
It also grows in partial shade and full sun areas of 
forest clearings. 

Encephalartos voiensis shares some characters 
with E. en E. gratus Prain, E. hildebrand- 
tii A. & Bouché, and F. 


are all ihe in the same general vicinity (Fig. 1): 


tegulaneus, which 
and, to a lesser extent, it resembles FE. manikensis 
(Gilliland) Gilliland and its allies in Central Mozam- 
bique and in Zimbabwe. Encephalartos votensts 
shares different features with each of these taxa. 
For example, foliage characteristics are closest to 
E. bubalinus and (especially) E. hildebrandtii in 
that 1—5 spinose teeth are present on the basal 
curve of the superior margin of the median pinnae, 
and at least one spinose tooth is present near the 
basal portion of the inferior margin. Encephalartos 
voiensis differs from these two species by having 
subfalcate, very abruptly cuneate basally, very dark 
green to blue-green leaflets, and the transition to 


Annals of the 


936 


Missouri Botanical Garden 


A 


Volume 76, Number 3 
1989 


Notes 937 


reduced leaflets in the lower l6 of the rachis is 
abrupt rather than gradual. Encepha 


sis can be distinguished further from E. bubalinus 


lartos voien- 
because the former has median leaflets 4 (vs. 1— 
2) cm wide, 25-40 (vs. 10-20) em long, and 
gradually acuminate (vs. acute) apices. In general, 
in E. hildebrandtii, the median leaflet apices are 
in E. 


contrast, they are always pungent. Moreover, the 


either bifurcate or trifurcate; volensis, by 
median leaflets of E. hildebrandtii have up to nine 
spinose teeth along each margin; in £. votensis 
these teeth are reduced to barely discernible callose 
bumps. 

The mega- and microsporangiate strobili of Æ. 
volensis resemble those of E. bubalinus, E. gratus, 
E. hildebrandtii, and E. tegulaneus with respect 
to overall size, shape, and color. There are, how- 
ever, significant differences in mega- and microspo- 
rophyll characters. As in vegetative characters, £F. 
votensts shares individual characters with each of 
the aforementioned species, but none that is com- 
mon to all except perhaps the presence of inferior 
and sagittal crest tubercles. For example, only E. 
gratus and E. voiensis have a rectangular to trap- 
This median lobe is 
5-12 mm long in £. gratus but only 2 
in K. 


unique abaxial tuberculate facet that replaces the 


ezoidal adaxial median lobe. 
4 mm long 
volensis. Encephalartos tegulaneus has a 


abaxial angle of the pedicel of its megasporophylls. 
but here 
usually does not extend the 


This feature also occurs in E. voiensis, 
he facet is narrower, 
full length of the pedicel, is barely discernible in 
herbarium material, and is nontuberculate. The a 

and adaxial margins of the bulla of the megaspo- 
rophylls of £F. 


tuberculate and without ridges, whereas those of 


votensis and E. hildebrandtii are 
E. bubalinus, E. gratus, and E. tegulaneus always 
have some distinct ridges. The overall appearance 
of the megasporophylls of E. voiensis in shape and 
arrangement of the bulla facets is most similar to 
those of E. hildebrandtii. Compared with the pre- 
viously discussed taxa, the lateral lobes of the bulla 
of E. 


and appear laterally compressed in the ab-adaxial 


voiensis are uniquely smaller and thinner 


ane. 
The deltoid to cuneate microsporophylls of £F. 
voiensis are more similar to those of E. bubalinus 


than to the oblong microsporophylls of £. gratus, 


and E. 


voiensis and E. 


E. hildebrandtii, In addition, 


only E. 


tegulaneus. 
bubalinus have their mi- 
erosporophylls inserted at a right angle to the stro- 
bilus axis at pollen shedding, whereas in the other 
species they are either ascendant as in £. 
and E. hildebrandtii or descendant as in E. te- 


gratus 


gulaneus. Encephalartos voiensis has a median 
facet on the ab- and adaxial bulla a these 
median facets are not present in £. gratus. 

Encephalartos voiensis exhibits little bl 
except that some leaflet characters appear to be 
associated with habitat differences. The leaflets of 
plants growing in deep shade are thinner in texture 
and barely subfalcate, and they generally have 
fewer and smaller marginal spinose teeth. Only 
shade plants exhibit these features, and they would 
most likely change under exposure to higher light 
intensities, as Stevenson et al. (1986) observed in 
Ceratozamia latifolia under otherwise similar con- 
ditions. 

Even though the relationship of Æ. votensis is 
not completely clear, it is our opinion, based on 
vegetative morphology, that Æ. 
closely related to E. hildebrandtii. 


vOlensis is most 
This is because 
in the cycads, in general, vegetative characters are 
more useful in separating species as, for example, 


980). 


Macrozamia 


in Ceratozamia (Stevenson et al., Dioon 
(Sabato De Luca, 1985), 
(Johnson, 1959). There are several reasons for this. 
First, vegetative characters are always present and 


and 


often exhibit more obvious macroscopic diversity 
than do reproductive characters. Secondly, most 
data concerning reproductive characters are either 
fragmentary or nonexistent, and herbarium ma- 
terial is highly unreliable because of the vast amount 
of distortion of size and shape induced by drying. 
Thirdly, because all cycads are dioecious, we think 
it is important to establish vegetative characters 
that can be correlated with reproductive characters 
to establish a common denominator. In the case of 
the species discussed in this work, microsporoph yll 
and megasporophyll characters do help corroborate 
the species ‘delimitations indicated by vegetative 
Finally. 
changes in vegetative morphology between species 
of different habitats and. o 
same time exhibiting only subtle differences in re- 


characters. cycads often exhibit extreme 
r localities while at the 


productive morphology. These vegetative changes 


— 

FIGURE 2 Encephalartos voiensis. A, B. Leaf and leaflet characters. — A. Leaf with characteristic abrupt transition 
to reduced m rg in lower portion. Scale bar = : dun Subfalcate leaflets of holotype with spinose teeth. 
Scale bar — ;.. End view of megasporophylls. S e bar = 5 cm. —D. Adaxial view of megasporophyll of 


the holotype. PPA bar = ] cm. 


—E. End view of a ed Scale bar = | cm. 


938 Annals of the 
Missouri Botanical Garden 


are stable and remain so even when the species ŒE. hildebrandtii (Fig. 1) occurs at the top and on 
are brought together and cultivated through suc- the sides near the top of coastal cliffs and on the 
cessive generations in a uniform environment. sides of river valleys. With the appropriate habitats 
The high incidence of locally endemic species isolated from each other, there is the opportunity 
of Encephalartos in Central Africa appears to be for locally endemic taxa to differentiate. Encepha- 
related to habitat preference and physical geog- — lartos voiensis fits this pattern because it also oc- 
raphy in this region. That is, in general, Encepha- curs on steep slopes of isolated mountains sur- 
lartos here prefers steep wooded slopes, and this rounded by savanna. As more areas of Central 
habitat occurs in isolated areas surrounded by sa- Africa are explored for Encephalartos populations, 
vanna in which this genus does not occur. Even more new taxa will certainly be discovered. 


Keys To East CENTRAL AFRICAN SPECIES OF ENCEPHALARTOS 


KEY TO VEGETATIVE PLANTS 
E. hildebrandtii 


la. Median leaflet apic es bi- or trifurc 
lb. Median leaflet apices exin pointed, ending in a spikelike tip. 
2a. Median leaflets subfalca 
2b. Median leaflets linear to qa 
3a. Median leaflets linear, narrower than 2 cm . E. bubalinus 
3b. Median leaflets lanceolate or oblanceolate, iy than 2 c 
4a. Median leaflets oblanceolate, margins exed, distinc a striate below oo — < < < —— E. tegulaneus 
4b. Median leaflets lanceolate, margins flat. aid striate below J 


E. voiensis 


KEY TO MEGASPORANGIATE PLANTS 

la. Microsporaphylis oblong, either ascendant or descendant at pollen shedding. 
2a. Bulla puberulent 
2b. Bulla glabrous. 


3a. Microsporophylls ascendant | 


. E. gratus 


E. hildebrandtii 
E 


3b. Microsporophylls descendant tegulaneus 

lb HR rosporophylls deltoid to cuneate, at t right angles to strobilus axis at pollen shedding. 
. Lateral ridges of bulla acute E. bubalinus 
i Lateral ridges of bulla subacute t to obtuse ... E. voiensis 


KEY TO MEGASPORANGIATE PLANTS 
Median lobe of bulla well developed, 7-12 mm long, bulla puberulent . E. gratus 
lb aee lobe of bulla absent or if anes shorter than 5 mm, bulla glabrous or nearly so. 
edian lobe of bulla 2-4 mm lor 

A ae » of pulla abse 

subtriangular, abaxial angle of pedicel absent 
3b. Bulla vb abaxial angle of pedicel present. 

xial m argin of bulla with GENE ridges 

les 


Ha Mrd margin of bulla with tubercles een pe d es E. hildebrandtii 


— 
mr 


E. voiensis 


E. tegulaneus 


E. bubalinus 


Cycadacée nouvelle de Congo-Kinshasa. Bull. Jard. 


This work was financially supported in part by 
7 Bot. Etat a 40 


NSF Grants BSR-86707049 and BSR-8796279 


MELVILLE, R. 1957. Ence € nialari in Central Africa. 
» DWS. ns B 
to | Kew Bull. s 237- 
s SABATO, S. & P. DE Tdi 1985. Evolutionary trends 
LITERATURE CITED , y 
in Dion reed Amer. J. Bot. 72: 1353-1363. 
DEVRED, ^ 1959. Une Cycadacée nouvelle du Congo — STEVENSON, D., S. SaBato & M. VazouEz-TonnEs. 1986. 
5 fus ephalarto marunguensis Devred. Bull. A new rid E Ceratozamia re from 
x, 91: 103-108. Veracruz, Mexico with comments on spec rela- 
DYER R, VR & L VERDOORN. 1969. V pes man- VEA habitats, and "e rav SD V es in c er 
ikensis and its near allies. Kirkia 7: 58. atozamia. Brittonia 38: 26. 
HEENAN, D. 1977. Some observations on d cycads of 
u : >, y i no 
Ceritral Africa, 1. Linn.. Soc. Bot 74.9 Aldo Moretti and | aolo De Luca, Dipartimento 
JOHNSON, L. . The families of cycads and A di Biologia Vegetale, Università di Napoli, 223 
a o srl Proc. Linn. Soc. New South via Foria, 80139 Napoli, Italy; Jean Pierre Scla- 
Wales 84: 6 vo, Villa La Finca, Plateau du Mont Boron, 


LAVRAN & k E 9 nce o. " : mue s 
Ren J. 00DE. 1985. Encephalartos tur- 06300 Nice, France; and Dennis Wm. Stevenson, 
neri (Cycadaceae), a new species from Mozambique. . . . 

Cia de oe te ]l=14 Vew York Botanical Garden, Bronx, New York 
MararssE, F. 1969. Encephalartos schmitzii Malaisse; 10458, U.S.A 


A NEW SPECIES OF 
PERSEA (LAURACEAE) 
FROM SURINAM, WITH 
A DISCUSSION OF ITS 
POSITION WITHIN 
THE GENUS 


In a shipment of Lauraceae recently received 


— 


from the University of Surinam, Paramaribo, 
encountered a specimen of Persea that turned out 
to represent a new species, differing from all other 
neotropical Persea species by its deciduous and 


strongly unequal tepals. 


Persea julianae van der Werff, sp. TYPE: 


Surinam: Wilhelmina Mountains, Juliana Peak, 


nov. 


small tree, 7 m tall, flowers very pale yellow, 
axes of inflorescence reddish, Schulz 10234 


(holotype, MO: isotype, BBS). Figure 1. 


A congeneribus tepalis deciduis et valde inaequalibus 


differt 


Tree, 7 m tall. Very young shoots and buds 
densely appressed-pubescent, the twigs soon be- 


10-14 


the 


coming glabrous. Leaves alternate, elliptic, 
3:9-5. Gm, Æ 


tips acute or shortly acuminate, the upper surface 


coriaceous, the base acute, 


glabrous, the lower surface with scattered ap- 
pressed hairs, the midvein clearly impressed and 
the lateral veins weakly impressed on upper sur- 
face, both of these vein types clearly raised on 
lower surface; tertiary venation weakly raised on 
lower surface. Petioles 1-1.5 cm long, glabrous, 
canaliculate. Inflorescences 4-6 cm long, in axils 
of deciduous bracts or leaves, paniculate, ap- 
pressed-pubescent, the axes reddish (fide collector). 
Pedicels ca. 4 mm long. Flowers very pale yellow. 
Tepals 6, strongly unequal, united at the very base 
and falling off in old flowers as a unit, with stamens 
and staminodia attached. Outer tepals broadly ovate, 
ca. 2 mm long, appressed-pubescent outside, gla- 
brous inside. Inner tepals elliptic, ca. 6 mm long, 
appressed-pubescent on both surfaces. Fertile sta- 
mens 6, these representing the outer 2 whorls, ca. 
5 mm long, the filament slender, ca. 4 mm long, 
4-celled, 


the cells introrse. Stamens of the third whorl about 


densely pubescent, the anther glabrous, 


as long as the outer stamens, the filament similarly 
pubescent, the anther as wide as the filament and 


devoid of anther cells; glands present, sub-basal. 


Outer 9 stamens with widened bases. Staminodia 


(whorl IV) ca. 2.5 mm long, widened at base, 
filament densely pubescent, inside of the anther 
glabrous except for an apical tuft of hairs, the 
outside sericeous, the base cordate. Ovary ellipsoid, 
ca. 1 mm long. glabrous or with few hairs near the 
apex; style ca. 2.5 mm long, glabrous. Fruit un- 
known; tepals deciduous in very early fruiting stage. 


The distinctive characters of P. julianae make 
it difficult to place in the most recent infrageneric 
classification. In her revision of the neotropical 
species of Persea, Kopp (1966) recognized two 
subgenera. Subgenus Persea is characterized by 
its equal tepals that are pubescent on inner and 
outer surfaces, usually puberulous ovary, and te- 
pals deciduous at the fruiting stage. This subgenus 
includes the type species of Persea, P. americana 
Mill., 


Subgenus Eriodaphne is 


and three other Central American species 
characterized by its 
strongly unequal tepals with the outer tepals in- 
ternally glabrous, usually glabrous ovary, and per- 
sistent tepals in the fruiting stage. This subgenus 
includes about 70 neotropical species and is further 
divided by Kopp (1966) into four sections. Section 
Mutisaea includes the species with a corymbiform 
inflorescence; section Hexanthera includes species 
with only six fertile stamens (representing the out- 
ermost whorls), section lurataea includes species 
with a (sometimes slightly) pubescent ovary; and 
section Eriodaphne includes the remaining species, 
with a glabrous ovary. The species with partly or 
entirely two-celled stamens belong to section £r- 
iodaphne. Thus, the two subgenera can be readily 
identified in flower (tepals equal vs. unequal) and 
fruit (tepals deciduous vs. persistent). 

e usefulness of this classification of the neo- 
tropical Persea species is undercut by several species 
with characters of both subgenera. Several Central 
American species (for example P. albida Koster- 
Allen, and P. 
Werff) have equal tepals, which are persistent in 


mans, P. rigens sylvatica van der 
the fruiting stage. These species were included in 
section Eriodaphne by Kopp, but the equal tepals 


ANN. MISSOURI Bor. GARD. 76: 939-94]. 1989. 


940 Annals of the 
Missouri Botanical Garden 


FIGURE 1. Persea julianae. — A. Habit. — B. Flower. — C. Detail of flowers showing two outer tepals (small), each 
with a fertile stamen (whorl I) and a tall staminode (whorl III), and one inner tepal (long) with a fertile stamen (whorl 
II) and a short, sagittate staminode (whorl IV). — D. Staminode of whorl II, with sub-basal glands. — E. Stamen of 
whorl L/II. —F. Pistil.— 6G. Staminode of whorl IV. —H. Old flower, showing dehiscence of tepals and stamens. 


do not support this placement. Kopp's (1966) in- P. hexanthera Kopp. These species are compared 

frageneric classification is weakened further by the in Table 1. 

unequal tepals deciduous in the fruiting stage of Persea julianae, named after the Queen Mother 

?, julianae. of the Netherlands, is only known from the type 
If it were not for the deciduous tepals, P. ju- collection from the Juliana Peak in Surinam at an 

lianae would be immediately placed in subg. Ær- elevation of 1,050 m. It was collected in a 20-m- 

iodaphne section Hexanthera, which is repre- — tall forest, rooting in a thick humus layer on sandy 


sented in Amazonian Brazil and Venezuela by P. clay. 
fastigiata Kopp, P. pseudofasciculata Kopp, and Kostermans (1936) recorded three species of 


Volume 76, Number 3 
1989 


Notes 941 


and P. hexanthera, based on MO type specimens, with exception of P. 


' 


TABLE 1. 
hexanthera, where data are based on the protologue and the collection Bernardi s.n., annotated by Kopp. 


Selected characters of Persea julianae, P. pseudofasciculata, P. fastigiata 


Lengths of 


Lengths of 
Inner Tepals 


Length of 
Inflorescence 


Lengths of 


Twigs 


Petioles 


Pedicels 


Pubescence 


Tepals 


solid 


6 mm 4 mm 1.0-1.5 cm 


sparse, appressed 


14-14 leaves 


deciduous 


P. julianae 


E 


2.5 cm 


3 mm 
3 mm 


persistent 


P. pseudofasciculata 


sparse, appressed 1.0-1.5 em 
9 


longer than leaves 


persistent 


P. fastigiata 


mm .5 em hollow 


8.5 mm 


sparse, appressed 


longer than leaves 


persistent 


hexanthera 


P. 


Persea from Surinam, P. americana Miller, P. 
benthamiana Meissner, and P. coriacea Koster- 
mans. As noted, the strongly unequal tepals, gla- 
brous ovary, and number of fertile stamens sepa- 
rate P. julianae from the cultivated P. americana. 
Specimens treated by Kostermans (1936) as P. 
benthamiana were included by Kopp (1966) under 
P. nivea Mez and are characterized by having nine 
fertile stamens, persistent tepals, and glaucous 
whitish lower leaf surfaces. 

Persea coriacea is an enigmatic species known 
only from the type collection, from the Hendrik 
Peak in the ma Chain, not far from the Juliana 
Peak. This species is described as having subequal 
tepals + 1.75 mm long and nine fertile stamens; 
these details separate it clearly from P. julianae. 
Kostermans (1936) also mentioned that the fruit 
was subtended by a small, flat cupule with an entire 
margin, a character not found in any other Persea 
species. This was probably one reason why Ko 
(1966) excluded P. coriacea from Persea without 
suggesting an alternative generic placement. The 

MO isotype of P. coriacea does not solve these 
difficulties. The label states that flower buds and 
fruits were collected. Presence of floral buds and 
fruits on the same specimen is very unusual for 
Lauraceae and is often the result of a disease in 
which galls, developed from buds, are mistaken for 
fruits. The buds on the MO isotype are diseased, 
which explains Kostermans's observation that fre- 

ently a staminode of the fifth whorl is present. 
The papillose lower leaf surface is against its in- 
clusion in Persea. Papillose lower leaf surfaces are 
common but not restricted to Aniba, but P. co- 
riacea does not match any Aniba species | know. 
I regard P. coriacea as a name based on a mon- 
strosity, different from the other Persea species 
reported from Surinam; for correct placement, ad- 
ditional specimens are needed. 


I thank Dra. M. Werkhoven for making the 
specimen available. J. Myers drew the illustration. 


LITERATURE CITED 


Korr, L. E. 1966. A taxonomic revision of the genus 
ersea in the Western Hemisphere. Mem. New York 
120. 


Bot. Gard. 14 
KosTERMANS, .G. H. 1936. Lauraceae. Ín: A. Pulle 
(e 2: 244-336 


ditor), Flora of Surinam 2: 2 
—Henk van der Werff, Missouri Botanical Gar- 
den. P.O. Box 299, St. Louis, Missouri 63160- 
0299. U.S.A 


THE PUBLICATIONS OF 
EDGAR ANDERSON: 
ADDITIONS 


Eisendrath (1972) published a bibliography of 


the writings of Edgar Anderson, botanist at the 
Missouri Botanical Garden and Professor of Botany 
at Washington University for most of his career, 
spanning the years from 1922 until his death in 
1969. For biographical information on Anderson, 
see Finan (1972). 

The Missouri Botanical Garden Library is gath- 
ering for binding and cataloging the collected works 


'The fol- 


of the curatorial staff, past and present. 


lowing additions to the Eisendrath bibliography of 


Edgar Anderson have turned up: 


1934. A chromosomal interchange in maize. in- 
volving the attachment to the nucleolus. 
Naturalist 68: 345-350 
Chromosomes involved in a series of inter- 
changes in maize. Amer. Naturalist 68: 
440-445. [With I. W. Clokey. ] 
Chromosomal interchanges in maize. Ge- 
netics 20: 70-83 

Christmas Eve thoughts from an herb gar- 
den. News Bulletin, New York unit of the 
Herb Society of America [2?]: [1]. 


Amer. 


1 934. 


1935. 


1946. 


ANN. MISSOURI Bor. GARD. 76: 942. 1989. 


. A nice quiet evening with a potato. Missouri 
Bot. Gard. Bull. 43(April): 50-53. (Re- 
printed as pp. 39-42 in C. B. Heiser, 1969. 
Nightshades, the Paradoxical Plants. W. 
H. Freeman and Company, San Francisco.) 
Islands of tension. Landscape 15: 7-8. (Re- 
printed in Arnoldia 48(3): 28-31. 1988.) 
Review of R. H. Platt, 1965, The Great 
Forest. Prentice-Hall, Engle- 
wood Cliffs, New Jersey. Landscape 10: : 


1966. 


1967. 


American 


LITERATURE CITED 

EISENDRATH, E. 1972. The oe of Edgar An- 
derson. Ann. Missouri Bot. Gard. 59: 346-361. 

FINAN, J. J. 1972. Edgar ne 1897-1969. Ann. 
Missouri Bot. Gard. 59: 325-345. 


— Sally Erickson, Volunteer, Missouri Botanical 
Garden Library, P.O. Box 299, St. Louis, Mis- 
souri 63166, U.S.A. 


BOOK REVIEW 


Flora of Australia, Volume 19. Myrtaceae— Fu- 
calyptus, Angophora. By G. M. Chippendale, 
including an introduction and appendix, with 
contributions by D. J. Chippendale & S. G. 
M. Carr, A. S. George, K. D. Hill, L. A. S. 
Johnson, and P. J. Lang. Australian Govern- 
ment Publishing Service, Canberra. 1988. xvi 
+ 543 pp., 111 figs. (38 color), 564 maps. 
ISBN 0-644-05866-8. Paper, price unknown. 


This recently published volume of the Flora of 
Australia is the first of three that will cover that 


arge group of trees and shrubs, the Myrtaceae. It 
is beautifully illustrated with numerous line draw- 
ings and seven pages of color photographs, is print- 
ed on high-quality paper, and has a good paperback 
binding. The volume includes a family description 
by A. S. George and treatments by G. M. Chip- 
pendale of the diverse genus Fucalyptus L Hér. 
(513 species) and the much smaller Angophora 
Cav. (7 species). George calculates that there are 
about 75 genera and over 1,500 species of Myr- 
taceae in Australia. Thus, Fucalyptus accounts for 
about one third of the family there. As stated in 
"more than any other group, 


e 


the introduction, 
these plants [the eucalypts ] give a distinctive char- 
Of course, 


acter to the Australian landscape. 
species of Fucalyptus are also planted around the 
world for ornamental and forestry purposes. Chip- 
pendale's contribution is very welcome indeed. 
An informative introduction relates some of the 
taxonomic history of Fucalyptus. The last com- 
plete account was that of Blakely, 4 Key to the 
Eucalvpts (1934), with updated editions appearing 
in 1955 and 1965. Until the late 1950s taxonomy 
was based on macroscopic characters of buds, fruits, 
and anthers, but since then a variety of anatomical 
and inflorescence features have been investigated 
by various workers. Presently, studies of Kucalyp- 
tus are appearing at an explosive rate. A quick 


look at the 18 issues of the dustralian Journal of 


Botany for the period 1984-1986 shows that not 
one issue was without at least one article with 
Eucalyptus in the title and that 59 articles on 
Eucalyptus appeared, 18 of which were system- 
atic. Thus, Chippendale's treatment is necessarily 
a synthesis of knowledge of Fucalyptus as it was 
in the mid 1980s, and it will soon be out-of-date. 
In fact, 24 species published while the book was 
being typeset could not be included. But consid- 


ering the history of Fucalyptus, we cannot expect 
another comprehensive work soon. 

Chippendale briefly reviews some of the features 
that have been used in systematic studies of Fu- 
calyptus, viz., stamens and anthers, cotyledons, 
operculum, ovules, leaf waxes, oil glands, tri- 
chomes, and stigma and style morphology. How- 
ever, his key to species depends on “readily-ob- 
servable characters of habit, bark, leaf arrangement 
and form, inflorescence, buds and fruit. ^ 

Near the beginning of the volume are color 
photographs illustrating types of bark, habit, and 
various species in flower. Labeling the photographs 
of the various bark types would have made them 
more useful. 

There are 60 pages of keys for Eucalyptus, 
first one dividing the species into 20 groups and 
then a key to the species in each group. For sys- 
tematic treatment the species are divided into 92 
series, but these have no close relationship to the 
20 groups of species in the key. A description of 
each series is provided with an accompanying ci- 
tation of type and notes on size, distribution, and 
distinguishing characteristics. There is no key to 
or synopsis of the series, but they are divided into 
groups in the introduction of the genus under the 
discussion of some of the systematically important 
characters. For each species are provided: a de- 
scription of ca. 8 lines; citation of types, synonyms, 
and published illustrations; a figure illustrating buds 
and fruits; a short list of representative specimens; 
and notes on distribution and distinguishing char- 
acteristics. Distribution maps for all species are 
found on 38 pages at the end of the systematic 
treatment. 

he style of the keys is very brief. Usually only 
one character comparison is made in each couplet 
and the alternatives can sometimes seem rather 
“adult 


“adult leaves 


arbitrary, e.g.. on page 8, lines 8 and 9 
leaves more than 12 cm long” versus * 
less than 12 cm long”; or lines | and 2 on page 
9 —“leaves green" versus "leaves glaucous or grey 
green or yellow green." The very first character 
used in the key is bark type, and bark characters 
are utilized at other points in the key. | experi- 
mented by attempting to “identify” three species 
I know and was able to run them through the key 
without problem. | was less successful with iden- 
tified specimens of three other species | did not 
know, even when I allowed myself knowledge of 
the bark by looking it up in the descriptions. In 


ANN. MISSOURI Bor. GARD. 76: 943-944. 1989. 


944 


Annals of the 
Missouri Botanical Garden 


my quick look at the koye [i ran into one contra- 
diction: if one takes lead “3:” on page 7, “oper- 
culum + equal to or longer A hypanthium,” 
" in group 9 on page 


one 
may later come to lead *100:7 
31, "operculum + equal in length to, or shorter 
than, hypanthium.’ 

In an appendix, Chippendale describes as new 


» 


or lectotypifies several series, and nine new species 
included in ie flora are Dx published by D. 
J. Carr & S. G. M. Carr S. Johnson & K. 
D. Hill, and es J. Lang. 


Chippendale's impressive treatment of Kucalyp- 
tus is sure to be a vital reference for botanists, 
foresters, and horticulturists. It obviously repre- 
sents many years of careful study and deep fa- 
miliarity with the genus. I am very happy that it 
has been published and recommend it to anyone 
with an interest in Eucalyptus. 


L. R. Landrum, Department of Botany, Ari- 


zona State University, Tempe, Arizona 85287, 
U.S.A 


Volume 76, Number 2, pp. 365-625 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN was published on 
989 


June 22, 


Works on Orchids 
from the Missouri Botanical Garden 


Orchids of Panama 

L. 0. Williams & P. H. Allen 

A facsimile of the Orchidaceae, Flora of Panama, which originally appeared in four separate issues of the Annals of the 
Missouri Botanical Garden from 1946 to 1949. A checklist of Panama's 800 species of orchids has been pue by 
R. L. Dressler and appears as an appendix. viii + 484 + xxvi pp., illustrated. 1980. $20.00. 


Icones Pleurothallidinarum I: Systematies of the Pleurothallidinae RE 

Carlyle A. Luer 

The 29 genera of this subtribe, which contains about 4,000 species, are discussed and illustrated. The introduction 

contains a detailed discussion of pleurothallid morphology and a key to the genera. 81 pp., color frontispiece, illustrated. 
$7.50. 


Icones Pleurothallidinarum Il: Systematics of Masdevallia (Orchidaceae) 

Carlyle A. Luer 

Masdevallia contains e 350 species. This overview is at the infragenetic. level and includes 29 pates eite a 
represen tati ach infrageneric taxon recognized. In Menit ut 


species listed. 63 pp. estates, color e: 1986. $6.5 


leones Pleurothallidinarum III: Systematics of Pleurothallis (Orchidaceae) 

Carlyle A. Luer 

This work provides a guide to be: genus Pleurothallis with keys to the dedit. keys to the sections and su bsections, 
and lists of representative species for each infrageneric taxon. A black and white illustration of a representative species 
of each genus, section, and subsection i is included. 109 pp., color fronsispiooë, illustrated. 1986. $8.50. 


Icones Pleurothallidinarum IV: Systematics of Acostaea, Condylago, | 

and Porroglossum erica: | 

Carlyle A, Luer | 

The 33 taxa included in these pleurothallid genera are net Apo riii. dor each has dora an red mole — 

labellum. In each, a stimulus by the pollinator causes the lip to rise quickly, temporarily trapping the pollinator between 
column, and petals. Escape requires the pollinator to brush past the stigma, r prelium, “a anther. giua 

illustrated; 1987. $8.00. 


Thesaurus Dueb 1&2 
Carlyle A. Luer with Rodrigo Escobar R. German translations by Pri» H 
Thesaurus Dracularum is a 80-odd illustrated : 
a popular m h of the genus Denel Sach. of the Lan seo is ated 
rd à watercolor painting accompanied. vd Uv deseripti riptions, discussions, and distributions in both English and German, with - 
stribution maps and black and white line nées In ing E format. Standing g orders yos. The first. two of 
six fascicles, $40.00 each, postpaid. AY 


Ih deu Tou ae A > a. 


ee ^. E X 3 E ; 
To place an order, send check or mo: 3: dd $1 50 for - 
ney order in U.S. funds, suite to U.S. bank; U. shipments a | 
bei EN and : 50 for each additional = non-U.S. shipments: add $2.50 for one book, and $.50 for each additional T 
hould be prepaid; a $1.00 fee will be added to orders requiring invoices. No ical Garden are made until rd 
Payment is Ecol Mail form with Y check or monet. en, Aaa Jr cur Botanical TRTE i F 2: 


> Deparment Eleven Missuri Botanical Garden 
i . p.0. Box 299- : 
loco AMETE DS y Louis, MO 63166:0299, USA. 
ase send the publications circled above to: .—. A Lae. 
Ne MÀ VERS ds. Dr D Pama en 


Name LIN 
de > AE Mate E AUCH ee LA Ae "e added tt ul. 


‘Postal Code. Gay 768) s * 


Css Ane Sean To Css Verr Noe 


CONTENTS 


Steyermark Recollections (No author—organized by G. Rogers) 
Plant Taxa Described by Julian A. Steyermark Mary Susan Taylor 
Systematics of Zapoteca (Leguminosae) Héctor M. Hernández 
Pollen Morphology of the Melanthiaceae and Its Systematic Implications CSS 
Takahashi & Shoichi Kawano 

A New Subgenus and Five New Species in Pantona 1 (Passifloraceae) from South 
America Linda K. Escobar xd 

A Revision of Mesoamerican Psychotria Subgenus Psychotria (Rubiaceae), Part III: Species 

... 48-61 and Dp te Clement W. Hamilton 
NOTES d Rice | 
Notes on Araceae boda S. Bunting AR 
Hosta jonesii (Liliaceae/Funkiaceae), a New Species from Korea Myong Gi Chung E : 
A New Species of AOUR (Apocynaceae) and Notes on the Genus Alwyn H. A 
Gentry 
Chromosome Number in Walleria (Tecophilaeaceae) Peter Goldblatt & pr a 


Manning : DRA T PATH TI 


Co L: xt JT 


New ; ies in TAE Barry Hammel ecco 
p New Species of Colnipis rise from the Venezuelan Guayana Leslie R. 
 Landrum hera 
Chromosome Number in Plenaogermun and Strelitzia and the fond Chromosome : 
s. Nerbini in Strelitziaceae (Zingiberal John C. Manning & Peter Goldblatt ..... 
E l is (Zamiaceae), a ae East Central African Species in the E. hilde- : 
‘brandi gren. | _ Aldo Moretti, Paolo De EAS Ane Pierre Sclavo & Dennis — 
AES Wm. Stevenson ........ —— 
; : A New Species of Persea (Lauraceae) from » Surinam, With a Discussion of Its P 


E . Within the Genus Henk van der : nner 
E Publications of Edgar Anderson: Additions | o Enden RE: 
| Book Review = “ei exe i e; 


Cover ilustracion. Solanum iiaracaD'Ary & Rekotrty, by Joho Myers: 


Annals 
ofthe | 
a 
Botani 


* AS 4 à 
wo ji f ; 
Vr ; E 
A N FK " 1 » 
f ; H. 
A I ; d 
| ail 
i 7; | te 
J a 41 
P \ » , 
\ j 3 
Ff 
LA f 
| XA N : 
| Le y y 
» 3j 4 / i 
f MWERA 
q } i f 
y. cum “El f i / 
4 ~ MOE FN I » 
| AE A y Ve PANA 
AA. oth «Qu Wa. was 
\ Í 
, Je OEA 
Vr : / x X 
i RES A: ‘ad 4 j Y 7 S 
ba BA c o E \ j 


shes ot ¡a P 
he l ? SS "s 
pa A NT 
R [EL 
H “3 
e t 
y kia 


Volume 76 
Number 4 


> KS 


Volume 76, Number 4 
Winter 1989 


Annals of the 
Missouri Botanical Garden 


The Annals, published quarterly, contains papers, primarily in systematic botany, con- 
tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the 
Garden will also be accepted. Authors should write the Editor for information concerning 
arrangements for publishing in the ANNALS. Instructions to Authors are printed in the back 


of the last issue of each volume. 


Editorial Committee 


- George K. Rogers 
Editor, Missouri Botanical andes 


Amy Scheuler 
Editorial Assistant, 
Missouri Botanical Garden 


Glenda Nau 
Magdalen Lampe 
Publications Staff 


Marshall R. Crosby 


Missouri Botanical Garden 


Gerrit Davidse 
Missouri Botanical Garden 


John D. Dwyer 
Missouri Botanical Garden & 
Saint Louis University 


Peter Goldblatt 


Missouri Botanical Garden 


Dale E. Johnson 


Missouri Botanical Garden 


Henk van der Werff 


Missouri Botanical Garden 


For subscription information contact Department 


. - Eleven, P.O. Box 299, St. Louis, MO 63166. Sub- 


= scription price is $75 per volume U.S., $80 Canada 
~ and Mexico, $100 all other countries. Four i issues 
= Pe volume. 


O Missouri Botanical Garden 1989 — 


The ANNALS or THE MISSOURI Bor EA 
(ISSN 0026-6493) is published quarterly Y 
Missouri Botanical Garden, 2345 Tower Grove 


. enue, St. Louis, MO 63110. Second class 


paid at St. Louis, MO and additional mailing 
POSTMASTER: Send address changes tO ANN. 
THE Missouri BOTANICAL GARDEN, Department 
Eleven, P.O. Box 299, St. Louis, MO 63166. 


NNALS 0f 


A A RRA 


PS X 


Volume 76 Annals 
Number 4 of the 
1989 Missouri 


Botanical 


Garden 


FLORA OF THE Julian A. Steyermarkt, Bruce K. Holst,? 
VENEZÜELAN CUAYANA==VIT. 8 tokitara 
CONTRIBUTIONS TO THE 

FLORA OF THE CERRO 

ARACAMUNI, VENEZUELA: 


NA 


ABSTRACT 


A preliminary checklist of the vascular plant flora of Cerro Aracamuni, an isolated sandstone massif of the Roraima 
Shield in southern Territorio Federal Amazonas, Venezuela, is presented. The list is based on collections from five 
P , 


ons 
made. The new ims include /lex aracamuniana (Aquifoliaceae); Philodendron roraimae subsp. aracamuniense 
(Araceae); Pochota aracamuniana (Bombacaceae); Navia carnevalii, N. iosothrix, N. patria, N. platyphylla 
terramarae (Bromeliaceae); Hexapterella steyermarkii (Burmanniaceae); Symbolanthus aracamuniensis cas 
naceae); Licaria trinervis (Lauraceae); Tococa liesneri (Melastomata ceae); Sauvagesia guianensis subsp. araca- 
muniensis (Ochnaceae); dices amunia liesneri (Orchidaceae); Piper mosaicum (Piperaceae); Rapatea aracamuniana 
(Rapateaceae); Cephalodendron aracamuniensis, Coussarea liesneri, Neblinathamnus aracamunianus, Remijia 
reducta (Rubiaceae); Bonnetia liesneri, Ternstroemia guanchezii, T. maguirei (Theaceae); Abolboda scabrida, 
Xyris aracamunae, X. liesneri, and X. prolificans (Xyridaceae). The three new combinations are: Paepalanthus 
sulcatus (Eriocaulaceae); Senefeldera yutajensis (Euphorbiaceae); and Persea perseiphylla (Lauraceae). 


>? 


Of the sandstone massifs of the Roraima Shield, Thus its neighbors to the north and northeast, Cerro 
Cerro de la Neblina, on the Venezuelan-Brazilian Avispa and Cerro Aracamuni, have emerged as 
border, to date has yielded the highest number of — especially interesting for botanical exploration. The 
endemic genera and species of vascular plants. first botanical collections from the summits of Cerro 


' We thank Paul Berry, Germán Carnevali, and George Rogers for carefully reviewing the manuscript, and Luther 
Raechal for assistance in preparing the text. John Myers prepared several of the plates. Part of this work was 


ipsia and Latin iria of Aracamunia were done by Bruno Manara. 
sc ni d n, P.O. Box 299, St. Louis, Missouri, 63166-0299, U.S.A. 
Ste ark died on October 15, 1988. For li on him and a record of much of his work see the 


J 
x 


Annals Ed 76. : 


ANN. Missouri Bor. GARD. 76: 945-992. 1989. 


946 


Annals of the 
Missouri Botanical Garden 


Avispa, and apparently from Cerro Aracamuni, 
were made by Felix Cardona Puig and G.C.K. 
& E. Dunsterville on a trip with the Venezuelan 
Border Commission in 1972. Coordinates used by 
them for what they called the ““Aracamuni section 
of Cerro Avispa” (01*30'N, 65?*51'W) match very 
closely with those from the most recent expedition 
to Cerro Aracamuni. Charles Brewer Carias gath- 
ered several plants during a stop to the summit by 
helicopter in 1983. Small gatherings of plants from 
the base and slopes of Aracamuni were made on 
two other occasions, in 1959 from a 900-m-tall 
granitic outlier of Cerro Aracamuni by John Wur- 
dack and Lincoln Adderley, and from an eastern 
escarpment of Aracamuni at 750 m by Otto Huber 
and Ernesto Medina in 1981. 

Cerro Aracamuni is the largest of the table 
mountains closest to Cerro de la Neblina, so ex- 
ploration of its summit was considered essential to 
help verify that the high degree of endemism re- 
ported from Neblina was not an artifact of incom- 
plete collecting. Although the summit of Cerro Ara- 
camuni attains an elevation of only 1,500 m, con- 
trasted with. 3,040 m on Neblina, most of the 
summit of Neblina lies between 1,800 and 2,200 m. 

In 1987, the Terramar Foundation of Venezuela 
provided Ronald Liesner of the Missouri Botanical 
Garden and Germán Carnevali and Francisco De- 
lascio of the Herbario Nacional de Venezuela the 
opportunity to explore the summit and slopes of 
Cerro Aracamuni between October 15 and No- 
vember 3, 1987. They gathered a total of 1,018 
collections (including some bryophytes and fungi, 
which are not considered in this paper) from the 
summit and slopes. 

Cerro Aracamuni lies near the southern tip of 
Territorio Federal Amazonas, Venezuela, in Dept 
Rio Negro, approximately 125-140 km northeast 
of Cerro de la Neblina (Fig. 1). It is a small to 
medium-sized massif for the Guayana region, rath- 
er long and narrow with a north-to-south orienta- 
tion, extending from 01?24'N, 65°47'W to 
01°32'N, 65?49'W. Vertical sandstone escarp- 
ments flank some of the sides, and a few rock 
outcrops appear on the upper edges. From its sum- 
mit, streams drain into the Matapire and Siapa 
rivers on the east, north, and south, and into the 
Rio Yatúa on the west. Several kilometers of low- 
land rainforest extend on the Rio Siapa side to the 
base of the massif. The large rapids, Raudal and 
Salto Gallineta, are prominent topographic features 
on the igneous outcrops along the Rio Siapa. Most 
of the massif is believed to be of Roraima sandstone. 

A close relationship with the flora of Cerro de 
la Neblina is expected on Cerro Aracamuni due to 


the proximity of the two massifs. Four of the genera 
previously known only from Cerro de la Neblina, 
Neblinanthera, Celiantha, Cephalodendron, and 
Neblinathamnus, are now known from Cerro Ara- 
camuni as well. Species (and subspecies) previously 
known only from Neblina, and now recorded from 
Aracamuni, are: Sobralia speciosa C. Schweinf., 
Endlicheria vinotincta Allen, Spathelia nebli- 
naensis R. Cowan & Briz., Diacidia glaucifolia 
(Maguire) W. R. Anderson, Philacra auriculata 
Dwyer, Tyleria silvana Maguire, Maieta nebli- 
nensis Wurd., Neblinanthera cumbrensis Wurd., 
Tococa pachystachya Wurd., Orthaea wurdackii 
Maguire, Steyerm. & Luteyn, Celiantha bella Ma- 
guire & Steyerm., Faramea boomii Steyerm., Fer- 
dinandusa neblinensis Steyerm., and Pagameop- 
sis maguirei subsp. neblinensis Steyerm. 

Two additions to the Venezuelan flora from the 
Aracamuni expeditions are Tyleria silvana Ma- 
guire, previously known from the Brazilian side of 
Cerro de la Neblina and Serra Pirapucü, and an 
as of yet unidentified species of the genus Jacques- 
huberia, previously known only from Amazonian 
Brazil and Colombia. 


VEGETATION OF THE SUMMIT 


The summit is covered mostly by 0.5—1-m-high, 
dense savanna (non-Poaceae) and low shrub vege- 
tation, and small forested areas occur in depres- 
sions and along streams. The savanna is inhabited 
mostly by Eriocaulaceae, Rapateaceae, Bromeli- 
aceae, Xyridaceae, Liliaceae, and Sarraceniaceae. 
Common shrubs and small trees of the savanna 
include Poecilandra retusa, Pagameopsis ma- 
guirei subsp. neblinensis, Bonnetia tristyla, Di- 
stictella monophylla, Emmotum glabrum, and 
Gongylolepis huachamacari. The patches of for- 
est and gallery forest include Neblinanthera cum- 
brensis, Aegiphila roraimensis, Clusia sp., Dia- 
cidia glaucifolia, Phyllanthus vaccintifolius, 
Psychotria duricoria, P. tapajozensis, and Ty- 
leria silvana. 

The remainder of this paper is divided into two 
sections, descriptions of new taxa and presentation 
of new combinations, and the preliminary checklist 
of the flora of Cerro Aracamuni. 


New TAXA AND COMBINATIONS 
FROM CERRO ARACAMUNI 
AQUIFOLIACEAE— contributed by Julian Steyermark 


Ilex aracamuniana Steyerm., sp. nov. TYPE: 
Venezuela. Territorio Federal Amazonas: Dept. 
Rio Negro, Cerro Aracamuni, summit, Proa 


Volume 76, Number 4 Steyermark, Holst & Collaborators 947 
1989 Venezuelan Guayana Flora— VII 
73 72 71 70 69 68 67 66 65 64 €3 62 61 eo 
puer: Er PUE | = an 1 4 4 -— eed 
12 7 | 1 | 12° 
EN d ` | 
H © c 2 11 
= Le 7 
ARE 
10 10 
9 3 9: 
8 je 
r Estado Bolivar hkz 
e 
C— + 5 e 
Territorio Federal 
Amazonas . 
| 4 
| Km e 
o 100 200 300 
= — 2 
a A 
| 1 
68 67 66 65 64 63 62 61 60 


FIGURE 1. 


Camp, medium height, semi-open forest, 
01°32'N, 65%49'W, 1,400 m, 26 Oct. 1987, 
R. Liesner & G. Carnevali 22473 (holotype, 
MO; isotype, VEN). Figure 2. 


r 6-10-metralis; foliorum laminis lanceolato- vel 


atropunctatis, decurrentibus, nervis lateralibus 
utroque latere 5-7 subtus impressis haud (warn ve- 
natione tertiaria tenuiter reticulatis; petiolis 6-12 m 


longis; apy asa foem mine a solitaria xillari lateralique; 
pedicellis foemineis matu ongis; calycis 
lobis breviter aioe ularibus apice acutis vel acuminatis; 
fructu subgloboso maturo 8-10 x 8-10 mm; preen 
quinque trigonis 6 mm longis dorsaliter unisulcatis 


ree 6-10 m tall, glabrous throughout. Leaf 
blades coriaceous, lance- to oblong-elliptic, obtusely 
acute at apex, acute at base, 6.5— cm long, 
(1.7-)2-3.5(-4) cm wide, dark punctate below, 
strongly decurrent on the petiole; midrib narrowly 
sulcate above, elevated below; lateral nerves 5-7 
each side, slightly sulcate above, impressed but not 
strongly conspicuous below, ascending at 45-50%, 
inconspicuously anastomosing 2-4 mm from the 
margin; tertiary venation subimpressed and grossly 


Map of Venezuela with détail of southern Territorio Federal Amazonas. 


reticulate above, finely reticulate below with im- 
pressed venation. Petioles 6-12 mm long, winged 
by the decurrent leaf bases. Pistillate inflorescence 
solitary, axillary and lateral; pedicels in post-an- 
thesis 7-11 mm long, 13-16 mm long in fruit. 
Calyx lobes in anthesis triangular, acuminate or 
slenderly acute, in fruit broadly ovate and abruptly 
acute at apex. Fruit subglobose at maturity, 8-10 
x 8-10 mm; pyrenes 5, 6 mm long, trigonal, 
unisulcate dorsally. Stigmatic lobes in post-anthesis 


prominent, discoid-capitate, 1 mm high, 1 mm 
diam. 
Paratypes. | VENEZUELA. TERRITORIO FEDERAL 


AMAZONAS: Dept. Rio Negro, Cerro Aracamuni, summit, 
Popa Camp, edge of tepui in ravine, 01?26'N, 65°42'W, 
1,550 m, 19 Oct. 1987, R. Liesner & F. Delascio 22181 
(MO, VEN); R. Liesner & F. Delascio 22060 (MO, 
VEN). 


This species is related to Ilex sipapoana Edwin 
and I. huachamacariana Edwin. It differs from 
the former in the longer mature pedicels; promi- 
nently decurrent leaf bases; and leaf blades with 
an obtusely acute apex, less conspicuous reticu- 
lation of the abaxial tertiary venation, and fainter 


948 Annals of the 
Missouri Botanical Garden 


uuo, 


uiuo. 


x30 

FIGURE 2. Ilex aracamuniana. Ws Habit. — B. Immature fruit. —C. Mature fruit. — D, E. pua stem. — F. 

eaf tip, upper surface. — G. Leaf tip, lower surface. — H. Detail of upper leaf surface. — I, J, K. Seed. — L. Detail 
of petiole. (A, C-L, from fon & bono 2247 3; B, from Liesner & Delascio 22181.) 
Rio Negro, Cerro Aracamuni, Proa Camp, in 
medium height semi-open forest, 01?32'N, 
65?49'W, 1,400 m, 26 Oct. 1987, R. Liesner 
& G. Carnevali 22510 (holotype, MO). Fig- 


ure 3. 


lateral nerves of the lower surface. The new species 
differs from /. huachamacariana by having longer 
petioles and broader leaf blades not retuse at the 


apex. 


ARACEAE— contributed by George Bunting A 1 typica foliorum lamina aliquantum minore 
Philodendron roraimae K. Krause subsp. ara- ee angustiore, petiolo tereti 
É : (lide aeria a abhorr 
camuniense Bunting, subsp. nov. TYPE: Ven- 


ezuela. Territorio Federal Amazonas: Dept. Climbing herb. Stem to 5 m long, terminal part 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 949 


Venezuelan Guayana Flora—VIl 


FIGURE 3. 


hidden by mass of coarsely fibrous cataphyll re- 
mains. Cataphyll broad, to 24 cm long, marcescent. 
Petiole stout, terete, 50-57(-68?) cm long in vivo 
with many lines of small gray dots. Leaf blade 
coriaceous though easily fracturing along veins, 
ovate (narrowly auk rok in juvenile shoots), 44— 
54 cm long, 2 cm wide -1.6 times 
longer than hu apex obtuse and abruptly acu- 
minate, 1-1.5 cm long; posterior lobes introrse, 
rounded at apex or somewhat angular but very 


Philodendron roraimae subsp. aracamuniense (Liesner & Carnevali 22510). 


blunt, slightly overlapping and enclosing a sub- 
rhombic space 5 cm long, or separated by a + 
mitriform sinus 14-18 cm long; margins undulate; 
midrib wide and prominent, primary lateral veins 
adaxially sulcate and abaxially raised, ca. 6-7 per 
side arising at angles of ca. 55-655; lesser veins 
all manifest and abaxially raised; posterior lobes 
with 2 primary lateral veins on each side of basal 
ribs, the latter nude in sinus for ca. 2-3.5 cm. 
Inflorescence solitary. Peduncle 4.5 cm long. Spathe 


Annals of the 


950 
Missouri Botanical Garden 

v " 

pU 

E [f 
o 
9 
N 

M 
FIGURE 4. Pochota aracamuniana. — A. Habit of sins —B. Androecium.— C. Staminal tube.— D, 
Anther.—G. Detail of style base, ovary summit. — H. Ovary and base of style. — I. Fruit. —J. Detail of lower bat 
» 


surface. (A-H, J, from Liesner & Carnevali 22559; I, fror Liesner & Carnevali 22577 


stout, 15-17 cm long, not constricted, outside with 
tube red and limb yellowish. Spadix 12.5-14.5 cm 
long, pistillate part 4 cm long and 2-2.5 cm thick, 
staminate part 8.5-10 cm long with sterile part to 
4.5 cm and fertile part to 5.5 cm long. Ovary 3- 
5-locular, locules each with 6-8 or more ovules 
apparently superposed from bottom to top. 

arat ue ME ELA. TERRITORIO FEDERAL 


Par 
AMAZONA o Negro, Cerro Aracamuni, Popa 
Camp, savanna ih. small to large patches of forest and 


stream, 01?26'N, 65?47'W, 1,550 m, 16 Oct. 1987, R. 
Liesner & F. Delascio 21956 (MO). 


The characters that separate the two subspecies 
of P. roraimae are few, yet the specimens seem 
quite distinct at first glance, chiefly because of the 
proportionally narrower leaf blade of subsp. ara- 
camuniense. Even the juvenile leaves of both taxa 
have a similar, narrow blade outline. An important 
difference may prove to be petiole shape, said to 
be terete in the Aracamuni collections; in the ma- 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 951 
| 


Venezuelan Guayana Flora— VII 


14mm 


FIGI . 
& Delasci io id 


terial of subsp. roraimae from the Gran Sabana, 
it is rounded abaxially, while the adaxial surface 
has a median, longitudinal, obtuse angle (ridge) and 
two plane faces. 

This is the common Philodendron growing on 
the summit of Cerro Aracamuni, climbing up tree 
trunks. Its relation to P. nebulense Bunting (de- 
scribed from Cerro de la Neblina) remains to be 
determined when fertile material of the latter be- 
comes available. 


BOMBACACEAE — contributed by Julian Steyermark 


Pochota aracamuniana Steyerm., sp. nov. TYPE: 

Venezuela. Territorio Federal Amazonas: Dept. 

Rio Negro, Cerro Aracamuni, summit, Proa 

amp, semi-open forest at edge of forest, 

01°32'N, 65°49'W, 1,400 m, 28 Oct. 1987, 

R. Liesner & G. Carnevali 22559 (holotype, 
MO; isotype, VEN). Figure 4. 


rbor 3-6-metralis; foliis (1-)3-foliolatis, foliolis ellip- 
tico-oblongis vel lanceolato-ellipticis apice obtusis vel sub- 


Navi xa carnevalit. —A. Habit. — B. Flower. 
2.) 


—C. Anther.— D. Sepal. — 


E. Floral bract. (From Liesner 


i Pri minute mucronatis e acutis vel subacutis, ma- 
joribus 6-10.5 cm longis 2-3.7 cm latis subtus 
atrolepidotis petiolis 2 m longis, petiolulis 4-8 
(713) mm longis; petalis 8-11 cm longis 7-15 mm latis; 

tubo staminali 10-11 mm longo praeter basin versus 
minute papillatum glabro; filamentis 6- s; stylo 
basi lepidoto parte inferiore 4.2 cm nan parte 
superiore 4 cm glabro 


Tree 3-6 m tall. Leaves (1 -)3-foliolate, leaflets 
coriaceous, elliptic-oblong to lance-elliptic, obtuse 
or subobtuse at apex with a minute mucro, acute 
to subacute at base, 6-10.5 cm long, 2-3.7 cm 
wide, densely dark brown lepidote beneath; midrib 
sulcate above, elevated below; lateral nerves im- 
pressed and manifest above, conspicuously im- 
pressed below, anastomosing 2-3 mm from margin; 
tertiary venation impressed above, subreticulate 
below. Petioles 1.5-2 cm long; petiolules 4-8(-13) 
mm long. Pedicel 2-3.5 cm long. Receptacle dark 
glandular. Calyx campanulate, 5-8 mm long (dried), 
to 12 mm (living), 10-12 mm wide at summit in 
anthesis, subundulate, minutely mucronate at sum- 


952 


Annals of the 
Missouri Botanical Garden 


mit, dark glandular-lepidote without, sericeous 
within. Petals linear-ligulate, 8-11 cm long, 7- 

mm wide, canescent-lepidote on both sides. Sta- 
mens ca. 110, in 5 principal phalanges. Staminal 
tube 10-11 mm long, glabrous except for minute 
papillation basally; filaments 6-7.5 cm long; an- 
thers 1-2.5 mm long. Style 8.2 cm long, sparsely 
dark glandular-lepidote in lower 4.2 cm, lepidote 
at very base, glabrous in the upper 4 cm. Ovary 
ovoid-deltoid, 4 x 3.5 mm, glabrous below, lepi- 
dote above. 
rounded at summit, 6 cm long, 2.5 cm wide at 


Fruit narrowly obovate, truncate- 


summit, minutely dark lepidote. 


Paratypes. | VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: Dept. Río Negro, Cerro Aracamuni, ro 
Proa Camp, edge of savanna, 01°32'N, 65?49'W, 1,400 
m, 28 um 1987 (ft), R. Liesner & G. Carnevali n 
(MO, VEN); Cerro Aracamuni, ue Popa Camp, sa- 
vanna at EN of forest, 01?26 °47'W, 1,550 m, 
19 Oct , R. Liesner & F. CMS 22174 (MO, 
V EN) 

This species is related to Pochota rob ynsii Stey- 
erm., but differs by having broader petals; generally 
obtuse, elliptic leaflets with a mucronulate apex; 
and conspicuous secondary nerves and tertiary ve- 
nation. 


BROMELIACEAE—contributed by Lyman B. Smith & 
Julian Steyermark 


Navia carnevalii Lyman B. Smith & Steyerm., 
sp. nov. TYPE: Venezuela. Territorio Federal 


987, R. Liesner 
& F. Delascio 22322 (holotype, US; isotypes, 
MO, VEN). Figure 5. 


N. umbratile Lyman B. Smith, cui valde affinis, 
on laminis subintegris glabris, bracteis pd an- 
guste oblongis vel oblongo-lanceolatis glabris differt 


Plant shortly caulescent, the stem variable in 
length, 1-6 cm long. Leaf sheaths brown, 1.5-2 
cm long, 5-10 mm wide, many ribbed. Leaf blades 
white at base, suberect, linear, attenuate, not con- 
tracted at base, coriaceous, 9-21 cm long, 3-5 
mm wide, glabrous above, 
uously lepidote below, margins thickened, subin- 


minutely inconspic- 


volute, subentire. Inflorescence 2-3-flowered, en- 
closed within the bracts, narrowly lanceolate, 2.5 
cm long, 2-3 cm wide. Outer bracts foliaceous, 
linear, to 6 cm long, 1-2 mm wide. Floral bracts 
membranous, narrowly oblong or oblong-lanceo- 
late, acuminate, about equaling the sepals, 13-14 
m wide. Sepals free, keeled, con- 
duplicate, lanceolate, 12-13 mm long, 3 mm wide, 


mm lon 


glabrous, obtusely narrowed to a slightly incurved, 
slightly mucronate tip. Corolla white, 20 mm long, 
with a narrow cylindric tube, 12 mm long, the 
lobes lanceolate, subacute, 6-7 mm long, 2-2.5 
mm wide. Ovary superior. 

The subentire leaves and the narrowly oblong 
or oblong-lanceolate, acuminate floral bracts dis- 
tinguish this species from N. umbratilis Lyman B. 
Smith of Cerro Yutaje. 


Navia lit d B. Smith. & Steyerm., 
: Venezuela. Territorio Federal 
egro, Cerro Araca- 
muni, riverbank in shade of forest, Quebrada 
Camp, in area of rapids flowing over laja, 
01?24'N, 65?38'N, 600 m, 20 Oct. 1987, R. 
Liesner & F. Delascio 22203 (holotype, US; 
isotypes, MO, VEN). Figure 6. 


speciebus ies adhuc i: a M Hae 
inflorescentiae parvi globosae aes eae pri- 
mariae valde re re a haud foliaceae, Fidis mini- 
morum lateralium verisimiliter o cucullatorum dif- 
fert 


Plant caulescent with a short thickened stem 5- 
7 cm long, in a dense spreading rosette. Leaf sheaths 
oblong-ovate, slightly broader than and merging 
into the lea 13-20 mm long, 13-15 mm 
wide, many-nerved. Leaf ge divaricately 
spreading, with a median stripe, linear, long-atten- 
uate, 15-23 cm long, 10-13 mm wide, minutely 


lades, 


serrulate throughout with teeth ca. 0.2 mm long, 
minutely and sparsely lepidote on both surfaces, 
especially beneath. Infructescence simple, sessile, 
subglobose-ovoid, many-flowered, 1.5-1.8 cm high, 
1.5-1.7 cm diam., ferruginous-tomentellose. Outer 
bracts few, not foliaceous, strongly reduced, 2.5- 
3 cm long, 4-5 mm wide, serrulate. Smaller lateral 
sepals apparently free, cucullate. Ovary superior. 
Capsule broadly ovate, 6.5 mm long; seeds blackish 
brown, 0.8 mm long, verruculose, costate, finely 
reticulate. 


The strongly reduced, nonfoliaceous primary 
bracts, together with the smaller, free, cucullate 
lateral sepals and ferruginous tomentose infructes- 
cence distinguish this species. The specific name 
refers to the rusty-colored hairs of the infructes- 
cence. 


Navia patria Lyman B. Smith & Steyerm., sp. 
nov. TYPE: Venezuela. Territorio Federal Ama- 
zonas: Dept. Rio Negro, Cerro Aracamuni, 
summit, Popa Camp, crevice of bluff at edge, 
savanna with small to large patches of forest 


and stream, 1?26'N, 65%47'W, 1,550 m, 17 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora— VII 


6.5mm 


Fic Navia 2m — A. Habit. - 
Liesner ei yos 22203.) 


Oct. 1987, R. Liesner & F. Delascio 22041 
(holotype, US; isotypes, MO, VEN). Figure 7. 
crispa Lyman B. Smith, cui valde affinis, foliorum 


laminis haud crispatis, bracteis primariis paulo reductis 
basi pallidis, bracteis florigeris acutis, multo majoribus 


Plant in dense spreading rosette, subacaules- 
cent. Leaf sheaths brown, membranous, 15-17 
mm long, 15-17 mm wide at base. Leaf blades 
widely spreading, with a broad white median stripe, 
linear, acuminate, 11-16 cm long, 8- 10 mm wide, 


- B. Fruit, lateral view. — C. Fruit, view from above. - 


- D, E. Seed. (From 


densely serrulate, not crisped on margins, both 
surfaces pilosulous, especially beneath. Inflores- 
cence sessile, subglobose, M M bipinnate from 
many short spikes, 2.5 c , 3.0-4 cm diam. 
Outer bracts foliaceous, edis de reduced, sparse- 
ly puberulous without, strongly carinate, entire. 
Floral bracts broadly lanceolate, acute, somewhat 
longer than the sepals, 15 mm long, 5 mm wide 
below the middle, entire, glabrous. Sepals lanceo- 
late, acuminate, the outer two carinate, 13 mm 
long, ca. 3 mm wide, entire, glabrous. Corolla white, 
10 mm long. Ovary superior. 


954 Annals of the 
Missouri Botanical Garden 
E 
E 
e 
B | I D 
e 
FIGURE 7. Navia patria. — A. Habit. — B. Sepal. — C. Floral bract. — D. Leaf. (From Liesner & Delascio 22041.) 


This species differs from N. crispa Lyman B. 
Smith in having the outer primary bracts pale and 
only slightly reduced basally, the floral bracts larg- 
er, and the leaf margins noncrispate. 


Navia platyphylla Lyman B. Smith & Steyerm., 
sp. nov. TYPE: Venezuela. Territorio Federal 
Amazonas: Dept. Rio Negro, Cerro Araca- 
muni, summit, Popa Camp, steep slope of ra- 
vine forest at edge of tepui, 01?26'N, 65°47'W, 
1,550 m, 17 Oct. 1987, R. Liesner & F. 
Delascio 22062 (holotype, US; isotypes, MO, 
US). Figure 8. 


AN - diffusa Lyman B. Smith, cui valde affinis, foliis 
latioribus spinis longioribus, inflorescentia meia innata 
; floribus juvenilibus et alteris reflexis diffe 


Terrestrial plant, 2 m tall when fruiting. Sheaths 
not seen. Leaf blades coriaceous, linear-ligulate, 
attenuate, 1.3-1.5 m long, 4-4.5 cm wide, strong- 
ly spinose in the lower 14-1% with black, broad- 
based spines 3 mm long, obscurely and sparsely 
lepidote beneath, mostly glabrous above. Inflores- 
cence laxly 4-pinnate, 2 m tall, longer axes 9-14 
cm long, the ultimate axes 3-11 cm long, glabrous, 
the flowers 3-4 mm apart. Primary bracts broadly 
triangular, long-attenuate, shorter than the sterile 


Volume 76, Number 4 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora— VII 


FIGURE 8. Navia platyphylla. — A. Leaf and portion of inflorescence. —B. Flowers, past anthesis. — C. Floral 
bract. — D. Sepal. — E. Pistil. (From Liesner & Delascio 22062.) 


bases of the main axes, 3.5-4.5 cm long, scarious 
margined. Bracts subtending the secondary axes 
broadly ovate, abruptly attenuate, 3-7 mm long. 
racts broadly suborbi- 
cular-deltoid, subamplexicual, acuminate, 3-5 mm 
long, about % length of the sepals. Sepals oblong- 
ovate or ovate, subobtuse, 4-7 mm long, the pos- 
terior strongly carinate. Petals (past anthesis) 9 
m long. Ovary superior, ovoid, 4 mm long. 


This species differs from N. diffusa by having 


reflexed flowers (even when young), larger sepals, 


and much broader leaves with longer, stouter spines. 
The four-pinnate inflorescences are also charac- 
teristic. It is distinguished from N. hechtioides 
Lyman B. Smith by having shorter sepals, the 
posterior ones nonalate. From NV. brocchinioides 
Lyman B. Smith, it is differentiated by the non- 
digitate, solitary, shorter spikes. 


Navia terramarae Lyman B. Smith & Steyerm., 
sp. nov. TYPE: Venezuela. Territorio Federal 
Amazonas: Dept. Rio Negro, Cerro Araca- 


956 


Annals of th 
Missouri Pu" Garden 


| | 
AAS 
WN SN, | S á 


ANS 


D 


FIGURE 9. Navia terramarae. —A. Habit. — B. 
Fruit. —C. Sepal.— D. Floral bract. (From Liesner & 
Delascio 21998, 22599.) 


muni, summit, Popa Camp, savanna with small 
to large patches of forest and stream, stream 
bank, 01?26'N, 65?47'W, 1,550 m, 16 Oct. 
1987, R. Liesner & F. Delascio 21998 (ho- 
lotype, US; isotypes, MO, VEN). Figure 9. 


A. N. liesneri Lyman B. Smith, Steyerm., & H. Rob- 


inson, cui valde affinis, foliorum laminis interioribus ad 


o 
_ obscure. serrulatis laxe minuteque obscure serru- 
latis, et ab ambabus caulibus SA ramosis subter- 
raneis, foliis or omnino Pai differt 


Caulescent plant with elongated, simple or di- 
chotomously branched stems 10-20 cm long, 0.7- 
cm wide. Leaf sheaths brown, membranous, 
2-6 mm wide, striate. Leaves crowded in a dense 
rosette, the older ones strongly reflexed, the interior 
leaves white basally; leaf blades narrowly linear- 
triangular, 4.5-6 cm long, 1.5-2 mm wide, gla- 
brous, terminating in a slender apiculum, laxly and 
minutely spinulose, the spinules ascending, 0.2— 
0.25 mm long, 1-1.5 mm apart. Inflorescence 


simple, sessile, glomerate. Outer bracts linear, 1.7— 
m long, minutely serrulate. Floral bract uni- 
formly membranous, ca. 6-striate, broadly oblong- 
lanceolate, acute, 10-12 mm long, 5.5 mm wide 
at base. Sepals free, linear-oblanceolate, obtusely 
acute, 8.5-9 mm long, 1-2 mm wide. Ovary su- 
perior. Capsule lance-oblong, slightly narrowed at 
summit, 9 mm long, 3 mm wide at middle 


This species is closely related to the recently 
described Navia liesneri and N. delascionis. The 
interior leaves are white at the base as in N. 
lascionis, but the more conspicuous teeth resemble 
those of V. liesneri. The leaf blades are narrower 
than in either N. liesneri or N. delascionis. The 
lower margins of the leaf blades are shortly white 


cose-ciliate, wher 
blades are completely glabrous. 


Par itype. VENE EZUELA. TERRITORIO , FEDERAL 


87, R. Liesner & G. Carnevali 22599 
(MO. US. d 


Th species is named for the Terramar 
amdanon of Venezuela in recognition of the in- 
valuable assistance provided on the expedition to 
Cerro Aracamuni. 


BURMANNIACEAE— contributed by Paul J. M. & H. 
aas 


Studying the material available of Burmanni- 
aceae for Steyermark's Flora of the Venezuelan 
Guayana, we came across an undescribed species 
of the saprophytic genus Hexapterella. We name 
it after Julian Steyermark to honor his lifelong 
dedication to the Venezuelan flora. 


Hexapterella steyermarkii P. Maas & H. Maas, 
sp. nov. TYPE: Venezuela. Territorio Federal 
Amazonas: Dept. Rio Negro, Cerro Araca- 
muni, summit, Proa Camp, 01?32'N, 
65?49' W, 1,400 m, 29 Oct. 1987 (fl, fr), R. 
Liesner & G. Carnevali 22622 (holotype, 
MO; isotype, U). Figure 10. 


Herba saprophytica. Flores hypocrateriformes sex-cos- 


aque dissimilia. Antherae sessiles 
tubum floralem non excedentes 

Saprophytic herbs 12-21 cm high. Rhizomes 
unknown. Stems lavender, unbranched, or some- 
times branched at the base of the plant, upper part 
of stem and inflorescence papillate (papillae ca. 
0.02 mm long). Leaves narrowly triangular to tri- 


Volume 76, Number 4 Steyermark, Holst & Collaborators 957 
1989 Venezuelan Guayana Flora— VII 


FIGURE 10. Hexapterella steyermarkii. —A, B. Habit. 
Lea 


—F Inner side of flower showing outer and inner tepals, and stamens. — C. Detail of stamen. — 


+ outer tepal 


inner tepal ( i 


ww p'O 


C. Old flower with dehiscing capsule. — D. ag 
d. (B, 


from Maguire et al. 42251; G, from Steyermark 105202; A, C, D, E, F, and H, from Liesner & Carman 2 ) 


angular-ovate, 2-4 mm long, 0.8-2 mm wide, base 
auriculate, apex acuminate. Inflorescence a bifur- 
cate or single cyme, cincinni 2-6-flowered, 8-25 
mm long, flowers 1.8-11 mm apart, or plant hav- 
ing a solitary, terminal flower. Bracts narrowly 


triangular to triangular-ovate, 1.8-3.5 mm long, 
0.6-1.9 mm wide, base often auriculate, apex acu- 
minate. Pedicels 2-4(-5.5) mm long. Flowers pale 
purple to lavender, or white with purple to lavender 


lobes, (4.5-)6.5-9.5(-11) mm long. Tepals per- 


958 


Annals of the 
Missouri Botanical Garden 


sistent, papillate on both sides. Outer tepals 3, 
narrowly triangular(-ovate), 3.0-4.3 mm long, 
0.8-1.4 mm wide, margins involute. Inner tepals 
3, obcordate to spathulate, 0.6-1.3 mm long, 0.5- 
mm wide. Floral tube 2.5-4.9 mm long, 0.8- 
1.5 mm diam., 6-ribbed. Thecae sessile. Style 3.5- 
4.6 mm long, branches 0.6-0.9 mm long. Ovary 
(transversely) obovoid-ellipsoid, 1.2-2.7 mm long, 
-2.5 mm diam., upper part 1-locular, basal part 
3-locular. Capsule globose to transversely broadly 
ellipsoid, 1.5-3.2 mm long, 2-3.5 mm diam., 
crowned by the persistent perianth, dehiscing by 
transverse slits and irregularly withering of the wall 
between the ribs. Seeds irregularly globose to ovoid, 
0.3-0.5 x 0.2-0.3 mm 


Distribution. Known only from the Venezue- 
lan Guayana (Territorio Federal Amazonas and Es- 
tado Bolivar); (470-)1,100-1,800 m; often along 
rivers or on dry rock ledges. 


P VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: Cerro de la Neblina, Rio Yatúa, slopes of 
Cañon Grande below Cumbre Camp, 1,100-1,800 m, 26 
Nov. 1957 (fl, fr), B. Maguire et al. dn (NY); Cumbre 
del Cerro Autana, 1,2 ; , 21-22 Sep. 1971 
(fl, fr), J. Steyermark 105202 (UL. VEN). BOLIVAR: Chi- 
manta Massif, along Rio Tirica (Rio Apapurén) just above 

echiné-Merü, 410 m, 16 Jan. 1955, J. Steyermark & 
J. Wurdack 113 (VEN). 


Hexapterella steyermarkii is only the second 
species described in this genus. The other species, 
H. gentianoides Urban, is a fairly common sap- 
rophyte in Guyana and Surinam, and has been 
collected once in Venezuela—at Cerro de la Ne- 
blina, between 1,100 and 1,800 m. Hexapterella 
steyermarkii is known from three collections in 
Territorio Federal Amazonas (Cerros Aracamuni, 
Neblina, and Autana) and one from Estado Bolivar 


(Chimantá Massif). 


The two species of Hexapterella can be distin- 
guished by the following key: 
la. Anthers on long filaments slightly exceeding the 
floral tube; upper part of perianth caducous; 
inner tepals narrowly ovate-triangular 
. gentianoides 
lb. Anthers sessile, not exceeding the floral a 
perianth persistent on fruit; inner ug obco 
date oui 


ERIOCAULACEAE—- contributed by Nancy Hensold 


Paepalanthus sulcatus Hensold, nom. et stat. 
nov. Paepalanthus stegolepoides var. acu- 


talis Moldenke, Phytologia 15: 463. 1968. 


Selected specimens examined. VENEZUELA. TERRI- 
TORIO FEDERAL AMAZONAS: Dept. Rio Negro, Cerro Ara- 
camuni, summit, Popa Camp, O1°26'N, 65°47'W, 1,550 


m, 15 Oct. 1987, R. Liesner & F. Delascio 21940 (MO, 
VEN). 


EUPHORBIACEAE—contributed by Grady Webster 


Senefeldera yutajensis (Jabl.) Webster, comb. 
nov. Sapium yutajense Jabl., Mem. New York 
Bot. Gard. 17: 184. 1967, fig. 24. TYPE: 
Venezuela. Territorio Federal Amazonas: Cano 
Yutaje, B. Maguire & C. Maguire 35261 
(holotype, NY). 


Jablonski noted that the leaf venation and inflo- 
rescence of this species suggested Senefeldera, but 
e placed it in Sapium because of the staminate 
flower with a two-lobed calyx and two connate 
stamens. However, the branched inflorescence, ca- 
ducous stipules, and especially the non-arillate seeds 
of the type collection are quite incompatible with 
Sapium. The small stamen number is shared with 
Senefelderopsis, but that genus has unbranched 
inflorescences and leaves with the veinlets not 
prominulous. Surprisingly, there is a resemblance 
to Mabea, especially to Mabea pohliana (Benth.) 
Muell. Arg. in sect. Apodae Pax & Hoffman 
(Pflanzenr. 147. V: 39. 1912). That species of 
Mabea has a branched inflorescence, an androe- 
cium of only two or three stamens, and an indu- 
mentum of dendritic hairs. Yet Mabea pohliana 
differs by having serrulate, short-petiolate leaves 
without prominulous venation, discrete staminate 
sepals and stamens, pedicellate pistillate flowers, 
and an elongated stylar column. In its subsessile 
pistillate flowers, elongated stylar column, and en- 
tire leaves with prominulous venation, Senefeldera 
yutajensis agrees with other species of Senefeld- 
era, and this seems the best generic placement 
despite the indumentum of dendritic hairs. Along 
with Senefeldera triandra Pax & Hoffman (Pflanz- 
enr. 147. XIV: 55. 1919) and possibly Mabea 
paraguensis Muell. Arg. (Fl. Brasil 11(2): 527 
1874), Senefeldera yutajensis may represent a 
phyletic branch intermediate between the genera 


Senefeldera and Mabea 


Additional Ta examined. VENEZUELA. TERRI- 
TORIO FEDERAL A : Dept. Rio Negro, Cerro Ara- 
camuni, ue Proa oe 01932: N, 65?49' W, 1,400 
m, 29 Oct. 1987, R. Liesner & G. Carnevali 22625 
(DAV, MO, VEN). 


GENTIANACEAE— contributed by Julian Steyermark 


Symbolanthus aracamuniensis Steyerm., sp. 
nov. TYPE: Venezuela. Territorio Federal Ama- 
zonas: Dept. Rio Negro, Cerro Aracamuni, 

mmit, Popa Camp, savanna, 01?26'N, 
65°47'W, 1,550 m, 17 Oct. 1987, R. Liesner 


Volume 76, Number 4 


Steyermark, Holst & Collaborators 959 


1989 Venezuelan Guayana Flora— VII 
C 
FIGURE 11. Symbolanthus aracamuniensis. —A. Habit of branch. — B, C. Anther. — D. Stamen. — E. Pistil. — 
F. Seed. —G. Cross section of stem. (From Liesner & Delascio 22079.) 


& F. Delascio 22079 (holotype, MO; isotype, 
VEN). Figure 11. 


. huachamacarensis et S. yaviensis calyce valde 
reducto, 7-8 mm longo, lobis 5-6 mm longis differt. 


Glabrous herb, 0.3-0.8 m tall. Stem 4-angled, 
3-3.5 mm diam. Leaves short-petiolate, blades 
lance- to oblong-elliptic, the apex acute, the de- 
current base acute to cuneate, silvery abaxially, 
the larger lower and middle blades 4—6.5 cm long, 
1.5-2.2 cm wide, faintly plinerved above, obso- 
2-7 mm long. Inflo- 
rescence terminal, cymosely 3-4-flowered; pedicel 
slender, 0.5-2 cm long. Calyx 7-8 mm long, 8- 


mm wide, 5-lobed; lobes ovate, obtuse, nearly 


letely nerved below; petiole 


divided to the base, 5-6 mm long, 4 mm wide, 
scarious-margined. Corolla lavender-pink, pink, or 
whitish, infundibuliform, 5-5.5 cm long; tube 3- 
3.8 cm long, 3-3.5 
wide at summit; lobes 5, broadly ovate, 2.2-2.5 
cm long, 2-2.2 cm wide, broadly curved above to 


mm wide at base, 2-3 cm 


an obtuse mucronulate apex. Stamens 5, subequal, 
as high as or higher than the orifice of the corolla. 
Filaments coiling, attached to corolla tube 5 mm 
above its base, 4 cm long; anthers suboblong, dor- 
sifixed, lance-oblong, sagittate at base, narrowed 


with a dorsal basal enlargement. Capsule oblong, 
13 mm long, 6 mm wide. Seeds quadrangular, 


960 


Annals of the 
Missouri Botanical Garden 


Uu 
OR Y 
( y! 4 


UN uU 
NU Wl TRU ITA 
A IO 


\ 
E 
tas 


à 


1 MM 
EN 
At. 


P 10977 
A Tix Mie 2 


"HH Lett? 
UTIMS: 


FIGURE 12. Licaria trinervis. — A. Habit of branch. — B. Flowers. — C. Part of inflorescence. — D. Leaf. — E, 
Ventral and dorsal views of stamen.—G. Ovary.—H, I. Ventral and dorsal views of tepal.—J. Staminode. (From 


Liesner & Delascio 22161.) 


broader than long, 0.2 mm long, 0.3 mm wide, 
depressed at the summit, the surface plane with a 
microscopic cellular appearance. 


This species differs from 5. yaviensis Steyerm. 
and S. huachamarensis Steyerm. in its much 
shorter calyx. Other distinctive characters of the 
new species are the anthers with revolute, aristate 
tips and a dorsal enlargement at the junction of 
the anther with the filament. 


LAURACEAE— contributed by Henk van der Werff 


AY 
ATAR 
SANS AR 
3 
Sx 


BEALS P 
EY Tr 

R = hh 
pe EE. D 

S NI > = 


"d Ns 
Assen 77 


TA VIVAS 
27 Te 


NON; 
so 
x 
es 
SS 


RS 


EX DC ao 
xo EVA 
RS 

SS 


Y INS 
AN - 


Y 


F. 


Licaria trinervis van der Werff, sp. nov. TYPE: 
Venezuela. Territorio Federal Amazonas: Dept. 
Rio Negro, summit of Cerro Aracamuni, 
01?26'N, 65°47'W, 1,550 m, 19 Oct. 1987, 
R. Liesner & F. Delascio 22161 (holotype, 
MO; isotypes, HBC, NY, VEN). Figure 12. 


Frutex vel arbor parva. Ramuli teretes, juvenales fusco- 
tomentosi, velustiores glabri. Folia alterna, 4-10 x 1. 
3 cm, anguste elliptica, fortiter chartacea, basi apiceque 
acuta, super glabra (juvenalissima sparse tomentosa), sub- 
tus fusco-tomentosa (vetera glabrescentia). Nervi utroque 
costae latere 4-5, marginem versus arcuati et connati; 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora— VII 


super immersi, subtus elevati, preci pue costa et nervi 
basales. Petioli 0.5-1 cm, fusco-tomentosi, glabrescentes. 
snip AFA axillares, fusco-tomentosae, paniculatae, 
folis breviores, 2-3 c uds ongae. Pedicelli ad 1 mm longi. 
Flores fusco- ionis ntosi, ,ca. l.5 mm longi, 
ca. 2.2 mm lati; stamina dere tan obtecti, solum api- 
cibus expositis. Tepala 6, aequalia, fusco-tomentosa, in- 
curvata, ca. |l mm jw a et 1.2 mm lata. Stamina 3, ca. 
1 mm longa, 2- eue filamentis "liberis. pubescentibus, 
basi 2 glandulis praeditis; locellis parvis, fere apicalibus. 
Staminodia 6, 0.5-0.6 mm longa, basibus pubescentia. 
Ovarium glabrum, dus. ca. 0.6 mm longum; stylo 
gracili, ca. 0.6 mm longo. Fructus ignotus. 


Shrubs or small trees, to 5 m tall. Twigs terete, 
rather slender, brown tomentose when young, gla- 
brescent with age. Leaves alternate, 4-10 x 
3 cm, narrowly elliptic, evenly distributed along 
the twigs, base acute, apex acuminate, firmly char- 
taceous, upper surface glabrous or loosely tomen- 
tose along the midrib when very young, lower sur- 
young, 
wearing off with age and the oldest leaves occa- 
sionally glabrous; lateral veins 4—5 pairs, the basal 
pair much more strongly developed than the others, 
all laterals arching upwards and becoming loop- 
connected; veins immersed above, raised below (es- 
pecially the midvein and the basal laterals). Petioles 

.9-1 cm long, flattened above, brown tomentose 


face brown tomentose when tomentum 


when young, glabrescent with age. Inflorescence 
in axils of leaves or deciduous bracts, brown to- 
mentose, paniculate, shorter than subtending leaves, 
2-3 cm long. Pedicels to 1 mm long, brown to- 
mentose. Flowers brown tomentose, depressed-glo- 
bose, 1.5 mm long, ca. 2.2 mm wide, the stamens 
nearly enclosed by the tepals with only their tips 
exposed. Tepals 6, equal, incurved, ca. 1 mm long, 
1.2 mm wide, brown tomentose. Stamens 3, ca. 1 
mm long; the filaments free, with 2 glands near 
the base, pubescent, about as wide as the anthers; 
anther cells small, ar T opening towards 
the tip of the anther. ia 6, representing 
the two outer whorls, ca. ries m long, their 
bases pubescent. Ovary glabrous, ellipsoid, ca. 0.6 
mm long; style slender, ca. 0.6 mm long. Fruit 
unknown 


atypes. | VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: Dept. Río Negro, summit of Cerro Aracamuni, 
Proa Camp, 01*32'N, 65%49"W, 1,400 m, 25 Oct. 1987, 
R. Liesner & G. Carnevali 22451 (MO, VEN), 2 Nov. 
1987, R. Liesner & G. Carnevali 227 14 (MO, VEN). 
Licaria trinervis belongs to subg. Licaria, as 
defined by Kurz (1983). It does not seem to have 
close relatives within this subgenus; among its strik- 
ing characters are the brown tomentose pubescence 
on the young parts, depressed-globose flower shape, 
strongly developed basal pair of lateral veins on 
the leaves. Another distinction is that no other 


Licaria species in the Venezuelan Guayana is known 
from the tepui summits, although L. macrophylla 
(A.C. Smith) Kostermans is known from slopes of 
Cerro Huachamacari at 600 m, L. tomentosa van 
der Werff occurs on the slopes of the Chimantá 


Massif. 


Persea perseiphylla (Allen) van der Werff, comb. 
nov. Ocotea perseiphylla Allen, Mem. New 
York Bot. Gard. 10: 109. 1964 


This species was known to Allen only from the 
holotype (Schultes & Lopez 10103, US), a spec- 
imen in young bud. Characters necessary for cor- 
rect generic placement were not discernible at this 
young stage, and Allen mistakenly placed the species 
in Ocotea. Fruiting specimens show clearly that 
this species belongs in Persea. Inspection of the 
holotype also showed that the tepals of the buds 
are unequal, with the outer three shorter than the 
inner ones, a character found in most neotropical 
Persea species but never in Ocotea. Hence Ocotea 
perseiphylla is transferred to Persea. Because 
open flowers have not yet been collected, P. per- 
seiphylla cannot be placed in any of the sections 
of subg. Eriodaphne Nees; the general resemblance 
to P. maguirei L. Kopp and P. croatii van der 
Werff suggests that it belongs in sect. Aurataea 
Kopp. It has nine four-celled stamens and therefore 
does not belong to sect. Hexanthera Kopp. 

Additional 21: ns examined. | VENEZUELA. 
TERRITORIO FEDERAL AMAZONAS: Dept. Rio N 


Aracamuni ind opa p, 01?26'N, 65°47'W 
1,550 m, 18 Oct. 1987 (fr), " Liesner & F. Delascio 
22099 ); 19 Oct. 1987 (fr), R. Liesner De 


lascio 22169 (MO); ip sur de la Sierra de Unturán, 
cabeceras del Rio Mavaca, 01°32'N, 65?11'W, 550-650 
m, 26 Feb. 1981 (fr), F. Fais 809 (MO, TFAV). 


MELASTOMATACEAE— contributed by John J. Wur- 
dack 


Tococa liesneri Wurd., sp. nov. TYPE: Vene- 
zuela. Territorio Federal Amazonas: Dept. Rio 
Negro, Cerro Aracamuni, summit, Proa Camp, 
savanna with small to large patches of forest, 
"d 65°49'W, 1,400 m, 1 Nov. 1987, 

. Liesner & G. Carnevali 22700 (holotype, 

T 502486; isotype, US). 


T. obovata Gleason affinis, foliorum laminis (3-nervatis) 
et petiolis minoribus, calycis m exterioribus vix em- 
inentibus, petalis wee dn differ 


Shrub 0.4-0.5 m tall, the terete branchlets mod- 
erately puberulous with dendritic hairs ca. 0.2 mm 
long and sparsely glandular-setose with smooth hairs 
1-2 mm long. Petioles 0.5-1 mm long; blade 1.5- 


962 


Annals of the 
Missouri Botanical Garden 


2.5 x 0.8-1.4 cm, ovate-oblong, apex rounded- 
acute, base cordulate, subcoriaceous and entire, 
ciliate with loosely appressed and in part gland- 
tipped hairs, above sparsely loose-strigose with 
smooth hairs 1.5-2.5 mm long, 
loose-strigulose with gland-tipped hairs on the pri- 


below sparsely 


mary veins and very sparsely puberulous with short 
dendritic hairs but esetulose on the surface, 3-nerved 
(excluding the tenuous incomplete inframarginals), 
with obscure lax venule reticulation. Flowers 5- 
merous, solitary or ternate at branchlet ends, ped- 
icels 2-7 mm long and (as the hypanthium) mod- 
erately dendritic-puberulous and glandular-setu- 
lose, the persistent bracteoles 2-2.5 x 0.7 mm 
and usually inserted ca. 0.5 mm below the ca. 5- 
5.4-mm-long hypanthium; calyx tube ca. 0.6 mm 
long, the oblong interior lobes ca. 3-3.5 x 2 mm, 
the glandular-setulose external teeth not or barely 
projecting (to 0.2 mm); torus glabrous within. Pet- 
als 10-11 x 8-9 mm, obovate-oblong with round- 
ed-truncate apex, pink and glabrous. Stamens es- 
sentially isomorphic, glabrous; filaments ca. 6.5- 
7 mm long; anthers ca. 5 x 0.6 x 
subulate with ventrally inclined pore and with short 
blunt-basal spur. Stigma not expanded; style ca. 
11-12 x 0. -celled and ca. 3 


mm inferior, the truncate glabrous apical cone 1.4- 


0.5 mm, oblong- 


o mm; ovary 


1.5 mm long. 


Paratype. VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: Dept. Río Negro, Cerro Aracamuni, summit, 
Popa pes savanna, 01?26'N, 65?47'W, 1,550 m, 16 
Oct. 1 , R. Liesner & F. Delascio 22007 (MO, US). 


Tococa obovata has distinctly five-nerved leaf 
blades measuring mostly 4-8 x 2-5 cm, petioles 
0.3-0.7 cm long, external cayx teeth projecting 
1-2 mm, and petals 15-21 
calyx lobes are similarly Fo and both species 
have glabrous petals. In general vegetative aspect, 


7-9 mm; its interior 


T. liesneri resembles the typical subspecies of T. 
bolivarensis Gleason, which has broadly deltoid 
calyx lobes ca. 0.5 mm long and densely pruinose- 
granulose petals. The collection Pires & Marinho 
15708 (US, in fruit, from Brazil. Amazonas: alto 
Rio Negro, Sào Gabriel da Cachoeira) bears a strong 
resemblance to T. liesneri except for completely 
eglandular vegetative setae. 


OCHNACEAE— contributed by Julian Steyermark 


Sauvagesia guianensis (Eichl.) Sastre Bd 
aracamuniensis Steyerm., subsp. nov. TYPE: 
Venezuela. Territorio Federal Paid Dept. 
Rio Negro, Cerro Aracamuni, summit, Popa 

amp, savanna, 01?26'N, 65°47'W, 1,550 
m, 17 Oct. 1987, R. Liesner & F. Delascio 
22075 (holotype, MO; isotype, VEN). 


A subsp. guianense Sip aa laminis suborbicu 
laribus apice late rotundatis, foliorum glandulis kapis 
alibus utroque latere 1-3 adpressis recedit: a su uai- 
quinimensis petalis apice rotundatis, rl lami 
a subsp. sipapoensis atque subspeciebus alus foliorum 
glandulis marginalibus paucis differt. 


Subligneous plant 0.5-1 m tall. Leaves linear- 
oblanceolate, 9-10 mm long, 1.8-2 mm wide, 
remotely appressed serrulate with 1-3 appressed 
glandular teeth on uppermost 1.5-2 mm of mar- 
gins, obtusely acute at apex with a blunt, glandular 
protuberance; lateral nerve depressions obliquely 
ascending. Pedicels 2 mm long, flowers hidden in 
the upper leaves. Petals pink, obovate, rounded at 
apex, 6.5 mm long, 3.8-4 mm wide at the middle. 
Staminodia 9-10 mm long, the lamina suborbi- 
cular, 0.3-0.5 x 0.3-0.5 mm, stipitate portion 
0.6-0.7 mm long. 


This subspecies is distinguished from subsp. gui- 
anense by having the leaf margins remotely ap- 
pressed serrulate with 1—3 glandular teeth on each 
side of the uppermost 1.5-2 mm of the margins, 
and the leaf apex with a blunt, glandular protu- 
berance. From subsp. sipapoensis Steyerm. it dif- 
fers by the relatively few appressed glandular teeth 
on each of the uppermost margins and the subor- 
bicular rounded lamina of the staminode. It differs 
from subsp. guaiquinimensis Steyerm., which has 
a similar suborbicular rounded lamina, in the 
rounded petals. 


ORCHIDACEAE— contributed by Germán Carnevali 
& Ivon Ramirez 


The generic revision of the Spiranthinae by L.A. 
Garay (1982) formed the basis for a better un- 
derstanding of the spiranthoid complex. A reeval- 
uation of this group for a treatment of the Orchi- 
daceae from the Venezuelan Guayana has revealed 
the presence of a very distinctive new species that 
cannot be assigned to any of the genera delimited 
in Garay's revision. 


Aracamunia liesneri Carnevali & I. Ramirez, 
gen. et sp. nov. TYPE: Venezuela. Territorio 
Federal Amazonas: Dept. Rio Negro, Cerro 
Aracamuni, summit, Popa Camp, on moss cov- 

ered streambank in almost closed forest, 

01?26'N, 65°47'W, 1,550 m, 16 Oct. 1987, 
R. Liesner & F. Delascio 21988 (holotype, 
VEN; isotype, AMES, MO). Figure 13. 


Plantae terrestres, grac iles, minutae (vix 4-7 cm altae); 


libus e rhizomate brevi decumbenti vix erectis; racemo 


Volume 76, Number 4 Steyermark, Holst & Collaborators 963 
1989 Venezuelan Guayana Flora— VII 


elongato remote paucibracteato, apice laxe spicato; flo- 


a ungue tubo 
deinde ittat 


elliptico; rostello valido, oblongo, apice truncato, apice 
structura globulari et papillosa praedito. Anthera ellip- 
soidea. Pollinia lineari-clavata, viscidio parvulo, rotundo 
affixa. Ovarium subcilindricum, sessile, haud tortum 


Small terrestrial or muscicolous erect herbs, 4— 
7 cm high. Roots cylindric, 3-6, fleshy, coarse, 
pubescent, up to 3 cm long. Stems 5-15 mm long, 
ca. 3 mm thick, totally enveloped by the leaf sheaths, 
terete. Leaves glabrous, thinly subfleshy, marces- 
cent, 7-11 per rosette, 4-5 fresh at flowering, 
sheaths imbricating and invaginating the stem, lam- 
ina flat or concave, margins slightly undulate, el- 
Ed or a Ps acute or shortly acuminate, Detail of PH dos ^ C Leaf 
5-9 mm long, 3-4 mm wide, basally contracted with ligulelike structure at leal sheath. A Flower, ped- 
into a channeled pseudopetiole 1-1.5 mm long and  icellate ovary, and floral bract. — E. Perianth segments 
wide, this merging into the sheath. Sheath ovate F. Labellum. — G. Upper side view of ih with anther 
to ovate-elliptic, ca. 5 mm long, 4 mm wide when retracted. — H. Lower side view of column showing stig- 
flattened, with 1(2) erect, liguliform structures con- matic surface, (From Liesner & Delascio 21988.) 
nected at the center of the sheath base and pro- 
truding from it, these 3.8-4.5 mm long, 1-1.2 
mm wide, rigid, + expanded apically where cov- 
ered by multicellular short-pedunculate, globose 
hairs ca. 0.05 mm long. Inflorescence a terminal 
raceme, 35-45 mm long, laxly 3-6-flowered, fer- 
tile in the upper !4, covered (except for the bracts) 
with sparse, cylindric to subcapitate, unicellular 
hairs ca. 0.1-0.2 mm long. Peduncle terete, 21— 
28 mm long, triarticulate, each articulation sub- 
tended by one glabrous, lanceolate, long-acuminate 
bract, this basally enveloping and apically spread- 
ing, the lowermost bracts 6.5-7 mm long, 3 mm 
wide, the apical bracts 5 mm long, 2 mm wide. 
Rachis 12-17 mm long, straight or slightly frac- 
tiflex. Floral bracts 3-7 mm long, 2-2.5 mm wide, 
identical to the peduncle bracts but somewhat 
smaller, much longer than the pedicellate ovary. 
Flowers tubular, not resupinate, white, 7— m 
long including the ovary, forming an acute a 
with the rachis. Ovary subcylindrical, 2-2.2 m 
long, 1-1.2 mm thick. Sepals fused in the basal 
34, forming a cylindric, 5-5.5-mm-long tube around 
the column, petals, and labellum, the free parts 
concave, triangular-oblong, rounded apically, 2- 
mm long, 0.9-1.1 mm wide, minutely glan- 
dular-papillose without; dorsal sepal partially ad- This new genus seems to belong to the Eury- 


Aracamunia Menem M Habit. — B. 
he 


nate to back of column in the basal Y within. 
Petals membranaceous, connivent with the dorsal 
sepal along their whole length, uninervate, linear- 
spathulate, ca. 5 mm long, 0.7 mm wide near apex, 
0.1-0.2 mm wide in the attenuated basal half. 
Labellum connivent with column margins, mem- 
branaceous, trinervate, 5.5-6 mm long, ca. 1 mm 
wide near apex, narrowly oblong in general outline, 
somewhat dilated in the apical !4, apex obtuse, 
thickened and finely papillose-ciliate, base sagittate 
with 2 retrorse, linear-oblong, apically rounded and 
thickened lobes 1.3-1.5 mm long, 0.4 mm wide; 
labellum attached to ovary apex by a relatively 
long, linear-oblong claw, 0.9-1.2 mm long, 
mm wide, this adnate to the sepaline tube. Column 
erect, in subcylindric, 4.5 mm long, 0.7- 

mm wide at base. Anther ellipsoid, bilocular, 
obtuse at on extremes, 2 mm long. Rostellum 
oblong, 1.8 mm long, ca. 1 mm wide, apically 
truncate and bearing a globular, papillose structure 
0.4 mm long and wide. Clinandrium margins nar- 
row, membranaceous, + erect and slightly enclos- 
ing the anther. Stigmatic surface ventral, papillose, 
transversely elliptic, 0.8 mm wide 


964 


Annals of the 
Missouri Botanical Garden 


styles Wawra complex, with which it shares the 
small vegetative habit, nonresupinate flowers, and 
sepals connate for at least part of their length. 
However, Aracamunia differs from the other gen- 
era of the KEurystyles complex in its terrestrial 
habit, eciliate leaves and bracts, presence of lig- 
ulelike structures, noncapitate inflorescences, and 
ecristate sepals connate for about % of their length 
(vs. up to 4% their length). Within the complex, it 
seems to be closer to Eurystyles as defined by 
Burns-Balogh et al. (1985), but in Aracamunia 
the stigmas are confluent as in Pseudoeurystyles 
Hoehne. 

The most distinctive feature of Aracamunia is 
the presence of the ligular, apparently glandular 
structures that originate from the bases of the leaf 
sheaths and persist even after disintegration of the 
leaf blades and sheaths. These ligular structures 
are erect, rigid, solitary, or (rarely) in pairs on each 
leaf sheath. They show traces of vascular axes, 
with phloem extending farther apically than the 
xylem, suggesting a glandular function. The apices 


age dentate, or expanded into a suborbicular 
he multicellular hairs of the liguliform 
structures are denser in younger more apical por- 
tions, with each of the hair cells containing a large 
nucleus proportionate to the size of the cells. 
he ligular structures have vascular connections 
with the leaf sheaths and the stem. Further study 
is required to determine if they are glandular or 
perhaps have an assimilative function. 


Etymology. Generic name for Cerro Araca- 
muni where the type species was collected, in the 
Federal Territory of Amazonas in southern Ven- 
ezuela. The specific epithet honors Ronald Liesner 
who collected the type material. 


PIPERACEAE— contributed by Julian Steyermark 


Piper mosaicum Steyerm., sp. nov. TYPE: Ven- 
ezuela. Territorio Federal Amazonas: Dept. 
Rio Negro, Cerro Aracamuni, summit, Popa 
Camp, savanna with small to large patches of 
forest and stream, 01?26'N, 65?47'W, 1,550 
m, 16 Oct. 1987, R. Liesner & F. Delascio 
21968 (holotype, MO; isotype, VEN). Figure 
14 


tex ] metralis; ramis retrorse Maier foliis 
elliptico- T apice acute acuminatis, basi aequilat- 
eraliter s sutis vel iE pacem mucronatis 8.5- 14 
cm ae 25 4 cm latis, conspicue reticulatis argenteo- 
maculatis ubique rugulosis, praeter squamas farinaceas 
lepidotas glabris, subtus nervis et costa media adpresso- 
pubescent tibus, squamis farinaceis onustis; totis pinnatim 
venosis, nervis lateralibus utroque latere 5-8; petiolis 2- 


7 mm longis adpresso- puberulentis, sin medium vagina 
tis; spicis erectis 17 mm longis 5 m bond sell 
4.5 mm longis, breviter retrorse Sube 


Shrub 1 m tall; stems nodose, minutely and 
retrorsely subappressed-pubescent, 3-4 mm thick. 
eaf blades elliptic-lanceolate, acuminate at the 
minutely mucronate apex, equilaterally acute to 
subacute at base, drying gray-silvery and blackish, 
reticulate-rugulose both sides, 8.5-14 cm long, 
2.5-4 cm wide, upper surface glabrous except for 
farinaceous lepidote scales, the darker areas with 
more minute scales, lower surface paler, covered 
with a dense farinaceous lepidote layer with pale 
stellulate trichomes along the midnerve, pinnately 
nerved throughout to the upper !4, the lateral 
nerves 5-8 each side; midnerve and lateral nerves 
on lower surface appressed-pubescent. Petioles 2— 
7 mm long, minutely appressed-puberulent, the 
petiolar sheath vaginate in lower half. Spikes erect, 
17 mm long, 5 mm wide, peduncle 4.5 mm long, 
shortly retrorsely puberulous. Stigmas sessile, 
spreading over the summit of the fruit. Fruit de- 
pressed-truncate at summit, suborbicular, subquad- 
rangular. 


This species differs from Piper gentryi Steyerm. 
by having mottled, smaller leaf blades; shorter pet- 
ioles with sheaths that are longer, winged; re- 
trorsely pubescent stems and peduncles; and fewer 
lateral nerves. It is distinguished from P. holtii 
Trel. & Yuncker by having shorter peduncles and 
spikes and smaller, mottled leaf blades that lack 
the sulcate nerves on upper leaf surface and lack 
conspicuously elevated nerves of the lower surface. 


RAPATEACEAE— contributed by Julian Steyermark 


Rapatea aracamuniana Steyerm., sp. nov. TYPE: 

enezuela. Territorio Federal Amazonas: Dept. 

Rio Negro, Cerro Aracamuni, slope, Quebrada 

Camp, area of rapids flowing over laja, river 

bank in sun, more common in shade, 01?24'N, 

65?38'W, 600 m, 20 Oct. 1987, R. Liesner 

& F. Delascio 22197 (holotype, MO; isotype, 
VEN). Figure 15. 


Planta herbacea; foliorum laminis supra omnino minute 
dense papillato-verruculosis, su 


18 mm mene pace 10 mm longis; filamentis ciliolatis. 


Herb. Leaves petiolate, sheaths 11—13 cm long, 
6-8 cm wide, conspicuously many-costate, densely 
papillate; leaf blades ligulate-lanceolate, falcately 
acuminate at apex, inequilaterally acute at base, 
36-66 cm long, 5-7.4 cm wide at the middle, 
upper surface densely verruculose-papillate, lower 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 965 


Venezuelan Guayana Flora— VII 


FIGURE 14. Piper 


surface. — E. Detail of stem node. — F. Stem. 


surface with brown glandular punctae regularly 
distributed between smaller punctate glands, midrib 
conspicuously impressed on upper surface, elevat- 
ed on lower surface, bordered by 55-60 finely 
elevated pe nerves on each side. Petiole 3- 
7.5 cm 
idea shallowly hemispherical, 4.5-6.5 c 

long, 8- cm wide, enveloped by 2 broadly 
deltoid involucral prid these cordate at base and 
acute at apex, 7— m long, 9-10 cm wide, 
minutely brown papillate- verruculose without, mi- 


, 8-9 mm wide. Inflorescence on 


nutely pale papillate within. Spikelets numerous, 
pedicellate; pedicels 10—13 mm long, 1 mm wide, 
broadened to 1.5 mm wide at summit. Spikelets 
numerous, (excluding sepals) 23-25 mm long, 
mm wide, ca. 12-bracteolate. Bracteoles graduate, 


cen 


mosaicum. — A. Habit of branch. — 


ail of spike. — C. —D. tip, lower 


Det k nther Leaf tip, 
—G, H. Details i ene leaf surface. (vani pr & Delascio 21968.) 


the lowest 17-18 mm long, 3 mm wide, % length 
of spikelet, linear-lanceolate, acute, scarious-mar- 
gined; the innermost (uppermost) 22-23 mm long, 
3-4 mm wide, narrowly triangular-lanceolate, acu- 
minate, minutely puberulent without in upper por- 
tion, minutely ciliolate. Sepals with free expanded 
portion broadly lanceolate, acuminate, 12 mm long, 
4 mm wide at base, membranous, 5-nerved, the 
lower portion attached to corolla tube for 15 mm 
long and 4 mm wide, hyaline throughout. Corolla 
yellow, 3 cm long; tube 12 mm long, 2.5-3 mm 
wide; laminate portion rhomboid, abruptly acute, 
17 mm long, 9 mm wide at the widest portion, 
narrowed to claw 9 mm long, 2 mm wide. Anthers 
linear, bilobate at base, 10 mm long, 0.5 mm wide, 
prolonged at apex into a linear-oblong gland 1.5- 


966 Annals of the 
Missouri Botanical Garden 
o 
3 
(d 3 
MAN ll 
| WI "P 
EMI I Il fos 
| | || q 
Y HI f i E 
T i] 
nl ! | 
MI 
nT 
A | niyi | 
i 
I 
HII 
ul 
il 
i 
€ | 
O 
iD TOI 
Vl 
^d HA 
Il 
Hii 
lll 
JJ] | 
All 
| 
Y 
| O 
Han 3 
LH] i 
BESNNI | 
uli H 
Rss 15. dE ea aracamuniana. — A. Leaf. —B. Spikelet with open flower. — C. Inflorescence. Sis es 
rescence with bracts. —E. Fruit. —F. Corolla in bud. —G. Flower with corolla opened. —H. Petal.— I. Stamen. — J. 


Anther de 1d. Tk “Detail of upper leaf surface. — L. Detail of lower leaf surface. (From Liesner & Delascio 22197. ) 


1.7 mm long, 0.4-0.5 mm wide. Filaments 4 mm 
long, 0.5 mm wide, pilosulous-ciliolate in the up- 
permost 2 mm. Style glabrous, 28 mm long, 0.3 
mm wide, curved at apex, narrowed to a conical 
apex. 


This species is related to Rapatea steyermarkii 
Maguire of extreme southeastern Venezuela in Es- 
tado Bolivar. Although similar in inflorescence, in- 
volucral bracts, and leaf shape and size, R. ara- 
camuniana has filaments that are ciliolate (vs. 
glabrous in R. steyermarkii). The anthers, lowest 


bracteoles, and spikelets are longer in R. araca- 
muniana. The upper and lower leaf surfaces of R. 
steyermarkii are densely brown pubescent on the 
lower surface with elongate, crisp-villosulous, loose 
multicellular hairs, whereas in R. aracamuniana 
the lower surface has only brown glandular dots 
between smaller, pale punctations. In R. steyer- 
markii the upper leaf surface has short, divaricate, 
pointed hairs irregularly scattered on the nerves 
in addition to a minutely papillate surface, whereas 
the upper surface in R. aracamuniana is densely 
verruculose-papillate throughout. 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora— VII 


Since Rapatea steyermarkii was originally de- 
scribed without flowers, the following description 
(taken from a specimen of Holst & Liesner 2847 
from Río Acanan, Estado Bolivar) is added to pro- 
vide details lacking from the original description. 


RAPATEA STEYERMARKII 


pala! ar libera) 19-20 mm longis; antheris 5 mm 
longis, 0.7 mm latis; filamentis 3 mm longis omnino gla- 
bris; stylo 15 mm longo glabro. 


RUBIACEAE—contributed by Julian Steyermark 


Cephalodendron aracamuniensis Steyerm., 
sp. nov. TYPE: Venezuela. Territorio Federal 
Amazonas: Dept. Rio Negro, Cerro Araca- 
muni, summit, Popa Camp, ravine forest at 
edge of tepui, 01?26'N, 65?47"W, 1,550 m, 
17 Oct. 1987, R. Liesner & F. Delascio 

22064 (holotype, MO; isotype, VEN). 


Arbor 4-6- metal ramulis cenae hirsutis pilis diva- 


M c Re d a 2-3 m 
o dense antrorse adpres 


nse pubescenti pilis 
adpressis retrorsis 0.5 mm longis munito; antheris 7 mm 


longis 


Tree 4-6 m tall, stems of the branches ferru- 
ginous-brown hirsute with spreading hairs 2-3 mm 
long. Stipules at the stem apex prominent, oblong- 
obovate, rounded at apex, 5.5-7 cm long, 3-3.5 
cm wide, densely ferruginous-brown hirsutulous. 
Leaf blades subcoriaceous, both sides rugulose, 
broadly obovate, abruptly acute at apex, rounded- 
subcordate at base, 32-35 cm long, 16-26 cm 
wide at or near the base, lower portion of upper 
surface hirsute with erect to ascending ferruginous 
hairs 2-3 mm long, hecoming more shortly pu- 


bescent in age, lower surfa ly villous-hirsute 
with ferruginous-brown hairs, the midrib and main 
lateral nerves with denser, loosely spreading to 
subretrorse hairs 2-3 mm long. Lateral nerves 16- 

each side, arcuate-ascending at 35-40*, in 
conspicuously anastomosing near the margin, el- 
evated below, impressed above; tertiary venation 
conspicuously rugose on lower surface. Inflores- 
cence capitate, globose, at anthesis 3 cm long, 3.5 
cm wide, up to 4 cm long and 4.5 cm wide at post- 
anthesis, enveloped at anthesis by 2 suborbicular 
rounded bracts 3 cm long, these 3.5-4 cm wide, 


densely ferruginous hirsute. Peduncle 9-29 cm 
long, 5-6 mm wide, densely ferruginous hirsute 
with loosely spreading hairs 2-3 mm long. Bract 
subtending base of hypanthium lanceolate, acute 
5 mm long, 1.5 mm wide. Hypanthium turbinate- 
oblong, in anthesis 6 mm long, 4.5 mm wide, 
densely ferruginous hirsute with ascending hairs 
2.5-3 mm Calyx and hypanthium 19-20 
mm long in anthesis. Calyx tubular-cylindric, 15 
mm long, 5-6 mm wide, the tube in anthesis 13 


long. 


mm long, 3.5 mm wide, appressed-pubescent with 
hairs 1-2.5 mm long; calyx lobes 5, unequal, 4 
lobes broadly suborbicular, rounded, 1.5 mm long, 
2 mm wide, the 5th lobe broadly subtriangular- 
ovate with a 0.5-0.6 mm long caudate appendage, 
densely short-sericeous within. Corolla white, 
subsalverform, 35-37 mm long, the tube 27 mm 
ong, 3 mm wide in the upper 15 mm, 1.5 mm 
wide in the lower 10-12 mm, densely pubescent 
without, with appressed hairs 0.5 mm long; corolla 
lobes 8 mm long, 2 mm wide, densely antrorsely 
pubescent without, glabrous within in the basal 2. 

0.5-0.7 


the adjacent 11 mm, glabrous in the uppermost 


mm, pilose with spreading hairs mm in 
12 mm. Stamens inserted in the upper ?4 of the 
corolla tube; anthers linear, 7 mm long, 0.5 mm 
wide, slightly bilobate-rounded at base; filaments 1 
mm long, glabrous, inserted 10 mm below base of 
corolla je Style 


glabrous. Fruits not seen. 


8.5 mm long, glabrous. Disk 


type. | VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: Dept. Río Negro, Cerro Aracamuni, summit, 
Proa Camp, ravines near oe of tepui, 01°32'N, 65°49'W, 
1,400 m, 31 Oct. 1 , R. Liesner & G. Carnevali 
22682 (MO, VEN). 


Cephalodendron was known until now as a 
monotypic genus endemic to the summit of Cerro 
de la Neblina in Venezuela. The new taxon differs 
from C. globosum Steyerm. in the longer, spread- 
ing-hirsute pubescence of stems, peduncles, peti- 
oles, and lower leaf surface; longer calyx, corolla, 
and anthers; and cylindric calyx tube 314-342 times 
longer than broad. 


Coussarea liesneri Steyerm., sp. nov. TYPE: Ven- 
ezuela. Territorio Federal Amazonas: Dept. 
Rio Negro, Cerro Aracamuni, slope forest, 
Quebrada Camp, 01?24'N, 65°38'W, 600 m, 
2] Oct. 1987, R. Liesner & F. Delascio 
22262 (holotype, VEN; isotype, MO). 


Arbor 8-metralis; foliis oblongo- pinus apice abrupte 
acuminatis, acumine bd ad 1.5 cm producto, basi 
utis 22-3 pud cm Be ubique glabris, 
nervis lateralibus que i tere 10-12; petiolis 1.5-2 

cm longis; inflorescentia Mri. pedunculo incluso 8- 


£ 
e 


968 


Annals of the 
Missouri Botanical Garden 


10 cm longa basin versus 6 cm lata; pedunculo 4-4.5 
cm longo puberulento; cuyos 4-lobato profunde campa- 
nulato 2.5 mm lo lato extus minute 
puberulento, lobis inaequ 
apice obtusis 0.5 mm longis puberulento-ciliatis; hypanthio 
minute puberulento, staminibus basi corollae insertis. 


Tree 8 m tall. Leaf blades oblong-elliptic, 22- 
33 cm long, 7-14 cm wide, glabrous throughout, 
abruptly acuminate at apex (the acumen up to 1.5 
cm long), acute at base; principal lateral nerves 
10-12 each side, elevated below, slightly elevated 
or impressed above; tertiary venation reticulate and 
elevated below, inconspicuous above. Petiole 1.5- 
2 cm long, 2.5-3.5 mm wide, glabrous. Inflores- 
cence terminal, paniculately branched, 8-10 cm 
long including the peduncle, 6 cm broad at the 
base, with 3 main divaricate tiers of lateral axes. 
Flowering portion of inflorescence 6 cm long with 
7-10 axes, the lower 4 proximate, 1-2 cm long 
and branched into 3 secondary axes bearing capitu- 
liform clusters 7-10 mm broad, each containing 
5-10 sessile flowers; the other tiers of axes shorter, 
1-3 cm long; rachis and axes minutely puberulent. 
Peduncle 4-4.5 cm long, 2-2.5 mm wide, puber- 
ulent. Calyx 4-lobed, deeply campanulate, 2.5 mm 
long, 2 mm wide at summit, narrowed below, mi- 
nutely puberulent without, sparsely puberulent at 
the summit margin, glabrous within; lobes unequal, 
squarrose, sublanceolate, obtuse at apex, 0.5 mm 
ong, 0.2 mm wide, puberulent-ciliate; hypanthium 
turbinate, 1.5 mm long, ] mm wide at summit, 
minutely puberulent without. Corolla cylindric, api- 
cally narrowed, 6 mm long, 1.5 mm wide (in bud); 
corolla lobes 3 mm long (in bud). Stamens inserted 
at base of corolla tube; anthers 2.5 mm long. 


This species is related to Coussarea rudgeoides 
Rusby of Bolivia and Peru, from which it differs 
in the denticulately lobed calyx with pubescent, 
squarrose lobes; longer peduncle; longer calyx tube 
exceeding the hypanthium; and insertion of the 
filaments at the base of the corolla tube. In C. 
rudgeoides the filaments are attached in the lower 
1⁄4 of the corolla tube, and the calyx tube is about 
as long as the hypanthium. 


Nahi; 1 


Steyerm., sp. 
nov. TYPE: Venezuela. Territorio Federal Ama- 
zonas: Dept. Rio 
summit, Popa Camp, savanna at edge of tepui, 
01?26'N, 65?47'W, 1,550 m, 19 Oct. 1987, 
R. Liesner & F. Delascio 22153 (holotype, 
MO; isotype, VEN). 


egro, Cerro Aracamuni, 


Suffrutex 0.5-metralis; foliis diera elliptico- oblongis 
e am vel subacutis majoribus -4.2 cm longis 
1.3-2.2 cm latis subtus praeter ae foveolatas minute 


albido- pubescentes glabris; calycis lobis spathulato-linear- 
ibus apicem versus dilatatis apice subacutis 4 mm longis 
superne 0.5-0.7 mm latis extus albido-hispidulis, margi- 
nibus hispido-ciliatis; corollae tubo 16 mm longo extus 
glabro intus m antheras minute papilloso- -puberulen- 
toali , lobis ovato- oblongis apice late rotundatis 
5 mm To 3 mm lat 


Suffruticose plant 0.5 m tall. Stems slender, 
densely introrsely hirtellous. Leaves ternate, ellip- 
tic-oblong, mainly obtuse to subacute at apex, acute 
at base, the larger and principal leaf blades 3.5- 
4.2 cm long, 1.3-2.2 cm wide, glabrous above 
except for sparse, appressed hairs in the sulcate 
midnerve, paler and mainly glabrous below except 
for depressed, minute, white pubescent, foveolate 
areas within the reticulate venation; principal lat- 
eral nerves 4-6 each side, inconspicuous to ob- 
solete above, the midnerve and lateral nerves gla- 
brescent or with sparse, appressed trichomes 0.2- 
0.5 mm long; tertiary venation obsolete above, 
finely reticulate below and impressed, the areoles 
in depressed foveolate areas with circles of minute 
white trichomes. Petiole 2-3 mm long, moderately 
to densely appressed-pubescent with pale hairs. 
Inflorescence terminal or axillary, the flowers sol- 
itary or in pairs. Pedicels 1-2 mm long. Hypan- 
thium turbinate, 2 mm long, 1.5 mm wide, densely 
gray hirtellous. 
broadened near the subacute apex, subacute at 
base, 4 mm long, 0.5-0.7 mm wide in upper half, 


Calyx lobes linear-spathulate, 


hispidulous without, with gray-white pubescence, 
glabrous within, the margins hispid-ciliate. Corolla 
tube red, greenish white on lobes, 16 mm long, 
slenderly salverform, the tube 12 mm long, 2 mm 
wide, glabrous without, minutely papillose-puber- 
ulent within for 3 mm in the zone of the anthers, 
elsewhere glabrous; corolla lobes 5, ovate-oblong, 
rounded at apex, 5 mm long, 3 mm wide. Anthers 
linear, 2 mm long, inserted 2% way up tube; fila- 
ments free for 1 mm, attached 3 mm to tube 4 
mm above its base. Style 10 mm long, glabrous; 
stigmas 2 mm long. 


The genus Veblinathamnus was originally de- 
scribed by Steyermark (1963) with two species 
known, N. argyreus from Venezuela on Cerro de 
la Neblina and N. glabratus from adjacent north- 
ern Brazil on Serra Pirapucü. This third species 
differs from the other two in its shorter corolla; 
larger leaf blades with subacute or obtuse apices; 
and shorter, spathulate-linear, subacute calyx lobes, 
which are more dilated at the apex, dorsally pu- 
bescent, and more abundantly ciliolate. 


Remijia reducta Steyerm., sp. nov. TYPE: Ven- 


ezuela. Territorio Federal Amazonas: Dept. 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora— VII 


Rio Negro, Cerro Aracamuni, summit, Proa 
Camp, savanna near edge of tepui, 01?32'N, 
65?49'W, 1,400 m, 25 Oct. 1987, R. Liesner 
& G. Carnevali 22407 (holotype, MO; iso- 
type, VEN). Figure 16. 


Frutex 0.5- 1-metralis; ramulis juvenilibus dense hir- 
sutis pilis antrorsis; foliis parvis revolutis lanceolato-ellip- 
ticis vel elliptico-oblongis apice obtusis vel subacutis, basi 
4-10 cm longis 2-3 cm latis, supra tenuiter rugosis 


12 mm longis 2-3 mm latis; antheris 3.5 mm dics 
capsulis maturis 13-18(-32) mm longis; seminibus 5- 
6.5 mm ve corpusculo suborbiculari 3 mm longo, alis 
1-2 mm lat 


Shrub 0.5-1 m tall. Stems antrorsely densely 
hirsute with appressed to ascending ferruginous 
hairs; terminal stipule ovate-oblong to broadly ovate, 
obtuse or rounded, 10-17 mm long, 5-14 mm 
wide, densely appressed-hirsute with elongate fer- 
ruginous-brown hairs 1.5-2 mm long. Internodes 
3-6 mm long. Leaf blades coriaceous, drying dark 
above and ferruginous-brown below, revolute, lance- 
elliptic to elliptic-oblong, obtuse to subacute at apex, 
acute at base, 4-10 cm long, 2-3 cm wide, ru- 
gulose or bullate and glabrous above at maturity, 
densely hirsutulous with subappressed to ascending 
hairs in early stages, densely tomentose below, 
midrib and lateral nerves with densely appressed, 
crisp, longer hairs; lateral nerves 10-12 each side, 
slightly to deeply sulcate above, finely and minutely 
reticulate and elevated below. Petioles 2-9 mm 
long, 1-2 mm wide, densely appressed-hirsute. In- 
ternodes 3-6 mm long. Inflorescence greatly ab- 
breviated, lateral, the flowers solitary, one on each 
side. Peduncle at anthesis 1-2 mm long, in fruit 
5-7 mm long, densely and antrorsely ferruginous 
hirsute. Hypanthium at anthesis 2-3 mm long, in 
fruit cylindric to oblanceolate and elongating to 9 
mm long, densely antrorsely ferruginous-hirsute. 
Calyx 1-2 m 
broadly deltoid lobes, glabrous within. Disk Panis 


mm 


ong, subtruncate or with minute, 


the calyx. Corolla white, salverform, tube 1 
long, 3 mm wide above, 2.5-3 mm wide an the 
middle, densely buff-brown hirsute with antrorsely 
appressed hairs 33 mm long, glabrous within; lobes 
5, recurved, linear-lanceolate, acutish, 10-12 mm 
long, 2-3 mm wide, densely antrorse-hirsute with- 
out, glabrous within. Stamens attached to tube 
slightly below the middle; anthers linear, 3.5 mm 
long, 0.7 mm wide; filaments 4 mm long, attached 


6.5 mm above base of tube, glabrous. Style 3 mm 
long, glabrous; stigmas linear, 2 mm long. Capsule 
at maturity obovate or broadly oblong, 13-18(-32) 
mm long; valves conspicuously osseous, bifid at 
apex, densely brown hirsute without, glabrescent, 
8-13 mm wide; seeds few, wings pale buff-brown, 
body pale brown, finely cellular on surface, oval 
or subovate, rounded or subtruncate at one or both 
ends, often more narrowed at the other end, 5- 
6.5 mm long, 2.7-3 mm wide; wings 1-2 mm 
wide. 


Paratype. | VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: Dept. Rio Negro, 
Popa Camp, low savanna, 01?26'N, i 
17 Oct. 1987, R. Liesner & F. e ied 22045 (MO, 
VEN) 

This species differs from other species of Re- 
mijia by having solitary flowers. It is further dis- 
tinguished by its small leaf blades, short petioles, 
densely ferruginous tomentose lower leaf surface 
with strongly reticulate tertiary venation, finely 
rugulose upper leaf surface, and relatively small 
s 


THEACEAE— contributed by Julian Steyermark 
(Bonnetia) and Brian Boom (Ternstroemia) 


Bonnetia liesneri Steyerm., sp. nov. TYPE: Ven- 
Dept. 


Rio Negro, Cerro Aracamuni, summit, Popa 


ezuela. Territorio Federal Amazonas: 
Camp, savanna with small to large patches of 
forest and stream, 01?26'N, 65?47'W, 1,550 
m, 18 Oct. 1987, R. Liesner & F. Delascio 
22105 (holotype, MO; isotype, VEN). Figure 
17 


Suffrutex 0.5-metralis; foliis anguste ei Qc vel 


ulis autem sub m een inulato-rotun 

1 longi atis; sepalis ae elliptico- 
bud. aristato- oblandi apice rotundatis 11-12 mm 
longis 3-4 mm latis; style simplice vix 3- un 8 mm 
longo. 


Subshrub 0.5 m tall with spreading branches. 
Leaves clustered at the tips of the branches, nar- 
rowly oblanceolate to narrowly elliptic, seemingly 
subacute or obtusish at apex, but under 20 x mag- 
nification rounded with an emarginulate summit, 
attenuate to the base, 18-30 mm long, 4-8 mm 
wide, glabrous throughout, leaf margin microscop- 
ically repand, cartilaginous, shortly 1-2 mm pet- 
iolate; midrib slightly elevated below, obsolete above; 
lateral nerves and tertiary venation obsolete. Flow- 
ers solitary near the summits of the branches. Pe- 
duncle ancipital, ebracteolate, mainly 13-15 mm 
long (sometimes much longer), glabrous. Sepals 


Annals of the 


Remijia dig ta. — 
Fru 


y.—E. Seed. —F. 


narrowly elliptic-lanceolate to ovate, aristate-acu- 
minate at apex, acute at base, pars mm a 
3-4 mm wide at the middle, entire. Stamens nu- 
merous; anthers suborbicular, ghi longer dion 
broad, 0.7 mm long, mm Style 8 m 
long, undivided, merely shallowly 3 lobulate at is 
apex. Ovary (post-anthesis) ovoid, 5 mm long, 3 
mm wide. Fruit ovoid, 3-celled, tapering to the 
persistent style, 10 mm long 


This species is most similar to Bonnetia hube- 
riana Steyerm. of the Chimanta Massif, Estado 
Bolivar, but differs by having undivided, shallowly 
three-lobulate style; larger petals; larger, aristate- 


970 
Missouri Botanical Garden 
C 
Ein rat A. Habit of branch. — B. Corolla. — C. Corolla opened. — D. Pistil with calyx cut 
)7.) 


t. — G. Detail of lower leaf surface. (From Liesner & Carnevali 22407 


acuminate, entire sepals; and  microscopically 
rounded emarginulate leaf apex. 


Ternstroemia guanchezii Boom, sp. nov. TYPE: 
usais onn Federal Amazonas: Dept. 
tures, matorral ribereno has- 
ta an bajo n en la margen izquierda 
del cano “Cabeza de Manteco,” afluente del 
Rio Autana, en el “Raudal Manteco,” 04*52'N, 
67°27'W, 100-120 m, 10 Nov. 1984, F. 
Guánchez & E. Melgueiro 3447 (holotype, 

NY; isotype, TFAV). Figure 18 
Frutex altus, ramulis oppositis vel suboppositis 
teretiusculis ravidis; foliis oblanceolatis vel angustis ellip- 


Volume 76, Number 4 


Steyermark, Holst & Collaborators 


1989 Venezuelan Guayana Flora— VII 
w 
O 
3 
o 
a 
3 
3 
D E F 
FIGURE 17. Bonnetia liesneri. —A. Habit of branch. — B. Flower. — C. Fru 


G, H. Stamen.— I. Lower leaf surface. —J. Leaf 


ticis 5-7(-8.5) x 1-1.5(-3) cm aliquantum asymmetricis 
apice acutis vel acuminatis basi acuminatis vel apenas: 
petiolis 0.2-0. 5(- 1) mm longis, co 
infra prominula, venis obscuris a floribus idi 
ulatis, p ad 10 mm longo a vel pen 
dulo, bracteolis 2 suboppositis cymbiformibus 

l- 


oglanduloso- a, et tribus intimis eglan- 
dosis- integris dh. lobis 5 liberis ovatis vel lanceolatis 
3.5-4 5 mm s obtusis marginibus integris, 
teu ca. 24 liberis me ws basi xis, filamentis 
0.3-1 mm longis, antheris i 
eas introrsus eee stigmate et 

m longis, ovario 1 x 1(-1.2) mm 2-loculari, utroque 
bh 2-ov ju fructibus incognitis 


— D. Sepal-—E. Petal: —F.. Pistil.— 
.) 


apex. (From o & EMT 22105 


Shrub 2-3 m tall; branchlets opposite to subop- 
posite, terete, grayish. Leaves clustered within the 
apical 3(-5) cm of branchlets, coriaceous, glabrous, 
oblanceolate to narrowly elliptic 5-7(-8.5) x 1- 
1.5(-3) cm, slightly asymmetrical, apex acute to 
acuminate (to rounded), base acuminate to atten- 
uate, petiole 0.2-0.5(-1) mm long, midrib cana- 
liculate above, evident the entire length, prominent 
below, the veins obscure on both surfaces. Flowers 
fasciculate, axillary among leaf bases, up to 7 per 
cluster, peduncles to 10 mm long, suberect to 
pendulous; bracteoles 2, cymbiform, subopposite, 
unequal in size, 2.5-3 x 1.5-2 mm, apex acute, 
black gland-tipped (the gland itself tipped by sev- 


972 Annals of the 
Missouri Botanical Garden 


FIGURE 18. Ternstroemia guanchezii. —A. Habit. — B. Flower, longitudinal view with two calyx lobes and one 
ps removed, showing stamens and summit of gynoecium. — C. Gynoecium. — D. Stamen. (From Liesner & Delascio 
22230.) 


Volume 76, Number 4 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora— VII 


eral eglandular cilia), the margin black glandular- 
serrulate; calyx lobes 5, free, cymbiform, 3.5-4 
x mm, the margin scattered black glan- 
dular-serrulate on outer lobes, entire on inner 3 
lobes; corolla of 5 free petals, these ovate to lan- 
ceolate, 3.5-4 x 


entire; androecium of ca. 24 uniseriate stamens, 


1-1.5 mm, apex obtuse, margin 


free from corolla; filaments of varying lengths, 0.3— 
] mm long; anthers 0.6-1 mm long, basifixed, 
thecae parallel, dehiscence introrse; gynoecium with 
terete stigma and style, 2.5 mm long, ovary | x 
1(-1.2) mm, 2-locular with 2 pendulous ovules per 


locule, the ovules 0.5-0.6 x 0.1-0.2 mm. Fruits 
unknown. 
Paratype. VENEZUELA. TERRITORIO FEDERAL 


AMAZONAS: Dept. Río Negro, Cerro Aracamuni, slope, 
Quebrada Camp, in area of rapids F over laja, 
01°24'N, 65°38'W, 600 m, 20 Oct. , R. Liesner 
& F. Delascio 22236 (MO, NY, a 


This distinctive species, with graceful, subop- 
posite to opposite branchlets and small, fasciculate 
flowers, is known from lower to middle elevations 
of the western Venezuelan Guayana in forests along 
streams running through igneous rocks known lo- 
cally as “lajas.” The new species is named in honor 
of Francisco Guanchez, former director of the Her- 
bario TFAV in Puerto Ayacucho, who first col- 
lected this species in 1984. 


Ternstroemia maguirei Boom, sp. nov. TYPE: 
Venezuela. Territorio Federal Amazonas: Cer- 
ro de la Neblina, south slope of Cumbre Camp, 
caño toward Cañon Grande, occasional on ex- 
posed rock outcrop, 1,500 m, 16 Jan. 1954, 
B. Maguire, J. Wurdack & G. Bunting 37347 
(holotype, NY; isotype, VEN). Figure 19. 


rutex vel arbor 1-6 m altus, 6 cm diametro, ramulis 
fasciculatis angularibus ravido-fuscis; foliis ellipticis ad 
ovatis vel obovatis Me 5-)10-14 x (3.8-)4.5-6.7 mm 
aliquantum asymmetricis apice acutis vel rotundis basi 
acutis vel rotundatis pe (5-)8-13(-15) mm longo, 

15 


(-19)-jugis utrinque vix manifestis, lam 

punctata; floribus geminatis subterminalibus, pedunculo 
11-38(-50) mm longo; bracteolis duabus suboppositis 
cymbiformibus 5.5-6 x 3.5-4 mm rd cutis, margin e 


atroglanduloso-serrulatis, pd lobis : 
tis (5-)7-8.5 x (6.5-)8-9 m arem pan aag mar- 
glanduloso- Son eeey wie intimis fere eglan- 
duloso-integris; corollae lobis 5 basi per ca. 3 mm connatis, 
is li ¿ 5 x 3.5 mm apice acuminatis 

s ca. 37 uniserialis basifixis 


longis, thecis parallelis 1-1.5 mm longis introrsus dehis- 
centibus, connectivo ultra thecas producto, stigmatibus et 
stylis teretibus 8 mm longis; ovario biloculari, ovulis 0 

0.2 mm; fructibus laevibus perlate ovoideis 2-3- icut 


ibus 15 x 16 mm, stylo persistenti ca. 5 mm longo, 
parietibus 2.5-3 mm crassis; seminalibus 16, 8-9 x 4.5 


Shrub or small tree 1-6 m tall, 6 cm diam., 
branchlets fasciculate, angular in cross-section, 
grayish brown. Leaves distributed along branchlets, 
coriaceous, elliptic to ovate or obovate, (7.5-)10— 
14 x bs 8-)4.5-6.7 mm, slightly asymmetrical, 
apex te to rounded, base acute to rounded, 
sS (5- )8-13(-15) mm long, midrib canalicu- 
ate above, evident the entire length, prominent 
below, the veins 10-15(-19) pairs, scarcely visible 
above and below, blade below with scattered dark 
glandular punctations. Flowers subterminal, paired, 
peduncles 11-38(-50) mm long; bracteoles 2, 
cymbiform, subopposite, unequal in size, 5.5-6 x 
3.5-4 mm, apex acute, black gland-tipped, the 
margin black glandular-serrulate; calyx lobes 5, 
free, widely depressed ovate, (5-)7-8.5 x (6.5-)8- 
9 mm, the margin scattered black glandular-ser- 
rulate on outer 2 lobes, entire to slightly glandular 
on inner 3 lobes; corolla of 5 petals, these fused 
basally for ca. 3 mm with lanceolate lobes ca. 5 
x 3.5 mm, apex acuminate, margin entire; an- 
droecium of ca. 37 uniseriate stamens, these free 
from corolla, connate basally; filaments of varying 
lengths, 0.8-1 mm long; anthers 2 mm long, ba- 
sifixed, thecae parallel with apiculate connective, 
this 1-1.5 m 
cium with terete stigma and style, 8 mm long, 
ovary 2.5-3.5 
ovules per locule, the ovules 0.4 x 


m long, dehiscence introrse; gynoe- 


mm, 2-locular with 8 pendulous 

.2 mm. Fruits 
15 x 16 mm, 
persistent style 5 mm long, fruit wall 2.5-3 mm 
thick, 2-3-locular, 16-seeded, the seeds 8-9 x 
4.5 mm 


smooth, widely depressed-ovoid, 


Paratypes. VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: Dept. Río Negro, Cerro de la Neblina, Canon 
EL 3S mbre Camp, 1,200-2,200 m, 25 
7, B. Maguire et al. 42510 D VEN); Cerro 
de eblina, Camp m NNE of Pico egy 
el 40"N, 65°58'5 oW, 1,450 m, 27 Feb. 1985, M. 
Nee 31158 (NY, VEN); Dept. hys Negro, Cerro pa: 
camuni, due 2n 01°26'N, 65°47'W, summit, 1,550 
n, 16 Oct. R. Liesner € T Delascio 21985, 
same bci, * rn 1987, Liesner & Delascio 22117, 

, R. Liesner & F. Delascio 22158 (MO, 
NY, irn Aracamuni, 
65°49'W, Sait. 1,400 
& G. Carnevali 22506 (MO, VEN); 
escarpment along escarpment above Caleira, 1,400 m, 
20 Nov. 1950, B. Maguire et al. 29570 (NY); Cerro 
Duida, Caño F S N slopes and ridges, 2,000 
3,000 m, 23 Nov. 1950, B. Maguire | 29660 (NY); 
al SE de la MU de Culebra, , 65?45'W, 
800-900 m, 6 Feb. 1982, J. Steye Ain et al. 126125 
(MO, NY, V 


~ 


974 Annals of the 
Missouri Botanical Garden 


DX 


19. Ternstroemia maguirei. — A. Habit, Showing: mature fruit. —B, "n in flowers. —C. Flower, 
longitudinal view with two calyx lobes and Sas petals remove cium. — D. Gynoecium. — 
E. Stamen. (A, from Maguire et al. 37347; B-E, from [ripe et al. 42510.) 


Volume 76, Number 4 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora—VIl 


This new species is notable for its inflorescence 
of strictly one pair of flowers and its relatively 
large, smooth capsular fruits that are apparently 
nondehiscent. It is restricted to elevations above 

m on sandstone in southern Venezuelan Gua- 
yana (Cerros Neblina, Aracamuni, and Duida). It 
is named in honor of Bassett Maguire, Senior Sci- 
entist Emeritus of the New York Botanical Garden 
and inveterate botanical explorer of the Guayana 
Highland. Maguire and colleagues first. collected 
this species in 1954, and he first directed my at- 
tention to Ternstroemia as a genus in need of 
taxonomic study. 


XYRIDACEAE— contributed by Robert Kral (VDB) 


Abolboda scabrida Kral, sp. nov. TYPE: Vene- 
zuela. Territorio Federal Amazonas: Dept. Rio 
Negro, Cerro Aracamuni, summit, Proa Camp, 
medium height, semi-open forest, 01?32'N, 
65?49' W, 1,400 m, 28 Oct. 1987, R. Liesner 
& G. Carnevali 22598 (holotype, VEN; iso- 
types, MO, VDB). Figure 20. 


Herba perennis densicaespitosa, praeter inflorescen- 
tiam et ini ers et scabridorugosa. Radices spon- 
glosae. ascendentes, usque ad 2-3 cm 
longi, d. ed O obtecti. 
ola extima rosularum praesertim va inalia, a coi 
re oria jw e erecta vel e oan , leviter 
excu m longa, ad ba ld ca. 5. -costata, 
ee 2-3-costa ‘ask pin ae 0.5-1 

, transversim 


ca. 1 mm longis. Anth erae E e ca 
flamenta ca. a mm longa. 

d ub. clavatae, l r 
2.5 mm longa, obscure trilobata, valvis ad a inflexis 
et HEA Semina mmetrice obovoidea, 0. 
mm longa, deba ut valde 12-14-costata, fer 
cina. 


Perennial, densely cespitose herbs, all but the 
bases and inflorescence scabrid and scabrid-rugose. 
Roots spongy. Stems short, ascending, up to 2-3 
cm long, the bases covered by old leaves. Outer 
leaves of a rosette for the most part sheath, nar- 
rowly triangular; principal foliage leaves erect to 
spreading, slightly excurved, 3-6 cm long, dilated 
at base, ca. 5-costate, then abruptly contracted 
above, 2-3 costate; blades 0.5-1 mm wide, pale 
green, somewhat compressed, transversely irreg- 
ularly rugose and tuberculate; apices gradually to 
abruptly narrowed, thickened, at tip obtuse or ex- 
curved-mucronate; margin slightly thickened to very 
thickened, scabrid; upper surface flat to somewhat 
convex, the lower surface concave to flat or prom- 
inently 2—3-costate. Scape solitary, scabrous, 6— 
10 cm high, 0.5-0.7 mm wide, at base somewhat 
compressed to tricostate and sulcate, terete toward 
apex. Scape sheaths erect, linear-lanceolate, sub- 
ulate, convex, slightly striate, unequal, in 2 pairs, 
the lower pair basal, the upper pair ca. 2 way up 
from scape base, the outer sheaths 1-1.5 cm long, 
34— 56 as long as the inner sheaths. Spike narrowly 
turbinate, 1.1-1.4 cm long, few-flowered (flowers 
usually 4); bracts 2 pairs, subdecussate, lanceolate, 
thickened-subulate, 1-1.4 cm long, convex, cari- 
nate from middle to tip, scabridulous, obtuse at tip, 
medially green with margins broadly scarious-bor- 
dered. Sepals 2, lateral, subopposite, lanceolate, 
inequilateral, ca. ong, curvate, narrowly 
acute, sharply and scabridulously carinate from 
middle to tip. Corolla blue, ca. 1 cm long. Stami- 
nodia bibrachiate, the branches flat, narrowly ob- 
long, ca. 1 mm long. Anthers oblong, ca. 1 mm 
long; filaments ca. 1.5 mm long. Stylar appendages 
3, recurved, claviform, 1 appendage reduced. Cap- 
sule obovoid, ca. 2.5 mm long, obscurely trilobed, 
the valves inflexed and erose at apex. Seeds broadly 
asymmetrically obovoid, ca. 0.5 mm long, longi- 
tudinally strongly 12-14-ribbed, coal black. 


Paratype. | VENEZUELA. TERRITORIO FEDERAL 
AMAZONAS: on Rio Negro, Cerro Aracamuni, summit, 
Popa Camp, on edge of tepui in eas savanna, 01?26'N, 
65°47'W, 1,550 m, 17 Oct. , R. Liesner & F. 
Delascio 2207 1 (MO, VDB, a 

This species is related to 4. acicularis Idrobo 
& Smith, particularly var. granularis Maguire, 
but the foliage and scapes are rugose and scabrid 
(vs. smooth or granular), the sepal keels are sca- 
bridulous (vs. smooth or granular), the petals are 
blue (vs. white), and the seeds are smaller. 


Xyris aracamunae Kral, sp. nov. TYPE: Vene- 
zuela. Territorio Federal Amazonas: Dept. Rio 


976 Annals of the 
Missouri Botanical Garden 


FIGURE 20. Abolboda scabrida. —a. Habit. —b. Leaf tip. — c. Sector of leafblade, upper side. — d. Sector of leaf 
blade, lower side. —e. Leaf.— f. Spike and upper scape.— g. Involucral bract, side view (left), upper side (right). — 
h. Lateral sepal, outer side (left), side view (right). —i. Scape sheath pair, side view. —j. Style, showing appendages 
and stigma head. —k. Stamen. —1. Staminode. — m. Capsule and style base. — n. Seed. (From Liesner & Carnevali 
22598.) 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 977 


Venezuelan Guayana Flora— VII 


Negro, Cerro Aracamuni, summit, Proa Camp, 
savanna with small to large patches of forest, 
in savanna, 01?32'N, 65?49'W, 1,400 m, 25 
Oct. 1987, R. Liesner & G. Carnevali 22452 
(holotype, VEN; isotypes, MO, VDB). Figure 
21 


Herba perennis, densicaespitosa. Radices graciles. Caas 
i cta bs levite 


cicostatae; apices gradatim contracti, angus ste incurvo- 
acuti, incrassati, ventraliter Si. n leviter in- 
crassatae, dense albotomentos a bas apicem 
ventraliter sulcatae; vaginae ias: convexae, 


sae, eripere braste omens fir 


iin iue vel ovatae, 3-63 tiene cb ha d 
tilibus breviores et in eas urere co infimis ad 
i retrorse albohirsutae; bracteae o 
ovatae, 7-9 mm longae, humiliter « hah . Se- 
pala lateralia bera, x een lanceolata, ca. 6 

ala carinalis fambusciliats. lata, 
n obovatae 


firma. Lamin ol ul 5 mm a 
luteolae, fer grae. Staminodia bibrachiata, brachiis 
longe penicillatis. Antherae oblongae-lanceola 


O 
Im 
= 
= 
A 
77) 
dà 


a, 
leitet: spiraliter mulucostata; ad apicem pallide 
et obtuse apic ula 


Densely cespitose perennial herb. Roots slender. 
Stems short. Leaves stiff, erect to slightly spread- 
ing, narrowly linear, 30-55 cm long, strongly flex- 
uous and twisted; blades slightly compressed, in 
cross section elliptic, 1-2 times longer than the 
sheaths, 2-2.5 mm wide, distinctly few-nerved; 
apices gradually narrowed, narrowly incurved- 
acute, thickened, ventrally sulcate; margins some- 
what thickened, densely white-tomentose, the inner 
edge sulcate from base to tip; sheaths much elon- 
gated, convex, dull brown, papillose-rugulose, very 
open, gradually narrowing from base to tip, elig- 
ulate, the edges long-ciliate with pale, villous hairs. 
Scape sheaths with blades as in foliage leaves. 
Scapes linear, somewhat compressed, twisted, 4— 
7 dm high, densely white tomentose, in cross sec- 
tion elliptic, apically subterete, prominently sev- 
eral-costate. Spikes broadly obovoid to turbinate, 
obtuse, several-flowered, the bracts convex, firm, 


red-brown, spirally imbricate, entire; sterile bracts 
several, triangular to ovate, 3-6 mm long, sub- 
acute, shorter than the fertile bracts and grading 
gradually into them, the lowest bracts retrorsely 
white-hirsute at the acute apex; fertile bracts ob- 
long-ovate, 7-9 mm long, narrowly rounded api- 
cally; dorsal area distinct, triangular, 4-5 mm long, 
low-costate. Lateral sepals free, subequilateral, lan- 
ceolate, ca. 6 mm long, curved, acute; keel fim 
briociliate, broad, firm. Petal blades obovate, ca 
mm long, yellow, nearly entire. Stamin ae 
2-branched, the branches long-penicillate. Anthers 
oblong-lanceoloid, 2.5 mm long, strongly emargin- 
ate and sagittate; filaments ca. 1 mm long. Capsule 
ellipsoid, acuminate, ca. 4 mm long; placentation 
axile. Seeds numerous, cylindric-ellipsoid, ca. 
long, translucid, pale amber, longitudinally 
spirally multiribbed, at apex with a pale and obtuse 


— 


apiculus. 

Paratype. VENEZUELA. TERRITORIO FEDERA 
AMAZONAS: Dept. Río Negro, Cerro Aracamuni, Si 
i Yes e low savanna, 01*26/ 47'W, 1,550 

1, 18€ , R. Liesner & F. Delascia Y 2100 (MO. 
“DB. tg 
Xyris liesneri Kral, sp. nov. TYPE: Venezuela. 


Territorio Federal Amazonas: Dept. Rio Ne- 
gro, Cerro Aracamuni, summit, Proa Camp, 
savanna with small to large patches of forest 
especially along streams, in ravines and near 
edge of tepui, 01°32'N, 65?49'W, 1,400 m, 
25 Oct. 1987, R. Liesner & G. Carnevali 
22086 (holotype, VEN; isotypes, MO, VDB) 
Figure 22. 

a perennis dense caespitosa, caulibus multis, ram- 

ete 


ific Miri ramificationibus usque ad 3 cm longis, prae 
margines scaporum et apices foliorum glabra. Radices 


laminae planae 
vel fe rrugineae, inconspicue mute vosae, praete 
; gradati im c ste curvato-acuti, 


r apices 


atroferrugineis, nitidis; va 
ferrugineis, ciliolatis, later multicostatis, ferrugineis, 
basin versus fuscatis vel atro-ferrugineis, le in 
laminas gradatim convergentibus infimis gradatim 
ia aciebus integris. Vaginae 
iedio ad apicem valde r 
um ciliatis. Scapi d flexuosi, 20-30 cm longi, leviter 
i, ad apicem in sectione transversali 
bicostati, costis piloso-ciliatis, 


rugineis, spiraliter bihiak convexis; 
, decussatae, ovatae, carinatae, 
bracteae fertiles paucae, obovatae, ca. 4 mm longae; 


978 Annals of the 
Missouri Botanical Garden 


icm 


FIGURE 21. Xyris aracamunae. —a. Habit. —b. Leaf tip.— c. Sector of leafblade, midblade.— d. Leaf base.— 
e. Spike. — f. Fertile bract. —g. Lateral sepal. —h. Petal blade, stamen, staminode, and enlarged beard hair. — i. Ovary 
and stylar apparatus. —j. Median sector of capsule, offsets showing placentation, septation. —k. Seed. (From Liesner 
& Carnevali 22452.) 


Volume 76, Number 4 Steyermark, Holst & Collaborators 979 
1989 Venezuelan Guayana Flora—VIl 


FIGURE 22. Xyris liesneri. —a. aa —b. me Dp — C. d of leaf at E sheath junction. — d. Leaf base. — 


e. Spike, 1 Ter tile bract.—g. Sector of upper scape.— ateral sepal. —i. Petal blade and stamen (top); 
apex (left); staminode and beard te (below) —j. Pa valve outline, also mne placenta and funiculi. — k. 
(From Liesner & Carnevali 22086. 


=~ 


stylar 
Seed 


980 


Annals of the 
Missouri Botanical Garden 


apicem versus papillosae; area dorsalis i inconspicue ellip- 
tica obr io indistincte 


c 
5 
d 
c 
p 
a 
S. 
5 
o 


valida, usque ad 
ata id eds late obo- 

MU es prominenter laceratae. 

Antherae Sein iat 
itta 


5mm 
asali; s. cylindraceo- -ellip- 

mm ha. pallide et obtuse apiculata, atro- 

brunnea, a obsc e multinervosae. 


soidea, 


Dense cespitose perennial herb, the stems many, 
branching, the branches to 3 cm long, smooth 
except for leaf tips and scape edges. Roots slender. 
Principal foliage leaves narrowly linear-gladiate, 
10-20 cm long, spreading flabellately, longer than 
the scape sheaths; blades flat, 1-2 mm wide, com- 
pressed, yellow-green to reddish brown, inconspic- 
uously multinerved, entire except at tips; tips grad- 
ually narrowed, narrowly curved-acute, long-ciliate, 
the hairs reddish brown; margins with thickened, 
dark red-brown, shining edges; sheaths carinate 
the carinas dark red-brown, ciliolate, the sides mul- 
ticostate, red-brown, dull brown to dark red-brown 
toward base, the margins gradually narrowed into 
the blades, below gradually dilating, entire. Scape 
sheaths multicostate, strongly red-carinate from 
middle to tip, with short, red-ciliate blades. Scapes 
slender, flexuous, 20-30 cm long, slightly twisted, 
somewhat compressed, at tip in cross section nar- 
rowly elliptic, ancipital, bicostate, the costas pilose- 
ciliate, the hairs brownish. Spikes ellipsoid, 7-8 
mm long, obtuse, few-flowered, the bracts entire, 
nearly uniform, red-brown, spirally imbricate, con- 
vex; sterile bracts 4, decussate, ovate, carinate, 
ca. 3.5 mm long; fertile bracts few, obovate, ca. 
4 mm long, papillose toward tip; dorsal area in- 
conspicuous, elliptic, dark brown, slightly de- 
pressed, medially indistinctly 1-nerved. Lateral se- 
pals free, strongly inequilateral, lanceolate, 5 mm 
long, acute; keel strong to strongly ciliate up to 
the tip. Petal blades broadly obovate, ca. 5 mm 
long, yellow, strongly lacerate. Anthers lance-ob- 
long, 2 mm long, emarginate and sagittate, longer 
than ie EUR Staminodia with 2 long-peni- 
cillat apsule ellipsoid, ca. 2.5 mm long, 
dee the ea basal; seeds cylindric-ellip- 
, 121.3 mm long, with a pale and obtuse apic- 
E dark brown, longitudinally spirally multi- 


VENEZUELA. TERRITORIO FEDE 
AMAZONAS: Dept. Río Nn. Cerro Aracamuni, summit, 
Proa Camp, savanna with small to large patches of forest, 
m along streams, in savanna, 01°32'N, 
1,400 m, 25 Oct. 1987, R. Liesner & G. 
22454 (MO, VDB, VEN). 


, 
Carnevali 


This is a near relative of Xyris frondosa Ma- 
guire & Smith, a common high-altitude species of 
the tepuis of Estado Bolivar, particularly in habit 
and general leaf and spike dimensions. However, 
it is somewhat shorter-stemmed, its leaves are en- 
tire except at the very tips, and their margins more 
wirelike and thickened. The scapes, rather than 
being thick-edged and entire, are ancipital, the two 
costas thin, broad, and distinctly pilose-ciliate. 


Xyris prolificans Kral, sp. nov. TYPE: Venezuela. 
Territorio Federal Amazonas: Dept. Río Ne- 
die slope of Cerro Aracamuni, Quebrada 

mp, ea of rapids flowing over laja, 

01°24'N. 6538" W. 600 m, 20 Oct. 1987, 

R. Liesner & F. Delascio 22200 (holotype, 
VEN; isotypes, MO, VDB). Figure 23. 


erba perennis caespitosa, tenella, glabra, 
sulata, rhizomatosa. a 
flexuosa, saepe s 
veternarum rosularum foliorum prodeuntia internodiis ex- 
positis. Radices graciles. Folia principalia rosularu 
iter vel valde flabellate expansa, linea ri-lane eolata 
cm Ma gini 
valde compressae, 0.8-1.5 
longiores longitudinaliter indistincte nervosae, olivaceae, 
a medio ad apicem gra ua contractae; apices anguste 
et acute incurvati; margines integri, tenues; vaginae car- 
ae, tenues, lateribus longitudinale multicostatis, 


3-7 


ina 

marginibus in laminas gradatim convergentibus, apicibus 
li gua anguste. acuta -2 m longa ornatis, infimae 
gr i dilat „a p longe ciliatae. 


axae, apertae, tenues, ad apicem cuspidatae. Scapi sub- 
teretes, filiformes, spiraliter torti, 10-25 cm alti, ca. 0.4 
0.5 1 i 


O 


ali 
ussata satan pari. 
3 


oblongo-ovatae, 5 
area dorsalis viridi mm longa, ssa ndi d 
medigindistincte costatis. Sepala later 
teralia, curvata 
lata, a basi ad dion usque integra, a medio ad apicem 
usque indistincte lacerata. Laminae petalorum obovatae, 
3.5 mm longae, luteolae, late rotundatae, valde erosae 
Staminodia bibrachiata, brachiis dense longe penicillatis. 
Antherae ore i lanceolatae, ca. 
ginatae et sagittatae; filamenta ca. 0.5 mm longa. Capsula 
matura non visa sed ovario placentationibus marginalibus 
et funiculis elongatis, praedito. 


B. 


mm longae, emar- 


Cespitose, slender, smooth, lax, rosulate, rhi- 
zomatous herb. izomes numerous, slender, 
strongly zig-zag, usually branching, short-leaved, 
from the axils of older rosette leaves and with the 
internodes exposed. Roots slender. Principal ro- 
sette leaves slightly to very flabellately spreading, 
lance-linear, 3-7 cm long, longer than the scape 
sheaths; blades flat, strongly compressed, 0.8-1.5 


mm wide, 1—3 times longer than the sheaths, lon- 


Volume 76, Number 4 Steyermark, Holst & Collaborators 981 
1989 Venezuelan Guayana Flora— VII 


IMN 


FIGURE 23. Xyris prolificans. —a. Habit with inset showing outline of mature spike.—b. Leaf tip.—c. Leaf 
blade-sheath junction. —d. Leaf base.—e. Dry spike.—f. Fertile bract.—g. Lateral sepal.—h. Petal blade and 
stamen. — i. Staminode. —j. Ovary and style.—k. Capsule valve (construction) showing placentation. (From Liesner 


& Delascio 22200.) 


982 


Annals of the 
Missouri Botanical Garden 


gitudinally indistinctly nerved, olivaceous, gradu- 
ally narrowed from middle to tip; tips narrowly 
incurved-acute; margins entire, thin; sheaths car- 
inate, thin, the sides longitudinally multicostate, 
the margins gradually narrowed, at apex producing 
a narrowly acute ligule 1.5-2 mm long, gradually 
dilating below, long-ciliate at base. Scape sheaths 
lax, open, slender and thin, cuspidate. Scapes sub- 
terete, filiform, spirally twisted, 10-25 cm high, 
ca. 0.4-0.5 mm thick, olivaceous, striate. Spikes 
lanceoloid, drying turbinate, 7-9 mm long, few- 
flowered, the entire bracts with broad, scarious pale 
brown borders; sterile bracts ca. 6, subdecussate, 
ovate, the lower pair carinate, shorter than the 
inner pairs, 2.5-3 mm long; fertile bracts oblong- 
ovate, 5-6 mm long, convex, rounded; dorsal areas 
green, ca. 2 mm long, lanceolate-elliptic, medially 
with an indistinct rib. Lateral sepals free, inequi- 
lateral, curvate, 3-3.5 mm long, acute, the keel 
broad, entire from base to middle, from middle to 
apex low-lacerate. Petal blades obovate, 3-3.5 mm 
long, yellow, broadly rounded, strongly erose. 
Staminodia bibrachiate, the branches densely long- 
penicillate. Anthers oblong-lanceolate, ca. 1 mm 
long, emarginate and sagittate; filaments ca. 0.5 
mm long. Mature capsule not seen, but the ovary 
with marginal placentation and elongate funicles. 


The growth habit of this species is somewhat 
unusual. From the limited material available one 
concludes that rosettes produce either scapes or 
rhizomal buds from older axillae. The rhizomes are 
elongate, zig-zag, and with short, evenly separated 
leaves. The offshoots frequently branch and ulti- 
mately produce new rosettes at the tips, the whole 
proliferating product presumably forming extensive 
clones. Mature seeds would be necessary to estab- 
lish the relationship of this to others of the section 

ris 


PRELIMINARY CHECKLIST OF 
THE VASCULAR PLANT FLORA 
OF CERRO ARACAMUNI 


The checklist has been arranged into two groups: 
pteridophytes and seed plants. Within each group, 
families are arranged alphabetically and are divided 
up into the four main areas of collection in the 
following order: Summit, East Escarpment, Que- 
brada Camp, and Granitic Dome. Within each of 
the collecting areas, genus and species are listed 
alphabetically. Each species is followed by the name 
of the principal collector and number. 

Details of the camp and collecting locations, 
along with a list of the co-collectors with their 
number ranges and dates of collection, are given 


SUMMIT 
The Dunsterville collections give bg coordinates: 
01°30'N Lat., 65%51'W Long., 1,510 


G.C.K. & E. Dunsterville s.n., Dec. 1972 


Brewer Carias gives only elevation, 1,500 m. 
Charles Brewer Carias s.n., 13 Dec. 1983 


The 1987 expedition had two summit camps located 
at extreme ends of the massif: Popa Camp, situated at 
01°26'N Lat., 65?47'W Long., 1, 530 m, € p aa 
situated at 01%32'N Lat., 65?49'W Long., 
Carnevali & Santana 2362-2457, Popa Camp, 27-30 
Oct. 1987 
Carnevali & d 
ov. 
Liesner & Dalasi 21934-22182, Popa Camp, 15-19 
Oct. 
Liesner E M 22407-22 
( 3 Nov. 1987 


456-2584, Proa Camp, 30 Oct.- 


2730, Proa Camp, 25 


EAST ESCARPMENT 
,65?48'W Long., 750 m. Habitats here 


consisted of nan thickets, exposed rock, and low dense 
Orest. 
Huber & Medina 5892-5942, 10 Feb. 1981 


QUEBRADA CAMP 
01°24'N Lat., 65*38'W Long., 500-600 m. Located 
th 


along a stream slightly south and east of the summit in 
low open forest on a smooth, rocky outcrop near rapids. 
Liesner & Delascio 22183-22270, 20-21 Oct. 1987 
Liesner & Carnevali 2227 122406, 21-24 Oct. 1987 


GRANITIC DOME 


Collections from this site were made by Wurdack and 
Adderley from a 900-m granitic dome with low herbaceous 
vegetation on "pun rock and scrub forest. The dome is 
on the right side of Río Siapa, just below the rapids, 
Raudal Callineta. 


Wurdack & Adderley 43538-43572, 21 July 1959 


PTERIDOPHYTA 
Adiantaceae — Alan R. Smith (UC) 


Summit 


Pterozonium scopulinum Lellinger — Liesner 22569 


Aspleniaceae — John Mickel (UC), Elaphoglossum; Alan 

R. Smith (UC), Asplenium 

Summit: 

Asplenium serra Langsd. € Fischer —Liesner 22727 

dd lis Si plumosum (Fée) T. Moore — Liesner 
2 907, 22 90 

Ean eae produc ba ee 
22009, 22555, 22615, 22616, 2266 

Quebrada camp: 

Elaphoglossum glabellum J. Smith—Liesner 22277 

Elaphoglossum obovatum Mickel— Liesner 22343. 


Volume 76, Number 4 


Steyermark, Holst & Collaborators 983 


Venezuelan Guayana Flora— VII 


Cyatheaceae — Alan R. Smith (UC) 


Summit: 
Cyathea demissa (C. Morton) Lellinger — Liesner 
21997, 22601 


P Anus platylepis (Hook.) Domin — Liesner 22482 


Cy athec aff. williamsii (Maxon) Domin — Liesner 
21966, 22014, 22660 


Quebrada camp: 
C yathea macrosora (Baker) Domin 2 eee nae (Win- 
) A. R. Smith — Liesner 22 


Davalliaceae — Alan R. Smith (UC) 
Sum 
O ded articulata (Sw.) C. Presl — Liesner ~ 
Oleandra duidae A. C. Smith— Liesner 2272 


Dennstaedtiaceae — Alan R. Smith (UC) 


Summit 

Lindsaea sp. —Liesner 21978 

Lindsaea stenomeris Kramer — Liesn 

Lindsaea stricta (Sw.) S s var. pa er 
ramer — Liesner 22539, 22591 

L indsaea um ta (Sw.) Sea var. stricta — Liesner 


Pe camp: 
Lindsaea cyclophylla Kramer — Liesner 22264 


Gleicheniaceae — Alan R. Smith (UC) 


Summit 


Dicranopteris flexuosa (Schrader) L. Underw. — Lies- 
ner 21962 


Dicranopteris pectinata (Willd.) L. Underw. — Liesner 
22567 


Quebrada camp: 
Dic iir eM pectinata (Willd.) L. Underw. — Liesner 
2230 


Grammitidaceae — L. E. Bishop (UC); Alan R. Smith (UC) 


Sum 

C m —— (Desv.) L. Bishop — Liesner 22620 

Ceradenia deed yeri (Baker) L. Bishop— Liesner 

270 

C Viadon a (Mett.) L. ee 

Cochlidium connellii (C. H. Wright 
Liesner 22576, 2263 

Cochlidium serrulatum (Sw.) L. Bishop— Liesner 
21994, 22092, 22574 

d tepuiense (A. C. Smith) L. Bishop — Lies- 


- Liesner 22724 
A. C. Smith — 


47 
GIyphotaenium campbellii (Baker) Copel. — Liesner 
22 22685 


Crit bry ae (Maxon) F. Seymour — Liesner 
22; , 22619 
Grammitis melanosticta (Kunze) F. Seymour — Lies- 
ner 2267 
Grammitis taenifolia (Jenman) Proctor — Liesner 
22509 
Grammitis aff. williamsii (Maxon) Lellinger — Liesner 
22662 
Quebrada camp: 
Cochlidium serrulatum (Sw.) L. Bishop— Liesner 
22299, 22301 
Hymenophyllaceae — Alan R. Smith (UC) 


Summit: 
Hymenophyllum elegans Sprengel — Liesner 22552 


Hymenophyllum fendlerianum Sturm in C. Martius — 
Liesner 22668 

Hymenophyllum ferax Bosch — Liesner 22524, 22526 

Hymenophyllum karstenianum Sturm — Liesner 
22528 

Trichomanes crinitum Sw. —Liesner 21972, 22566, 


Trichomanes cf. pelluc ens Kunze — Liesner 22548 
Trichomanes sprucei Baker — Liesner 22434, 22435, 
22551 


Quebrada camp: 
Trichomanes ankersii Parker ex Hook. & Grev.— 


Liesner 22398 
Hymenophyllopsidaceae — Alan R. Smith (UC) 


Summit: 

Hymenophyllopsis hymenophylloides L. D. Gómez — 
Liesner 22568 

Hymenophyllopsis superba Lellinger — Liesner 22432 


Lycopodiaceae — Alan R. Smith (UC) 


Summit: 
Lycopodium funiforme Cham. ex Spring — Liesner 
22721 


Lycopodium iuliforme L. Underw. & Lloyd — Liesner 
22175, 22491 


Quebrada camp: 
— Liesner. 22201 
D ann linifolium L.—Liesner 22351A 


Lycopodium 


ee R. Smith (UC) 
Quebrada camp 


Metaxya rostrata (H.B.K.) C. Presl — Liesner 22361 


Polypodiaceae — Alan R. Smith (UC) 


mmit: 
Polypodium laevigatum Cav. — Liesner 22691 
Polypodium triseriale Sw.— Liesner 22083 
Quebrada camp: 
Microgramma baldwinii Brade — Liesner 22297 


Schizaeaceae — Alan R. Smith (UC) 


Summi 

Schizaea — s (M. Vahl) Sw. 

mos aea poeppigiana Sturm in C. 
2171 


Liesner 22658 
Martius — Liesner 


— camp: 
tachys pennula (Sw.) on — Liesner 22184A 


Actino 
Schizaea elegans (M. Vahl) S Liesner 22259, 
22290 


Schizaea stricta Lellinger (mixed collection with Ac- 
84 


tinostachys pennula)— Liesner 221 


Selaginellaceae— Alan R. Smith (UC) 


Summit: 
Selaginella rhodostachya Baker — Liesner 22553 
Tectariaceae— Alan R. Smith (UC) 


Quebrada cam 
Triplophyllum di caladas (Fée) Holttum — Liesner 
22260, 2235 


SEED PLANTS 
Annonaceae — Paul J. M. Maas (U), 


Anaxagorea 


984 Annals of the 
Missouri Botanical Garden 


Quebrada Quebrada camp: 
Anaxagorea brevipes s a 22398A Anthurium a, EE subsp. rionegrense 
Duguetia sp. — Lies 228 unting 
Apocynaceae — Bruce did pa Odontadenia; A. Hereropsis Aaa var. T Bunting — Liesner 
Leeuwenberg (WAG), dado 21926 
Friedrich Markgraf (Z), Parahancornia; James Za- Heteropsis presan Schott. var. spruceana — Lies- 
rucchi (MO), misc.; G. Morillo (VEN), Mandeville ner 22832 
ics ndron englerianum m subsp. duidae 
Sumn Steyerm.) Bunting — Liesner 22406 
As Serm Ime sp: pea 22042, 22167 Philodendron maguirei ae Liesner 22243. 
Couma rigida Muell. — Liesner 22696 er also from Cerro Arauicaua and Cerro de la 


oe se parse ma Woodson — Liesner 
Philode on cf. maroae Bunting—Liesner 22332. 


Mandevilla aracamunensis Morillo — Brewer Car- he determination is in question because of the pau- 
las s.n. (type) city of material and the lack of data concerning the 
East escarpment: uad shape; until now d maroae vnd been known 
Mandevilla annularüfolia Woodson. — Huber 5895 only from lowland s 
Parahancornia krukovii Monach. — Huber 5928 Sc hismatoglottis i aane Banag & fear — 
192. A rath 


Parahancornia negroensis Monach. —Huber 5911 


parently th record outside of liva 
Quebrada camp: The specimen is identical to other material except 
Couma n Muell. Arg. Pry 22400 for the greater size of the leaf blade (45 cm long, 
Galactophora sp.— Liesner 22403 32.5 cm wide). 
Lacmellea gs ner 22 206 Stenospermation cf. ulei K. Krause — Liesner 22320. 
Mandevilla spp. —Liesner 22216, 22321 Specimen sterile but oli a to this 
Odontadenia neglecta Woodson — Liesner 22296 widespread species with nodding inflorescence 


Prestonia sp. — Liesner 22279 
p dip dai n C. Martius ex Muell. — Arecaceae — Andrew Henderson (NY) 
Arg. (?) — Liesner 55 


Summit: 
Aquilone — Julian icd (MO) Euterpe montis-duidae Burret ex Gleason — Liesner 
Summit: 22120 
Ilex aracamuniana Steyerm.— Liesner 22060 (para- Quebrada camp: 
type), 22181 (paratype), 22413 (type Bactris sp. (sterile) — Liesner 22256 
Illex retusa Klotzsch — Liesner 22109, 22043 Bactris simplicifrons C. Martius— Liesner 22283 
llex tateana nd — Liesner 22533 actris cf. simplic d C. Martius— Liesner 22258 


Iriartella setigera (C. Martius) H. A. Wendl. — Liesner 
22261 


East escarpmen 


llex divaricata C Martius ex Reisseck — Huber 5896 


A ae—G s. ing f - y ae 
raceae— George 5, Bunting Asclepiadaceae — Gilberto Morillo (VEN) 
Summit: , 
: : oon Summit 
Anthurium acm Steyerm. —Liesner 22024, c 99491) 99759 
22039. 22684 : Blepharodon sp.— Liesner 22420, 22752 
pin Matelea neblinae Morillo —Liesner 22164, 22503, 
Phälodendron 73 K. Krause ae ptarianum 99795 
79 í 
erm.) Bunting — Liesner 22672, 22720. Sterile 
pi feni specimen ms slender, exce stem A ; ‘ 
ES lca . ? S — Asteraceae — John Pruski (NY); Victor Badillo (M Y) 
Philodendron roraimae K. Krause subsp. aracamu- Sun 
nse Bunting — Liesner 22502, 22510, 21956 TuS huachamacari Maguire € Wurd. 
Philodendron tatei K. Krause subsp. iore hlorum Liesner 21960 
(Bunting) Bunting —Liesner 22178A, 22487, Ime ria idi ps rd & Wurd.) King & H. 
22704. Described originally from collections from m— Liesner 21983 
the Gran Sabana, similar material has subsequently Mikania lucida H. L. Blake f. lucida — Liesner 22449, 
been collected in Territorio Federal Amazonas, as 
well as some plants nearly intermediate between this uS haeta Mum His (H. L. Blake) Maguire & 
and subsp. tatei that demonstrate the close rela- 'urd. — Liesner 2208 
tionship of these taxa. Four collections from Cerro TN 
racamunl appea represent subsp. melano- Past escar pment 
PP P P Gongylolepis martiana (Baker) Suia & Cuatre- 
chlorum, which has a much narrower leaf bla c 
: casas — Huber edina 592 
than subsp. tatei. Of these, Liesner & Delascio 
Erga Imeria serratifolia V. Badillo Har ie (type) 
221784, taken from an immature plant, has ovate 
EM campestris Maguire & Wurd.— Huber 
lea des ee at base in contrast to the 
usually elliptic blades with obtuse or acute bas 
relevance of this variation is unknown. Quebrada camp: 
c o e E nebulense Bunting — Liesner a (Praxelis) asperulaceum Baker — Liesner 
2437. The t collection is from Cerro de la 2190 


Neblina at l, 500 1 m. 
Stenospermation ulei K. Krause — Liesner 22676 Bignoniaceae — Alwyn Gentry (MO) 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora— VII 


Summit: 

Digomphia densicoma ( 
Liesner 22106 

Distictella monophylla Sandw. —Liesner 21954 


C. Martius ex DC. 


— 


Pilger — 


East escarpm 
Distictella Ae Sandw.— Huber 5892 


Quebrada camp: 
Pleonotoma jasminifolium (H.B.K.) Miers — Liesner 
22246 


Schlegelia spruceana Schumann — Liesner 22399 
Bombacaceae — Julian Steyermark (MO) 


Summit: 

Pochota aracamuniana Steyerm.—Liesner 22174 
(paratype) 22559 (type), 22577 (paratype) 

Pochota cf. ewelii Steyerm. — Liesner 22444, 22664 

Quebrada camp: 

Pochota sp.— Liesner 22390 


Bromeliaceae — Lyman B. Smith (US), with Julian Stey- 
ermark (MO) 


Summit: 
Brocchinia acuminata Lyman B. Smith — Liesner 
22027 
Brocchinia hitchcockii Lyman B. Smith — Liesner 

0 
Brocchinia micrantha (Baker) Mez (ovaries variable 
in form) — Liesner 22000 
Brocchinia vestita Lyman B. Smith — Liesner 22028 
I HE pec — A 22706 
a Lyman B. Smith — Liesner 22057, 


s 
Lindmania aff. Lp illima Lyman B. Smith, (leaf spines 
large)— Lies 22440 


Nas aps c ner 22093 


Navia patria d B. Smith & Steyerm. — Liesner 
22041 (ty 

Navia platyphylla Lyman B. Smith & Steyerm. — 
Liesner 22002 (type) 


ea terramarae Tons B. Smith. & Steyerm. — 
Liesner 21998 (type), 22599 (paratype 
Tillandsia confinis Lyman B. Smi th var. caudata Ly- 
man B. Smith — Liesner 22446 
Tillandsia ook iR nee B. Smith var. confinis — 
5 


Liesner 22019, 
Pillans rude | um var. spiculosa — Liesner 
, 22442 
Vriesea eapitulige ra om ) Lyman B. Smith & Pit- 
dr. — Liesne 678 


ten 

Vriesea duidae (L yman B. Smith) Gouda (det. G. Car- 
nevali)—Carnevali 2460, 25 

Vriesea incurva (Griseb.) Read — Liesner 22085 


Quebrada camp: 

Aechmea brevicollis Lyman B. Smith — Liesner 22345 

Aechmea politi Lyman B. Smith (leaf blades atten- 
uate)— Liesner 22363 

Ananas parguazensis Camargo & Lyman B. Smith — 
Liesner 22394 

Brocchinia paniculata Schult. f. (7) — 

an B. Smith & Steyerm. 


Liesner 22319 
Navia carnevalii Lym: - Lies- 
ner 22322 (ty ype) 
Vavia aa Lyman B. Smith & Steyerm. — Lies- 
ner 22203 (type) 
Pitcairnia patentiflora Lyman B. Smith — Liesner 
2198 


Tillandsia paraensis Mez —Liesner 22303 


Vriesea socialis Lyman B. Smith — Liesner 22294 

Granitic dome: 

Brocchinia acuminata Lyman B. Smith — Wurdack 
435 

Ur tatei Lyman B. Smith — Wurdack 43563, 
435 

s patentiflora var. armata Lyman B. Smith — 
Wurdack 4 

Tillandsia confinis Lyman B. Smith — Wurdack 43501, 
43562 


Vriesea fibrosa Lyman B. Smith — Wurdack 43569 
Vriesea platynema Gaudich. var. platynema — Wur- 
dack 43568 


Burmanniaceae — Paul J. M. & H. Maas (U) 


Summit: 

Dictyostega orobanchoides (Hook. U^ Mens subsp. oro- 
banchoides — Liesner 22004 

Hexapterella ME P. Maas & H. Maas- 
Liesner 22622 


, 2242 


Quebrada camp: 
de paie (Nutt.) Barnhart ex Small — Liesner 


iu Du ega orobanchoides (Hook.) Miers subsp. par- 
viflor li 1.) Snelders & P. Maas— Liesner 
22 253, 402 


ee Daly (NY) 


Sumn 


Not determined to genus — Liesner 22072, 22465, 
22626 


Bis camp: 
Dacryodes glabra (Steyerm.) Cuatrec. — Liesner 22187 


Caesalpiniaceae — Richard S. Cowan (PERTH); Rupert 
Barneby (NY) 


East escarpmer 

add ua e pm Sprague ex Sandw.— Huber 
5926 

Jacqueshuber .— Huber 


Macrolobium de olor Benth 


m 5917 
Quebrada camp: 
Macrolobium sp.— Liesner 22196, 2 
Chl lx 


Sun 
Hedman vel aff. gentryi D'Arcy E Liesner — Lies- 
r 22438, 22472, 22517, 2253 


N 
w 
wn 
=J 


Carol Todzia (TEX) 


Chrysobalanaceae — Ghillean Prance (K) 
Quebrada cc 
Hirtella punctidaka Ducke 
Hirtella schultesii Prance- 


Liesner 22295 
Liesner 22360 


Clusiaceae — Bassett Maguire (NY); Barry Hammel (MO) 
Summit 
Cas aff. lopezii Maguire — Liesner "ie 22575 
Clusia wat hiorii d sner 221 22717 


Clusia pusilla Steyer - Liesner 2: 

Clusia sessilis Klotzsch ex Engler — Liesner. 22021, 
85 

Eder ye obovatus Spruce ex oo & 
na—Liesner 22111, 22543, 226 


East escarpment: 


Clusia sp.— Huber 5922 


986 


Annals of the 
Missouri Botanical Garden 


Granitic dome: 
Clusia pusilla Steyerm. subsp. orinocensis Maguire — 
Wurdack 43555, 43556, 43557 

Quebrada camp: 

Clusia columnaris En E Liesner 22225, 22233 

Clusia aff. gratula Ma Liesner 22344 

Clusia aff. penduliflora Engler —Liesner 22298 
Cunoniaceae 


Sum 

us spp.— Liesner 21935, 22098, 22429, 
22641 

Cycadaceae 


Viri opis camp: 
s Liesner 22367 


— 
= 


Cyclanthaceae — Barry Hammel (MO) 
Summit: 

Sphaeradenia rubescens Harling—Liesner 22447, 
22587 


Quebrada camp: 
Dicranopygium imeriense Harling — Liesner 22309 
Cyperaceae — Gerrit Davidse (MO) 
Summit: 
Didymiandrum stellatum (Boeckeler) Gilly — Liesner 
22015 


Rhynchocladium steyermarkii (T. Koyama) T. Ko- 
yama — Liesner 21941 

Quebrada camp: 

Calyptrocarya glomerulata (Brongn.) Urban — Lies- 
ner 22364 

Cephalocarpus confertus Gilly — Liesner 22333 

Hypolytrum pulchrum (Rudge) Pfeiffer — Liesner 
2288 


Mapania ar rar a Koyama & Steyerm. ex T. 

Ko asia 22241 

n c micrococca T. Koyama — Liesner 22193 

Rhynchospora rigidifolia (Gilly) T. O bou 
planifolia T. Koyama — Liesner 22224, 


£g 
= 
Hd 
5 


ae — Julian Steyermark (MO) 
Sum 
Cyrilla. pm L.- Liesner 
73 


Carnevali 2365; 


hose nutans Planchon — Carnevali 2548; Liesner 
22711 

Dilleniaceae — Klaus Kubitzki (HBG); Ronald Liesner (MO) 

East escarpme 

Doliocarpus savannarum Sandw.— Huber 5902 

Quebrada camp: 


Doliocarpus savannarum Sandw.—Liesner 22 
22276 


185, 
Dioscoreaceae 


mit: 
d Nos —Liesner 22415, 
10 


22436, 22484, 
22541, 
Droseraceae — Bruce Holst (MO) 
Summit: 
Drosera Klotzsch ex Diels) Maguire & Laun- 


don— Liesner 22687, 22126 


Quebrada camp: 
Drosera roraimae o un ex Diels) Maguire & Laun- 
don— Liesner 22 
dee MM (MO) 
Summit: 
Sloanea steyermarku C. E. Smith subsp. pera 
(Steyerm.) Steyerm. Sterile, but probably to 


ferred here. —Liesner & Carnevali 22722 


Ericaceae — James Luteyn (NY) 


Sum 


Cav iuc) callista J. D. Smith, s.l. — Liesner 22536 


show an interesting transition m the type collec- 
tion at Neblina with a short racemose inflorescence, 
to recent Neblina collections whore the inflorescence 
ery short raceme, to 


em to be equal. The een is pk 

stc e m Cerro de la d 

Satyria carnosiflora Lanj.— Liesr r 2252 

dein. Ae cordifolium Benth. ae 21979, 
22563, 22686 

Thibaudia cupatensis H. E. Huber 
y 22579 


- Liesner 21946, 


Quebrada camp: 
Rica. callista | Donn. Sm., 
Satyr h.) B 
32323 


s.l. — Liesner 22272 
sath & Hook.— Liesner 


Thibaudia cupatensis H. E. Huber — Liesner 22189 
e INQUE Hensold (MO) 

Sun ] 
e sulcatus Hensold — Liesner 21940 
Syngonanthus phelpsae Moldenke — Liesner 22443 
East escarpment: 
Syngonanthus anomalus (Koern.) Ruhl — Huber 5934 
Quebrada camp: 
Paepalanthus capillaceus Klotzsch ex Koern. — 

22308 


ner 22 


Lies- 


Syngonanthus biformis (N.E. Br.) Gleason — Liesner 
22199 


Euphorbiaceae — Michael Huft (MO); Grady Webster 
me 


Sun 

oa ed (Tul.) Muell. Arg. var. oblonga — 
Liesner 226 

Phyllanthus i Jabl. —Liesner 21947, 
22032 


anadenus 


Phyllanthus vacciniifolius (Muell. Arg.) Muell. Arg. — 
984 


Liesner 2 
Sue cd yutajensis (Jabl.) Webster — Liesner 
22625 


Senefelderopsis sipapoensis Jabl. —Liesner 22466, 
22475 


Quebrada camp: 

Pera bicolor (Klotzsch) Muell. Arg. —Liesner 22238 
Granitic dor 

Phy lana lt Jabl.— Wurdack 43550 
Phyllanthus subapicaulis Jabl.— Wurdack 43549 


Fabaceae — James Zarucchi (MO) 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 987 


Venezuelan Guayana Flora— VII 


| ali camp: 

Clito sp.—Liesner 22404 

nsa sp.— Liesner 22234 

Swartzia sp.— Liesner 22271 
Flacourtiaceae — Julian Steyermark (MO) 


East escarpmen 
Laetia coriacea pm ex Benth.- 


—Huber 5920 

Quebrada camp 

Hi atda c. Mart 

Laetia coriacea Spruce ex Bent 
48 


ius— Liesner 22300 
h.—Liesner 22284, 


223 
odd t var. subulifera (Sandw.) Monach. — 
Liesner 22380 


Gentianaceae — Paul J. M. Maas (U), in part 


Celiantha bella Maguire & Steyerm. — Liesner 22126 
anthus aracamuniensis Steyerm. — Liesner 


eo ^ E Benth. — Liesner 22082, 


22585, 


East esc arpment 


Irlbachia alata (Aubl. ) P. Maas subsp. alata — Huber 
5940 


Quebrada camp 
Irlbachia alata À Aubl. ) P. Maas subsp. alata — Liesner 
22305 


Tachia grandifolia Mag. & Weaver var. orientalis 
Maguire & Weaver — Liesner 22254, 223 

Tapeinostemon Vin i Benth. —Liesner 22082 
Voyria aphylla (Jacq.) Pers. — Liesner 223: 
Voyr ria (?) sp. — Liesner 99952 

Granitic dom 

Eua ine as (H.B.K.) Gilg — Wurdack 

3552 
-Wurdack 43554 


Chalonanihis uliginosus Griseb. 
Gesneriaceae 


'ummit: 
Not determined to genus —Liesner 22525, 22530, 
22679 


anes camp: 
Not ao to genus — Liesner 22324, 22346 
Codonanthe sp. — Liesner 22389 
Humiriaceae — Ronald Liesner (MO) 
Sum 
rod balsamifera oe Standley — Liesner 
226 


22410, 22589, 


Icacinaceae — Richard A. Howard (A) 


Summit: 
d idus a ex Miers — Carnevali 2583; 
Liesner 21953, 22070 


Lauraceae — Henk van der Werff (MO) 


Sum 
dude y Allen — Liesner 21967, 22065. 
e type of this species is cn Cerro Sipapo, 
ritorio Federal Amazon Aniba vinnamonifore 
has > been reporte fon lowlands in Estados 
Aragua, and Táchira. Distribution and habit 
(a pun or small tree fro r d Ara 
camuni, large timber tree inn the other localities) 


suggest that tions oi > Territorio Federal 
Amazonas a not belong in 20 cinnamomiflora; 
additional collections are needed to solve this prob- 
em. 

Endlicheria vinotincta Allen — Liesner 22008 (9), 
22414 (8), 22292 (8). On the summit and slopes of 
Cerro Aracamuni, 600-1,500 m. Previously known 
from two collections from Cerro Neblina. Liesner & 
Carnevali 224 14 and 22292 2 are the first staminate 


rff — Mae 22161 (type), 
22451 (paratype), 227 14 (par 

Persea perseiphylla (Allen) van re Werf — Liesner 
22099, 22169 


pie dien. camp: 
niba sp. —Liesner 22374 


Lentibulariaceae — Peter Taylor (K) 
Summit 


Genlisea guianensis N.E 

Genlisea repens Beuj.- 

Utricularia alpina Jac 

Utricularia amethystina A. St. Hil.- 
0 


[; 


. Br. — Liesner 22488 
Liesner 22573 

q.--Lesner 2197] 

Liesner 22051, 


Liesner 21950 
21992, 


Utricularia humboldtii Schomb. - 

Utricula jamesoniana | Oliver — Liesner 
2201 

Utricularia neottioides A. St. Hil. — Liesner 21995 

Liesner 2 22565 

P 'sner 21989 


Utricularia triloba Benj. Lena s í 


East escarpment: 
Genlisea pygmaea A. St. —Huber 5935 
Genlisea repens Benj. — Huber 5936 


— 
— 
mt + 
-— 


Quebrada cl 
Genlisea pygm St. 
Utricularia canal Oliver 


E 


Liesner 22195 
Liesner 22370 

Granitic dom 

Utricularia humboldtii Schomb. — Wurdack 43560 


Liliaceae — Robert Cruden (IA) 


Summit 
Tofieldia schomburgkiana Oliver — Liesner 22469 
Loganiaceae — Rupert Barneby (NY) 


Quebrada camp: 


Strychnos guianensis (Aublet) C. Martius — Liesner 
2405 


Loranthaceae 


Summit 
Not determined to genus — Liesner 


22] 
22630, 22455, 22544, 22561, 2262 


55. 22170, 
8, 22729 
Quebrada camp: 


Not emo to genus — Liesner 22084, 22191, 
393 


22293, 

Lythraceae — Alicia Lourteig (P) 
Summit: 

Cuphea annulata Koehne 


East escarpment 


pment: 
Cuphea annulata Koehne 


Liesner 21948 


Huber 5906 


988 Annals of the 
Missouri Botanical Garden 


Quebrada camp: Pachyloma huberioides (Naudin) Triana — Huber 5903 
Cuphea annulata Koehne — Liesner 22207 
Quebrada cc 
Malpighiaceae — William R. Anderson (MICH) Adelobotrys rotundifolia Triana — Liesner 22378 
Adelobotrys saxosa Wurd. — Liesner 22285 


Summit: 
Banisteriopsis a Gates — Liesner 22162 
Byrsonima amoena Cuatrec. — Liesner 22422, 22578 


"llidemia heteroneura (DC.) fon — Liesner 22395 
Clidemia novemnervia (DC.) Triana — Liesner 22306 


Diacidia pee ed (Magi e) W. R. Anderson — Macairea thyrsiflora DC.— Liesner 22188 nm 
Maieta poepp igii C. Martius ex Cogn. — Liesner 22831 
Liesner 21963 À 
Miconia sp.— Liesner 22379 
East escarpment Miconia cautis Wurd. Tm 22242, 22795 
Blepharandra m (Benth.) Griseb. — Huber Miconia marginata Triana — Liesner 22213 
5 Miconia rugosa Triana — Liesner 22 37] 
Heteropterys huberi W. R. Anderson — Huber 5893 Miconia splendens (Sw.) Griseb. — Liesner 22758 


Miconia cf. traillii Cogn. — Liesner 22830 


Marantaceae — Lennart Andersson (GB), /schnosiphon 
" (GB) inc aid sane mac rosperma (C. Martius) C. Martius — 
26 


Quebrada camp: Liesner 22326 

Ischnosiphon surumuensis Loesn. — Liesner 22255 Buchs lena huberioides (Naudin) Triana — Liesner 

Ischnosiphon puberulus Loesn. — Liesner 22377 22215 

"nd sp. — Liesner 22249, 22327 [Note: this Tococa macrophysca Spruce ex Triana — Liesner 
w species will be described soon in the Annals.] 22286 

Melastomataceae — John J. Wurdack (US) Granitic dome 

Summi Adelobotrys saxosa Wurd. — Wurdack 43543 

jo l AM Miconia ciliata (Rich.) DC. — Wurdack 43544 

Clidemia tepuiensis Wurd. — Liesner 22070 ficonia ciliata (Rüch.) 


e i: cf. (ur ee poo subsp. neblinensis Mimosaceae — Rupert Barneby (NY) 
— Liesner 22423, 22699, 22730. The 


Summit: 
camuni collections, mostly i in young bud, differ from 


Abarema arenaria (Ducke) — Liesner 22113 


surfaces with very attenuated pubescence and very Quebrada camp: 

faint secondary nerves, obscure pedicels only 0.5 Calliandra sp. — Liesner 22304 

l mm long, and smaller flowers (petals ca. 8 x 5.5 Inga sp.— Lies 11 

mm, anthers 3.5-4.5 mm long with basal spur l- Zygia basijuga (Ducke) Barneby & Grimes — Liesner 
1.3 mm long). I am reluctant to desc ribe a new 22269 


ne ' À l Monimiaceae 
Macairea theresiae Cogn. — Liesner 21949. Other- 
wise known from lowland white sand savannas of Quebrada camp: 
the middle and lower Amazon and Rio Negro basins Mollinedia sp. —Liesner 22270 


in Brazil, the closest locality being the Rio Araca 
(fide Renner Mss. 


Moraceae — C. C. Berg (BG) 


Maieta neblinensis Wurd.— Liesner 22068, 22583. Summit: 
Otherwise known only from the upper slope forests Coussapoa sp. —Liesner 22180 
of Cerro Neblina in Venezuela and Brazil, 650- Ficus sp. —Liesner 22160 
300 m. 
Miconia curta (Gleason) Wurd. — Liesner 22688 Myrsinaceae-John Pipoly 


Miconia dodecandra (Desr.) Cogn. — Liesner 22166 


Summit: 
Miconia roraimensis Ule— Liesner 22150, 22523, 
22663 


n y qe to E Liesner 21936, 22416, 
86, 2272 
Neblinanthera cumbrensis Wurd. —Liesner 21937, 


22657. Otherwise known from the upper slopes of Bos escarpment: 
cera ia in Venezuela and Brazil, 1,300- Cybianthus deltatus Pipoly — Huber 5910 
1,800 n 


Quebrada car 


eee pusilla (Gleason) Wurd.— Liesner 21986, Not ea ml a to genus — Liesner 22273 
987, : 


rwise restricted to the summit 
a upper slopes of Duida and Neblina, often a quasi- Myrtaceae — Bruce Holst (MO) 


rheoph 


, Summit 
ias aidas Gleason — Liesner 22627 f s pee . 
p. : "iden > : we Blepharocalyx d Cie (Kiaerskov) Landrum — Lies- 
Tibouchina dissitiflora Wurd.— Liesner 22080 E 
E : iris nnd ner 22061, 22588. Known previously in Venezuela 
Tococa liesneri Wurd.— Liesner 22007 (paratype), 

22700 (type) only from Isl sla Margarita in — Nueva Esparta, 


and northeast Estado Boliv 

Eugenia sp. — Liesner 22680 

Marlierea summa McVaugh var. summa — Liesner 
22002, 22419 


Tococa pachystachya Wurd.— Liesner 21938. Oth- 
erwise known only from cloud iren of i upper 
slopes of Cerro Neblina, 1,450- On 


East escarpment: Myrcia bolivarensis (Steyerm.) McVaugh — Liesner 
Macairea lanata Gleason subsp. lanata — Huber 5897 22581 
Macairea thyrsiflora DC.— Huber 5931 i rcia revolutifolia McVaugh — Liesner 22097, 


Meriania urceolata Triana — Huber 5901 2702 


Volume 76, Number 4 
1989 


Steyermark, Holst & Collaborators 
Venezuelan Guayana Flora— VII 


Quebrada camp: 
por sp. — Liesner 22330 
Myrcia revolutifolia McVaugh — Liesner 22274 


Granitic dom 
Myrcia eet (Poiret) DC. — Wurdack 43559 
Nyctaginaceae — Julian Steyermark (MO) 


East escarpment: 
Neea obovata Spruce ex Heimerl — Huber 5900 


Ochnaceae — Julian Steyermark (MO) 


Summit: 
¡diera auriculata Dwyer — Liesner 22168. Known 
reviously only from Cerro de la Neblina. 

Ho ud. retusa Tul. — Liesner 21942, 22631 

Sauvagesia piden (Eichler) Sastre subsp. ara- 

1. — Liesner. 22075 (type) 

Tyleria silvana Maguire — Liesner 22006. This species 
was originally described from the upper slopes of 

e table mountain in north- 


a . The collection of Liesner 

& Delara 22006 iom the summit of Cerro Ara- 
1,550 m marks the frst 
report of this The Cerro 
Aracamuni specimens differ from typical Tyleria 
silvana by having fimbriate intermediate append- 
es between the staminodes 5.5 r mm 


g mm vs. 2.5 

and primary bracts of the inflorescence 20- 30 mm 
292 

VS a 


wm 
O 
=> 
[77 
RE 
[a] 
a 
> 
A 
£ 
e 
= 
UN 


glandular- -scarious 
typica al of T. silvana end agrees in all other char- 
ers 


East escarpment 
Sauvagesia falcisepala Sastre — Huber 5918 (type) 


Quebrada camp: 
Ouratea ramosissima Maguire & Steyerm. — Liesner 
22217 


Granitic dom 


Sauvagesia longifolia Eichler — Wurdack 43545 
Olacaceae 


oe camp 
é determined t to genus— Liesner 22358 
Orchidaceae — Germán Carnevali € Ivón Ramirez (MO) 


Summit: 

Aracamunia — ri Carnevali & I. Ramirez — Liesner 
21988 (ty 

Bifrenaria LE C. Schweinf. p aa 2401; 

sterville s.n.; s sner 22176 8 
rs stricta C. Schweinf. Carnevali 2363, 2388, 
; Liesner 22052, 22421, 

"s e hy D Reichb. f. say nie 2515 
ner 22439 

Elleanthus sp. evali 2525; Liesner 22431 

Epidendrum dendrobioides Thunb. — Liesner 22054 

Epidendrum elongatum Jacq. — Carnevali 249 

Epidendrum noc disi Jacq. — Carnevali 2571; Lies- 
ner 22017, 2257 

isi Lm Garay — Carnevali 2551 

Epid rum ramosum Jacq. 

i ien ium hernandii Dario acaevali 2480; 

Liesner 22172, 22483 


; Lies- 


dd 


Cr ae subrepens Hoehne — Carnevali 2459; 
Liesner 22096 


Leaner 22049 
einf. & Garay) Gar- 


noc biloba cima 
cns; americana (C. Schw 


¡— aes 2202: 


N 
Ww 


Jacquiniella globosa (Jacq.) Schltr. — Carnevali 2452 


esne 
Lepanthes dude nsis Ames € C. Schweinf. — Carne- 
vali 2 
mir nin amazonica Schltr. — Carnevali 2482; Lies- 
ner 22163 


Maxillaria aff. arachnites Reichb. f. — Liesner 22036 

Maxillaria mapiriensis (Kranzl.) L. O. Williams — 
Carnevali 2422 

ea a Lindley 
2249: 


Oc pelai 
22602 


- Liesner 21970, 


sp. —Carnevali 2550; Liesner 22597, 


Octomeria cordilabia C. Schweinf. — Carnevali 2439, 
2569; Liesner 21965, 22089 

Octomeria erosilabia C. Schweinf. — Liesner 22600 

Ornithidium aureum Poeppig ex Endl.— Carnevali 
2423, 2504; Liesner 22445 


Pleurothallis a " o hweinf. — Carnevali 2533; 


Liesner 22462, 225 
ieri rn splurirace emosa Garay — Carnevali 2402 
2486; sner 22522 
Prescottia stachyoides (Sw.) Lindley — np 22460 
Psilochilus dusenianus Kranz aray & n 


Carneva 
speciosa C. 


sterv.— 
Sobralia 
Dunsterville s.n 
Sobralia yauaperye ensis Barb. Rodr. — Carnevali 2420, 
2474; Liesner 22152, 22467 
Stelis tristyla Lindley — Carnevali 2526; Liesner 22692 
Trichosalpinz Janata (Griseb.) Luer — Carnevali 2437, 
2541, 2554; Liesner 22128 
Pic duin obliguipetalus (Ames & C. Schweinf.) 
uer — Carnevali 2549 
Vanilla wrightii Reichb. f.— Carnevali 2403; Liesner 
22456 


Quebrada camp: 

Catasetum ochraceum Lindley, vel sp. aff. — Liesner 
22349 

Dichaea hystricina Reichb. f. Li esner 22396 

Epidendrum nocturnum Jacq. 

Epistephium subrepens Hoehne 

Mss iria grobyoides Garay € Dunsterv. — Liesner 


i nasuta Reic 
Octomeria 

Palmorchis 
Sobralia nee Lindley 


jJesner 222 
Liesner 22385 


Liesner 22240 


ib. f. — Liesner 22289 
82 


Granitic dome: 


Epidendrum ramosum Jacq. — Wurdack 43570, 
43571 


Passifloraceae — Julian Steyermark (MO) 
Summit: 
Passiflora sclerophy a Harms— Liesner 22053, 
22474, 22564, 227 
o M (MO) 
Sum 
P suecia duidana Trel. —Liesner 22707. This col- 
lection from the summit at the Proa Camp has some 


990 


Annals of the 
Missouri Botanical Garden 


ee as much as 4 x 4 cm and suborbicular and 
roun summit, 2s most of same collection 
number has smaller leaves ls arrowed apex (ob- 
use) from a broadl 
a hernandijfalia | (M. VERD A. Dietr.- 
ner 22518, 226 


Pepa ami tenella UM ) A. Dietr. —Liesner 22527, 
718 


- Lies- 


E lemaense Yuncker — Liesner 22580. Previously 
known ei us eastern. Venezuelan Guayana of 
Estado Boliv 

Piper mosaicum Steyerm.- Liesner 21968 (type) 

Piper poiteanum Kunth — Liesner 22425 


Quebrada camp: 

Piper xr um Trel. & Yuncker — Liesner 22368. 
Previously known from the eastern Venezuelan 
yuayana. 

Piper Francoville anum C. DC. 

Piper cf. pubive aginatum Steyeri 

Piper cf. jauaense Steyern 


Poaceae — Gerrit Davidse (MO) 


oa a 22266 
— Liesner ¿ 


Lie iesner 22214 


22353 


Summit 

Ic hnanthus a addio re Black € Fróes ex Black 
& Pires — , 22637 

Ichnanthus Fete din i eauv. lisse er 2191 

Myrioc Viger ROME Swallen — Liesner 22040, 

22005, 2267 

Myriocladus i M Swallen — Liesner 22634 

Myriocladus simplex Swallen — Liesner 22634A 

Myriocladus virgatus Swallen — Liesner 
22182, 22497, 22501 

Panicum chnoodes Trin. — Liesner 22441 

Panicum eligulatum N.E. Br.- Liesner 22107, 22642 

ss tepuianum Davidse & Zuloaga— Liesner 


226 


22118, 


East escarpmen 
Myriocladus Ke pom a 594.2 
Paspalum petilum Chase — Huber 


Quebrada camp: 
Andropogon bicornis L.— Liesner 22235 
Andropogon leucostachyus H.B.K.— Liesner 22237 
Ichnanthus epheme ic cin Black € Fróes ex Black 
Pires — Liesner 

Ichnanthus bius m -Liesner 22375 
Olyra micrantha H B.K.— Liesner 22267 
j ji en — Liesner 22209 

(Steudel) Calder & Söder- 
om — Liesne 239 
Hos petrosa es ) Chase — Liesner 22230 
Granitic dom 
Olyra mic i (H.B.K.) Swallen — Wurdack 43558 
Oly ra wurdackii Swallen — Wurdack 43540 
Panicum nervosum (Lam.) Swallen — Wurdack 43541, 

3542 


Polygalaceae — John J. Wurdack (US) 


Fast escarpment: 
Securidaca retusa Benth. 


Huber 5894 
boxed — Julian Steyermark (MO) 


Summ 

Sa xofriderici ia compressa Maguire — Liesner 22055 
Stegolepis celiae Maguire — Liesner 22030, ngo 
Stegolepis grandis Maguire — Liesner 21944 


Stegolepis linearis Gleason — Liesner 22058. 7th col- 
lection. Known iously from Cerro Duida, Avispa. 


East escarpme 
CORE ae pe (Ducke) Maguire — 
Huber 5905 


Monotrema xyrido ides Gleason — Huber 5939 
Saxofridericia spongiosa Maguire — Huber 5924 
Schoenocephalium lieri Maguire — Huber 5923 


Quebrada camp: 
apatea aracamuniana Steyerm. — Liesner 22197 


ype 
d pe 00): 
Rapatea spruceana Koern. —Liesner 22226 


Granitic dom 
Saxofridericia spongiosa Maguire — Wurdack 43538 
Rubiaceae — Julian Steyermark (MO) 
Summit: 
Md aracamuniensis Steyerm. — Liesner 
22064 (type), 22682 (paratype) 
Gleasonia duidana Standley var. duidana — Liesner 
22418 
Veblinathamnus aracamunianus Steyerm. — Liesner 
22153 (type 
dle Si montana Gleason & Standley — Liesner 
2629 
Pagameopss maguiret Steyerm. subsp. neblinensis 
Steve ya r 21951. Pre- 


aah j n from Cerro de la Neblina. 
Perama i hotoma var. scaposa (Gleason & Standley) 


Steyerm. — Liesner 22624 
Psyc horria campylopoda Standley — Liesner 22173, 
2266: 


99 


Pía ceratantha Standley — Liesner 225: 

Psychotria coussarioides Standley — Liesner 22667 

ae duricoria Pepe i Steyerm. var. duri- 

cc Liesner 2199 

Psychorria tapajozensis el 
23 


3; 22€ 


21996, 


Psclotr teputensis (Standley) Steyerm. — Liesner 


22 


Re reducta. Steyerm.—Liesner 22045 (para- 


type), 22407 (type) 

East escarpment: 

Pagamea vel aff. a Benth. subsp. caudata (H. 
E. Huber) iia Ee d 5916 

Pagamea coriac 


pant . Benth. — Huber 5915 
Retiniphyllum truncatum ; Muell. Arg. 


er 591 
Huber 5899 


Quebrada cam 

Chomelia NE Benth. — Liesner 22369 
Coussarea liesneri Steyerm. — Liesner 22262 (type) 
Miramar homi Steyerm. — Liesner 22350. Previ- 


reviously known from Cerro de la Neblina. 
ý Hei coriacea Spruce ex Benth. var. coriacea — 
2 


ne 
E nitidella (Muell. Arg.) Standley — Liesner 
22329 
—_— dienot Beat & Endl. var. dichot- 
jesner JS, 222 ph 
[NES a mal .—Liesner 22383 
Psyc hotria deflexa DC. subsp. elation (Muell. Arg.) 
Steyerm. — Liesner 2235 
Psychotria iodotricha Muell. oe subsp. todo- 
icha—Liesner 22248, 


D, 222. 


Volume 76, Number 4 Steyermark, Holst & Collaborators 991 
1989 Venezuelan Guayana Flora— VII 
Psychotria medusula Muell. Arg. — Liesner 22362 Theaceae — Brian Boom (NY), Ternstroemia; Anna 


Psychotria cf. paupertina Standley & Steyerm. 
Liesner 22 

d podocephala (Muell. 
Liesner 22268, 22373 


Bode spadicea (Pittier) Standley & Steyerm. 


Arg.) Standley 


DI 


jiesner 22 
i cueca ocn var. sororiella (Muell. 
Steyerm.—Liesner 22257, 22370 
Remijia roraimae (Benth.) Schumann var. adpressa 
Steyerm. —Liesner 22251, 22291, 22335 
Retiniphyllum truncatum Muell. Arg. — Liesner 22229 
Rudgea stipulacea (DC.) Steyerm. — Liesner 22244 


Arg.) 


> oon y LO: 


ho ard 


Isertia r ex Schumann — Wurdack 43553 


Retiniphyllum bci orum (Benth.) N.E. Br. var. laxi- 
florum — Wurdack 43551 
Rutaceae — Julian Steyermark (MO) 
Sur 
. — Liesner 


Spathelia neblinaensis R. Cowan & Briz 
. Known previously only from Neblina. 
Sapindaceae 
Summit: 
Toulicia sp.—Liesner 22477, 22683 
d camp 
Not lied to genus — Liesner 22334 


K) 


~ 


pe — Terence D. Pennington 


Summ 


Mic "opui suborbicularis Aubrév. — Liesner 21939, 
22179, 22413 


East esc arpment 


Pradosia sc ec PR (A.DC.) Cronq. — Huber 
5927 

Quebrada camp: 

Pradosia schomburgkiana (A.DC : Cronq. subsp. 


schomburgkiana — Liesner 2221 


Sarraceniaceae — Julian Steyermark (MO) 


umm 

Heliamphora tatei Gleason var. neblinae (Maguire) 
eyer parva oe Steyerm. — Liesner 

22119, 22643, 226 


Simaroubaceae 


Quebrada camp: 
Picramnia sp. — Liesner 22384 


Smilacaceae 


Summit: 
Smilax sp. —Liesner 22022, 22560 


pad camp: 


Smilax sp. —Liesner 22250, 2228 
Solanaceae 
Summit: 


Cestrum sp. —Liesner 22505 
Styracaceae — Reinaldo Monteiro (HRCB) 


Summit: 
dl sp. —Liesner 22681 
tus Steyerm. —Liesner 22151, 


Wei eltzman (US), Frezierea, Julian Steyermark (MO), 
Bonnetia, Neotatea 
Sum 
Arc wm ibd p du ae —Liesner 22123 
Bonnetia liesneri Steyerm.— Liesner 22105 > (type) 
bir cin tristyla Gleason subsp. nervosa Steyerm. — 
Lies dim 22029 
peep trinata Weitzman — Liesner 
Veotatea lon Em (Gleason) Maguire — Liesner 22110 
T. o JUS ki tees ner 21957, 
65, od 248 


~ 
® 


22154 


oa y 441« T, de 
Te puce maguir Bo ae 
2117, 22158, 22506 (all ibit: 
recen pungens Gleason — Liesner 22618 


21985, 


(, 24 


East esc arpm 

Archytaea angustifolia Maguire — Huber 5943 

Quebrada camp: 

Ternstroemia guanchezii B. Boom 
(paratype 


-Liesner 22236 


Granitic dome: 
Bonnetia paniculata Spruce — Wurdack 43548 


Verbenaceae — Julian Steyermark (MO 


<~ 


Summit: 
Ee roraimensis Moldenke —Liesner 21958, 
59 


Quebrada cam] 
Not pe E to genus — Liesner 22265 


neds Robert Kral (VDB) 


Moli grandis Griseb. var. grandis — Liesner 
1955. Thi 


a toward var. 


is mostly in ios to 
northern Brazil north into the Guianas, westward 
ety Venezuelan Guayana into southeastern Co- 


lor 
Atl scabrida Kral — Liesner 22071 (paratype), 
8 (t 


T 
fva ris MAE Kral — Liesner 22100 (paratype), 
22452 (type) 
Xyris od Nees 
Xyris liesneri Kral- 
(para 


type) 
yris witsenioides Oliver 


Liesner 21943, 22701 
22080 (type), 22454 


- Liesner 


- Liesner 22025 
ast escarpmen 


Huber 5912 


I 
Abolbada i: Griseb. var. grandis 


Quebrada cam 
S DPA grandis Griseb. — Liesner 22194 

Xyris e > Steyerm. — Liesner 22339 

Xyris fallax Ma lo —Liesner 22221 

Xvris p ans Kral — Liesner pe (type) 

Yyris spruceana dede — Lone 22219. Locally 
abundant in mostly low- 


western Colombia. This example from 600 m elevation 
s from one of the higher sites known, probably 


Annals of the 
Missouri Botanical Garden 


migrating upward along edges of gallery forest in 
the lajas. 
initic don 
Abolboda pa Griseb. var. 
guire — Wurdack 43539 
poma lopezii (Lyman B. Smith) Steyerm. & 
erry subsp. lopezii — Wurdack 43564 
The first set of the Wurdack & Adderley col- 
lections is deposited at NY, with a second set at 
VEN. The first set of the Huber & Medina col- 
lections is deposited at VEN with duplicates at NY. 
he collections of Liesner have the first set at VEN, 
the second at MO, and the third at the institutions 
of the specialists that provided identifications. The 


guayanensis Ma- 


Carnevali, Dunsterville, and Brewer Carias collec- 
tions are deposited at VEN 


LITERATURE CITED 


DURNS-BALOGH,. P. H. ROBINSON & M. S. Foster. 1985. 


anew genus from Paraguay. Brittonia 37: 154-162. 

GARAY, 982. A generic revision of the Spiran- 
thinae. Bot. Mus. Leaflt. 28: 277-425. 

Kurz, H. 1983. pasan ct einiger Gattung- 
en neotropischer Lauraceen und Revision der Gat- 
tung Licaria (Lauraceae). Thesis, Univ. Hamburg. 

STEYERMARK, J. A. 196: ubiaceae. In: aguire 
(editor), Botany of the Ang Highland V. Mem. 
New York Bot. Gard. 


COMPOSITAE OF THE 
GUAYANA HIGHLAND—IL. 
NOVELTIES IN 
GONGYLOLEPIS AND 
STENOPADUS (MUTISIEAE) 


John F. Pruski? 


ABSTRACT 


Described from the Guayana Highland are: Gongylolepis Ad Vea from Serra Aracá, Brazil, and from Sierra 


de Unturán and Cerro de la Neblina, Venezuela; 
megacephalus from Mount Ayanganna, Guyana. 


species of both genera is given. Stomatochaeta eolv eei is transferred to Stenopadus and 


Stenopadus aracaensis fro 


m Serra Aracá, Brazil; and Stenopadus 
xon are discussed and a key to 
the combination Stenopadus 


colveei is proposed. Additionally, five species from the Guayana Highland are newly documented for the flora of 
razil. 


The shrubby, homogamous, generally large- 
headed Mutisieae are a striking element of the 
Guayana Highland flora and are typically found 
on tepui summits. Among Guayanan taxa of its 
size—57 of the 6l currently recognized species 
(93%) are endemic to Guayana — the Mutisieae are 
unparalleled in their endemism level (Maguire, 
1956; Pruski, 1989). The woody, hermaphroditic 
members of the tribe Mutisieae from Guayana fall 
into two main generic groups. The first group (sub- 
tribe Mutisiinae sensu Cabrera, 1977) consists of 
six genera having homogamous capitula composed 
of don flowers and includes Gongylole pis 

, with one species described herein. The 
nud group (subtribe Gochnatiinae sensu Ca- 
brera, 1977) contains four genera, which have only 
actinomorphic flowers within their capitula. The 
second group was previously represented in Gua- 
yana only by Quelchia N. E. Brown and Steno- 
padus S. F. Blake, but has been redefined to admit 
two new genera, equivalent in part to species for- 
merly placed in Stenopadus s.l. The newer genera 
are Stomatochaeta (S. F. Blake) Maguire & Wur- 
dack (1957), which was raised from subgeneric 
level within Stenopadus, and Chimantaea Maguire 
et al. (1957), which was described as new. All 


subsequent authors (Aristeguieta, 1964, 1968; 
Steyermark, 1980, 1984; Pruski, 1989) have rec- 
ognized Stomatochaeta and Chimantaea, the ge- 
neric status of which is justified by floral mor- 
phology. 

In this paper new species of Gongylolepis and 
Stenopadus are described preliminary to my treat- 
ment of the Compositae for the Flora of the Ven- 
ezuelan Guayana by Julian Steyermark and col- 
laborators, where the localities and complete 
synonymies of all our Mutisieae will be given. Also, 
fertile material of Stomatochaeta colveei, recently 
made available to me for study, shows that this 
species has the flower color and morphology typical 
of Stenopadus, to which it is herein transferred. 


GONGYLOLEPIS 


Gong ylolepis is a genus of 15 species with bi- 
labiate flowers and glabrous, reticulate, lanceolate 
to obovate leaves. Fourteen of these occur in the 
Guayana Highland and 11 are endemic to the 
Venezuelan Guayana. The nearest generic relative 
of dii us is the monotypic Eurydochus Ma- 
guire & Wurda 


leaves and eared but Eurydochus differs 


The two genera have similar 


! [ thank Rupert Barneby for help with the Latin diagnoses and for many helpful comments; Victor des: Rove 


Robinson, and an anonymous reviewer for their 


Huber, and [ain Prance 
financially supported by the Missouri Botanical Ga 
2 New Y 


York Botanical Garden, Bronx, 


r suggestions; Bruno Manara for drawing Figure 2; Hans Beck, Ot 
for permission to publish their photographs; and Don 
Stevens, and Muriel Weinerman for helping me prepare the figures. 


Black, Maria Del deas Marcia 


rium studies in Venezuela have been 


My herbar 


rden, Flora of the Venezuelan Guayana project. 


New York 10458-5126, U.S.A 


ANN. Missouni Bor. GARD. 76: 993-1003. 1989. 


Annals of the 
Missouri Botanical Garden 


FIGURE 1. 
of two capitula 
solitary capitulum, zygomorphic flowers, and the pubescent peduncle (Henderson et al 961). mu i H. 


Gong ylolepis oblanceolata. 


occurrence at the stem apex (Pran 


from Gongylolepis by deciduous (vs. persistent) 
pappus setae, by pubescent (vs. glabrous) phylla- 
ries, and by pubescent (vs. glabrous) achenes. Less 
closely related to Gong ylolepis are Glossarion Ma- 
guire & Wurdack and Duidaea S. F. Blake, which 
differ, respectively, in abaxially tomentose and one- 
nerved leaves. Described and keyed below is a new, 
predominantly monocephalous Gong ylolepis. It is 
most closely related to the other pedunculate, 
monocephalous species of the genus, these mostly 
confined to T. F. Amazonas, Venezuela. 


Gongylolepis oblanceolata Pruski, sp. nov 
YPE: Brazil. Amazonas: Plató da Serra es 
200 km N de Ba Mar. 1984, M. G. 
Silva et al. [co-collector names not given on 
type labels] 7/69 (holotype, INPA; isotypes, 
K, MO, NY, US). Figures 1, : 


rcelos, 


Gongylolepidi bracteatae et G. pedunculatae affinis, 
quibuscum capitulis pedunculatis foliisque aut petiolatis 
aut basin versus angustatis congruit, sed a priori pedun- 
culis ebracteatis et a secunda phyllariis magis numerosis 
et minoribus differt. 


A. Note the nearly glabrous, minutely bracteolate ceci and the 
nce et al. 


29014). Photo by G. Prance > the zm 


Branched shrubs 1-3 m tall; stems terete, striate, 
leafy and often lanate (especially in upper leaf axis) 
distally, glabrous and leafless proximally, inter- 
nodes to ca. | cm long, thereby much shorter than 
the leaves. Leaves simple, alternate, often apically 
clustered, ds or petiolate; blade coriaceous, 
oblanceolate, ca. -4.5 cm, narrowly 
cuneate or attenuate basally, often narrowed into 
a subpetiolar base, apically acute, obtuse or round- 
ed, margins entire, these not thickened, rarely re- 
volute, venation pinnate, reticulate, upper blade 
surface dark green, often puberulent when young, 
especially so along midrib, glabrous with age, lower 
blade surface light green, often pubescent when 
young, glabrous with age; petiole often represented 
only by a petioliform leaf base to 5 mm long, 
clasping and stout. Capitula generally solitary, ter- 
minal, pedunculate, homogamous, with ca. 14-31 
flowers, these perfect and fertile; peduncle often 
bracteolate, nearly glabrous to densely pubescent, 
ca. 3-8 cm ii inv Jolie re narrowly hemispheric, 
o X 2.5(3) cm, 5-6-seriate; phyl- 
necs ca. 25- "3 tole vie. graduated, erect or 


Volume 76, Number 4 
1989 


Pruski 
Compositae of the Guayana Highland—ll 


FIGURE 2. Corollas of representative species of Stomatochaeta and Stenopadus. —A. Stomatochaeta condensata 
54, NY.)— 


(Baker) Maguire & Wurd. (Drawn from Pruski 
(Drawn from the holotype, Huber 11374, VEN.)- 
from Pruski & Huber 3616, NY.)— 
29184, INPA.)—E. Stenopadus cc (S 


, B) the short corolla tube a 


corollas of Stenopadus are magenta 


sometimes slightly spreading, thickly coriaceous, 


entire, often somewhat purple; outer 
phyllaries triangular-ovate, ca. 2.5-6 x 2.5-5.5 


mm, apex broadly acute to rounded, basally round- 


glabrous, 


ed or cordate, sometimes decurrent onto peduncle; 
inner phyllaries elliptic-lanceolate to lanceolate, to 
ca. 2.5 cm 5 mm, apex commonly rounded; 
receptac le broadly dome-shaped, naked, aveolate, 


6-sided. Corollas bilabiate, 15— 


times pale yellow; tube 9— 
with upper portion broadened near sinuses of the 
2 lips, obscurely ca. 10-nerved; lips coiled; the 
outer, larger lip apically 3-dentate, to ca. 10— 
mm long when uncoiled, commonly coiled at apex 
or for more than half its length; the inner, lower 
lip 2-cleft, generally strongly coiled, thereby short- 
er than the outer lip; anthers exserted, linear, 
reddish, less commonly pale yellow, 10-12 m 
long, apically acuminate, prominently sagittate ba- 
sally; tails white, squamose, linear, 5-7 mm long, 
thereby more than half as long as the anthers, tails 
of adjacent anthers commonly completely connate 
or occasionally free basally; filaments yellowish or 
reddish, inserted well below bases of lobes; styles 
reddish above, yellowish below, branches spread- 
mm long, apically obtuse. 4chenes linear 
10-costate, 10-11 
long, brown, glabrous; pappus setae numerous, 


ing, ca. | 
or slightly fusiform, ca. 
linear, brown, ca. 


persistent, -]7 mm long, 


thereby longer than the achenes. 


& Steyern mark 1454 

. Stenopadus chimantensis Maguire, Steyerm. ur -( 
D. Stenopadus aracaensis . 
ruski. (Drawn from ue 12405, VEN.) Note in Stomatoc haeta 
nd the dun. stiffly. erect lobes. 
Stenopadus (C-E) the long corolla tube and the coiled or flexuous lobes as long 


B. Stomatochaeta acuminata Pruski. 
& W rawn 

ruski. (Drawn from the holotype, Prance 

e corollas of Sonato haeta are yellowish. Note in 

as or shorter than the tube. The 


Distribution. 
ra Aracá, Brazil, ridges of Sierra Unturán, Ven- 


Known from the plateau of Ser- 


ezuela, and low elevation slopes and canyon walls 
of Cerro de la Neblina, Venezuela. This species has 
) 700-1,250 m and is 


known to flower and fruit from February to July. 


been collected from ( -], 


Paratypes. BRAZIL. AMAZONAS: Municipio de Bar- 
celos: plató da Serra Aracá, parte SE da Serra Norte, 
095]'N, 63?22'W, 1,150-1,250 m, 14 Feb. 1984 EP 
Amaral 1557 (F, INPA, NY, US); plateau of Mic wn 
massif of Serra Arani, savanna, S E of north mountain, 
0951-57"N, 63%21-22'W, 1,20( abe 19 
T. Prance, L L. Pind J. J. "x 

M.€ 4. G Fess 
K, NY, vc " platô da Se rra m. 


. Tavares, 
205 (INPA, 
SE da Serra 


150 1,250 m, 13 Feb. 
Cisneros & M. G. Silva 22 (INPA 


masa Cañón eue. dune e Río Vae aie be- 


of the river, ca. 7 airline km ENE of Puerto Chimo, 0°50- 
: : j in m, 9-14 July 1984, 

Davidse & J. Miller 27303 (MY); Dept. Rio Negro, 
Neblina massif, Sanii ng midi at the mouth of Canón 
Grande, N of Puerto Chimo, 0°50'N, 66?06'W, 500 
100 m, 15 July 1984, C. e & J. Miller 27399 
(VEN); Sierra de Unturán, ridge i camp, 01°33'N, 
65°12'W, 1,150 m, 4 Feb. 1989, 4. Henderson, H. 
Beck, B Scott & L. Leigh 961 n VEN); Cerro de 
la Neb slope N of Rio Mawarinuma, above Puerto 
n. OSON, 66°05'W, 500-700 m, 25 Apr. 1984, 

Stein & 4. Gentry 1686 (VEN). 


Gongylolepis oblanceolata, most readily rec- 
ognized by its minutely bracteolate peduncles, is 


996 Annals of the 
Missouri Botanical Garden 


FIGURE 3. Stenopadus colveei showing a few-flowered, narrowly cylindric capitulum (Huber 12405). Photo by 
O. Huber. 


closely related to G. bracteata and G. peduncu- 
lata. These three species are linked by having 
pedunculate heads and large leaves that are either Other seemingly less closely related monocephalous 
petiolate or narrowed into a petioliform base. Gon- species of Gongylolepis generally have sessile heads 
gylolepis oblanceolata differs from G. bracteata and smaller, sessile leaves. 

in apically nonbracteate (vs. bracteate) peduncles 


and from G. pedunculata in smaller, more nu- 
merous (vs. larger and fewer) involucral bracts. 


Pruski 997 


Volume 76, Number 4 
19 Compositae of the Guayana Highland—ll 


oblanceolata 


colveel 


aracaénsis 


megacephalus 


y 


FIGURE 4. Distributions of Gongylolepis oblanceolata, Stenopadus colveei, S. aracaensis, and S. megacephalus 


KEY TO THE SPECIES OF GONGYLOLEPIS 
la. e Wei ipe monocephalous, rarely with two c Ie stem. 
2a. pro talked on a naked or leafy peduncle ca. 10 cm long 
aves with subparallel venation, 8.5-15 x 3.3-6 cm, sessile; capitula ca. be flowered, ca. 5.5 
NA jauaansis (Aristeg., Maguire & Wu cos A V. Badillo 
4-8.5 cm, sessile or petiolate; a -35-flowered. 
l cm or more ds e; 


m tall 
3b. bos pinnately e a= — x (1.5 
capitula taller th cm, outer and intermediate phyllaries ca. 
e mey es naked or le es bag qu che el 20(50) x 4-8.5 cm, basally cuneate, 
essentially sessile, wider than 5 1 
5a. Peduncles leafy, hers aa [o 1-3 large, leafy bracts; stem apex lightly 
pubescent . G. bracteata Maguire 
5b. diria naked, capitula not subtended by bracts; stem ften densel scent 
o flocc edunculata ib 
bra 


mm; peduncles minutely 


4b. Capitula shorter than 4.5 cm, phyllaries narrower than 
eolate; leaves 5-21 x 1.5-4.5 cm, basally attenuate, e or petiolate, narrower 
G. oi Pruski 


han 4 mm at base __........ 
r than 4 cm, 


c 


Subshrubs to 0.5 m tall, leaves often clustered at apex of n 
7 x 0.7-2 cm wide and at least 3 times as long as vie "M 


a Maguire . & Wurd. 


. fru 
6b. Shrubs or trees commonly 1.5-3(5) m tall, leaves not apically a leaves iru b res vate 
2.5-10 x 1.2-5 cm, less than 24 times e wide. 
o 10 x 4 cm 


oblanceolate, sessile, ca. 


e to petiolate, ca. 
boe and branches glabrous; leaves obovate, sessile, 
aberrima S. F. Blake 


aaa rre 
Tb. Young leaves and ding ie pilose; leaves oblanceolate to broadly elliptic, subsessile 
to En less than 
aves with ode ca. E mm long, secondary veins obviously pinnate; phyllarie 
edes tor x, wider than 5 mm. G. paruana ae 


rounded at ape 
8b. Leaves subsessile or with petioles shorter dui 3 mm, secondary veins subpalmate; 
G. erioclada S. F. Blake 


phyllaries acute to acuminate, narrower than 5 mm „u 
lb. rep ora ences branched. of few to many capitula, each of these pedunculate. 
s pubescent when young. 


10a. Captlescene leafy 
l 1 stem RAE commonly (12)15-20 cm long; corollas glabrous ...... 


llb. uu shorter than 13 cm; corollas hair 


G. yapacana Maguire 
benthamiana Schomb. 


998 


Annals of the 
Missouri Botanical Garden 


10b. 


ronem naked, no 
Cap 


t leafy. 
tulescence umbelliform, peduncles 3-6 cm long; leaves thickly coriaceous, narrower 


tian 3 .5 cm, margins often thickened and revolute; stem apex often densely pubescent 
G. 


ccose 


uachamacari n d 


floc 
12b. cow item corymbiform, peduncles 1-3 cm long; leaves thinly coriaceous, 3-6 c 
wide, margins slightly thickened, not revolute; stem apex generally lightly pubescent . 


erramarae Steyerm. 


9b. Stems glabrou 
3 


s of flowering stems subsessile o 


piles e naked, not leafy; leaves n petiolate, often prominently so so 
i ith 


s to ca. 1 cm long g; pappus « straw-colored. 


ers ves subsessile; ie als s asian plants of the Andes of Colombia and 
l 


Venezuela 


:olombiana (Cuatrec.) Cuatrec. 


15b. Leaves with petioles to ca. 1 cm fong; capitulescence umbelliform or — orm; 
m 


lants of Guayana, occurring 1,000-2,70 


I 
1 4b. i of floweri ing stems 


with petioles 1-3 cm long; pappus straw- ene to rose-co 
ts generally occurring ene 150(-750) m 


n to couplets 12) 
olored; 
PE G. martiana (Baker) Reis & Cuatrec. 


13b. € lpliulescence leafy or sometimes naked when in fruit; leaves sessile or aed petiola 
a. Ca 


apitulescence umbellifo 


huac nne Maguire 


16b. e corymbiform. 


as hairy, tube broadened; leaves 7.5-16.5 


x 2-5.5 em, upper surface RET 


icis heads rounded at the base, involucre ca. 3 cm tall, ca. 6-seriate; ac 
G 


m long, mostly eid ric 


6- 
17b. ON glabrous, tube narro 


moderately reticulate; heads narrowed at the ba 
6-8-seriate; achenes 8-15 mm long, frequently fusiform ............. 


STENOPADUS 


Described and keyed below are two new species 
of Stenopadus which, following the transfer of 
Stomatochaeta colveei to Stenopadus, raises to 
14 the number of species recognized in this genus. 
All are endemic to the Guayana Highland and eight 
species are restricted to Venezuelan Guayana. 
Stenopadus is closely related to Stomatochaeta, 
which was originally described as a subgenus of 
Stenopadus. The yellowish corollas of Stomato- 
chaeta (Fig. 2A, ABE with edi erect lobes much 
longer than the t ti h it from Stenopadus 
(Fig. 2C-E), which has magenta corollas with (at 
maturity) coiled or flexuous lobes about as long as 
or shorter than the tube. By these features generic 
status for the two taxa is justified (Maguire & 
Wurdack, 1957; Pruski, 1989). 

Stomatochaeta colveei resembles Stenopadus 
in a magenta corolla with tube longer than the 
lobes. It was originally (incorrectly) described and 
illustrated as having corolla lobes stiffly erect and 
was thus placed by Steyermark (1980) in Sto- 
matochaeta. Pruski (1989) noted it as possibly 
intermediate between these two genera, but he 
retained it within Stomatochaeta until flowering 


o 


material could be observed. Specimens examined 
by me during a recent visit to the Instituto Botánico 
in Caracas show that the corolla lobes of Stoma- 
tochaeta colveei are flexuous and shorter than the 
tube (Fig. 2E). Thus, this species fits well within 
Stenopadus as envisioned here, and the formal 
transfer of this species to Stenopadus is proposed 
below 


Stenopadus and Stomatochaeta now stand 


7. rnthanian Schomb. 
-8(9) cm, A ida surface 
se, involucre ca. 3-3.5 tall ca. 
G. polus e Maguire 


; leaves 12-27 x (2.5 


clearly apart from each other on the basis of flower 
morphology. Furthermore, Stenopadus, a genus 
of stately plants up to 25 m tall, is typically more 
robust than Stomatochaeta, plants of which are 
less than e m tall. Other genera from Guayana 

only actinomorphic flowers, and 
slightly less ied related to Stenopadus are Chi- 
mantaea, which has abaxially tomentose leaves and 
corolla lobes pilose when young, and Quelchia, 
which has small, uniflorous capitula. 


Stenopadus colveei (Steyerm.) Pruski, comb. 
nov. Stomatochaeta colveei Steyerm., Brit- 
tonia 32: 21. 1980. TYPE: Venezuela. Bolivar: 
cumbre Cerro Guaiquinima, sector SE, cerca 
del borde, 5%40'N, 63?26'W, 1,250 m, 26 
May 1978, Steyermark, Berry, Dunsterville 
& Dunsterville 117407 (holotype, VEN). Fig- 
ures 2E, 3, 4. 


This species is a shrub 1.5-2.5 m tall charac- 
terized by basally attenuate, oblanceolate leaves 
and by narrowly cylindric, 5-6-flowered capitula. 
It is endemic to Cerro Guaiquinima, where it occurs 
in rocky or sparsely covered shrubby chaparral 
from 900-1,350 m and flowers in December and 
May. 


Additional specimens examined. 
var: meseta del deg qe sector SE de la cumbre 
piae 63?25'W, 50 m, 26 Sep. 1985 (sterile), Hu- 

r & Medina Per: (VEN); cumbre de la meseta del 


a sector S, 
8 kr 


Vene i Rp Boli- 
E 


: Dec. ec rd TOMUS (AAU n.v., HBG n.v., MY, 
N). 


, TFAV n.v., US n.v., VEN 


Volume 76, Number 4 


Pruski 999 
Compositae of the Guayana Highland—ll 


1989 
NY NEGATIVE 
No. 12531 

3 F: AA 
“Ee /e 
ZZ / m t. 
x 2 le : 2, g T 
Tis = qn 

sE KP, 

Va av? " d 
ES d 
3 5 a f f 
i 3 
1 E INDOL O Ls t Le 
PA | da 
|_UMICATA ] 
FicuRE 5. Holotype of Stenopadus aracaensis (Prance et al. 29184, INPA). 


Stenopadus aracaénsis Pruski, sp. nov. TYPE: 

Brazil. Amazonas: plateau of northern massif 

of Serra Aracá, N part of N slope, 0%51- 

dp 63°21-22'W, 1,400 m, 20 Feb. 1984, 

. T. Prance, I. L. Amaral et al. 29184 

Hof INPA; isotypes, K, MO, NY, US). 
Figures 2D, 4, 5. 


Ste i is m ed = et cum illa corollae lobulis 
tubo multo brevioribus congruens, sed capitulis haud s a 
itatis ad maximum 25 di 35-70) flosculosis divers 


Tress 10 m tall, 30 cm diam.; stems terete, 
striate, glabrous, leafy distally, leafless proximally, 
internodes to ca. 1.5 cm long, thereby shorter than 
the leaves. Leaves simple, alternate, often apically 


1000 


Annals 


of the 
Missouri Botanical Garden 


NY  necarve 


No. 12532 


Vl lett a ol 547106 
lus megacephaluz Fila, 


de? Dohu Prost, 11938 


im 


FIGURE 6. 


clustered, petiolate; b/ade rigid-coriaceous, gla- 
brous, broadly oblanceolate or obovate, ca. (4)6- 
13.5 x (1.5)2.5-4.7 cm, narrowly cuneate or 
attenuate basally, apically rounded, margins entire 
thickened, somewhat revolute, venation pinnate 
reticulate or third-order veins often obscure, upper 


34AL 


, 


u 
o 


wpeg punog yo, MIN pn p 


THE NEW YORK BOTANI GARDEN GUIANA EXPLORATIONS 
PLANTS OF BRITISH GUIANA 
UPPE I RIVER E 

Ne 45100 

f, f et 

He maf des la Me as Plaga et ibu 

Shrub to 2 m; le st k, stiff, rrittle, 

i ve, lig} eneeth, mer tl lute, 

re € r 
t ow for - 
MA gan 


Holotype of Stenopadus megacephalus (Tillett et al. 45100, NY) 


blade surface dark green, lower blade surface pale 
green; petiole ca. 0.9-1.5 cm long, thin and non- 
clasping. Capitula solitary, terminal, sessile and 
thus nonstipitate, homogamous, with 21-25 flow- 
ers, these d and fertile; involucre cylindric, 
n, 7—8-seriate; phyllaries 


Gee 


ca 


Volume 76, Number 4 
1989 


Pruski 1001 


Compositae of the Guayana Highland—ll 


ca. 40-45, imbricate, graduated, tightly appressed 
and rigidly erect, coriaceous, glabrous, entire; out- 
er phyllaries slightly carinulate, triangular-ovate, 
ca. 4-8 mm long and broad, apex broadly acute 
or obtuse, not de nem onto stem; inner ip epe 
elliptic-lanceolate to lanceolate, ca. 3 cm x 

5 mm, apex commonly rounded; receptac le weakly 
paleaceous, paleae linear, ca. 2.5 cm 
colored. Corollas actinomorphic, tubular, strongly 
5-cleft, ca. 19-22 mm long, magenta, glabrous; 
tube 14—16.5 mm long, narrow and much longer 
than the lobes, weakly ca. 15-nerved; lobes either 
flexuous and only apically coiled or strongly coiled 
nearly to base, ca. 5-6 mm long, generally 3(-5)- 
nerved and with an obvious central nerve; anthers 


ong, cream- 


exserted, linear, cream-colored but prominently 
purple-spotted, 8.5-9 mm long, apically acumin- 
ate, basally sagittate; tails white, smooth, linear, 
mm long, tails of adjacent anthers connate; 
filaments yellowish or reddish, inserted within the 
tube; styles obscurely branched, magenta, glabrous 
but asperulous apically, to 4 cm long. Achenes 
(immature) obconic, ca. 10-costate, ca. 6 mm long, 
olivaceous, glabrous; pappus setae numerous, lin- 
ear, cream-colored, ca. 13-19 mm long, thereby 
longer than the achenes. 
Distribution. 
lection made on the plateau of Serra Aracá, Brazil 
at 1,400 m. Flowers in February. 


Known only from the type col- 


Stenopadus aracaensis, collected on a recent 
Projeto Flora Amazónica expedition, is most closely 
related to S. kunhardtii. Both species have three- 
nerved corolla lobes that are much shorter than 
the corolla tube and thereby closely resemble one 
another. However, S. aracaensis clearly differs 
from S. kunhardtii by nonstipitate (vs. stipitate) 
heads, smaller basally attenuate (vs. larger basally 
cuneate) leaves, and capitula that contain 25 or 
fewer (vs. 35-70) flowers. Also closely related is 
S. huachamacari, but this differs from the two 

mer species by two-nerved (vs. three-nerved) 
corolla lobes that are about as long as (vs. much 
shorter than) the corolla tube. 


Stenopadus megacephalus Pruski, sp. nov. 
YPE: Guyana: upper Mazaruni River basin, 
Mount Ayanganna, shoulder of E flank, above 
Thompson Camp, 1,418-1,585 m, 11 Aug. 
1960, S. S. Tillett, C. L. Tillett & R. Boyan 
45100 (holotype, NY). Figures 4, 6. 

sericeo, etiam in monte Ayanganna obvio, 


nis S. 
d foliis M et glaberrimis (nec petiolatis et infra 
sericeis) absta 


Shrubs 2 m tall; stems terete, nonstriate, thick, 
glabrous, leafy distally, leafless proximally, inter- 
nodes to ca. 1.5 cm long, thereby shorter than the 
leaves. Leaves simple, alternate, clustered toward 
end of branches, subsessile; blade rigidly and thick- 
ly coriaceous, glabrous, oblanceolate or obovate, 
8.5-19 x 
narrowed into a petiolariform base, apically round- 
ed, margins entire, thickened, slightly revolute, 
venation pinnate, obscurely reticulate with third- 


3-8.5 cm, cuneate basally and mostly 


order veins often immersed, upper blade surface 
dark green, lower blade surface lighter green; pet- 
ioliform base stout, ca. 5 mm long and broad, not 
clasping the stem. Capitula (well past maturity 
with flower parts mostly fragmentary) solitary, ter- 
minal, sessile or very shortly pedunculate, homog- 
amous, with many (ca. 40 or more) flowers, these 
perfect and fertile; involucre hemispheric, ca. 5 
X 4.5 cm, ca. 5-seriate; phyllaries ca. 40, im- 
bricate, graduated, erect, coriaceous, glabrous, en- 
tire; outer phyllaries broadly triangular-ovate, ca. 
3-1.5 cm, apex broadly acute or 

obtuse; inner phyllaries lanceolate, ca. 4.5 cm X 
5-7 mm, apex acuminate to rounded; receptacle 
seemingly naked. Corollas actinomorphic, tubular, 
strongly 5-cleft, ca. 13 mm long, magenta, gla- 
brous or weakly puberulent internally near sinuses 
of lobes; tube ca. 8-10 mm long, broadened near 
sinuses of lobes, weakly ca. 10-nerved; lobes some- 
times flexuous but more commonly strongly coiled 
nearly to base, ca. 2-6 mm long, seemingly 
2-nerved; anthers partly exserted, linear, cream- 
colored, ca. 13 mm long, apically acuminate, ba- 
sally sagittate; tails cream-colored, smooth, linear, 
3-4 mm long, those of adjacent anthers ap- 
parently appressed but not connate; filaments 
mostly disintegrated, seemingly inserted within the 
corolla tube; styles not seen. Achenes broadly ob- 
x 2.5 
mm, light brown, glabrous; pappus setae numer- 
ous, linear, light brown, tips mostly broken off, but 


conic, with ca. 4 or 5 main angles, ca. 5.5 


intact setae to 15 mm long, thereby much longer 
than the achenes. 


Distribution. Known only from the holotype, 
collected on Mount Ayanganna, Guyana, where 
the new species is frequent in dense scrubland 
between 1,418 and 1,585 m. Known to be in fruit 
in August. 


Stenopadus megacephalus is closely related to 
S. sericeus, the only other species of Stenopadus 
reported from Mount Ayanganna. However, the 
leaves of S. sericeus are clearly petiolate and are 
sericeous below, whereas those of 5. megacephalus 
are subsessile and completely glabrous. Also closely 


1002 Annals of the 
Missouri Botanical Garden 


campestris, which has flexuous three-nerved (vs 


related are S. chimantensis, which has wider leaves 
strongly coiled two-nerved) corolla lobes. 


and smaller heads, and the distantly disjunct 5. 


KEY TO THE SPECIES OF STENOPADUS 
iud bons a to corolla tube 1-4(10) mm below base of lobes. 


la. 
ssile or brevipetiolate with petioles shorter than 7 mm 
"Hd eaves seta than 11 cm; capitula with up to 25 florets; involueres. een than 3 em; corolla 
lobes 3-nerve 'ampestris Maguire & Wurd. 
3b. Leaves to 19 cm long; capitula with more than 30 florets; involucres n 5 un tall; corolla lobes 
2-nerved megacephalus Pruski 
2b. Leaves with petioles 8-20(35) mm long. 
4a. Outer phyllaries acuminate, inner ones attenuate; capitula often 2-4 n dia TE eapite ocu 
onicus Maguire & Wurd. 
4b heu ai obtuse to acute, inner ones usually acute or gbh, never attenuate; capitula 
Sa. ‘Corolla lobes 2-nerved, about as long as the tube S. huachamacari Maguire 
5b. Corolla lobes 3-nerved, about 21% to 3 times shorter than the tube. 
Phyllaries not pui involucre eR leaves 6-13.5 cm long, basally attenuate; 
mare with up t flowers; corollas 19-22 mm long; anthers (molading tails) 
. 11-12 mm ks tails of adjacent is onnate s s 5. aracaensis Pruski 
6b. ua phyllaries decurrent onto peduncle, the = one stipitate and the eas 
campanulate; leaves 9-21 cm long, basally cun s0; capitula t 
flowered; corollas (20)28-35 mm long; anthers (including tails) e 16 m 
tails of adjacent anthers connate, but easily d able — s gene Maguire 
lb. Anther filaments attached to corolla tube at the sinuses of the lobes. 
7a. Mature leaves sericeous below 5. sericeus Maguire & Aristeg. 


Tb. Mature leaves glabrous or puberulent below. 
8a. Capitula even cylindric, 5-6-flowered; leaves narrower than 2.3 cm 


S. colveei (Steyerm.) Pruski 


8b. ao he ere) to een qm ca. 12- 100-flowered; leaves wider than 3 cm 
r phyllaries keeled, s S. connellii (N. E. Brown) S. F. Blake 


9b. Du je not Lor. not shiny. 
10a. Leaves to 35 cm long ~S. colombianus Cuatrec. & Steyerm. 
10b. Da shorter than 26 e 
11a. Outer phyllaries ca. " 2 cm long, elliptic-lanceolate, apically obtuse to rounded 
ee Maguire & Wurd. 


p 


l Outer phyllaries shorter than 1 cm, triangular, A 
2a. Leaves shorter than 6 cm, rotund with rounded Ae pm yen DNE 


ensis Aristeg. 
at 


12b. Leaves commonly longer than 6 cm, elliptic to ovate vi. attenuate to 
bases and apices. 
Leaves subsessile, basally cuneate to obtuse; petioles shorter than 7 
mm; young leaves sometimes sericeous 
chimantensis Maguire, Steyerm. & Wurd. 
13b. Leaves ara basally attenuate to narrowly acute; Didi ca. 
m long; leaves glabrous or puberulent, due sericeous io 


S. inm. S. F. Blake 


(Amaral 1628; Prance et al. 28975; Prance & 

land of Guyana and/or Venezuela, the following Guedes 29600; Rosa & Lira 2250; Tavares & 
species are here recorded for Brazil, mostly from — Silva 36). 
Serra Aracá: Fupatorium (Guayania) bulbosum Aristeg. (Pi- 
poly et al. 6664; originally determined as Gua ya- 
nia by H. Robinson). King & Robinson (1987, p. 
412) listed it as restricted to Venezuela. 


Known previously only from the Guayana High- 


Calea abelioides S. F. Blake (Prance et al. 
9810, 9876, 21555; Tavares et al. 141). Th 
first three collections cited above were reported 
uen by re Sete aes due Fupatorium (C ip oa B. L. 
var. [subsp. in protologue] neblinensis Maguire & 

Robinson (Prance et al. ; Rosa & Lira 
Wurd., a taxon not known to occur in Brazil. 
2341). This species is not muni from Brazil on 


Calea lucidivenia Gleason & S. F. Blake p. 387 of King & Robinson (1987). 


Volume 76, Number 4 
1989 


Pruski 1003 


Compositae of the Guayana Highland—ll 


Mikania lucida S. F. Blake (4maral 1578, 
1619; Prance et al. 20058, 29021; Tavares & 
Silva 63; Tavares et al. 120). The provisional 
determination of M. cf. lucida based on Prance 
et al. 20058 from near Auaris, Brazil and reported 
on p. 123 of Huber et al. (1984) is herein con- 
firmed. King & Robinson (1987, p. 425) listed it 


as restricted to Venezuela. 


LITERATURE CITED 


ARISTEGUIETA, L. 1964. Compositae. Flora de Vene- 
uela 10(2): gs 941. 
19 s especies nuevas de pu rur] 
Tribu (Mutisieae). Acta Bot. Venez. 3(1-4): 39-44. 
pr AL. 1977. graph pr ii P 
039-1066 in V. H. Heywood, J. B. Harborne & 
B. L. Turner (editors), The Bioboer and Chemistry 
of the Compositae. Academic Press, mus ork 

HUBER, O., J. A. STEYERMARK, C. T. PRANCE & C. ALES. 
1984. Tho vegetation of the Serra Parima, Ven- 
ezuela-Brazil: some results o Brit- 


tonia 36: 104-139. 


—] 
~J 


Kınc, R. M. € H. Rosinson. 1987. 
brun Monographs Syst. Bot 
Gard. 22: 1-581. 

MAGUIRE, B. oe dep ota bei and evo- 
lution patterns amon ae of the Mis 
Highland of Venezuela. Proc. poss Phil. Soc. 100: 


The genera of the 
. Missouri Bot. 


J. J. Wurpack. 1957. Compositae. /n: B. 
Maguire, J. J. Wurdack & collaborators, The Botany 
of the Guayana Highland — Part II. Mem. New York 
Bot. Gard. 9(3): 366-392. 


. A. STEYERMARK & J. J. Wurpack. 1957. 
Colipositas. In: B. Maguire, J. A. Steyermark, J. 
Wurdack & collaborators, Botany f the Chimanta 


o 
Massif — I. Gran Sabana, Vene veli. Mem. New York 
Bot. Gard. 9(3): 425-439. 
PRUSKI, J. F. 1989. Notes on the Compositae of the 
Guayana Highland-l. A new species of Stomato- 
chaeta and the reduction of Guaicaia to Glossarion 
ME Mutisieae). Nina 41: 35-40. 
STEYERMARK, J. A w species from the summit 
of Cerro Guaiquinima, ala Brittonia 32: 17- 


f 84. Flora of the i pit es 
Ann. NE Bot. Gard. 71: 29 


CHROMOSOME NUMBERS 
IN COMPOSITAE, XVI: 
EUPATORIEAE IT' 


Harold Robinson, A. Michael Powell,’ 
Gerald D. Carr,* Robert M. King,? and 
James F. Weedin’ 


ABSTRACT 


An additional 159 po reports are provided for the tribe Eupatorieae with 13 
180 genera and over 450 
t of the subtribes, including the Adenostemmatinae (with first report 
of Sciadocephala), Eupatoriinae, Gyptidinae, most of the Rectatibas and Alomiinae, Liatrina 
Praxelinae. Th i 


species reports. Numbers own include 94 of the 


A primary base of x — lo is confirmed for most 


Hartw rightia), Fleischmanniinae, Critoniinae, and 


ew generic and 60 new 
of the more than 2,000 species. 


e (with first report of 


of n — 9 is consistent for four additional 


species in Ac ritopappus of the Ageratinae, independent of that base number à in the Brickellia group of the Alomiinae. 


Numerous pora a a 
numbers of x l 95 are found in new report 
umbers si n = ca. dl 26 are noted in two spec 


added for the Central American polyploid complex in Fleischmannia. n higher base 
s ui the subtribes Neomirandeinae, Mikan 
Kaunea of the Oxylobinae with correction of o 

ower y credited to K. DRE p ‘eeu of the two bases, x = 10a 
are ditd: and dee in Bartlettina and the Critoniine genus Eupatoriastrum are discussed. 


nae, and a 


nd 16, in the Hebecliniinae 


This paper continues a series dealing with chro- 
mosome numbers of Compositae (Raven et al., 
1960; Raven & Kyhos, 1961; Ornduff et al., 1963, 
1967; Payne et al., 1964; Solbrig et al., 1964, 
1969, 1972; Anderson et al., 1974; Powell et al., 
1974, 1975; King et al., 1976; Tomb et al., 1978; 
Robinson et al., 1981, 1985) and is the second 
dealing with the tribe Eupatorieae (King et al., 
1976). New reports are provided for 13 genera 
and 59 species (marked by ** 
in Appendix). 

The new reports in this paper are based primarily 
on material collected by R. M. King and counted 
by A. M. Powell and J. F. Weedin (not coded) and 


G. D. Carr (C). The chromosome counts have been 


and * respectively 


made from aceto-carmine or aceto-orcein squashes 
of microsporocytes in meiosis. Voucher specimens 
of the King collections are in US and a second set 
is in MO 

The previous paper in the series dealing with 
the Eupatorieae summarized the numbers then 
known from 327 of the approximately 2,000 species 


and 70 of the 180 genera now known in the tribe. 
The chromosome data were summarized again with 
few additions in the survey of the tribe by King & 
Robinson (1987). Studies subsequent to that of 
King et al. (1976) have added reports of ten ad- 
ditional genera: Urolepis n = 10 (Turner et al., 
1979; Waisman & Rozenblum, 1986); Hatsch- 
bachiella n = 10 (Waisman & Rozenblum, 1986); 
Lorentzianthus n = 10 (Bernardello, 
Alomia n — 10 (Mabry et al., 1981; B 
Turner, 1986; Sundberg et al., 1986); Amolinia 
n — 10 (Strother, 1983); Flyriella n = 10 (Baker 
Sí Turner, 1986; Sundberg et al., 1986); Eryth- 
16, Eupatoriastrum n = 16; Kyr- 
steniopsis n — 10; and Piptothrix n — ca. 17 
(Sundberg et al., 1986). 

The present paper adds chromosome counts for 
the 13 additional genera: Bahianthus n — 10; 


radenia n — 


Bejaranoa n — 10; Bishovia n — ca. 10; Cron- 
quistianthus n — ca. 10; Fleischmanniopsis n — 
10; Goyazianthus n — 20; Platypodanthera n — 
10; Pseudokyrsteniopsis n — 10; Sciadocephala 


' We thank Dr. Richard Wunderlin of the University of South Florida for collecting buds of Hartwrightia and 
Dr. Caloway Dodson of the Missouri Botanical Garden for collecting buds of Sciadocephala. We also thank Susan 
Richardson of the Smithsonian Institution for growing specimens from seed. Part of the work was supported by 


Nat ional Science Foundation 


grant #DEB 77-13457 to R. M. 


Wn 


illa nt of Botany, National Museum of Natural History, Shonin Institution, Washington, D.C. 20560, 


3 r of Biology, Sul Ross State University, Alpine, Texas 79832, U.S.A. 


‘ Department of Botany, University of Hawaii at 


Manoa, Honolulu, 


Hawaii 96822, U.S.A 


* Division of Science, Community College of Aurora, Aurora, Colorado 80011, U.S.A. 


ANN. Missouni Bor. GARD. 76: 1004-1011. 1989. 


Volume 76, Number 4 
1989 


Robinson et al. 1005 
Chromosome Numbers in Compositae, XVI 


n — 10; and Trichogonia n — 10. The new counts 
of n — ca. 20 and ca. 26 for Kaunea seem to be 
the first reasonably representative figures for the 
genus. Chromosome numbers are now known from 
94 of the 180 genera. 

A few previously reported numbers are based 
on vouchers needing corrections of identification. 
The name Ageratina aschenborniana (Schauer) 
K. & R. used in earlier papers is a synonym of A. 
bustamenta, and Ophryosporus origanoides 
[Meyen & Walp.] Hieron. is a synonym of O. 
heptanthus. The plant reported by Coleman (1968: 
235, Brazil: Coleman 295) as Ageratum cony- 
zoides L. is Barrosoa betoniciforme (DC.) K. & R. 

The general pattern of chromosome numbers in 
the tribe was first presented in King et al. (1976). 
Thirteen of the 18 subtribes had a base of x — 10 
or clearly derived numbers, such as x — 15. The 
remaining five subtribes show higher base numbers 
with no remnant of x = 10. As elaborated from 
the chart in King & Robinson (1987), the subtribes 
have the following numbers with primary bases in 
boldface: Adenostemmatinae 5, 10; Eupatori- 
inae 10, 15, 20; Disynaphinae 10; Gyptidinae 
10; Ageratinae 9 (Acritopappus), 10, 11, 12 
(Microspermum- Piqueria-Stevia group), 15, 20, 
25; Trichocoroninae 15; Ayapaninae 10; Alo- 
miinae 9 (Brickellia group), 10; Liatrinae 10, 
20, 30; Fleischmanniinae 4 (Fleischmannia 
microstemon), 10, 20, 30; Critoniinae 10, 20; 
Praxelinae 10, 20, 29; Hebecliniinae 10, 16; 
Neomirandeinae 17, 20, 24, 25; Mikaniinae 16, 
17, 18, 19, 20; Oxylobinae 16, 17, ca. 20, 51; 
Hofmeisterinae 18, 19; Oaxacaniinae 18. The 
pattern has not been altered by any of the more 
recent reports, and it is actually greatly reinforced 
by numerous new chromosome counts in the sub- 
tribes Ageratinae, Fleischmanninae, Hebecliniinae, 
and Oxylobinae. The only questionable additions 
since 1977 are in the Critoniinae and Hebeclini- 
inae. Questionable counts have often been over- 
turned by subsequent examination of vouchers, as 
in the case discussed under Kaunea in the Oxy- 
lobinae, or by reexamination of the species. The 
patterns seem well established, and deviations should 
be carefully rechecked. 

The significant new reports are best reviewed in 
the order of the subtribes established in King & 
Robinson (1987). 


ADENOSTEMMATINAE 


The new report of n = 10 for Sciadocephala 
is the first for that genus and the first in the subtribe 
outside of Adenostemma. Evidently both genera 


retain the basic number of the tribe. There has 
been no further evidence in the subtribe for the 
low n = 5 reported by Turner & Irwin (1960) 
from a plant now determined as 4. involucratum. 
Our new report for that species is n = . The 
two genera of the subtribe with the anomalous 
knob-formed pappus have now been counted, but 
the remaining genus Gymnocoronis, which lacks 
such a pappus, is still unstudied. 


EUPATORIINAE 


The Waisman € Rozenblum (1986) count for 
Hatschbachiella completes the survey of genera 
in the subtribe. All four genera show a base of x 
= 10, with Eupatorium (including Eupatoriadel- 
phus) showing some triploids and tetraploids. 


GYPTIDINAE 


The new generic reports for Bahianthus, Be- 
jaranoa, Litothamnus, Platypodanthera, and 
Trichogonia, and the Urolepis counts (Turner et 
al., 1979; Waisman & Rozenblum, 1986) show 
that n = 10 is basic to the subtribe. The recent 
cytological study of Gyptis pinnatifida Cass. (as 
Eupatorium tanacetifolium Gillies ex Hook. & 
Arn. by Rozenblum et al., 1988) confirms the 
triploid condition based on n = 10 that is already 
known to occur in that genus and examines details 
of the apomixis in the species. 


AGERATINAE 


The many new counts of Acritopappus, based 
on four species, strongly confirm the occurrence 
of n = 9 first noted for one of the species by 
Coleman (1970) and mentioned by King et al. 
1976). The isolated occurrence of n = 9 in Ac- 
ritopappus is apparently consistent for the genus, 
with reports from both viscid-leaved and non-viscid- 
leaved members. Prior to the discovery of n = 9 
in Acritopappus, the number was known in the 
tribe from only Brickellia and its close relatives 
in the subtribe Alomiinae. 

ther reports of chromosome numbers of Age- 
ratinae since King et al. (1976) confirmed other 
known patterns. New counts of Ageratum spp. 
show a base of 10. The Dillon & Turner (1982) 


report of ca. n = 10 for Ascidiogyne sanchez- 


— 


vegae Cabrera conforms with the n — 10 already 
known for the type species 4. wurdackii Cuatr. 
This further indicates the difference between the 
latter genus and the members of the Piqueria 
relationship. New reports from Piqueria and Stevia 


and the Strother (1983) and Sundberg et al. (1986) 


1006 


Annals of the 
Missouri Botanical Garden 


reports for Microspermum debile Benth. and M. 
nummularüfolium Lagasca conform to the already 
known bases of n — 12 and n — 11 in that group. 
The South American species reported for Stevia 
all have n — 11, and none show the n — 12 found 
in shrubby members of the genus in Central Amer- 


ALOMIINAE 


he new generic counts for Goyazianthus, 
Planaltoa, and Pseudokyrsteniopsis; generic 
counts by others for Alomia, Flyriella (Baker & 
Turner, 1986; Sundberg et al., 1986) and Kyr- 
steniopsis (Sundberg et al., 1986); and the counts 
for all three species of Carminatia (Turner, 1988) 
extend our knowledge of elements of the subtribe 
having the base number of 10. The Turner report 
for his new species, Carminatia anomala, is here 
attributed to the earlier name for the species, C. 
alvarezii Rzed. & Calderón de Rzedowski (1987). 
Chromosome numbers of the genera could all be 
anticipated on the basis of their evident relation- 
ships within the subtribe. The number n — 9 that 
is notably present in the Alomiinae in Brickellia 
(King et al., 1976; Keil et al., 1988), Barroetea 
(Sundberg et al., 1986), Pleurocoronis, and their 
immediate relatives is unknown in any of the pri- 
marily South American genera of the subtribe. 


LIATRINAE 


The new chromosome count for Hartwrightia 
completes the survey of the six genera of this small 
subtribe from the southeastern United States. All 
members of the subtribe have n = 10 or polyploids 
of that number. 


FLEISCHMANNIINAE 


The numerous new reports for Fleischmannia 
firmly establish n — 10 as the base for most of the 
species. No further examples have been found of 
the n — 4 that is characteristic of the weedy annual 
F. microstemon (Cass.) K. & R. and that has been 
reported for F. hymenophyllum by Grashoff et al. 
(1972). 


The counts reported in the present study formed 
the basis for statements regarding the polyploid 
complex first noted by King & Robinson (1978, 
1987) in Central America. The geographically re- 
stricted Guatemalan species Fleischmannia bohl- 
manniana K. & R., F. deborabellae K. & R., 
and F. viscidipes (B. L. Robins.) K. & R. prove 
to be diploid, while the more widely distributed 
weedy species F. pratensis (Klatt) K. & R. and 


F. pycnocephala (Less.) K. & R. continue to show 
a consistently polyploid condition. 


CRITONIINAE 


The three new generic reports for the subtribe 
(Bishovia K. & R., Cronquistianthus K. & R., 
and Fleischmanniopsis) and the Lorentzianthus 
count of Bernardello (1986) show the base of n — 
10 that is characteristic of all previously known 
members of the group. The only serious suggestion 
of another base in the subtribe seems to involve 
the presently uncounted genus Eupatoriastrum, 
which is noted for its paleaceous receptacles. The 
species belonging to this genus have been associ- 
ated with a similarly paleaceous species, Fupato- 
rium corvi McVaugh (McVaugh, 1972) of Chia- 
pas, which has an n = 16 (King et al., 1976). The 
latter is now recognized as the genus Matudina of 
the subtribe Hebecliniinae, and shows no detailed 
structural evidence of immediate relationship to 
‘upatoriastrum. In contrast, structural features 
of Eupatoriastrum suggest close relationship to 
Koanophyllon, which has numerous reports of a 
base of n = 10 in this study and in previous studies 
based on Central and South American material 
(King et al., 1976; Keil et al., 1988). 

Therefor: it is surprising to see a report of n 
= l6 from Fupatoriastrum nelsonii Greenman 
(Sundberg et al., 1986). The voucher cited by 
Sundberg et al. from Coatepec, Veracruz (Her- 
nández 320) has been sought for study, but the 
specimen is not deposited at TEX as indicated. The 
actual repository, XAL, has not replied. A specimen 
of E. nelsonii (Hernández 355 TEX), collected 
from the same place, is correctly determined. The 
latter is the voucher for the chemical study by 
Bohlmann et al. (1985). It is possible that the n 
= 16 reported from Eupatoriastrum nelsonii is 
correct, and that the genus is a member of the 
Hebecliniinae rather than the Critoniinae. No char- 
acteristic of the genus precludes this. In fact, the 
form and texture of the corollas of Fupatoriastrum 
are rather like that of various members of the 
Hebeclinium group. Structural details indicate that 
placement within the Hebecliniinae, if correct, would 
be closer to Decachaeta than Matudina. Although 
the structural characters of the Hebecliniinae are 
less definite than those of most subtribes, there is 
no reason to believe that n = 16 has arisen more 
than once from x — 10. 

King et al. (1976) placed Peteravenia in the 
Hebeclinium group and reported chromosome 
numbers of n — 10 and n — ca. 17. The genus 
was transferred to the Critoniinae in King & Rob- 


Volume 76, Number 4 
1989 


Robinson et al. 1007 


Chromosome Numbers in Compositae, XVI 


inson (1987), where the report of n — ca. 17 was 
rejected from the genus as the voucher proved to 
be Ageratina subinclusa (Klatt) K. & R. Two new 
counts further establish n — 10 as the base number 
for Peteravenia. 


PRAXELINAE 


New reports for Chromolaena and Praxelis (Ap- 
pendix) and reports in recent literature (Sundberg 
et al., 1986) show the trend already known for 
high chromosome numbers and irregular meiosis 
in the subtribe. 


HEBECLINIINAE 


King et al. (1976) noted the Hebeclinium group 
for the presence of two distinct base numbers, x 
= 10 and x = 16. The members of the subtribe 
with n = 10 were Hebeclinium and the South 
American species of Bartlettina. In contrast, the 
Central American genera Erythradenia (Sundberg 
et al., 1986), Decachaeta, and Matudina, and the 
Central American species of Bartlettina showed n 
— 16. The geographical pattern seemed to be most- 
ly South America versus Central America, with 
Hebeclinium being basically South American. Sub- 
sequent reports have generally confirmed the pat- 
tern. Strother (1983) reported n — 16 for four 
Central American species, Bartlettina breedlovei 
K. € R., B. pansamalensis (B. L. Robins.) K. & 
R., B. pinabetensis (B. L. Robins.) K. & R., and 
B. tuerckheimii (Klatt) K. & R., and the same 
number was reported by Sundberg et al. (1986) 
for B. hintonii K. & R. In an apparent exception 
to the characteristic n — 16, two Central American 
species of Bartlettina tamaulipanum (B. Turner) 
K. & R. and Eupatorium xalapanum B. Turner 
(7 B. brevipetiolata (Schultz-Bip. ex Klatt) K. & 
R.) have been reported as n = 10 (Turner, 1985). 
The types have been seen, and they are clearly 
members of the Central American complex in the 
genus. The reports of n — 10 are greatly in need 
of confirmation. 

The only basically Central American element of 
the Hebecliniinae presently known with n = 1 
seems to be Amolinia heydeana (B. L. Robins.) 
K. & R. reported by Strother (1983). 


NEOMIRANDEINAE 


New reports for this monogeneric tribe include 
one species of each subgenus of Neomirandea. The 
reports of n = 17 for N. eximia agree with earlier 
counts. The reports of n = ca. 26 for N. angularis 
are the first of that number in the genus, but they 


offer limited consistency in a member of the typical 
subgenus of Neomirandea, which has previously 
been reported variously as n — 20, n — ca. 24, n 
— ca. 25 (King et al., 1976), and the species has 
recently been reported as n — 25 by Sundberg et 
al. (1986). 


OXYLOBINAE 


Reports since King et al. (1976) have provided 
an expected count of n — ca. 17 for Piptothrix 
(Sundberg et al., 1986) and supported previously 
known chromosome patterns of n — 16 for Oxy- 
lobus (Turner & Kerr, 1985) and n — 
Ageratina subgenera Ageratina, Neogreenella, 
and Klattiella (Jansen et al., 1984; Sundberg et 

al., 1986). Strother (1983) noted a minor Ss 


} 


17 in 


ture of n = 18 for Eupatorium subp 
(= Ageratina subinclusa (Klatt) K. & R. The 
present study, Jansen et al. (1984), and Waisman 
& Rozenblum, (1986) give further examples of the 
subgenus Andinia, i.e., Ageratina ampla (Benth.) 
, A. dendroides, A. pseudochilca, A. 
Dr. (Hieron.) K. & R., 4. angustifolia 
(H. B. K.) K. € R., 4. dee (Benth.) K. & 
Ry A baccharoides (H. B ) K. & R., and 4. 
elegans (H. B. K.) K. & E ue counts confirm 
the consistent n — ca. 40 and n — ca. 42 already 
seen by King et al. (1976) in the subgenus except 
for the higher polyploid n — ca. 90 count for 4. 
angustifolia by Waisman & Rozenblum (1986). 
The new count of n = ca. 20 for Kaunea ig- 
norata seems to reinforce the generally high num- 
ber 2n — near 45 found by Royce Oliver in root 
tips of that species (King & Robinson, 1980). The 
new report of n = ca. 26 for Kaunea lasioph- 
thalma conflicts with an earlier report of n — 10 
for Eupatorium lasiophthalmum Griseb. (Turner 
et al., 1979), but the voucher for the latter report 
has been examined and proves to be Lorentzian- 
thus viscidus, a species known to have an n — 
10. It seems that Kaunea characteristically has a 
chromosome number at the level of n = 20-26, 
which is consistent with higher 
known members of subtribe Oxylobinae. The higher 
numbers in Kaunea are reminiscent of those of 
another Andean member of the subtribe, 
tina subg. Andinia. Nevertheless, the numbers in 
the two groups do not seem to be the same, and 
if other aspects of the plants are any indication of 
relationship, the higher numbers evidently origi- 
nated separately. Structurally, Kaunea seems to 
be one of the most divergent elements of the Oxy- 
lobinae, and it is not now considered close to any 


numbers in other 


Agera- 


part of Ageratina. 


1008 


Annals of th 
Missouri Botanical Garden 


LITERATURE CITED 


, D. W. Kyuos, T. Mosqu A. M. 

L& Pi H. Raven. 1974. o num- 

dee in pese IX: Er and other 
Astereae. Amer. J. Bot. 61 

BAKER, M. M. & B. L. TURNER. “Taxonomy of 

300 


Flyriella (Asteraceae- Eupatorieae). “Sida 1 
317 


ANDERSON, 


BERNARDELLO, L. M. 1986. Numeros cromosómicos en 
Asteraceae Padre Darwiniana 27: 169-178. 
BOHLMANN, F., ERO & B. L. TURNER. 1985. Ger- 


mecranides em Eupatoriastrum nelsonii. Phy- 
1263-1266. 


COR. J. 1968. Chromosome numbers in Brazilian 

Compositae. Rhodora 70: 228-240. 
1970. Additonal chromosome numbers in 

Braailian Compositae. Rhodora 72: 94-99, 

DILLON, M. & B. L. TURNER. 1982. diu num- 
bers of Peruvian Compositae. Rhodora 

ED. J. L., M. W. BIERNER & D. K. ein TON. 
1972. bremen numbers in North and Central 

American bugs gun 24: 379-39 

JANSEN, R. K., T , S. Díaz- A & V. 
A. "FUNK. 1984. ipak n Com- 
positae de pne Caldasia 14: 7-20 

Kern, D. J., M. Luckow & D. J. PINKAVA. 1988. 
Chromosome rin es in Asteraceae from the United 
States, Mexico, the West Indies, and South America. 


~ 


Amer. J. Bot. 75: 652-6 
Kinc, R. M. & H. Rowinson 1978. Studies in the 
d rieae (Aster CLXIX. Two new species 


leisc ina from Guatemala. Phytologia 38: 
n» 420. 

—— 80. Studies in the Eupatorieae 
Pi C C II. A new genus Kaunea. Phytologia 
257-260 


The genera of the Eupa- 
torieae (Asteraceae) Monographs in ei Bot- 
any, Missouri Bot. Ga 

—. cl V. KYuos, A. y POWELL, P. H. RAVEN & 
H. INSON. 197 Chromosome numbers in 
mutis dt XIII: Eupatorieae. Ann. Missouri Bot 

Gard. 63: 862-888 


á: 7X 


Manny, T. J., B. N. TIMMERMAN, N. Hem & A. M. 
OWELL. 1981. Systematic implications s of the fla- 
vonoids and chromosomes of Flyriella (C = d 
Eupatorieae). Pl. Syst. Evol. 137: 275-280 
19 Tribe III. Eupatorieae. ie Con 
positarum oe Pugillus. Contr. Univ. Mich. 
igan Herb. 9: 08. 

ORNDUFF, R., T. uu IN, D. W. KvHos & P. H. RAVEN. 
1967. Chromosome numbers in Compositae, VI: 
Senecioneae E Amer. J. Bot. 54: 205-213 

,P.H. ven, D. W. KvHos & A. R. KRUCKE- 
BERG. 1963. Chromosome numbers in Compositae, 
Ill: vp qp Amer. J. Bot. 50: 131-13 

Payne, W. W., P. H. Raven € D. W. Kyuos. 
Chromosdtmie Ae tin in Compositae, IV: Am 
sieae. bras . Bot. 51: 419-424. 

PowkELL, A. M., D. W. KyHos & P. H. Raven. 1974. 


I n mbari in Compositae, X. Amer. J. 
Bot. 6 913. 


i 


—— € —— ———. 1975. Chromosome 
ls E IAN XI: Heleniéae: Amer. J. Bot. 
62: 10 
Raven, P. H. & D WY. KvHos. 1961. 


Chromosome 


numbers in Compositae, II: Helenieae. Amer. J. Bot. 
48: jos 8 


T SOLBRIG, D. W. Kynos & R. 


ro Chromosome numbers in "C ANE T p 
. Amer. ot. 47: pe 
RON. H., A. M. PowEL & J. F. 


WEEDIN. 1981. Chromosome num ne rs in Com- 
peste, € Helianthene: Smithsonian Contr. Bot. 
52: 1-28 


— — & —— Chro- 
mosome —Ó— in Compositae, XV: Liabeae. Ann. 
Missouri Bot. Gard. 72: 469-479. 

ROZENBLUM, E., S. MALDONADO & C. E. Waisman. 1988. 
Apomixis in E us aii (Composi- 
tae). An Aes . Bot. 

RZEDOWSKI, J. . CALDERÓN DE d E 1987. 
iniu N i arezii, una nueva especie mexicana 
de Compositae, Eupatorieae. Anales Esc. Nac. Ci. 

: 9-11. 


0. T. L. C. ANDERSON, D. W. doas P. H. 
RAVEN: 1969, 'hromosome numbers in Compos- 
itae, VII: cda gu r. J. Bot. 56: 348-353. 

——— RM . KvHos, "M Pow WELL & P. H. RAVEN. 
1972 o sustbene- in bru a VIII: 
Hen Amer. J. Bot. 59: 878. 

, L. C. ANDERSON, D. W gwa P. H. RAVEN 

& a ES DENBERG. 1964. Chonuesamh pu in 

ror E V: Astereae II. Amer. J. Bot 513- 

id] J. L. 1983. More chromosome studies in 
31224. 

. TURNER. 1986 

Uomo counts of Latin eel Compositae. 
Amer. J. Bot. 73: 33-38. 

Toms, A. S., K. L. CHAMBERS, D. W. Kyu A. M. 
POWELL & P. H. Raven. 1978. aa num- 
bers in the em XIV: Lactuceae. Amer. J. 
Bot. 65: 717-72 

TURNER, B. L. wo new species of Eupatorium 
(Asterac eae) from northeastern Mexico. Brittonia 37: 
373-377. 

————. 19 88. Taxonomy of Carminatia (Asteraceae, 
Eupatorieae). Pl. Syst. Evol. 160: 160-169. 
——— & H.S. Hen Chromosome numbers 
in the Compositae. Meiotic counts for fourteen 
pm of Brazilian Compositae. Rhodora 62: 122- 

126. 


& K Kerr. 1985. Revision of the genus 
Oxylobus pe -Eupatorieae). Pl. Syst. Evol. 

ols 78- el 
, L. UnBaATSCH & B. SIMPSON. 1979. 


= 


ae. Ame 173-1 

WAISMAN, C. ROZENBLUM. 1986. Estudios 
iure en Compositae. III. Darwiniana 27: 179- 

189. 


UD Aia in South American Composi- 
. Amer. J. Bo 


APPENDIX. Chromosome counts and vouchers for 
species of Eupatorieae. a generic reports are marked 
new species reports *. K num 
collection numbers. Carr counts are m 
counts are by Powell ne Weedin unless marked other- 


* Acritopappus confertus ( (Gardner) K. & R. =9, + 
|. BRAZIL. BAHIA: | km S of Andarai rae road to 
Mucugé. K8708 (C). 


Volume 76, Number 4 
1989 


Robinson et al. 1009 
Chromosome Numbers in Compositae, XVI 


ar hagei K. = 9, BRAZIL. BAHIA: 
S of yaer K8711 C). 


* Ac ropas ie — 9, BnaZIL. BAHIA: 52 km E 
of * . K8773 (C. and Royce pad 
*Acritopappus morii K. & R., type. n = 9. BRAZIL. 
BAHIA cugé. K8172. 
* 4c rtopappus morii. n = 9. BRAZIL. BAHIA: 9 km N of 
ucugé. K8768 


* Ac a proa K. & R. n = ca. 9. BRAZIL. 
BAHIA: 8 km S of Mucugé. K8749 (C 2). 

Adenostemma involucratum K. R. n = 10. BRAZIL. RIO 
DE JANEIRO: grown from seed collected at Botanical 
( 1. K9701. 

Adenostemma tro Cass. n = . BOLIVIA. 
COCHABAMBA: 33 km from Colomi. KT D. 
Ageratina ener ie (Klatt) K. & R. n = 17. 
ICA. SAN JOSÉ: La Palma. K9657 (C). 
Ageratina azangaroensis (Schultz-Bip.) K. & R. n — 


^7 


. COCHABAMBA: Challa. K7539. 


7 OLIVIA 
dgeratina az Mi, ea n= 17. "s COCHABAM- 
m Challa. K754 
d az on roensis. n = Hyi [TM LA PAZ: ca. 
30 km from La Paz. K74 
Ageratina xis d n — ca. 17. PERU. ANCASH: 
Catac. K9071 T 


Ageratina azangaro 2 B chromo- 
somes. Bo OLIVIA. COCHABAMBA: 4 km W of Epizana. 


JK. & R n= 17. 


Ageratina bustamenta (DC 
22 km W of San G ris- 


MALA. ALTA lign ca. 
tóbal Verapaz. K7352. 
Ageratina o n= 
km SE of Panajachel. K7? 
— hustamenta, n 
ENANGO: 16 k 


s ATEMALA. SOLOLA: 13 
as CHIMAL- 


d of Patzun. Km 


Aeeranina “bustamer nt EMALA 
QUEZALTENANGO: 14 La SE of e eS 
K 1259. 

Ageratina bustamenta. n = 17. ATEMALA. 


QUEZALTENANGO: 4 km SW of San Mateo. K7266. 
Ageratina tanta n = 51. GUATEMALA. CHIQUIMULA: 
14 km E of Ipala. K7383. 


i Ageratina dendroides (Sprenge l) K. ca. 40. 
Ec p 


& R. n= 


LOJA: 1 E of Loja. K7911. 
Ageratina matretiana (DC.) K. = 17. Gua- 
TEMALA. HUEHUETENANGO: 5 km N of Chiantla. 
K?7: 
Ageratina matretiana. n 17 ESTA HUEHUE- 
TENANGO 8 km E of C Made K731 
* Ageratina ple hile a (Benth.) K. & R. n = ~ 42. 
ECUADOR. AZUAY: 5 km S of Cumbe. K775 
K. € R. n= 10. Duci 


dia eran (Cabe 
5 km N EG salina pe 
diit astute n= 10. . GOIAS: 2-7 km 
ESE of Cristallina on the Ned | to y Belo ius 
Age ratum tii: n = 10. BRAZIL. MINAS GERAIS: 
a do Cipo 2 km wen Chapeu de Sol. K8363. 
n. being de n= 10. BRAZIL. GOIÁS: 5 km NE 
of Cristallina. K8960 (€ 
ie nd rugosum J. Coulter. n = 20 + 1-2 fragments. 
TEMALA. CHIMALTENANGO: 12 km WSW of Chi- 
ao K71 
* dristeguietia pseudoarbor a (Hieron.) K. 
. PERU NAS: mountains SE of € irm 
as. K9208 


end digne us ns(Don) K. & R. n — Lo ~ IVIA. 
: 1 km from Comarapa. K76 
dustrorupatorium chaparense (B. L. "udis A : 
n 1 OLIVIA. COCHABAMBA: 40 km from Co- 
marapa. 'K7660. 
yapana e (Lam. K. &R. n= f BOLIVIA. 
LA PAZ: 27 km from Chulumani. K74 

"+ Bahiani, prar (Baker) K. & R. n = ca. 12. 
uem ZIL. BAHIA: A 30 km ao S - Aai Na estrada 

vai T Jussiape. K81* 

** Bahianthus viscidus. n = ca. 10. BRAZIL. BAHIA: A 
3 km ao S de ur cugé. Na estrada que vai par 
Jussiape. K816 

** Bahianthus 7 n= 10. BRAZIL. BAHIA: Mucugé. 
K8169. 


lis: cian esi is (Hieron.) K. € R. n = ca. 20. 
B IA: Trecho Vitoria da Conquista BR 415 
Ilheus/ ‘Brumado Anagé km 260. K8032. 

** Bejaranoa balansae. n — 10. BOLIVIA. SANTA CRUZ: 

23 km E of Comarapa. K9654 (C). 


** Bishovia bolivensis K. & R., type. n= ca. 10 (possibly 


2n = 10, + 1, or more than 10,; chromosomes 
morphologically domat M IA. SANTA CRUZ: | 
km from Coma K76 


Brickellia argy role B. L. Re m n — 9. GUATEMALA. 
SE of Pongan hel. K7230. 


SOLOLA 
xc pon n= 9. Gu ATEMALA. JALAPA: Ca. 
W of San Luis. 384 
*Campulo linium megacephalum e. Marius s) K. & R. 
— 20-22. BRAZIL 2 km S of Cristallina. 
K8252. 
*Campuloclinium ed lage tea 2n = 
somes. BRAZIL. GOIÁS: 20 s y p Alto 
Goiás. K881: 


is 

C 'hromolaena arnottiana eis ) K. & R. n= 27-: 
univalents. BOLIVIA NTA CRUZ: 1 km from Co- 

marapa. K7029 
Chromolaena arnottiana. 2n = 8-9, + 10- = 28- 
2 ARGENTINA. CATAMARCA: Cà. pt b 4 of Ca- 
461 (C. 

Chromolaena collina (DC.) K. & R. n = 10. GUATEMALA. 
PAZ: ca. 24 km W of San Cristóbal Vera- 


7 


ES 


ien c 9. 
*C oon Qo ese bn :hultz-Bip. ex Baker 
.n- RAZIL. € : 12 km S of Alto Paraiso 
de Goiás. K887 l (C. 
*Chromolaena cryptantha. n = 10. BRAZIL. 
km N of Cristallina. K8944 (C). 
*Chromolaena pseudinsignis K. & R., type 
ably 10. BRAZIL. GOIÁS: Chapada dos Veadeiros, 18- 
9 km N of Alto Para 
*Chromolaena tunariensis (Hieron.) K. & R. 
1,. BOLIVIA. COCHABAMBA: Cercado. a 
Conocliniopsis prasiifolia (DC.) K. R. n= ca. 
AZIL. BAHIA: com entrada no km 20 da Rodovia 
Ihéus/ Uruçuca (BA262). K8012 
ad ipia prasiifolia. n — 10. BRAZIL. 
n E of Ma 


GOIAS: 25 
. n = prob- 
aiso. K8279 

=11, 
(C). 


BAHIA: 40 


acas. K8019. 
e sae prastifolia. n = 10. Piu BAHIA: 12 
km N of Rio Verde Pequeno. K8588 
ipis id coelestinum (L.) DC. n — io U.S.A. 
MA D: Montgomery P Potomac. K8430. 
= 10. PERU. 


C undis acuminata (H.B.K. 
AMAZO 4 km from Chachapoyas K9227 (C). 


C ritoniella iaa n= 1 e . CAJAMARCA: 4 km 


n Ignacio. K9300 


o 
aan es K. & R. n = ca. 


1010 


Annals of the 
Missouri Botanical Garden 


10. PERU. AMAZONAS: 8 km E of Chachapoyas. 
K9164 (C). 

* Disynaphia dl un DC.) K. & R. n= 10. BRAZIL. 
DISTRITO 


7 


RAL: vicinity of the U aa of Bra- 
silia campus. 7 

Fleisc eri hohinammnena K. & R., type. 10. 

ATEMALA. CHIMALTENANGO: ca. 9 bn NW of . Anti- 


e 
Fleischmannia bohlmanniana. n= 10. Gu nt CHI- 
ALTENANGO: 14 km NW of Patzun. K720 


Fleischmannia Mpeg lon n = o estis 
SOLOLA: 3 km SE o najachel. K 72 
Fleischmannia bohImanniana n = ue Es JATEMALA. 


SUCHITEPEQUEZ: 20 km N of Patulal. p 8. 
Fleischmannia bohimanniana, n = TEMALA. 
QUEZALTENANGO km S of Quezaltenango. K7255. 
Fleischmannia aves nn n . GUATEMALA. 
QUICHE: 18 km SE of Santa Cruz del Quiche. K 7287. 
Fleisc O bohlmanniana. n = GUATEMALA. 
APAN: n. the Pan American Highway, ca. 
an LS of Huehuetenango border. K7299 
Fleise os deborabellae K. € R. n = 10. Gua- 
TEMALA. ALTA VER E ^ ca. l6 km W of San Cris- 
tóbal Verapaz. sa 1346. 
Fleisc sele: d i L. Robins.) K. & R. 
. ECUADOR. ZAMORA-CHINCHIPE: 25 km W of Za. 
8. 


-l 
oz 


iora. 
Fleischmannia pratensis (Klatt) K. & R. n= 20. ¢ 
MALA. ALTA VERAPAZ: 5 km WNW of San Cristobal 
. K7336 


Fleischmannia praten nsis. n = 20. GUATEMALA. IZABAL: 
ca. 27 km NE of Gualan. pa 371. 
Fleisc hmannia pycnocephala (Less.) K. & R. n = ca. 


20. GUATEMALA. SACATEPEQUEZ: ca. 24 km N of 
Escuintla. K7185. 
gis mannia m ephala. n = 20. GU Pu Vue A. SA- 


ATEPEQUEZ: 7 km NW of ta K71 
Pa Ora po is n = 20. 
SOLOLA: 5 km SE of Panajachel. K 7223. 
Fleischmannia s ephala. n — 40. GUATEMALA. ALTA 
VERAPAZ: 2 km from Coban. K733 
Fleisc adi p ephala. n — 40. Gu SERM A. BAJA 
VERAPAZ: 1 S of Purulha. K7362. 
Fleisc — pyene haloides (B. L. Robins.) K. & 
. 90. Gu, ATEMALA. CHIMALTENANGO: 16 km 
of Patzun. K7205 
Fleischmannia p» phaloides (B. L. Robins.) K. & 
R. n = ca. GUATEMALA. TOTONICAPAN: 33 km 
San Cristóbal i uc. K7294. 
Fleisc mammis pycnocephaloides var. Meere (B. 
. Robins.) . n = 30. EMALA. TOTONI- 
APAN: 2 km NE of Tornicopan. 'K7270 
Fleisc imana r. glandulipes. n = 
30. GUATEMAL o 2 km NE « j£ Toto: 
nicapan. K727 
Fleischmannia sp. n = - 90 * uud univ alent. GUATEMALA. 
SUCHITEPEQUEZ: ca. 3 km W of Cocales. du irs 
* Fleisc hmannia viscidipes (B. L. Robins.) K. & R 
MALA. kp ce » l6 km N o San 
Cristóbal Totonic apan. K71 
*Fleischmannia viscidipes. n = 


UATEMALA. 


TO TONICAPAN: 


GUATEMALA. 


OTONICAPAN: 18 km NE of ace K7282. 

** Fleischmanniopsis leucocephala (Benth.) K. & R. n 

= GUATEMALA. ESCUINTLA: ca. 16 km SW of 
AM an. 


NA Fleischmazimopiis uña n = 10. GUATE- 


MALA. QUEZALTENANGO: 30 km S of Quezaltenango. 
7252. 


725 
**Goyazianthus tetrastichus (B. L. Robins.) K. & R. 
2n — 20, erratic, d spindles. BRAZIL. GOIÁS: 
12 km 5 of Alto Paraiso de Goiás. K8881 (C) 


*Graz zielia dimorpholepis Baker n= 10. BRAZIL. 


~e 


MINAS 
28 km due N of the bridge 
i. 


" 
EI 
= 


Grazielia intermedia (DC.) K. & R. n = 10. BRAZIL. 
GOIÁS: Chapada dos Veadeiros, 5 km S of Alto Pa- 
297 


** Hartwrightia Joktdana A. Gray. n = 10. U.S.A. 
'LORIDA: Highlan ds Co., Hanson, Wunderlin & 
Richardson 6420. 

Hebeclinium macrophyllum (L.) DC. = 10. BOLIVIA. 


km from Kd eae K743 437 
Isoc 'arpha oppositifolia ( = 10. GUATEMALA. 
EL PROGRESSO: city limits of EL Progresso. 1365. 
Kaunea ignorata (Hieron .) K. € R. n= ca. 20. BOLIVIA. 
ANT. one km from Comarapa on road to 
Cochabamba. K763 
* Kaunea lasiophthalma (Crise. )K.& R. 
OLIVIA. COCHABAMBA: 5 km W of Pojo. Cue 
lr ea M Hed (Gardner) K . n= 
10 BRAZIL. BAHIA: 2 km a W de Ubatá. 
4 B 


* Koanophyllon conglobatum. n — BRAZIL. MINAS 
GERAIS: NE of Montes latos: K8575 (C). 
Koanophyllon pittieri (Klatt) K. & R. n = ca. 30. Costa 
CA. PUNTARENAS: Monteverde. K9685 (C). 
dri a standleyi (B. 5 eae K. & R. 
MALA. ESCUINTI km N of Ese m 


27 km 


= 


*Koanoiyton sridi: n = 10. Gu, pcs e 
km N of perg K71 

** Litothamnus ellipticus K. &R.:; er Ain 
8.7 km N of Porto t E 

Mikania cordifolia (L.f) Willd. n = ba x NAME 
ESCUINTLA: 7 km N of Eset K717 


BAHIA: 


pa buius n — 19, ATEMALA. SUCHITE- 
P :10 km E of Maraton. K7244. 

Mikania cordifoli n — ca. 17. AVIA. COCHABAMBA: 
26 km from rap K765 


* ha goyazensis (B. L. Robins.) K. & R. n= ca. 
. BRAZIL. Go1Ás: Chapada na Veadeiros, 18-19 
n N of Alto ej vas 
us goyazensi = ]7. BRAZIL. GOlÁs: 20 km N 
Alto Paraiso " TA K8814 (C). 


Moni e .n = 19 + 1. GUATEMALA. 
ABAL: ca. 27 km NE of Gualan. K7370 
T Mikania periplocifolia Hook. 


& Arn 

RGENTINA. CATAMARCA: city linis: of a 
K9458 (C). 

* Mikania pohlii (Baker) K. & R. n = ca. 34. BRAZIL. 
GOIÁS: 2 km S of Cristallina. K82 

* Mikania pohlii. n = ca. 36. BRAZIL. FEDERAL DISTRICT: 
18 km from Brasilia. K8028 8 (C). 

* Mikania purpurascens (Baker) K. ı = ca. l 

valents, one bivalent larger and 

d two univalents 


m due N of the bridge 
5i 
* Mikania pyramidata J. D. Smith n = 18-20 + 5 or 


OS BAJA VERA PAZ: 4 km S of 
o K735 


Volume 76, Number 4 


Robinson et al. 1011 
Chromosome Numbers in Compositae, XVI 


Neomirandea e bu (B. L. Robins.) K. € R. n = ca. 
26. CosTA RICA. CARTAGO: ca. 15 km from Tapanti. 


did dcc ois gularis. n = ca. 26. Costa RICA. CAR- 
o: V olcá rA Turrialba. K9695 (E) 
Neomirandew eximia (B. L. Robins.) K. R. n = ca. 
17. Costa RICA. SAN JOSÉ: La Palma. K9656 (C). 
Neomirandes eximia. n= ca. 17. CosrA RICA. CARTAGO: 
15 km from the dam at Tapant. K9669 (C). 
RR angustifolius B. L. Robins. n = 10. 
BOLIVIA. SANTA CRUZ: 2 km from Comarapa. K 7638. 
b. osporus ehilca (H.B.K.) Hieron. n = 10 + one 
und frag. or univalent. PERU. ANCASH: along the 
roa to Olleros ca. 3 km generally ea B of the in- 
rsection with Peru highway #3 
“Ophryosporus ees (Schultz- Bip. ex Lu K. 
& R. 20. BOLIVIA. COCHABAMBA: 26 km from 
Tolata "KTS 593. 
*Peievavent schultzii (Schnittsp.) K. & R. n = 


GUATEMALA. ALTA VERAPAZ: 


ca. 10. 
2 km from Coban. 
í a. 
icr schultzit. n = 10. GUATEMALA. ALTA VERA- 
9 km W of San Cristóbal Verapaz. pudo 


Figuera. trinervia Cav. n = 24. GUATEMALA. TOTON 
CAPAN: 33 km N of San Cristóbal Taonia 
K7295 

Piqueria trinervia = . GUATEMALA. HUEHUE- 
TENANGO: ca. 13 km N of rA K7311 


inc lychnophoroides G. Barroso. n = 10. BRAZIL. 
12-20 km N of Alto P de Goiás. K8821 


(C. 
**Platypodanthera d (DC)K.& R. n= 
AZIL. BAHIA: n N of Livramento do ud. 
K8597 (C) 
Polyanthina nemorosa (Klatt 


LS R. n = 10-12 


(probably 10). BOLIVIA. LA PAZ: 7 km from Chulu- 
mani. K7432. 
Pra m Mii pur ) - E R. n = 32,. BRAZIL. 
ANEIRO: Rio de Jar o. Coun obtained from 
> geny mcd collected in pu nical garden. K9700. 
Praxelis "rinde (H.B.K.) Schultz-Bip. n = 40 uni- 
valent a ictic). BRAZIL. BAHIA: entre n ramento 


do Bra 1ado e Rio de Contas, a 5 km do primeiro. 
** Pseudokyrsteniopis porem K. & R = 10. 
ATEMALA. QUICHE: 29 km E of Aguacatan. KI 319. 
**Sciadocephala se huliz e-rhonhofiae Mattf. 
ADOR. LOS RÍOS: Río Palenque Biological Ste Station. 
C. Dodson 5513. 
* Stevia PATA Schultz-Bip. 7 l (one larger;). 
)LIVIA. COCHABAMBA: 48 tn A Tolata. K 7609. 
*Stevia boliviensis. n = 11. y m COCHABAMBA: 23 
m fro ie ci K764 
*Stevia bolivianas n= ll. Bornia COCHABAMBA: 23 
from Comarapa. K766 
*Stevia boliviensis. n=11. NN COCHABAMBA: 9 km 


Stevia caracasar GUATEMALA. CHIMAL- 
TENANGO: 5 km SE of Chimaltenango. K7194. 
TUM deos ensis Hieron. LIVIA. COCHA- 

BAMBA: 24 km from Tolata. = (587 


ds EA sis Hieron. n 11. ARGENTINA. CA- 
TAMARCA: ca. 45 km N of rs K9463 (C). 

*Stevia “ovata Rusby. n = 11. BOLIVIA. COCHABAMBA: 

n from Colomi. K7678. 

* Stevia os ee E Bertol. n 2. 
TOTONICAPAN: 14 km NE of Totonicapan. K 7278 

. n = 12. GUATEMALA. HUEHUE- 

1 


GUATEMALA. 


TE 
*Stevia samaipatensis B. L. Robins. B BOLIVIA. 


COCHABA : 71 km from Tolata. 'K76 
*Stevia bd sis. n — 11. BOLIVIA. COCHABAMBA: 
2 from Eon K7673. 


= 10 or 11 (if n = 10 then 
one larger n). Bau COCHABAMBA: 29 km from 
Cochabamba. K7674. 
* Stevia uci i n = 11. BOLIVIA. COCHABAMBA: 
ca. 60 km E of the bridge at Punata. K9641 (C). 
*Stevia santacruzensis Hieron. n = 11 + l univalent. 
OLIVIA. COCHABAMBA: 59 km from Tolata. K 7606. 
T sarensis B. L. Robins. n = 11. BOLIVIA, SANTA 
23 km E of Comarapa. K9655 (C). 
Siena an Rusby. n = BOLIVIA. COCHABAMBA: 
outskirts of Cochabamba. K7579 
* Stevia ia pod Hieron.? n = 11. BOLIVIA. COCHA- 
BAMB 6 km from Challa. K7541. 
* Stevia urcicifolia. ai in Thunb. n = 11. 
COCHABAMBA: val from Tolata. K7617 
RE yaconensis Da n = 11. BOLIVIA. coc TO 
. 60 km E of da bridge at Punata. K964 


BOLIVIA. 


eS iomaunndlies aa ^nsis (B. L. Robins.) H. Robins 
n = 10. BRAZIL. GOIÁS: ca. 1 km N of Crscallina: 
244 


r 


*Sy pir in. ud didi m (Gardner) B. L. Robins. 
1 


. BRAZIL. MINAS 
pln 7 (€). 
. BRAZIL. 


icm compressus. n — 
27 km NE of Montes a. 
“Symphyopappus o urcz. n = 
0 km of Muc cugé. K87! 
e eee i K. & R., type. 
SANTA CRUZ: 1 km from Comarapa. 
ke des e zielae K. & R. n= 10. hus 
f Sào Joào d'Alianca. a 
*Tric hogonia ee G. Barroso. n = 
20 km N of Alto Paraiso de NN 
*Tric hogonia salviifolia Gardner. 
o 


3 (C * 

pit oF IA. 
K76 
GOIÁS: 


gs AZIL. GOIÁS: 
K8812 (C). 


n 5W along saan from Escola Fazendaria. K8 
*Trichogonia salviifolia. n = 10 bivalents + two uni- 
valents or fragments. BRAZIL DISTRITO FEDERAL: 32 
= N i the bridge at t As en "ide in Brasilia on the 


road to Alto Paraiso. K82 
*Tric hogoni scottmorii K. & R. type. n = 10. BRAZIL. 
IA: 42 km a E de Maracas. K8018. 


“Trichogonia n (DC.) Schultz Bip. ex Baker. n = 
0, appears to be 20 bivalents. BRAZIL. MINAS GERAIS: 
"m ra do Cipó, 9-11 km beyond Rio Cipó. K8343. 


A GEOGRAPHICAL 
ANALYSIS OF THE 
FAMILY RANUNCULACEAE 


Svetlana N. Ziman! and 
arl S. Keener? 


ABSTRACT 


The Ranunculaceae Juss. are distributed worldwide. 
place in the vast territory of Eurasia during the Paleogen 


time. The western and eastern floristic centers in eastern Pes played the m 


differentiation (i.e., formation of most g 


primary differentiation of this family apparently m 


at least 50 million years ago 


The aim of this paper is to analyze the role of 
the Ranunculaceae with respect to floras around 
the world and to define more precisely some ques- 
tions concerning the origin and differentiation of 


e, there is no general study of the geo- 
graphical distribution of the Ranunculaceae as a 
whole, except those of Tamura (1966-1968), whose 
data on the presence of species in the various 
floristic regions of the earth were helpful in pre- 
paring this essay. Additianalys we took advantage 
of our own herba rium t 

from KW, LW, LWS, LE, TBI, TAD, BP, and 


others) and the observations of the senior author 


ai 
w the iterpaetaticd of the floristic re- 
gions tep provinces (Table 1 and map) by T 

tajan (1970, 1978) and the interpretation d en- 
in the sense of Tolmachev (1974). 
Furthermore, we have attempted to recognize local, 
disjunctive, and widespread endemic species within 
each province or region. 


GEOGRAPHICAL CHARACTERISTICS OF THE 
ANUNCULACEAE 


The Ranunculaceae comprise 55 genera and 
about 2,200 species (Takhtajan, 1987; Tamura, 
1966-1968; Ziman, 1985). About 20 genera’ and 
1,200 species are endemics (Table 2). Some en- 
demic genera, e.g., Beesia, Souliea, Kingdonia, 
Asteropyrum, and Xanthorhiza, are morphologi- 
cally isolated within the Ranunculaceae, and thus 
we were able to use their areas to discuss pecu- 
liarities of the distribution and the differentiation 
of the family as a whole. 

To date, five floristic regions, the Eastern Asiat- 
ic, Atlantic North American, Andean, Chile-Pa- 
tagonian, and Irano-Turanian, contain at least one 
endemic genus. 

Most genera of Ranunculaceae have disjunctive 
distributions. Some, e.g., Calathodes, Dichocar- 
pum, Urophysa, Semiaquilegia, Beesia, Souliea, 
and Kingdonia, occur within restricted geographic 
ranges of two to four provinces in one region. 
Another group, made up of Helleborus, Eranthis, 
Isopyrum, Leptopyrum, Paropyrum, Oxygra- 


à fog of Botany of the Academy of Sciences of the Ukrainian S.S.R., Repina 2, 252601, Kiev-GSP 1, U.S 


The Pennsylvania State University, : 
Pennsylvania 16802, U.S.A. 


Park, 


208 Mueller Laboratory, College of Science, Department of Biology, U s 


3 ee i dr re ferences are rei The nomenclature and circumscriptions of ranunculaceous genera follow 


, 20, 74, 87, 213, 264, 333, and o 


monographs 4 


thers. Authorships for plant taxa follow Index Kewensis, 
). 


Cherepanov (46), the monographs listed ‘aie. and regional floras (5 and others 


ANN. Missouni Bor. Garp. 76: 1012-1049. 1989. 


Volume 76, Number 4 
1989 


Ziman & Keener 1013 


Geographical Analysis of Ranunculaceae 


phis, Paraquilegia, and Callianthemum, is char- 
acterized by more or less widely disjunctive areas, 
occupying two to four regions of Eurasia. 

Areas of a third group, which includes Trollius, 
Actaea, Cimicifuga, Enemion, Pulsatilla, He- 
patica, Halerpestes, and Trautvetteria, are rep- 
resented in three to seven regions of Eurasia and 
North America. In many cases the disjunctive 
species of these genera are considered as relicts of 
the Tertiary temperate flora (Thorne, 1973). 

Species of Caltha and Myosurus are widely 
distributed in the Northern and Southern hemi- 
spheres. It is remarkable that they are widespread 
in temperate areas but absent in the tropics and 
subtropics. 

The distributional patterns of the remaining 18 
genera of Ranunculaceae are unbroken or nearly 
unbroken. Species of the majority of them occur 
in two to six regions of Eurasia (Aconitella and 
Ficaria), Sout 
America (Hamadryas), Africa (Knowltonia and 
Clematopsis), Eurasia, North Africa and North 
America (Nigella, Aconitum, Consolida, Aquile- 
gia, Adonis, Ceratocephalus, and Coptis). Del- 


Southeastern Asia (Varavelia), 


phinium occurs in 11 regions of Eurasia, Africa, 
North America, and South America; Thalictrum 
occurs in 16 regions on these continents. Species 
of Anemone, Clematis, and Ranunculus are wide- 
spread in both hemispheres in the floras of about 
30 regions. 


REGIONAL DISTRIBUTION OF THE 
RANUNCULACEAE 


In following Takhtajan (1978) we examined the 
distribution patterns of the Ranunculaceae in six 
floristic kingdoms, eight subkingdoms, 34 regions, 
and about 150 provinces. 


HOLARCTIC KINGDOM 


Location: territories of the Northern Hemi- 
sphere (more than half of the Earth), including 
Europe, extratropical Asia, North America, and 
North Africa. 

Climate: predominately temperate, partly sub- 
tropical, and cold 

Ranunculaceae: 49 genera, 2,000 species. 


Boreal Subkingdom 


This subkingdom includes four regions: Eastern 


Asiatic, Circumboreal, Atlantic North American, 
and Rocky Mountain. 

Location: extratropical Eurasia and North 
America. 


Ranunculaceae: 48 genera, 1,500 species. 


Eastern Asiatic Region 


Location: main part of continental China, north- 
ern India, Burma, and Nepal, almost all Japan and 
Korea, eastern U.S.S.R 

Climate: warm temperate to mostly temperate. 

Ranunculaceae: 38 genera, 900 species; 12 
endemic genera, 400 endemic species. 


Sikang Yunnan Province 


Location: highlands; main part of the Chinese 
provinces Szechwan, Kweichow, and Yunnan, also 
northeastern Burma and northern Laos. 

aracteristic vegetation: mountain conifer- 
ous, broad-leaved and rain forest, subalpine, and 
alpine formations. 

Ranunculaceae: 28 genera, 400 species (14, 
42, 48, 66, 81, 84, 109, 112, 199, 316). 

Related floras: Central Chinese (21/60) and 
Eastern Himalayan (19/70). 

Observations: 8 endemic genera, 180 endemic 
species. All endemic genera are isolated morpho- 
logically within the family. For example, Astero- 
pyrum (A. peltatum) represents the monotypic 
tribe Asteropyreae and differs from other species 
by having simple peltate leaves. One other genus, 
Souliea (S. vaginata), belongs to the tribe Cimi- 
cifugeae and is distinguished by a unique type of 

ollen grain which lacks a primary germination 
apparatus (171). Another endemic monotypic ge- 
nus, Kingdonia (K. uniflora), is exceptional by 
having open dichotomous foliar venation (42). This 
genus makes up the monogeneric tribe Kingdon- 
ieae, subfamily Kingdonioideae.* 
also Urophysa henryi, Beesia calthifolia, 
athodes palmata, C. polycarpa, and C. oxycarpa, 
occur in isolated mountain forest locations at 2,000— 
4,000 m of the Sikang Yunnan, Central Chinese, 
Eastern Himalayan, and some other provinces. 

Disjunct areas of species of Semiaquilegia (S. 
adoxodoides and S. manschurica) and Dichocar- 
pum (D. fargesii, D. dicarpon, and others) are 
widely separated in provinces in the west and the 
east of the Eastern Asiatic Region. These species 
are also mountain forest plants, restricted to shade 
locations. 


* Some 
family, Kingdoniaceae. 


systematists consider Kingdonia to belong to the Circaeasteraceae or else classified in its own monotypic 


1014 Annals of the 
Missouri Botanical Garden 


TABLE |. Summary of numbers of genera and species, including endemics, of the Ranunculaceae for the principal 
floristic regions of the world (after Takhtajan, 297). 


Total numbers Endemics 

Kindgoms, subkingdoms, regions, provinces Genera Species Genera Species 

I. HOLARCTIC KINGDOM 49 2,000 16 1,000 
BOREAL SUBKINGDOM 48 1,500 15 700 
Eastern Asiatic Region 38 900 12 400 
l. Sikang Yunnan Province 28 400 8 180 

2. Central Chinese Province 22 150 7 50 

3. North Chinese Provi 15 70 - 5 

4. Eastern Himalayan Province 23 200 6 90 

5. North Burmese Province 14 60 3 10 

6. Khasi-Manipur Province 8 25 5 

7. Wolkano-Bonin Province 2 5 
8. Tokaro-Okinawa Province 2 7 

. Taiwanian Province 9 40 2 5 

10. Manchurian Province 23 160 l 50 
11. Japanese-Korean Province 22 130 3 50 
12. Sakhalin- Hokkaido Province 17 80 l 20 
Circ zc al Region 30 600 - 200 
l; 10tsk- Kame hatka Province 18 90 - 15 

2. rd ern Siberian Province 15 70 - 5 

3. Middle a Province 18 100 20 

4. Western Siberian Province 15 ) 5 

5. Altai-Sayan Province 2 130 30 
Transbaikalian Province 19 100 15 

7. Eastern European Province 22 110 10 

8. Central European Province 24 160 - 20 

9. Balkan Province 23 140 40 
10. Atlantic European Province 22 110 20 
11. North European Province 15 50 — — 
12, Arctic Province 12 80 -- 5 
13. Caucasian Province 19 160 - 45 
14. Euxine Province 18 100 — 35 
15. Canadian Province I3 60 5 
Atlantic North American Region 20 120 3 25 
l. North American Prairies Province 12 40 l 

2. je -Gulf Coastal Plain Province 7 20 -- 5 
yalachian Province 20 100 3 20 

Rocky i ea Region 17 100 — 35 
. Sitka-Oregon Province 16 80 -— 20 

7 Rocky Mountain Province 16 70 = 15 
MADREAN SUBKINGDOM 7 130 - 50 
rds Region 17 130 — 50 
. Californian Province 16 100 — 25 

: Great Basin Province 11 40 — 5 

3. Sonoran Province 5 10 5 

4. Mexican Highlands aie 5 50 40 
ANCIENT MEDITERRANEAN SUBKINGDOM 30 500 l 250 
Saharo-Arabian Region 9 35 - 6 
Saharan Province 4 10 — 3 

ytlan- -Arabian Province 9 30 3 

Mare aronesian Reg 2 5 — l 
Mediterranean Regi on 18 130 — 65 
i icum Province 17 60 — 20 

2. Iberian Pu e 16 50 10 
3. East Mediterranean Province 13 50 15 


4. Southwest Mediterranean Province 9 25 — 5 


Volume 76, Number 4 Ziman & Keener 015 
1989 Geographical Analysis of Ranunculaceae 
TABLE |. Continued. 
Total numbers Endemics 
Kindgoms, subkingdoms, regions, provinces Genera Species Genera Species 
5. South Mediterranean Province 7 20 - l 
6. Southern Moroccan Province 5 10 — l 
7. Balearic Province 9 15 - 2 
8. Adriatic Province 5 15 — i 
9. Krym-Novorossiysk Province 10 30 2 
Trano-Turanian Region 28 400 l 180 
Anterior Asiatic Subregion 28 360 4 360 
1. Armeno-Iranian Province 24 220 — 30 
2. Hyrcanian Provinc 14 60 5 
3. a Anatolian Province 18 110 — 20 
4. Mesopotamian Prov 8 40 -— 5 
5. Turkestanian "eri 19 130 50 
6. Turanian Provi 10 50 — 
7. Western Himalaya an Provin 19 100 l 35 
8. Northern Baluc histanian Pasties 14 50 — 5 
Central Asiatic Subregion 23 200 - 60 
1. Dzungaro-Tien Shan Province 2] 130 = 40 
2. Central Tien Shan Province 19 7 25 
3. Mongolian Province 18 100 — 5 
4. Tibetan Pro 20 80 10 
II. PALAEOTROPICAL Ki on 11 100 — 50 
INDO-MALESIAN SUBKINGDOM 6 80 35 
iir Region 5 60 -— 30 
Malaccan Province 3 5 — 
Kalimantan Province 3 4 - — 
. Moluccan Province ] 2 — 
4. Bismarckian Province ] l - 
5. Philippinean Province > 15 6 
6. South Malesian Province 5 15 — 5 
7. Sumatran Province 5 10 5 
8. Sulawesian Province 2 15 12 
. Papuan Province 3 30 25 
ae Region 3 4 3 
Indo-Chinese Region 6 29 3 
Indian Region 6 10 3 
POLYNESIAN SUBKINGDOM 2 5 l 
Polynesian Region l 2 -— 
Hawaiian Region 2 5 l 
NEOCALEDONIAN SUBKINGDOM l ] 
Neocaledonian m 1 l — 
AFRICAN SUBKINGD 8 50 = 25 
Guineo-Congo Pim 7 40 — 20 
Sudano- Zambesian Region 8 30 10 
Karro-Namib Region l 3 - 
5t. Helena and Ascension Region l 1 - = 
MADAGASCAN SUBKINGDOM 4 10 - 2 
Madagascan Region 4 10 — 2 
III. Cape KINGDOM 7 30 - 10 
Cape Regio 7 30 10 
IV. Nun e AL KINGDOM 9 70 2 40 
Caribbean Region 3 20 - 5 
Guayana ii doc, Region 2 4 
Amazonian Reg l 2 — 
Brazilian Region. 2 5 - 
Andean Region 9 60 2 30 


1016 


Annals of the 
Missouri Botanical Garden 


TABLE 1. Continued. 


Total numbers Endemics 


Kindgoms, subkingdoms, regions, provinces Genera Species Genera Species 
V. HOLANTARCTIC KINGDOM 8 100 2 80 
Chile- Patagonian Region 7 50 2 25 
Fernandesian Region 
South Subantarctic Islands Region l 5 l 
Neozeylandic Region 5 60 55 
VI. AUSTRALIAN KINGDOM 5 50 30 
East Australian Region 5 45 30 
Southwest Australian Region 2 5 2 


Central Australian Region 


In the flora of the Sikang Yunnan Province, 
Aconitum has about 70 species of which 40 are 
endemics; Delphinium has 70 species and 50 en- 
demics; Clematis has 40 and 20; Anemone 20 
and 15; Trollius 10 and 8; and Aquilegia 8 and 
9 species. 

In this province occur species that are important 
in considering the origin and dispersion of their 
genera and the family Ranunculaceae as a whole. 
For example, Aconitum gymnandrum, which rep- 
resents the monotypic subg. Gymnaconitum, is 
found at about 4,000 m in Szechwan and in the 
Eastern Himalayan Province. Its annual habit is 
unique within Aconitum; this species is character- 
ized by having primitive flowers and is considered 
a Tertiary relict (181, 241, 284). Another prim- 
itive species, Delphinium aconitoides, also occurs 
in this province. The local endemic Clematis ran- 
unculoides, occurring there, represents the mono- 
typic sect. Ranunculoides, and its life form (ro- 
sette-forming perennial herb with tripartite leaves) 
is regarded (341, 342) as the most primitive within 
Clematis. 

There are three monotypic endemic sections of 
Anemone: sect. Rivularis (A. rivularis), sect. Ane- 
moclema (A. glaucifolia), and sect. Begoniifolia 
(A. begoniifolia). Moreover, the most primitive 
and simultaneously relict species of Trollius (T. 
yunnanensis) (64, 270) and Aquilegia (A. ecal- 

ta) in the Sikang Yunnan Province (272). 

The few species endemic to this province include 
Delphinium ceratophorum, Ranunculus platy- 
petalus, and Thalictrum delavayi; many other 
species are shared with adjacent provinces. 


Central Chinese Province 


Location: Plateau Guichou, Ridges Zinlin-Da- 
bashan and Nanlin, and the southern part of the 


Yangtze Basin (Kiangsu, Shensi, Hupeh, Anhwei, 
and Hunan provinces of Chin 

Characteristic eran RN forests, 
subalpine and alpine formations. 

Ranunculaceae: 22 genera, 150 species (58, 
81, 112, 198, 220, 330, 332). 

Related floras: Sikang Yunnan (21/60, e.g., 
Aconitum gymnandrum, Anemone rivularis, 
Aquilegia ecalcarata), Eastern Himalayan (18/ 
30, e.g., 


Anemone obtusiloba 


Trollius pumilus, Aquilegia ecalcarata, 


Observations: 7 elena genera, 50 endemic 
species. (The endemic genera, Asteropyrum, Uro- 
physa, Kingdonia, Souliea, Beesia, Calathodes, 
and Dichocarpum, are shared with the flora of 
Sikang Yunnan.) Most of the endemic species be- 
long to Aconitum, Delphinium, Clematis, and 
Thalictrum. Endemic Helleborus chinensis to- 
gether with H. thibeticus represent the endemic 
sect. Dicarpon. In contrast with the Sikang Yun- 
nan flora, some species of Pulsatilla, e.g., P. chi- 
nensis, and P. turczaninovii are distributed here. 


North Chinese Province 


Location: southeastern part of the Loess Plateau 
and North Chinese Plain. 

Characteristic vegetation: steppe formations and 
broad-leaved forests 

Ranunculaceae: 15 genera, 70 species (112, 
139, 198, 199, 221, 332) 

Related floras: Central Chinese (15/30, e.g. 
Trollius Del- 
phinium grandiflorum) and Manchurian (15/20, 


chinensis, Cimicifuga dahurica, 


e.g., Clematis brevicaudata, Thalictrum baica- 
lense). 
Observations: no endemic genera, but 5 endem- 


Volume 76, Number 4 
1989 


Ziman & Keener 1017 


Geographical Analysis of Ranunculaceae 


ic species (e.g., 4nemone changtungensis, Clem- 
atis nannophila). 


Eastern Himalayan Province 


Location: main part of eastern and central Ne- 
pal, northern India, and southeastern Tibet. 

Characteristic vegetation: mountain forests, 
subalpine and alpine formations. 

: 23 genera, 200 species (35, 
. 123, 179, 205, 244, 261, 276, 


nunculaceae 


D. 

Related flora: Sikang Yunnan (19/70, includ- 
ing endemic genera Souliea, Beesia, Calathodes, 
and Dichocarpum; endemic species, e.g., Del- 
phinium bonvalotii, Aconitum heterophyllum, 
Anemone rupicola, and Clematis connata; and 
disjunct relict species, e.g., Aconitum gymnan- 
drum, Anemone rivularis, and Aquilegia ecal- 
carata). 

Observations: 6 endemic genera, 190 endemic 
species. Four of the endemic genera are shared 
with Sikang Yunnan Province, whereas Archiclem- 
atis and Paroxygraphis are local only. Archiclem- 
atis alternata grows in central Nepal in bushy 
border habitats at 2,200 m; Paroxygraphis sik- 
kimensis is found in Sikkim in the alpine zone at 
4,000 m. There are two endemic, monotypic sec- 
tions of Aconitum: sect. Fletcherum (A. fletcher- 
anum) and sect. Alatospermum (A. novoluridum), 
also the endemic, monotypic Thalictrum—sect. 
Piuttia (T. rotundifolium). Most endemic species 
belong to Delphinium (in all about 50 species with 
about 30 endemics, e.g., D. nepalense), Aconitum 
(30 and 20, e.g., A. spicatum), Ranunculus (30 
and 10, e.g., R. hirtellus), and Thalictrum (20 
and 10, e.g., 7. elegans). In contrast with the 
Sikang Yunnan flora, this flora does consist of some 
species of Pulsatilla, Coptis, and Hepatica. 


North Burmese Province 


Location: between the Eastern Himalayan Prov- 
ince and Southwestern China. 

Characteristic vegetation: warm temperate high- 
mountain forest and meadow formations. 

Ranunculaceae: 14 genera, 60 species. 

Related floras: Sikang Yunnan and Central 
Chinese (14/50, e.g., Anemone vitifolia, A. riv- 
ularis, Cimicifuga foetida). 

servations: 3 endemic genera, Beesia, Sou- 

liea, and Dichocarpum, about 10 endemic species 
(e.g., Trollius yunnanensis, Clematis sikkimen- 
sis). 


Khasi-Manipur Province 


Location: Northeastern India Eee Plateau, 
Khasi, Naga, Manipur, and Patkai 

Characteristic vegetation: most like Km of the 
North Burmese Province. 

Ranunculaceae: 8 genera, 25 species. 

Related flora: Eastern Himalayan (8/20, e.g., 
Anemone rivularis, Clematis buchananiana). 

Observations: no en 
species (e.g., Trollius pumilus, Delphinium altis- 


emic genera, 5 endemic 
simum, Anemone elongata). 


Wolkano-Bonin Province 


Location: small islands in Southeastern Asia 
(Wolkano, Bonin, and others). 

Characteristic vegetation: 
tropical formations. 

Ranunculaceae: 2 genera, 5 species (215, 216, 


tropical and sub- 


Related flora: 


Clematis pierotii, Ranunculus japonicus). 


Japanese-Korean (2/5, e.g., 


Observation: no endemic genera or species. 


Tokaro-Okinawa Province 


Location: small islands in Southeastern Asia 
(Ryukyu, Tokara, Okinawa, Amami, and others). 

Characteristic vegetation: tropical and sub- 
tropical formations. 

Ranunculaceae: 2 genera, 7 species. 

Related flora: Japanese-Korean (2/7, e.g., 
Clematis mejeniana, Ranunculus ternatus). 

Observation: endemic species absent. 


Taiwanian Province 


Location: Taiwan and some adjacent islands, 
e.g., Sakisima. 
tropical and sub- 


Characteristic vegetation: 


tropical formations, also warm-temperate, high- 
mountain forests and meadows. 

Ranunculaceae: 9 genera, 40 species (216, 
331, 339). 

Related flora: Japanese-Korean (9/35, e.g., 
Anemone stolonifera, Clematis uncinata, Ranun- 
culus cantoniensis). 

bservations: 2 endemic genera, Calathodes 

and Dichocarpum, and 5 endemic species (e.g., 
Aconitum formosanum, Ranunculus vernyi). 


Manchurian Province 


Location: Soviet far east, Northeastern China, 
North Korea, and Northeastern Mongolia. The main 


landscape features are more or less level plains, 


1018 


Annals of the 


Missouri Botanical Garden 


++ +++ 


++ + + 


+ + 


+ 


+ 


+ 


+ 


+ 


c 


+ 


+ 


+ 


+ 


++ +++ 


+ 


+ 


++++ + 


+ + 


+++ +4 


++ + + 


piuopsuly 
wns dos] 
sispap«gy] 
vonoday 
sn10q34]2H 


uonuau7 
undap20Q2i(] 


ppyosuo) 
sisdojpuiaj7) 
s11Dui2]7) 
ngnfioiun?) 
snjpudo20]0427) 
Diy]v7) 

wumuay my?) 


$9P0Y1D]D7) 


UNPUOIP 
Djjonuoap 


Il 


OT 


suotday 


PIQUd*) 


"PHo^ əy} jo suolgal 3usuog ureur əy} ut I9PIJE]NIUNUBY əy} jo Plouad 3u1 Jo uonnqugistp aur 


'Z 318v 


1019 


Ziman & Keener 
Geographical Analysis of Ranunculaceae 


Volume 76, Number 4 


1989 


ugi[ernsny 1s?7— 6[ 
oipue|&azoaN — 8 I 
ueiuogejeq-opu) — 21 
ueapuy — 9] 
ueaqque-)— G 

ade) — $1 


— 


asauly’)-Opuy — z | 
UBISITPIA — [T 


uetsaquie7-ouepng — ()T 
O3UO7)-02Utn*) — 6 


UI JUNO JA ÁXIOM — b 
UBOLIQUY YON 3Qugnvy — € 
[8310qUIN II) —6 

JUBISY ul9]se;] — I 


suOIdal JHsUopg əy} jo suoneuztsap [£uonuoAuo:) 


+ 


de 
+ 
+ 
B 


+ 


ERA FAA - BG. +++ 


+ + 


DZVYLOYIUDY 
psAydos) 
smpo4 
D1121]2a]nn4] 
Wn J 
DINOS 
pisapinbpiwmag 
snjnoununy 
MIEL 
snidpn18Áxoapg 
uniKdoang 
nigapinbpapg 
sinydpnidÁx() 
sappy nal) 
DNIDIN 
DINIADADN 
snansoXqy 
DIYDÁNA] 
una1Adojdo'] 
umjpjado2ov'] 
Diu0j] nou x 


Il 


Ol 


6 


suolsay 


EI3uat) 


'penuguo? `g IYL 


1020 


Annals of the 
Missouri Botanical Garden 


but there are some mountainous chains (Sikhote- 
Alinj and Hingans). 

Characteristic vegetation: broad-leaved and co- 
niferous-broad-leaved forest, steppe, and meadow 
formations. 

Ranunculaceae: 23 genera, 160 species (25, 
108, 112, 158, 174, 326, 327, 328). 

Related floras: Japanese-Korean (18/40, e.g., 
Caltha membranacea, Actaea spicata, Enemion 
raddeanum, Thalictrum filamentosum) and 
Okhotsk-Kamchatkian (18/25). 
| endemic genus (Semiaquile- 
gia) and about 50 endemic species, most of which 
belong to Aconitum (40 species, with about 20 
endemics, e.g., 4. desoulavyi), Ranunculus (30 
and 5, e.g., R. tachiroei), and Thalictrum (15 
and 5, e.g., T. amurense). In contrast to the Sikang 


Observations: 


Yunnan and other western floras of the Eastern 
Asiatic Region, this flora includes species of Ene- 
mion, Hepatica, Coptis, and Trautvetteria and 
few species of Delphinium and Clemati 


Japanese-Korean Province 


Location: Honshu, Shikoku, Kyushu, southern 
Hokkaido, Iki, Cusima, Goto, additional small is- 
lands, and South Korea 

Characteristic vegetation: subtropical and tem- 
perate coniferous and broad-leaved forests in low- 
lands and mountains, also subalpine and alpine 
formations. 

Ranunculaceae: 22 genera, 130 species (208, 
216, 271, 299). 

Related floras: Manchurian (18/40) and Sa- 
khalin-Hokkaidian (16/25) 

Observations: 3 endemic genera, disjunct Semi- 
aquilegia and Dichocarpum, and local endemic 
Anemonopsis (4. macrophylla); these in many 
mountain forest locations in Honshu. Endemic 
species are about 50. There are endemic sections 
of Anemone (sect. Keiskea), Coptis (sect. Japo- 
nocoptis), Cimicifuga (sect. Pityrosperma), Pul- 
satilla (sect. Preonanthopsis), and Clematis (sects. 
Patentes and Paratragene). In contrast to the flora 
of the Manchurian Province, this flora consists of 
few species of Aconitum and lacks species of Del- 
phinium. 


Sakhalin-Hokkaido Province 


Location: South Sakhalin, North Hokkaido, 
South Kuril Islands (Kunashir, Shikotan, and Itu- 
rup). 

Characteristic vegetation: predominately co- 
niferous forests. 


Ranunculaceae: 17 genera, 80 species (209, 
216, 299, 325, 326, 327, 328). 

Related floras: Manchurian (16/40, e.g., Cal- 
tha fistulosa, Trollius macropetalus, Aconitum 
arcuatum), Japanese-Korean (16/25, e.g., Aco- 
nitum volubile, Anemone raddeana, Adonis amu- 
rensis, Trautvetteria japonica) and Okhotsk- 
Kamchatkian (14/20, e.g., Caltha sibirica, Clem- 
atis ochotensis). 

Observations: | endemic monotypic genus Mi- 
yakea (M. integrifolia) which is related to Pul- 
satilla and 20 endemic species (e.g., Aconitum 
miyabei, Anemone yesoensis, Pulsatilla sacha- 
linensis). In contrast to the Manchurian and Jap- 
anese-Korean floras, this flora has no temperate 
species of Eranthis, Enemion, and Hepatica. 


Circumboreal Region 


Location: main part of Europe (except extreme 
south), the Caucasus, Urals, Siberia, Kamchatka, 
North Sakhalin, North Kuril, the Aleutian Islands, 
Alaska, and northern Canada. 

Climate: temperate to cold. 

Ranunculaceae: 30 genera, 600 species. En- 
demic genera are absent; there are about ‘ 
endemic species. 


Okhotsk-Kamchatka Province 


Location: Kamchatka Peninsula, littoral zones 
along the Okhotsk Sea, North Sakhalin and North 
Kuril, Komandor and Aleutian Islands. 

Characteristic vegetation: predominately co- 
niferous forests. 

Ranunculaceae: 18 genera, 90 species (6, 18, 
29. 125, 160, 322, 325, 328) 

Related floras: most similar to the Manchurian 
(18/25, e.g., Caltha fistulosa, Coptis trifolia, Ac- 
onitum fischeri), Sakhalin-Hokkaidian (17/35, e.g., 
Trollius macropetalus, Cimicifuga simplex, Aqui- 
legia flabellata), Northeastern-Siberian (13/30, 
e.g., Trollius riederianus, Anemone debilis, Pul- 
satilla dahurica), Arctic (10/15, e.g., Caltha arc- 
tica, Trollius membranostylis, Anemone richard- 
sonil). 

Observations: 
nitum kurilense, Pulsatilla taraoi, 
recurvatus). Numbers of species are noteworthy 
in Ranunculus (25), Aconitum (10), 4nemone (10), 
and Thalictrum (10). 


15 endemic species (e.g., Aco- 
Ranunculus 


Northeastern Siberian Province 


Location: northeastern U.S.S.R. (eastern Ya- 
kutia, Magadan district; Verkhoyansk, Chersk y, and 


Volume 76, Number 4 
1989 


Ziman & Keener 1021 
Geographical Analysis of Ranunculaceae 


Suntar-Hayata ridges; Jana-Indigirka, Kolyma, 
Anadyrj, and Korjakya uplands). 

Characteristic vegetation: undersized and thin- 
ly growing mountain forests (of Larix), sparse groves 
of trees (e.g., Pinus), high-mountain tundras, alpine 
stone deserts. 

Ranunculaceae: 15 genera, 70 species (25, 
134, 135, 142, 151, 152, 154, 176). 

Related floras: Okhotsk-Kamchatkian (13/ 30), 
Manchurian (12/20, e.g., Actaea erythrocarpa, 
Aquilegia parviflora, Delphinium cheilanthum) 
and Arctic (12/30, e.g., Caltha arctica, Del- 
phinium brachycentrum, Anemone multiceps). 

Observations: 5 endemic species (e.g., Trollius 
chartosepalus, Pulsatilla magadanense, Ranun- 
culus jacuticus). There are some nemoral Tertiary 
relicts (Anemone dichotoma, Pulsatilla dahurica, 


Ranunculus repens; 135, 154, 308). 


Middle Siberian Province 


Location: North Siberia, between the Enisey and 
Lena rivers (mainly Middle Siberian and Aldan 
plateaus). 

Characteristic vegetation: taiga, composed of 
coniferous forests (predominately Larix sibirica). 

Ranunculaceae: 18 genera, 100 species (7, 25, 
142. 175, 231). 

Related flora: Northeastern Siberian (14/35, 
e.g., Cimicifuga simplex, Delphinium crassifo- 
lium, Aconitum macrorhynchum, Anemone re- 
flexa). 

Observations: 20 endemic species (e.g., Aco- 
nitum contractum, Ranunculus borealis, R. smir- 
novii) There are some nemoral Tertiary relicts 
(Anemone altaica, A. sylvestris, Thalictrum foe- 
tidum; 166, 195), Quaternary relicts (Trollius 
asiaticus, Aquilegia sibirica, Anemone reflexa, 

56) 


Pulsatilla patens; 96, 256). 


Western Siberian Province 


Location: North Siberia, between the Enisey and 
Obj rivers (Western Siberian Plain). 

Characteristic vegetation: taiga (e.g., Picea 
alba, Abies sibirica). 

Ranunculaceae: 15 genera, 60 species (25, 
168, 169). 

Related floras: Middle Siberian (14/30, e.g., 
Actaea erythrocarpa, Anemone sylvestris, Ra- 
nunculus gmelinii) and Eastern European (14/ 
10, e.g., Trollius europaeus, Aconitum septen- 
trionale, Delphinium elatum). 

Observations: 5 endemic species (e.g., Trollius 
kytmanovii, Anemone jenisseensis). 


Altai-Sayan Province 


Location: south of Western and Middle Siberian 
provinces (highlands of Altai, Sayan, Kuznetskyi 
Alatau, Tuva, and others). 

Characteristic vegetation: coniferous forests (of 
Larix, Abies, and Pinus); broad-leaved forests; 
subalpine and alpine, steppe, and tundra forma- 
tions. 

Ranunculaceae: 21 genera, 130 species (166, 
172, 173, 195, 231, 266, 309). 

Related floras: Northeastern Siberian (14/25, 
e.g., Aquilegia sibirica, Anemone dichotoma, 
Pulsatilla flavescens, Adonis sibirica), Transbai- 
kalian (19/50, e.g., Trollius lilacinus, Eranthis 
sibirica, Paraquilegia microphylla), and Mon- 
golian (18/35, e.g., Aquilegia glandulosa, Del- 
phinium crassifolium, Oxygraphis glacialis). 

Observations: 30 endemic species (e.g., Aco- 
nitum sajanense, Delphinium sajanense, Pulsa- 
tilla ajanensis, Callianthemum sajanense). Nem- 
oral Tertiary relicts include Actaea spicata, 
Cimicifuga foetida, Anemone sylvestris, Eranthis 
sibirica (47, 128, 167, 182); high-mountain Qua- 
ternary relicts include Trollius asiaticus, Anemone 
narcissiflora, Pulsatilla campanella (194, 232, 
256). 


Transbaikalian Province 


Location: Burjatia, southern parts of the Irkutsk 
and Chita districts (Khamar-Daban, Baikalyan and 
some other ridges; Vitim and Stanovoe plateaus; 
some depressions). 

Characteristic vegetation: coniferous forest, and 
steppe and high-mountain formations. 

Ranunculaceae: 19 genera, 100 species (142, 
230, 231, 267, 319, 324). 

Related floras: Altai-Sayan (19/50) and Mon- 
golian (18/40, e.g., Trollius ledebourii, Cimicif- 
uga dahurica, Clematis hexapetala). 

Observations: 15 endemic species (e.g., Aco- 
nitum montibaicalense, Aquilegia burjatica, Del- 
phinium korschinskyanum, Ranunculus ajanen- 
sis); large number of species of Ranunculus (20), 
Aconitum (15), Thalictrum (10), Pulsatilla (10), 
Aquilegia (8), and Delphinium (8). 


Eastern European Province 


Location: Southeastern Scandinavia and the Eu- 
ropean part of U.S.S.R. (mainly Russian Plain but 
also Crimea, Donetsk, Urals, and other highlands). 

Characteristic vegetation: coniferous and broad- 
leaved forest, forest-steppe and meadow forma- 
tions. 


1022 


Annals of the 
Missouri Botanical Garden 


Ranunculaceae: 22 genera, 110 species (8, 23, 
44, 96, 97, 269, 323). 

Related floras: Central European (21/70, e.g., 
Actaea spicata, Isopyrum thalictroides, Hepat- 
ica nobilis), Western Siberian (14/10), and Man- 
churian (15/15, e.g., Actaea erythrocarpa, 
Anemone reflexa, Thalictrum minus). 

Observations: 10 endemic species (e.g., Aco- 
nitum besserianum, Delphinium sergii, Anemone 
biarmiensis, Ranunculus crymaeus). There are 
disjunct relicts (e.g., Anemone nemorosa, Aconi- 
tum anthora, Isopyrum thalictroides; 156, 170). 
Isopyrum thalictroides is distributed in Central 
and Eastern Europe, and western Himalayas; Pul- 
satilla grandis occurs in the Alps, West Carpa- 
thians, Balkans, and Crimea, and most species of 
Helleborus (e.g., H. purpurascens) are distributed 
in Europe, but two of them (of sect. Dicarpon) 
occur in western China. 


Central European Province 


Location: Austria, F.R.G., G.D.R., Poland, and 
other countries of Central Europe, also western 
USSR 

Characteristic vegetation: broad-leaved for- 
ests, subalpine and alpine formations. 

Ranunculaceae: 24 genera, 160 species (19, 
25, 44, 49, 65, 79, 87, 119, 130, 221, 279, 
291). 

Related floras: Eastern European (21/70), Bal- 
kanian (23/80, e.g., Trollius europaeus, Helle- 
borus odorus, Aconitum paniculatum, Callian- 
themum coriandrifolium), and Western Siberian 
(17/20, e.g., Aconitum anthora, 
vestris). 

Observations: 20 endemic species (e. 


Anemone syl- 


4 Aco- 
nitum firmum, Aquilegia transsilvanica, Ranun- 
culus carpaticus). Most species belong to Aconi- 
tum (in all about 20 species, within them 10 
endemics), Aquilegia (7 and 5), Ranunculus (50 
and 10). Disjuncts: Callianthemum eMe 
lium, Aconitum moldavicum, and ot 


Balkan Province 


Location: main part of Yugoslavia, almost all 
Bulgaria, northern Greece, and the European part 
of Turkey. 

Characteristic vegetation: high-mountain for- 
mations of forests and meadows. 

Ranunculaceae: 23 genera, 140 species (44, 
51, 89, 133, ) 

Related floras: Central European (23/80) and 


Liguro-Tyrrhenian (17/25, e.g., Delphinium pe- 


regrinum, Anemone apennina, Pulsatilla alpi- 
na). 

Observations: 40 endemic species (e.g., Hel. 
leborus multifidus, H. cyclophyllus; bulb. FM 
of the Mediterranean Ranunculus sect. Ranun- 
culastrum, Ranunculus rumelicus, R. millii). Most 
species belong to Ranunculus (50), Pulsatilla (10), 
Aquilegia (8), and Helleborus (5 species). 


Atlantic European Province 


Location: Southwestern Scandinavia, Denmark, 
Belgium, Holland, Great Britain, Ireland, main part 
of France, northern F.R.G. and G.D.R., northern 
Spain. 

Characteristic vegetation: coniferous and most- 
ly broad-leaved forests, high-mountain formations 
with mixture of arctic, boreal, and nemoral ele- 
ments. 

Ranunculaceae: 22 genera, 110 species (44). 

Related floras: Central European (22/50, e.g., 
Ranunculus auricomus, R. platanifolius) and 
Arctic (12/20, e.g., Ranunculus sulphureus, R. 
hyperboreus). 

Observations: 20 endemic species (most of them 
restricted to dry rocky mountain places, e.g., Aqui- 
legia pyrenaica, Thalictrum tuberosum, Ranun- 
culus pyrenaeus, R. gramineus), including warmth- 
loving (e.g., Helleborus foetidus, H. niger, He- 
patica nobilis), cold-tolerant (e.g., Oxygraphis 
glacialis, Ranunculus pygmaeus), and xerophyt- 
ic (e.g., Nigella gallica, Consolida regalis, Ado- 


nis pyrenaica) elements 


North European Province 


Location: main part of Scandinavia, northwest- 
ern U.S.S.R. (except Arctica). 

Characteristic vegetation: coniferous forests (of 
Picea abies and Pinus sylvestris). 

o 15 genera, 50 species (25, 44, 

197, 207, 

Related s Atlantic and Central European. 

Observations: no endemic species; most genera 
are represented with one or two species, except 
Ranunculus (about 25 species). 


Arctic Province 


Location: extreme northern Eurasia and North 
America. 
Characteristic vegetation: tundra formations. 


Related floras: Eurasiatic and Canadian taiga 


Volume 76, Number 4 
1989 


Ziman & Keener 1023 


Geographical Analysis of Ranunculaceae 


(10/50, e.g., Aquilegia sibirica, Anemone syl- 
vestris, Ranunculus monophyllus), also hh. 
, €.g., Anemone narcissi- 
Thalictrum alpi- 


mountain zones (12 
flora, Oxygraphis glacialis, 
num). 

Observations: few endemic species. About 50 
species are widely distributed in Eurasia, and about 
30 species occur in Eurasia and North America 
(e.g., Delphinium chamissonis, Anemone rich- 
ardsonii, Ranunculus affinis). Hypoarctic ele- 
ments include Delphinium middendorffii, Ranun- 
culus gmelinii, and others); arctic elements include 
Caltha caespitosa, Ranunculus sabinii, R. punc- 
tatus, and others. 


Caucasian Province 


Location: Big and Little Caucasus mountains, 
Dagestan and Stavropolje hills and adjacent ter- 
ritories. 

Characteristic vegetation: coniferous and broad- 
leaved (Quercus and Fagus) forests, mountain- 
steppe, subalpine and alpine formations. 

Ranunculaceae: 19 genera, 160 species (90, 
101, 102, 103, 149, 150). 

Related floras: Euxinian (16/65, e.g., endem- 
ics Trollius ranunculinus, Helleborus caucasicus, 


Armeno-lranian (17/40, e.g., Delphinium foetid- 
um, Aconitum orientale, Pulsatilla albana, Ra- 
nunculus brutius) and Central European (17/10, 
, Anemone nemorosa, Ranunculus repens). 
Observations: 45 endemic species, with most 
occurring in subalpine and alpine zones (e.g., Pul- 
satilla aurea, Delphinium caucasicum, D. spe- 
ciosum, Ranunculus baidarae, R. acutilobus). 


Euxine Province 


cation: western part of the Caucasian Isthmus 
(Abkhasia, Adjaria, and adjacent territories). 
aracteristic vegetation: like the Caucasian 
Pos 
anunculaceae: 18 genera, 100 species (57, 
103, 149, ). 

Related flora: Caucasian (16/65, including 25 
endemic species). 

Observations: 35 endemic species. In contrast 
to the Caucasian flora, this flora does not possess 
annuals of Myosurus and Ceratocephalus but does 
include a few perennials of Anemone, Clematis, 
Diedropetala), and Ranun- 
culus sect. Ranunculastrum). 


Delphinium (sect. 


Canadian Province 


Location: Canada (Labrador, Hudson Bay, and 
Mackenzie districts) and Alaska. 

Characteristic vegetation: coniferous forests 
(mainly of Pinus, Picea, and Larix 

Ranunculaceae: 13 genera, 60 ined (26, 86, 
99, 100, 115, 245, 315). 

Related floras: Sitka-Oregonian (13/25, e.g., 
Aconitum columbianum, Aquilegia formosa, Ra- 
nunculus macounii), Appalachian (13/15, e.g., 
Actaea rubra, Aquilegia canadensis, Ranun- 
culus cymbalaria), and Arctic (13/10, e.g., Ac- 
onitum delphinifolium, Delphinium brachycen- 
trum, Anemone multifida 

Observations: few e species (e.g., Ra 
nunculus occidentalis, Thalictrum occidentalis). 


Atlantic North American Region 


Location: eastern North America (from the coast 
of the Atlantic Ocean to the Great Basin and from 
southern districts of Canada to the Gulf of Mexico). 

‘limate: mainly temperate. 

Ranunculaceae: 20 genera, 120 species (15a, 
32, 33, 55, 116, 132, 144-148, 223, 290, 305, 
315 


North American Prairies Province 


Location: vast treeless plain in the middle of 
North America (Saskatchewan and Manitoba prov- 
inces of Canada, Nebraska, lowa, Kansas, North 
and South Dakota, Oklahoma, and Texas states of 

SA) 

Characteristic vegetation: grass formations 
(mainly of Andropogon, Buchloe, Stipa, and other 
genera). 

Ranunculaceae: 12 genera, 40 species. 

Related flora: Appalachian (11/18, e.g., Aqui- 
legia canadensis, Tha- 
lictrum revolutum). 


Clematis virginiana, 
Observations: | endemic species (Clematis fre- 
montii). 


Atlantic-Gulf Coastal Plain Province 


Location: Coastal Plain territories in south- 


eastern U.S.A. (parts of North Carolina, Virginia, 
Florida, Alabama, Mississippi, Arkansas, and Lou- 
isiana). 

Characteristic vegetation: salt strands, dune, 
pine barrens, bog, marsh, beach and sea coast 
formations. 

Ranunculaceae: 7 genera, 20 species. 

Related flora: Appalachian (7/15, e.g., Actaea 
pachypoda, Clematis virginiana). 


1024 


Annals of the 
Missouri Botanical Garden 


Tha- 


Observations: few endemic species (e.g., 
lictrum cooleyi). 


Appalachian Province 


Location: southeastern and south-central Can- 
ada (Ontario, Manitoba, and Quebec) and north- 
eastern and central states of U.S.A. (Minnesota, 
Wisconsin, New York, Pennsylvania, Kentucky, 
Tennessee, and others). Mainly mountain territo- 
ries (Piedmont Uplands, Alleghany, Blue Ridge, 
Catskill, Cumberland, Green, White mountains, 
Ozark Plateau, and others). 

Characteristic vegetation: broad-leaved forests 
(Fagus, Quercus, 


of coniferous forests (Pinus, 


Acer), with less representation 
Picea, Abies, and 
other genera). 

Ranunculaceae: 20 genera, 100 species. 

Related floras: Canadian (13/15), Manchurian 
(16/5). 
Observations: 3 endemic genera, 20 endemic 
species. All endemic genera are monotypic and are 
isolated morphologically within the family, e.g., Hy- 
drastis (H. canadensis), Xanthorhiza (X. simpli- 
cissima) (stands apart from Coptis), and Anemo- 
nella (A. thalictroides). Endemic species belong 
mainly to Clematis (e.g., C. addisonii, C. 
tilis), Thalictrum (T. cooleyi, T. subrotundum, and 
others), and Ranunculus (e.g., R. harveyi, R. 
alleghaniensis). 


coac- 


Rocky Mountain Region 


Location: western North America (western Can- 
ada, also Washington, Oregon, Wyoming, Califor- 
nia, and other states). 

Climate: temperate to rather xerophytic. 

Ranunculaceae: 17 genera, 100 species (1, 54, 


55, 116, 122, 228, 259, 265, 315). 


Sitka-Oregon Province 


Location: western islands (e.g., Vancouver), main 
part of British Columbia, Washington, Oregon, and 
northeastern California (Selkirks, Gold Russell, and 
Cascade mountains). 

Characteristic predominately co- 
niferous forests (Picea, Pseudotsuga, Abies, and 


vegetation: 


others), subalpine and alpine formations. 

Ranunculaceae: 16 genera, 80 species (mainly 
Caltha, Coptis, inen Delphinium, and Ra- 
nunculus). 

Related flora: Rocky Mountain (15/60, e.g., 
Caltha leptosepala, Actaea arguta, Pulsatilla oc- 
cidentalis, Trautvetteria carolinensis). 

Observations: 20 endemic species (e.g., Caltha 


asarifolia, Coptis asplenifolia, Delphinium ore- 


ganum 
Rocky Mountain Province 


Location: southern W yoming, central Colorado, 
and northern New Mexico (Rocky Mountains). 

Characteristic vegetation: like the Sitka-Ore- 
gonian but more xerophytic. 

Ranunculaceae: 16 genera, 70 species (mainly 
Delphinium, about 20 species, and Ranunculus, 
about 30 species). 

Related floras: Sitka-Oregonian (15/60) and 
Californian (15/40, e.g., Aquilegia chrysantha, 
Enemion stipitatum, Delphinium andersonii). 
Del- 
phinium decorum, Ranunculus inamoenus, R. 
oresterus). In contrast to the Sitka-Oregonian flora, 
this flora includes some species of Enemion and 


Observations: 15 endemic species (e.g., 


few species of Caltha and Coptis. Trollius is ab- 
sent. 


Madrean Subkingdom 


This subkingdom includes only the Madrean Re- 
gion. 


Madrean Region 


Location: southwestern U.S.A. (California, Ne- 
vada, Utah, New Mexico, and other states) and 
almost all of Mexico. 

Climate: warm and hot, dry. 

Ranunculaceae: 17 genera, 130 species which 
are restricted to mountains and are absent in des- 
erts and chaparral formations (1, 73, 116, 214, 


273, 286, 306, 315). 


Californian Province 


Location: main part of California. 

Characteristic vegetation: predominately co- 
niferous (Pinus, Sequoia) and broad-leaved forest 
(Quercus and other genera), also subalpine and 
alpine formations and chaparral. 

Ranunculaceae: 16 genera, 100 species. 

Related flora: Rocky Mountain (16/40). 

Observations: about 25 endemic species, mostly 
Delphinium (e.g., D. californicum, D. luteum, D. 
uliginosum). In contrast to the Rocky Mountain 
flora, this flora includes many species of Delphin- 
ium (in all about 40) and several species of Aqui- 
legia, and Myosurus. 


Great Basin Province 


Location: Great Plain and Colorado Plateau 
(western parts of Washington, Oregon, and Cali- 


Volume 76, Number 4 


Ziman & Keener 1025 


Geographical Analysis of Ranunculaceae 


fornia, almost all of Nevada, Utah, northern parts 
of Arizona and New Mexico). 

Characteristic vegetation: predominately cha- 
parral (of Cercocarpus and others), few high-moun- 
tain formations. 

Ranunculaceae: 11 genera, 40 species. 

Related floras: Californian (11/30, e.g., Ac- 
taea arguta, Delphinium menziesii, Anemone 
globosa, A. drummondii). 

Observations: few endemic species (e.g., Del- 
phinium depauperatum, Anemone tuberosa). 


Sonoran Province 


Location: Southeastern California, partly Ari- 
zona, New Mexico, and Texas, central Mexico. 

Characteristic vegetation: desert formations. 

Ranunculaceae: 5 genera, about 10 species. 

Related flora: Great Basin (5/5, e.g., Del. 
phinium parishii, Clematis lasiantha). 

Observations: endemic species are Delphinium 
parryi, Anemone sphoenophylla, Clematis pau- 
ciflora, Myosurus sessilis, and Ranunculus an- 
dersonii. 


Province of Mexican Highlands 


Location: western and eastern Mexico (Sierra 
Madre Mountains, United Cordilleras). 

Characteristic vegetation: predominately rath- 
er xerophytic high-mountain formations. 

anunculaceae: 5 genera, 50 sp 

Related flora: Californian (all dd genera, but 
common species are absent). 

Observations: about 40 endemic 
which most belong to Thalictrum (e.g., T. pelta- 
tum, T. lanatum, T. pubigerum), also to Del- 
phinium (e.g., D. scopulorum) and Anemone (4. 


species, of 


mexicana, and others). 


Ancient Mediterranean Subkingdom 


This subkingdom includes four regions (Saharo- 
Arabian, Macaronesian, Mediterranean, and Irano- 
Turanian). 

Location: Southern Europe, Northern Africa, 
and Western Asia. 

Ranunculaceae: 30 genera, 500 species. 


Saharo-Arabian Region 


Location: extratropical Sahara, main parts of 
the Arabian and Sinai peninsulas and adjacent ter- 
ritories. 

Climate: warm-temperate and hot xerophytic. 

Ranunculaceae: 9 genera, 35 species (23, 29, 


191, 237, 292). 


Saharan Province 


Location: Sahara Desert from the Atlantic coast 
to Egypt 
Characteristic vegetation: predominately des- 
ert formations. 
anunculaceae: 4 genera, 10 species (e.g., Del- 
did bovei, Consolida deserti, Adonis den- 


Related flora: Egyptian-Arabian (4/6). 


Observations: few endemic species. 


Egyptian-Arabian Province 


Location: eastern Egypt, Arabia, and Sinai. 
Characteristic vegetation: predominately des- 


tions. 

ecies (mainly 
species of Ranunculus, Consolida, and Delphin- 
tum). 

Related floras: Saharan (4/6, e.g., Adonis den- 
tata, Nigella deserti). 

Observations: few endemic species (e.g., Ni- 
gella assyriaca, Adonis palaestina). 


Macaronesian Region 


Location: Azores, Madeira, Canary and Cape 
Verde islands. 

Ranunculaceae: 2 genera: Clematis (5 species) 
and Ranunculus (30 species). 


Mediterranean Region 


This region includes nine provinces. 
Location: southern Europe, northern Africa, and 
western Asia Minor. 
Climate: mainly subtropical and rather arid. 
unculaceae: 18 genera, 130 species (23, 


44, 51. 191, 200, 237, 238) 


Liguro-Tyrrhenian Province 


Location: southern France, southwestern Italy, 
islands in the Tyrrhenian and Ligurian seas (e.g., 
Corso, Sicily, Sardinia). 

Characteristic vegetation: evergreen forest and 
shrub formations. 

Ranunculaceae: 17 genera, 60 spec 

Related floras: d (16/25, e.g., Pulsatilla 
rubra, Clematis cirrhosa, Hepatica nobilis) and 
Balkanian (15/20, e.g., Helleborus lividus, Aco- 
nitum anthora, Pulsatilla alpina). 

Observations: about 20 endemic species, most 

of which are high-mountain representatives of 
Aquilegia (e.g., A. bernardii, A. litardieri) and 
Ranunculus (e.g., R. cordigerus, R. rupestris). 
n contrast to the Balkanian flora, this flora has 


1026 


Annals of the 
Missouri Botanical Garden 


no mesophytic species of Caltha, Trollius, Actaea, 
or Cimicifuga. Like other Mediterranean floras, 
the Liguro-Tyrrhenian flora includes a large num- 
ber of annuals (about 20 species, e.g., Delphinium 
halteratum, D. pictum, Nigella damascena). 


Iberian Province 


Location: main parts of Pyrenees Peninsula 
(including Cantabrian Mountains, Sierra Nevada, 
Central Cordilleras). 

Characteristic like the Liguro- 
Tyrrhenian, but also steppe formations. 

Ranunculaceae: 16 genera, 50 spec 

Related flora: Liguro- Tyrrhenian (16/25) 

Observations: about 10 endemic species, mostly 
Aconitum (e.g., A. nevadense), Delphinium (e.g., 
D. nevadense, D. obcordatum), and Ranunculus 


, R. gramineus, R. abnormis). This flora dif- 


vegetation: 


fers from the former by lacking Eranthis, pos- 
sessing fewer species of Helleborus and Aquilegia, 
and having more species of Anemone and Pulsa- 
ti 


East Mediterranean Province 


Location: about all of Greece, west of Asia Mi- 
nor, and adjacent parts of Syria, Lebanon, and 
Israel. 

Characteristic vegetation: predominately shrub 
formations (Arbutus, Spartium, and others), partly 
semideserts. 

Ranunculaceae: 13 genera, about 50 species. 

Related flora: Mesopotamian (8/25, e.g., Con- 
solida regalis, Adonis annuus 

Observations: about 15 enden: species (e.g., 
Nigella cretica, N. doerfleri, Consolida helles- 
pontica, Myosurus heldreichii, Ranunculus cre- 
ticus). Almost all endemics and half of the ra- 
nunculaceous species are annuals. In contrast to 
the previously mentioned floras, this flora has no 
species of Aconitum, Pulsatilla, or Hepatica but 
possesses species of Aconitella. 


Southwest Mediterranean Province 


Location: southern Pyrenees Peninsula, also 
northwestern Africa (mainly Morocco). 

Characteristic vegetation: coniferous and broad- 
leaved forests (mainly cut down), shrub and grass 
formations. 

Ranunculaceae: 9 genera, about 25 species. 

Related floras: South Mediterranean (7 / 15, e.g., 
Anemone coronaria, Consolida orientalis, Del- 
phinium staphisagria). 


Observations: few endemic species (e.g., Clem- 
atis campaniflora, Delphinium sylvaticum). Some 
species (e.g., Aconitum lamarckii, Helleborus foe- 
tidus) show temperate Eurasiatic affinities. 


South Mediterranean Province 


Location: mainly northern Algeria and Tunis. 

Characteristic vegetation: hke the Southwest 
Mediterranean. 

Ranunculaceae: 7 genera, about 20 species. 

Related flora: Southwest Mediterranean (7/15). 

Observations: Virtually no endemic species. 


Southern Moroccan Province 


Location: southern Morocco 
Characteristic vegetation: shrub and grass for- 
mations of a mixture of Mediterranean and Saharan 
elements. 
Ranunculaceae: 5 genera, about 10 species. 
lated floras: Southwest Mediterranean (5/5) 
and Saharan 
bservations: virtually no endemic species; 
species are predominately annuals (e.g., Consolida 
mauritanica, Ranunculus muricatus). 


Balearic Province 


Location: islands of Majorca, Minorca, and Pi- 
tius. 

Characteristic vegetation: similar to the Iberian 
Province. 

Ranunculaceae: 9 genera, 15 species. 

Related flora: Iberian (9/10, e.g., Helleborus 
lividus, Anemone coronaria, Delphinium sta- 
phisagria, Ranunculus bullatus). 

Observations: endemic species are only Clem- 


atis balearica and Ranunculus balearicus. 


Adriatic Province 


Location: coastal districts of Italy, Yugoslavia, 
and Albania. 

Characteristic vegetation: evergreen forest and 
shrub formations. 

Ranunculaceae: 7 genera, about 15 species. 

Related floras: Balkanian (5/10, e.g., Anem- 
one coronaria, Nigella damascena), also Liguro- 
Tyrrhenian (7/8). 

Observations: l endemic species, Ranunculus 
garganicus 


Krym-Novorossiysk Province 


Location: small territory of the South Crimea 
and the northeastern Caucasus 


Volume 76, Number 4 
1989 


Ziman & Keener 1027 
Geographical Analysis of Ranunculaceae 


Characteristic vegetation: similar to most of 
the Mediterranean provinces. 

Ranunculaceae: 10 genera, about 30 species 
(25, 44, 103, 258, 269). 

Related floras: Eastern European (10/25, e.g., 
Aquilegia vulgaris, Adonis flammea, Ranun- 
culus sceleratus). 

Observations: virtually no endemic species. 


Irano-Turanian Region 


This region includes 11 provinces. 

Location: vast area of Asia Minor and Anterior, 
Middle, and Central Asia. 

Climate: temperate, but mainly arid. 

Ranunculaceae: 28 genera, 400 species. One 
endemic genus (Alexeya), 180 endemic species. 


Anterior Asiatic Subregion 


Location: western Irano-Turanian Region. 
Ranunculaceae: 28 genera, 360 species, in- 
cluding about 150 endemic species. 


Armeno-lranian Province 


Location: Armenia, Azerbaijan, part of Turk- 
menia, Southeastern Turkey, Iran, Iraq, and west- 
ern Afghanistan. 

Characteristic vegetation: steppe, semidesert, 
and desert formations, scrubs, sparse growth of 
trees (e.g., Juniperus, Pistacia), also high-moun- 
tain formations. 

Ranunculaceae: 24 genera, 220 species (50, 
56, 57, 91, 92, 103, 149, 225, 260, 294). 

Related floras: Caucasian (19/40, e.g., Caltha 
polypetala, Helleborus orientalis, Delphinium 
foetidum, Aquilegia olympica), Hyrcanian (12/ 
30, e.g., Aconitella hohenackeri, Delphinium 
flexuosum), and especially Central Anatolian (18/ 


— 


50, Actaea spicata, Eranthis hyemalis, Aconitum 
orientale, Anemone blanda). 

Observations: about 30 endemic species, of 
which most belong to Ranunculus (in all about 80 
species— 10 endemic, e.g., 
nisii), Delphinium (35 and 10, e.g., D. ararati- 
cum, D. carduchorum), and Consolida (30 and 
hex. Es: olopetala). All these 
species are restricted to open limestone high-moun- 
tain locations. There are about 60 species of an- 


. crateris, R. sinte- 


armeniaca, C. 


nuals, mainly species of Vigella, Consolida, Ado- 
nis, and Myosurus. In contrast to the Caucasian 
and Hyrcanian flora, this flora consists of some 
species with boreal and nemoral affinities in the 
floras of Eastern Asia (e.g., Paropyrum anemo- 
noides, Isopyrum thalictroides, Actaea spicata). 


Hyrcanian Province 


Location: small southeastern part of the Cau- 
casian Isthmus (Talysh Chain). 

Characteristic vegetation: relict broad-leaved 
forests (mainly of Quercus castaneifolia and Zel- 
kova carpinifolia). 

Ranunculaceae: 
103, 260). 

Related flo 
phinium foetidum, Helleborus orientalis, Trollius 
ranunculinus) and Armeno-Iranian (12/30 


14 genera, 60 species (88, 


ras: Caucasian (13/40, e.g., Del- 


Observations: 5 endemic species (among them, 
Ranunculus hyrcanus, Anemone caucasica, and 
Delphinium ochroleucum). In contrast to the pre- 
vious floras, there are no species of Actaea, Ac- 
onitum, Pulsatilla; there are many species of Ni- 


gella and Consolida. 


Central Anatolian Province 


Location: northwestern Asia Minor (about half 
of the territory of Turkey, including the Anatolian 
Plateau). 

Characteristic vegetation; semiarid and arid 
grass formations. 

Ranunculaceae: 18 genera, 110 species (57). 

Related floras: Armeno-Iranian (18/50) and 
East Mediterranean (13/ 
entalis, Anemone blanda, Ranunculus asiaticus). 


,e.g., Thalictrum ori- 


Observations: about endemic species, of 
which most belong, like the Armeno-lranian en- 
demics, to Ranunculus (in all about 50 of which 
5 are endemics, e.g., R. demissus, R. fibrillosus), 
Delphinium (10 and 5, e.g., D. 
ilgarense), and Consolida (10 and 5, e.g., C. ra- 


veyi, C. thirkeana). 


venulosum, D. 


Mesopotamian Province 


Location: mainly Syria, Iran, Iraq, Israel. 

Characteristic vegetation: semidesert and des- 
ert formations. 

Ranunculaceae: 8 genera, 40 species (23, 57, 
229; 23 ls 292): 

Related floras: East Mediterranean (8/25) and 
Egyptian-Arabian (8/15, 


Adonis palaestina). 


e.g. Nigella deserti, 


Observations: few endemic species (e.g., Del- 
phinium chodatum). Most of species are annuals. 
Turkestanian Province 


Location: mountain ridges and highlands within 
Middle Asia (Paropamiz, Badkhyz, Karabylj, Ba- 
dakhshan, Gyssar, Karatau, Nuratau, Mogoltau). 


1028 


Annals of the 
Missouri Botanical Garden 


Characteristic vegetation: arid high-mountain 
forest, shrub and grass formations. 

Ranunculaceae: 19 genera, 130 species (25, 
39, 140, 141, 143, 163, 219). 

Related floras: Dzungaro-Tien Shanian (19/ 
50, e.g., endemics Ranunculus rubrocalyx, Del- 
phinium oreophylum, Aquilegia atrovinosa, Cal- 
lianthemum alatavicum) and Armeno-lranian (17 / 
20, e.g., Eranthis longistipitata, Delphinium 
semibarbatum, Clematis asplenifolia). 

Observations: about 50 endemic species, in- 
cluding Pulsatilla kostyczewii of the monotypic 
sect. Jostemon, also about 10 species of Delphin- 
ium (e.g., D. lipskyi, D. karategini), 25 species 
of Ranunculus (e.g., R. mogoltavicus, R. ko- 
marovii), 8 species of Aquilegia (e.g., A. vicaria, 
A. darwasi) and Anemone (A. eranthioides, A. 
seravshanica—in all 10). In contrast to the Ar- 
meno-Iranian flora, this flora includes only about 
10 annual species and lacks Caltha, Actaea, Hel- 
leborus, and Isopyrum. Besides that, this flora 
includes some alpine disjuncts (e.g., Trollius lila- 
cinus, Delphinium brunonianum, Paraquilegia 
microphylla) that are regarded as Tertiary relicts 


(218). 


Turanian Province 


Location: eastern Transcaucasia, Caspian low- 
land and vast plain territories of the middle asiatic 
republics of U.S.S.R. 

characteristic vegetation: desert, semidesert, 
and dry steppe formations. 

Ranunculaceae: 10 genera, 50 species (25, 39, 
94, 129, 162, 163, 226). 

Related floras: Turkestanian (9/25, e.g., 
Anemone gortschakovii, Clematis tangutica, 
Thalictrum isopyroides) and Armeno-Iranian (8/ 
15, e.g., annuals Consolida stocksiana, Cerato- 
cephalus falcatus, Adonis parviflora). 

Observations: no endemic species. 


Western Himalayan Province 


Location: northern parts of Afghanistan, Pa- 
kistan, India, and northwestern China (Nuristan, 
Vasiristan, Kashmir, Simla, and other highlands). 

Characteristic vegetation: subtropical forma- 
tions in low zones, temperate forest, subalpine and 
alpine formations in high zones. 

Ranunculaceae: 19 genera, 100 species (123, 
179, 180, 243, 257, 283, 299). 

Related floras: Eastern Himalayan (17/35, in- 
cluding endemics Caltha gowaniana, Trollius 
acaulis, Delphinium coeruleum, Thalictrum gla- 


reosum, and others) and Dzungaro-Tien Shanian 
(15/25, e.g., Trollius dschungaricus, Hepatica 
falconeri, Ranunculus brotherusii, R. popovii). 

Observations: 1 local endemic genus, Alexeya 
(A. vvedenskyi), which is closely related to Par- 
aquilegia and separated from the latter (222). 
Endemic species are about 35 (e.g., Aconitum 
heterophyllum, Clematis barbellata, Anemone te- 
trasepala). There are some disjunct relicts (e.g., 

nemone vitifolia, Actaea spicata, lsopyrum 
thalictroides), which are absent in other provincial 
floras of the Irano-Turanian Region. 


Northern Baluchistanian Province 


Location: parts of lran, Afghanistan, and Pa- 
kistan (Suleyman, Toba-Kakar, Kweta-Pishin, and 
other ridges). 

Characteristic vegetation: high-mountain arid 
formations. 

Ranunculaceae: 14 genera, about 50 species 
(91, 123, 225, 299). 

Related floras: Western Himalayan (12/20, 
e.g., Delphinium brunonianum, Oxygraphis in- 
volucratum, Anemone narcissiflora) and Armeno- 
Iranian (14/20, e.g., Consolida rugulosa, Clem- 
atis orientalis, Thalictrum isopyroides). 

Observations: about 5 endemic species (e.g., 
Delphinium incanum, Thalictrum afghanicus). 


Central Asiatic Subregion 


Location: vast territories from central Tien Shan 
to Big Hingan (part of territory of middle asiatic 
republics of U.S.S.R., Mongolia, Tibet, and North 
China). 

Climate: temperate but rather arid. 

Ranunculaceae: 23 genera, 200 species, in- 
cluding about 60 endemic species. 


Dzungaro-Tien Shan Province 


Location: a part of the territory of the Kazakh 
Republic (Talass, Dzungar, Tarbagatai, and other 
ridges, also the Kazakh small ridges). 

Characteristic vegetation: high-mountain forest 
and grass formations, also rocky desert and dry 
steppe formations. 

anunculaceae: 21 genera, 130 species (25, 
39, 94, 140, 163, 226). 

Related floras: Central Tien Shanian (19/60, 
including 25 endemic species, e.g., Trollius 
dschungaricus, Aconitum karakolicum, Delphin- 
ium turkestanicum, Ranunculus gelidus), Tur- 
kestanian (19/50), and Western Himalayan (15/ 
25). 


Volume 76, Number 4 


Ziman & Keener 1029 


Geographical Analysis of Ranunculaceae 


Observations: about 40 endemic species, of 
schanica), Aconitum (e.g., 
phinium (e.g., D. batalinü), Anemone (e.g., A 
almaatensis), and Ranunculus (e.g., R. 
anus). In contrast to the Turkestanian flora, this 


regeli- 


flora includes more species of Aconitum, Callian- 
themum, and Adonis but fewer species of Ranun- 


culus, Nigella, and Consolida. 


Central Tien Shan Province 


Location: west of the Dzungaro-Tien Shan Ter- 
skey and Kungey Alatau, also Kirgiz and Fergana 
ridges. 

Characteristic vegetation: like the Dzungaro- 
Tien Shanian, but mostly steppe formations. 

Ranunculaceae: 19 genera, 70 spec 

Related flora: Dzungaro-Tien Shanian (19/60). 

Observations: All 25 endemic species are shared 
with the Dzungaro-Tien Shanian flora. 


Mongolian Province 


Location: Mongolia, the Kashgarian and Tsai- 
dam hollows. 

Characteristic vegetation: in the northern out- 
lying districts (Khubsugul, Khentei, Khangai, and 
Mongolian Altai) predominately high-mountain for- 
est and grass formations, in the central and south- 
ern parts predominately steppe and desert for- 
mations. 

anunculaceae: 18 genera, about 100 species 
(105, 106, 108). 

Related floras: Transbaikalian (18/40) and AI- 
tai-Sayan (18/35 

Observations: few endemic species (e.g., Clem- 
atis intricata, Adonis mongolica, Ranunculus 
mongolicus). Several species belong to Aconitum, 
Clematis, and Ranunculus. 


Tibetan Province 


Location: Pamir, Tibetan, and Tsinkhai pla- 
teaus. 

Characteristic vegetation: predominately high- 
mountain stone tundra and desert formations. 

Ranunculaceae: 20 genera, about 80 species. 

Related floras: Turkestanian (16/30, e.g., 
Trollius micranthus, Callianthemum alatavicum, 
Pulsatilla kostyczewii) and Sikang Yunnanian (13/ 
15, e.g., Aconitum franchetii, Anemone imbri- 
cata, Delphinium albocoeruleum). 

Observations: 10 endemic species (e.g., Del- 
phinium propinquum, Ranunculus pamiri, Tha- 
lictrum ruthifolium). 


The flora of the Eastern Asiatic Region is an 
extraordinary mixture of tropical and temperate 
elements, and is considered (296, 297) as the most 
significant center of development of the Angio- 
sperms, possibly the area of origin. 

ithin this region the flora of the Sikang Yun- 
nan Province is extremely rich with Angiosperms; 
therefore, some Soviet plant geographers (159, 
38) regarded Southwestern China as the 
cradle of the modern temperate flora of Eurasia 
and considered the latter as derived from an ancient 
tropical flora. Other botanists (78, 107, 109) sug- 
gested that the primary development of the tem- 
perate mesophytic forest flora of Eurasia took place 
within the Eurasiatic tropical flora, possibly within 
Southwestern China, which has been the greatest 
refugium for this temperate flora. 

According to the paleobotanical data (131), since 
Late Cretaceous time the Sikang Yunnan flora 
was mainly a high-mountain one, which included 
many temperate elements. The Yunnan Plateau is 
one of the most ancient mountain systems, formed 
in the Late Cretaceous (Yanshan movement of the 
earth crust— 224), while the Himalayas are much 
more recent and belong to the alpine stage of 
mountain development (192). Therefore, the East- 
ern Himalayan flora is considered young (297). 
280) claim that 
significant refugium of 


Some plant geographers (13, 
the Manchurian flora was a 
the Tertiary warm temperate forest flora and sug- 
gest that the former coniferous-broad-leaved me- 
sophytic forests, in which temperate elements now 
predominate, migrated there during the Miocene 
and 
changed since. 

According to Maekawa (190), a warm temperate 
flora developed in Japan and Korea since the Oli- 


flora of Manchuria has scarcely 


gocene-Miocene. Tanai (304) claimed that the me- 
sophytic coniferous-broad-leaved forests that sur- 
vived there are quite similar to those in Manchuria. 

Our analysis of the role of the Ranunculaceae 
in the provincial floras of the Eastern Asiatic Re- 
gion led us to postulate that the initial differentia- 
tion of many groups of this family took place there 
in two diverse floristic centers, western and eastern 
ones. The former includes the Sikang Yunnan, 
Eastern Himalaya, and adjacent territories, the 
latter coincides with Manchuria, Japan, and Sa- 

alin. 

The western floristic center consists of ten en- 


Asteropyrum, Urophysa, Kingdonia, 
Beesia, Calathodes, Semiaquilegia, and Dicho- 


carpum, are distributed disjunctively in two to four 


1030 


Annals of the 
Missouri Botanical Garden 


provinces of eastern Asia and only Archiclematis 
and Paroxygraphis are local Eastern Himalayan 
endemics. 

Most of the endemic ranunculaceous species 
within the western floristic center of eastern Asia 
are high mountain plants, and in this area a number 
of species are shared with two or three adjacent 
provincial floras. There are 22 genera and about 
60 species ees about 20 endemics) in com- 
mon with the Sikang Yunnanian, Eastern Hima- 
layan, and Central Chinese provincial floras. These 
floras consist of the most primitive species of Trol- 
lius (T. yunnanensis), Anemone (A. rivularis), 
Clematis (C. ranunculoides), and Aquilegia (A. 
ecalcarata), and all of these genera include sec- 
tions endemic to the western part of eastern Asia. 
Compared with the eastern floristic center, the 
western one is home to species of the disjunctive 
Eurasiatic genera Helleborus, Paropyrum, Par- 
aquilegia, and Oxygraphis. 

All these facts confirm our opinion concerning 
the close connections of the floras of the previously 

entioned western provinces of eastern Asia. In 
i region of these western provinces during past 
geological periods a single high-mountain flora ex- 
isted which became a center of development for 
several genera of the Ranunculaceae (all ten en- 
demic genera, also Trollius, Aconitum, Aquilegia, 
Anemone, and Clematis). We do not consider the 
Ranunculaceae in the Eastern Himalayan Province 
as consisting of young species; rather, we support 
the opinion of van Steenis (289) about their antiq- 
uity. 

The eastern floristic center in eastern Asia con- 
sists of four endemic ranunculaceous genera and 
about 120 endemic species. Two endemic genera, 
Semiaquilegia and Dichocarpum, are shared with 
the western center and the other two genera, Ane- 
monopsis and Miyakea, are local endemics. 

Most endemic species of Ranunculaceae in the 
eastern part of eastern Asia are montane forest 
plants. There are 17 genera and about 25 species 
shared by Manchuria, Japan, and Sakhalin, but 
there are almost no endemic species shared by two 
or three provinces. These floras consist of endemic 
sections of Cimicifuga, Anemone, Pulsatilla, 
Clematis, and Coptis. In contrast to the western 
floristic center, in the eastern center occur species 
of the disjunctive Palearctic genera Enemion and 
Trautvetteria, comparatively few species of Del- 
phinium (absent in Japan and Sakhalin), and com- 
paratively few Clematis species (especially few in 
Manchuria). 

A comparison of the roles of the Ranunculaceae 
in the provincial floras of the western and eastern 


parts of eastern Asia indicated that only one-third 
of the ranunculaceous genera are common to 
both areas (Caltha, Trollius, Actaea, Cimicifuga, 
Aconitum, Delphinium, Aquilegia, Anemone, 
Pulsatilla, Adonis, Thalictrum, Ranunculus), and 
all of them are shared with the flora of the Cir- 
cumboreal Region. 

The facts obtained as a result of our analysis 
are probably best explained on the basis of the 
hypothesis that the Ranunculaceae probably orig- 
inated in the vast land mass of Eurasia, with east- 
ern Asia as a most significant center of secondary 
differentiation and conservation of many ranun- 
culaceous genera. There were two centers of for- 
mation for genera and species of Ranunculaceae, 
western and eastern, which developed rather in- 
dependently, though some groups migrated from 
one to the other. In our opinion, in the high-moun- 
tain flora of Sikang Yunnan, the Eastern Hima- 
layas, and adjacent territories the endemic genera 
Asteropyrum, Beesia, Calathodes, Souliea, Uro- 
physa, Kingdonia, Archiclematis, and Paroxy- 
graphis evolved, as did such genera as Trollius, 
Aquilegia, Anemone, and Clematis. Meanwhile, 
in the montane forest flora of Manchuria, Japan, 
and adjacent territories, the endemic genera Ane- 
monopsis, Miyakea, Semiaquilegia, and Dicho- 
carpum evolved. 

Plant geographers (77, 136, 166) regarded the 
Altai as the refugium of the Tertiary nemoral relicts 
and one of the important centers of development 
of the high-mountain Eurasiatic flora. They pro- 
posed that many ancient Eastern Asiatic species 
migrated westward through the Altai. Our data on 
the Ranunculaceae confirm the importance of the 
Altai flora in the development and distribution of 
many Eurasiatic groups of Angiosperms. Further- 
more, endemic genera and sections of the Ranun- 
culaceae in the Siberian provincial floras are ab- 
sent, and moreover, almost no species occur in all 
provincial floras of Siberia. These facts can be 
explained if one is to Ern that contacts of 
Siberian floras were not clos 

According to Tikhomirov (308) and Vasiljev 
(321), the origin of Siberian taiga floristic com- 
plexes are recent (Pleistocene), but Tolmachev (310) 
claimed that these complexes were more ancient 
(Early Pliocene). Our data support the latter opin- 
ion but show that the floristic elements of the Ra- 
nunculaceae of the northern Siberian taiga were 
derived from Eastern Asiatic ones. Consequently, 
some Siberian species of Trollius, Aconitum, Del- 
phinium, Anemone, Ranunculus, and Thalictrum 
are considered today as boreal or taiga elements, 

ut they are really related to the nemoral high- 


Volume 76, Number 4 


Ziman & Keener 1031 
Geographical Analysis of Ranunculaceae 


mountain elements of the more warmth-loving flo- 
ras of Manchuria and western China. 

Our analysis of the Ranunculaceae of the pro- 
vincial floras of Europe shows the presence of 25 
genera and about 250 species. Endemic ranun- 
culaceous genera are absent, but there are about 

5 endemic species restricted mainly to the Alps, 
Carpathians, Pyrenees, and other highlands, and 
include species of Aquilegia, Aconitum, Delphin- 
ium, Ranunculus, and Thalictrum. 

The distinctive features of the European pro- 
vincial floras include not only the presence within 
them of many common ranunculaceous species but 
also historical connections with the Siberian and 
especially with the Eastern Asiatic floras. Accord- 
ing to Popov (236), the Eastern Asiatic and Eu- 
ropean floras have the same age, but our data show 
that the latter are derivative from the former. In 
our opinion, therefore, the majority of the Tertiary 
ranunculaceous groups have migrated into Europe 
from the east during the Pliocene and became 
preserved there in the Alps, Carpathians, and other 
mountains. More recently, in these territories in- 
tensive radiation of new groups of Aconitum, Del- 
phinium, and Aquilegia took place. 

Our data from analysis of the role of the Ra- 
nunculaceae in arctic regions uphold the opinion 
of Tolmachev (312) and Budantsev (36) concern- 
ing the comparatively recent age of the arctic 
floristic complexes and their derivation from boreal 
and partly from nemoral complexes. Contrary to 
some plant geographers (95, 314, 320) who con- 
sider the territory of the Arctic Province as a 
floristic region or a special zone without division 
into regions and provinces, we follow Takhtajan's 
(2977) interpretation of this province. 

aucasus was a remarkable center in the 
formation of some high-mountain groups of An- 
giosperms in the Late Pliocene and Pleistocene (77, 
101, 102, 193, 293). These groups originated out 
of subtropical-forest species (complexes of the Pol. 
tava flora— 1 70), but some of them migrated there 
from the east, mainly from eastern Asia. 

We note also the presence within the modern 
Caucasian flora of 22 genera and about 230 species 
of Ranunculaceae (about 60 endemic). Almost all 
ranunculaceous genera in the Caucasian provincial 
floras are shared with those in the European and 
Siberian floras of the Circumboreal Region. Most 
species of these genera (Aquilegia, Aconitum, Del- 
phinium, Anemone, Pulsatilla, Clematis, and 
Thalictrum) occur in eastern Asia and Siberia. 

Some genera (e.g., Nigella, Consolida, Cera- 
tocephalus) occurring in the Caucasian floras are 
shared with the floras of the Mediterranean Region. 


In the flora of the Caucasus, there are some Eu- 
rasiatic disjuncts (e.g., Actaea spicata, Anemone 
nemorosa), which are generally accepted as Ter- 
tiary relicts. These data led us to hypothesize close 
connections among the ranunculaceous groups in 
Eurasia (including the Caucasus) during past pe- 
riods and to hypothesize possible migrations of their 
ancestral groups into the Caucasus from the east. 

The membership within the Ranunculaceae of 
large numbers of Caucasian endemics indicates lo- 
cal differentiation of this family in the high-moun- 
tain zones, especially within Delphinium and Ra- 
nunculus. Most of the Caucasian endemic species 
(e.g., Aquilegia colchica, Delphinium pyrami- 
datum, Anemone speciosa, Aconitum pubiceps, 
Ranunculus helenae) originated in the Western 
Caucasus (or Colkhis), which is considered to be a 
significant refugium of the Late Tertiary warm- 
temperate forest flora. Another large Tertiary flo- 
ristic refugium, which was in the Eastern Caucasus 
Hyrcan or Talysh), did not play a large role in 
the differentiation of the Ranunculaceae and hence 


— 


the Talysh endemic species of this family are not 
numerous. The evolution of the ranunculaceous 
flora of these two centers developed independently, 
and shared endemics as well as vicariant species 
of Ranunculaceae are almost absent. 

Some species of Ranunculaceae in the Caucasian 
floras are shared with the Mediterranean Region 
or are closely related to species there. They are 
predominately annual xerophytes (e.g., Nigella ar- 

ensis, Consolida ambigua, Ceratocephalus tes- 
ticulatus, Adonis aestivalis). The Caucasian ran- 
unculaceous species with connections in Anterior 
Asia are xerophytic and mainly annual (e.g., Del- 
phinium cyphoplectrum, Consolida divaricata, 

igella orientalis, Aconitella hohenackeri, Tha- 
lictrum sultabadense). At the same time, within 
the ranunculaceous species shared between the 
are high-moun- 
tain, mesophytic, mainly forest plants (e.g., Caltha 
polypetala, Trollius Aquileg 


caucasica, Anemone impexa, Pulsatilla RR 


Caucasus and Asia Minor, many 
ranunculinus, 


cea). 

All previously mentioned data led us to suggest 
close connections of the ranunculaceous groups in 
Eurasia (including the Caucasus) in past periods; 
possible migration into the Caucasus of some an- 
cestral groups of this family from the east through 
Asia Minor and some xerophytic groups from the 
Mediterranean Region; evolution of some groups 
within the Caucasian flora itse 

According to the literature, a flora of the Ap- 
palachian Province of the Atlantic North American 
Region is ancient (mainly Tertiary), whereas the 


1032 


Annals of the 
Missouri Botanical Garden 


flora of the North American Prairies and the At- 
lantic-Gulf Coastal Plain provinces are compara- 
tively recent and derivative. Our data support this 
conjecture, including close relationships of the bo- 
real floras of Eurasia with those of North America 
while, on the other hand, maintaining that there 
were relatively prolonged disconnections of the flo- 
ras in the western and eastern parts of both con- 
tinents (31, 98, 184). These floristic events were 
promoted by means of sea barriers (ancient Siberian 
Sea and the Straits of Mexico) and Miocene and 
more recent land bridges between Eurasia and North 
America through the Pacific and Atlantic oceans 
(135, 251). 

Seventeen ranunculaceous genera are shared by 
North American and Eurasian boreal floras. It is 
remarkable that more than half of them (Caltha, 
Actaea, Cimicifuga, Delphinium, 
Clematis, Anemone, Pulsatilla, Ranunculus, and 
Thalictrum) consist of sections with Eurasiatic— 
North American distributions. Although species in 
common to these floras are few (e.g., Anemone 


Aconitum, 


narcissiflora, Thalictrum alpinum, Ranunculus 
pedatifidus), pairs of corresponding or closely re- 
lated species are numerous (e.g., Actaea spicata- 
A. rubra, Trautvetteria japonica-T. grandis, 
Enemion raddeanum-E. biternatum). 

u ysis showed that the territories of the 
Appalachian and Rocky Mountain provinces were 
important centers of North American differentia- 
tion of the Ranunculaceae. Within the former, the 
Xanthorhiza, and Ane- 


=] 
E 


relict genera Hydrastis, 
monella evolved, and within both of them, some 
sections of Trollius, Aquilegia, Ranunculus, Tha- 
lictrum, Enemion, and Cimicifuga. Possibly, the 
two last-mentioned genera originated within these 
floras. Compared with the peculiarities of distri- 
bution of species of Hydrastis, Xanthorhiza, and 
Anemonella (they occur disjunctively through about 
ten states in the Appalachian province, e.g., Illinois, 
Tennessee, Virginia, and others), we regard the 
age of these genera (at least that of Enemion) as 
more recent than the Eastern Asiatic endemic gen- 
era (e.g., Beesia, Calathodes). 

A comparison of the distribution of the Ranun- 
culaceae in all four regions of the Boreal Subking- 
dom (Eastern Asiatic, Circumboreal, Atlantic North 
American, and Rocky Mountain) showed the con- 
siderable floristic role of this family in each. These 
data form the basis of our supposition that the 
development of the Ranunculaceae took place over 
a long period of time within the Tertiary warm- 
temperate flora which was in common to all cir- 
cumpolar territories of the Northern Hemisphere 
(31, 296). We regard western and eastern parts 


of eastern Asia and eastern North America (Ap- 
palachians) as significant refugia of this ancient 
flora. The Altai-Sayan id Alps, Carpathians, 
Pyrenees, Caucasus, and t ocky Mountains 
were, in our opinion, ee centers of the 


differentiation of some groups of the Ranuncula- 


eae. 

In the Madrean Region of the Madrean Sub- 
kingdom all ranunculaceous species are concen- 
trated on mountain slopes, and they are mainly 
members of the Californian and Mexican Highland 
provinces. These species are scanty in chaparral 
formations and absent in deserts. All genera of the 
Ranunculaceae are in common to those in both 
Eurasiatic and North American floras of the Boreal 
Subkingdom, but species in common are few and 
most of them are relict disjuncts. The mountains 
of California were one of the secondary centers of 
differentiation of Delphinium, and some groups of 
Thalictrum evolved in the Mexican Highlands. 

Our analysis of the Ranunculaceae within the 
Madrean Region upholds the suspected (251, 286) 
relationships of many northern Californian groups 
of species with those in the boreal floras in both 
western and eastern North America, and, more- 
over, with those in boreal Eurasia. Stebbins & 
Major (251) regarded Trautvetteria grandis, Del- 
phinium californicum, Clematis occidentalis, 
Anemone quinquefolia, and some other disjuncts 
as relicts of Tertiary deciduous mesophytic forests, 
although most endemic species of Delphinium have 
a more recent age, and our data confirm that 
hypothesis. It is remarkable that 15 genera of the 
Ranunculaceae are in common to the floras in the 
above-mentioned regions of the Boreal and Mad- 
rean subkingdoms. 

The modern flora of the regions of the Ancient 
Mediterranean Subkingdom is rather xerophytic, 
but the Ancient Mediterranean flora developed as 
a derivative and mainly migratory one from boreal 
and tropical mesophytic floras. Most of the Angio- 
sperms in the Mediterranean flora are related to 
corresponding Eastern Asiatic groups (Popov— 
245). 


Within the Mediterranean Region the floras of 
the Liguro- Tyrrhenian and Iberian provinces are 
quite rich with Ranunculaceae. They are similar 
to the Balkanian flora of the Circumboreal Region 
but much different from the rest of the Mediter- 
ranean provincial floras. At the same time, the 
flora of the East Mediterranean Province is related 
to the most xerophytic provincial floras of the Ir- 
ano-Turanian Region (e.g., Mesopotamian), and 
within the former about half of the species of Ra- 
nunculaceae are annuals. 


Volume 76, Number 4 


Ziman & Keener 1033 


Geographical Analysis of Ranunculaceae 


Our analysis of the Ranunculaceae within the 
Mediterranean floristic provinces showed their xe- 
rophytic features (presence of many annuals and 
bulb plants) and clear connections with mesophytic 
groups of this family in the floras of the Boreal 
Subkingdom, especially with those in the high- 
mountain floras of Southern Europe. 

Within the Irano-Turanian Region, most Ra- 
nunculaceae belong to the provincial floras of the 
Anterior Asiatic Subregion, while in the floras of 
the Central Asiatic Subregion there are relatively 
few Ranunculaceae. Most ranunculaceous repre- 
sentatives are concentrated in the flora of the Ar- 
meno-Iranian Province, which is surpassed world- 
wide for the richness of the Ranunculaceae only 
by the Sikang Yunnan Province. Nevertheless, the 
number of endemic species is remarkably low, es- 
pecially as compared with the flora of the Sikang 
Yunnan Province (30/220 vs. 180/400—see Ta- 
ble 1). Within Ranunculaceae predominate species 
with boreal and nemoral affinities predominate in 
the floras of eastern Asia and South Siberia. These 
phenomena confirm the close connections between 
the Armeno-Iranian, South Siberian, and Eastern 
Asiatic floras and might reflect that the ancient 
flora of Anterior Asia was once more mesophytic. 

From an analysis of the role of the Ranuncu- 
laceae in the provincial floras of the Irano-Turanian 
Region, we note that the highlands of Middle Asia 
(Pamiro-Alai, Tien Shan, and others) were sites of 
secondary formation of some groups of this family 
(more than 100 endemic species of Aconitum, 
Delphinium, Aquilegia, Anemone, and Ranun- 
culus). The floras of the Turkestanian, Dzungaro- 
Tien Shan, and Central Tien Shan provinces are 
very similar in terms of Ranunculaceae. This phe- 
nomenon reflects the autochthonous development 
of the Mountain Middle Asiatic flora on the base 
of the ancient boreal groups of Angiosperms (Ka- 
melin, 140, called them the Turkmenian and the 
Himalayan-Mediterranean branches). According to 
Pavlov (227) and Vykhodtsev (329), most of the 

iddle Asiatic mountain endemics are recent (Qua- 
ternary) and progressive, while Ovchinnikov (218) 
claimed that within them are some relict elements 
of nemoral complexes with certain Eastern Asiatic 
relatives (e.g., disjuncts Trollius lilacinus, Del- 
phinium brunonianum, Hepatica falconeri, Par- 
opyrum anemonoides). We agree therefore wit 
Kamelin (140) that the Mountain Middle Asiatic 
flora, including the floras of the Tien Shan, is a 
single unit. 

The Armeno-Iranian and Central Anatolian pro- 
vincial floras are similar to the flora of the Medi- 
terranean Region and are much different from the 


floras of Mountain Middle Asia 


annuals (species of Consolida, Nigella, and oth- 


. There are many 


ers), but almost no endemic species of the Ranun- 
culaceae. Species in common to the floras of Moun- 
tain Middle Asia, Armeno-Iranian, and Central 
Anatolian provinces are few, especially within 4c- 
onitum, Delphinium, Aquilegia, Anemone, and 
Pulsatilla. Meanwhile, the two last-mentioned flo- 
ras include some genera and many species shared 
with those in the Caucasian floras but absent in 
both the Middle Asiatic and Mediterranean floras 
(e.g., Caltha polypetala, Helleborus orientalis, 
Actaea spicata, Isopyrum thalictroides). It is re- 
markable that Caltha, Helleborus, 
Isopyrum are boreal mesophytes with many species 
in both the floras of the Eastern Asiatic and Cir- 
cumboreal regions. 

The floristic richness of the Ranunculaceae in 
the floras of the Western Himalayan and North 
Baluchistanian provinces is intermediate between 


Actaea, and 


the high-mountain floras of Middle and Eastern 
Asia. 

The Ranunculaceae in the Central Asiatic pro- 
vincial floras, both Mongolian and Tibetan, consist 
of few endemic species. The outlying northeastern 
districts are related to those in the Altai-Sayanian 
and West Chinese floras. 

We conclude that the flora of the Ancient Med- 
iterranean Subkingdom played a noteworthy role 
in the differentiation of the Ranunculaceae largely 
because of the evolution of annual (Vigella, Con- 
solida, and Aconitella) and bulb plants (species of 
Within the high- 


mountain floras, some groups of ranunculaceous 


Anemone and Ranunculus). 


endemic species recently evolved. 


PALAEOTROPICAL KINGDOM 


Location: territories of Southeastern Asia and 
Africa, also adjacent islands. 
Climate: mainly tropic 


Ranunculaceae: 11 genera, 100 species. 


Indo-Malesian Subkingdom 


This subkingdom includes four regions: Male- 
sian, Fijian, Indo-Chinese, and Indian. 

Location: Hindustan and Indo-China, also nu- 
merous islands in the Indian and Pacific oceans. 

Ranunculaceae: Ó genera, 80 species. 


Malesian Region 


Location: south of the Malacca Peninsula, the 
Malaya Archipelago, New Guinea, and adjacent 
little islands. 


1034 Annals of the 
Missouri Botanical Garden 
Ranunculaceae: 5 genera, 60 species (12, 51, Papuan Province 
70, 177, 203, 204, 287, 288, 289). 


Charasierisii vegetation: tropical forests in 
lowlands, also insular high-mountain formations. 


Malaccan Province 


Location: Malacca Peninsula and some little 
islands. 

Ranunculaceae: 3 genera, 5 species (Anemone 
sumatrana, Clematis smilacifolia, C. meyeniana, 
Naravelia dasyneura, N. laurifolia). 

elated VN South Malesian (3/5) and Ka- 
limantanian ( 

O EON no endemic species. 

Kalimantan, Moluccan, and Bismarckian 
Provinces 


Ranunculaceae: 3/4, 1/2, and 1/1. Endemic 
species are few (e.g., Ranunculus lowii). 


Philippinean Province 


Ranunculaceae: 5 genera, 15 species. 

Related floras: South Malesian (5/3, e.g., Na- 
ravelia laurifolia, Clematis lechenaultiana). 

Observations: 6 endemic species (e.g., Nara- 
velia pauciflora, Clematis antonii, Ranunculus 


philippinense, Thalictrum philippinense). 


South Malesian Province 


Ranunculaceae: 5 genera, 15 species. 

Related floras: Malaccan (3/5) and Philippi- 
nean (5/3). 

Observations: 5 endemic species (e.g., Clematis 
korthalsii, Ranunculus javanicus, Thalictrum ja- 
vanicum). 


Sumatran Province 


Ranunculaceae: 5 genera, 10 species. 

Related flora: South Malesian (5/8, e.g., com- 
mon endemics Ranunculus javanicus, Clematis 
javanica, and others). 


Observations: 5 endemic species (e.g., 4nemone 


angustiloba, Ranunculus sundaicus). 


Sulawesian Province 


Ranunculaceae: 2 genera, 15 species. 

Related floras: South Malesian (2/2, e.g., Ra- 
nunculus blumei). 

Observations: about 15 endemic species (e.g., 
Clematis molissima, Ranunculus celebicus, R. 


frigidurbis). 


Ranunculaceae: 3 genera, 30 species. 

Related flora: Sulawesian (2/2, e.g., Clematis 
pickeringii). 

Observations: about 30 endemic species (e.g., 
Clematis papuasica, C. i aati Ranun- 


R. brass 


culus bellus, 


Fijian Region 

Location: Fiji, New Hebrides, Samoa, Tonga, 
and adjacent small archipelagoes. 

Ranunculaceae: 3 genera, 4 species (Clematis 
pickeringii, Ranunculus perindutus, R. keysseri, 
Thalictrum papuanum — 

Related flora: Papuan (2/2, a 
pickeringii, Thalictrum papuant 

is region includes the Fijian nd Hebridean 


Clematis 


provinces. 


Indo-Chinese Region 


Location: peninsular Indo-China, adjacent con- 
tinental territories (Southeastern Bangladesh, 
Southeastern Burma, South China, Thailand, Laos, 
Kampuchia, and Vietnam), also Andaman and 
Hainan islands. 

Ranunculaceae: 6 genera, 25 species (21, 70, 
277, 299, 302). 

Related floras: Eastern Asiatic (6/10, e.g., 
Clematis acuminata, Ranunculus chinensis, Tha- 
lictrum punduanus) and Malesian (5/5, e.g., 
Clematis smilacifolia, Naravelia laurifolia). 

Jbservations: few endemic species (e.g., Clem- 
atis thaiana, Thalictrum siamense). 


Indian Region 


Location: peninsular Hindustan, Ganges Plain, 
South Foothills of the Himalayas, and some islands 
(e.g., Sri Lanka). 

Ranunculaceae: © genera, about 10 species 
(123, 262); all ranunculaceous species are con- 
centrated in the high-mountain flora of the Deccan 
Plateau, West and East Gaty ridges. 

Related floras: Eastern Himalayan (5/5, e.g., 
Anemone rivularis, Clematis nutans, C. grevi- 

aflora), Malesian (5/3, Anemone vitifolia, 
Naravelia laurifolia), Er dan (5/2, e.g., Del- 
phinium dasycaulon and Clematis grata). 

Observations: few endemic species (e.g., Clem- 
Naravelia Thalictrum 


atis triloba, zeylanica, 


dalzellii). 


Volume 76, Number 4 


Ziman & Keener 1035 
Geographical Analysis of Ranunculaceae 


Polynesian Subkingdom 


Location: many islands of Polynesia, Hawaii, 
and Micronesia. 

This subkingdom includes two regions— Poly- 
nesian and Hawaiian—and three provinces— Mi- 
cronesian, Polynesian, and Hawaiian. 

Ranunculaceae: 1 genus and 2 species (Ra- 
nunculus hawaiensis and R. mauensis— 45). 


Neocaledonian Subkingdom 


This subkingdom includes only the Neocaledo- 
nian Region and the Neocaledonian Province. 
Location: New Caledonia. 
Climate: tropical. 
anunculaceae: only Clematis pickeringú 


(110) 


African Subkingdom 


This subkingdom includes four regions— Gui- 
neo-Congo, Sudano-Zambesian, Karroo-Namib, and 
St. Helena and Ascension islands. 

Location: main part of the African continent to 
latitude 20°S, also tropical deserts in the south of 
the Arabian Peninsula and in Anterior Asia. 

Climate: tropical, but in highlands more or less 
temperate. 

Ranunculaceae: 8 genera, 50 species (3, 34, 
69, 72, 117, 121, 127, 206, 217, 242, 252, 
282, 318). The Ranunculaceae are absent in trop- 
ical deserts and semideserts; are scarce in tropical 
rainforests and arboreal woodlands in low places 
of northern, western, and central tropical Africa; 
and are important in tropical mountain and afro- 


alpine floras (117, 118). 


Guineo-Congo Region 


Location: territories adjacent to the Guinean 
Bay, including North Angola and the Congo Basin, 
and western Kenya, Tanzania, and Uganda. 

Characteristic vegetation: predominately trop- 
ical forests and savannas, also highland formations 
in the extreme east. 

Ranunculaceae: 7 genera, 40 species concen- 
trated in the Congo Province, but absent in the 
Guinea Province. 

Related floras: Sudano-Zambesian (6/ 15, e.g., 
Anemone thomsonii, Clematis simensis, Del- 
phinium leroyi, Thalictrum rhynchocarpum), 
Cape (7/10, e.g., Knowltonia transvaalensis, 
Delphinium leroyi, Clematopsis hombleyi, Ra- 
and Western Chinese (5/2, 


nunculus multifidus), 


e.g., Clematis grata and Delphinium dasycau- 
on). 

Observations: about 20 endemic species (e.g., 
Delphinium macrocentron, Clematis dolichopo- 
da, Thalictrum zernyi, Ranunculus aberdaricus). 
Most species belong to Clematis (about 15) and 
Ranunculus (about 20). Endemic species occur 


mainly in the upper belts of Elgon, Kilimanjaro, 
Aberdare, and other mountains at elevations of 
2,500-4,500 m 


Sudano-Zambesian Region 


Location: vast territories of western, central, 
eastern, and southern Africa, also territories in the 
Arabian Peninsula and southwestern Asia. 

Characteristic vegetation: predominately sa- 
vanna and desert formations, some high-mountain 
areas. 

This region includes the Sahelian, Nubo-Ara- 
bian, Omano-Rajastanian, Socotran, South Ara- 
bian, Sudanian, Somalo-Aethiopian, and Zambe- 
sian provinces. 

Ranunculaceae: 8 genera, about 30 species 
mostly in the two last-mentioned provinces. 

Related floras: Guineo-Congo (6/ 15) and Cape 
(6/5, e.g., Clematis welwitchii, Clematopsis sca- 
biosifolia, Thalictrum rhynchocarpum ). 

Observations: O endemic species 
Clematis | 


(e.g., 
Delphinium wellbyi, 


about 
inciso-dentata, 
Ranunculus oligocarpus). 
the floras of the Karroo-Namib Region and 
the region of the St. Helena and Ascension islands, 
Ranunculaceae are almost absent. 


Madagascan Subkingdom 


This subkingdom includes only the Madagascan 
Region. 

Location: Madagascar and adjacent islands (e.g., 
Komory, Seychelles, and others) to the east of the 
African continent. 


Madagascan Region 


Location: Madagascar. 

Characteristic vegetation: predominately trop- 
ical forests. 

Ranunculaceae: 4 genera, about 10 species 
(e.g., Clematis stanleyi, Clematopsis bojeri, Ra- 
nunculus madagascarensis, Knowltonia vesica- 
toria— l6, 252). 

Related floras: Sudano-Zambesian (4/2). 

Observations: few endemic species. 


1036 


Annals 
Sora ES Garden 


The Indo-Malesian Subkingdom is regarded as 
the greatest refugium of the Palaeotropical Angio- 
sperms in which were preserved many endemic 
and relict families and genera (10, 296, 337). 
There are few ranunculaceous representatives, and 
they occur mainly in the mountain flora of this 
subkingdom. Lam (177) and van Steenis (287) 
regarded Anemone, Ranunculus, and Thalictrum 
as examples of temperate genera that penetrated 
the Malesian tropical mountain flora from temper- 
ate eastern Asia through ancient mountain ridges. 
Our data do not challenge this idea. Possibly, the 
Ranunculaceae migrated through Malesia in the 
Southern Hemisphere by two different routes. The 
western route went through Sumatra, Kalimantan, 
and Java, and the eastern one passed through the 
Philippines, Molucca, Sulavesi, and New Guinea. 
Van Steenis (288) and Aubreville (9) supposed 
these groups of islands to be the distinct centers 
of development and dispersal of the Angiosperms: 
the former island group was related to the South- 
eastern Asiatic floristic center, while the latter group 
was home to part of the Australo-Papuan flora. 

The formation of uplands in New Guinea in the 
Late Oligocene limited north-to-south penetration 
by high-mountain Ranunculaceae (201, 202, 248). 
After that time some endemic groups of Clematis 
and Ranunculus evolved in the highlands of Ma- 
lesia. Areas of some species (e.g., Naravelia da- 
syneura, Anemone sumatrana, Clematis leche- 
naultiana) occur in the adjacent Indo-Chinese, 
Malesian, and Indian regions, while some other 
species (e.g., Anemone vitifolia, Clematis mey- 
eniana) occur disjunctively also in the highlands 
of West China, the Himalayas, and Japan. The 
most interesting area has Clematis pickeringii be- 
cause it includes Malesia, New Caledonia, and 
northeastern Australia. 

n the flora of the African Subkingdom only two 
genera are restricted to Africa and Madagascar: 
Clematopsis, which is related to Clematis, and 
The 
rest of the ranunculaceous genera (Anemone, 
Clematis, Delphinium, Nigella, Ranunculus, and 
Thalictrum) are shared by the floras of the Boreal 
and Ancient Mediterranean subkingdoms, and many 
species of them are related to the species of the 


Knowltonia, which is related to Anemone. 


latter subkingdom. For example, Anemone thom- 
sonit is distributed in the alpine and subalpine belts 
of Mount Kilimanjaro and a related species, 4. 
glaucifolia, grows in high mountains of western 
China. Clematis grata and Delphinium dasycau- 
lon are common to the floras of western China, 
the Himalayas, and in Hindustan. 


Our analysis of the distribution of the Ranun- 
culaceae in Africa confirms Hedberg's (117, 118) 
and Boughey's (27) suggested existence of the An- 
cient Tropical Mountain flora in Africa and the 
possible connections between the Afro-Alpine and 
Boreal floras of Africa and Eurasia in past periods. 
We suppose that the ranunculaceous representa- 
tives migrated into Africa from Eurasia by means 
of some ancient land bridges (e.g., Sunda, Arabian) 
that existed there until the Late Cretaceous and 
early Quaternary (246, 250, 335). Afterwards, 
formation of the mountain ridges in eastern Africa 
(dated as Miocene) promoted development of the 
high-mountain groups of Ranunculaceae and mi- 
gration of some of them into South Africa, where 
cold and temperate conditions existed until the 
Cretaceous (53). Few groups (e.g., species of Clem- 
atis, Clematopsis, and Knowltonia) migrated 
southward and secondarily became adapted to the 
tropical conditions of Africa 


CAPE KINGDOM 


Location: southern tip of Africa. 

This subkingdom includes only the Cape Region. 

Characteristic vegetation: predominately scle- 
rophyllous scrub and 

Ranunculaceae: 7 genera, about 30 species. 

Related floras: Guineo-Congo (7/10) and Su- 
dano-Zambesian (6/5). 

Observations: about 10 endemic species (e.g., 
Clematis brachiata, Knowltonia hirsuta, Ranun- 
Anemone 


forest formations. 


culus cooperi). Some endemics (e.g., 
capensis and 4. alchemillifolia) are related to 
endemics in the floras of the Eastern Asiatic Re- 
gion. 


The Cape flora is extraordinarily rich and dis- 
tinctive (297), but the Ranunculaceae in it are 
rather few and are related to those in the floras of 
the African and Boreal subkingdoms. 


NEOTROPICAL KINGDOM 


Location: southern Florida in North America, 
lowlands of Mexico, Central America, and South 
America (except the extreme south), also adjacent 
islands. 

Climate: tropical in lowlands, temperate in high- 


Pennatus 9 genera, 70 species. 

This kingdom includes five regions: Caribbean, 
Amazonian, Brazilian, Andean, and the Guayana 

ighlands 


Volume 76, Number 4 
1989 


Ziman & Keener 1037 
Geographical Analysis of Ranunculaceae 


Caribbean Region 


Location: extreme southern North America 

(tropical Florida), almost all Central America, and 

some islands mainly in the Caribbean Sea. 
Ranunculaceae: 3 genera, about 20 species (2, 


67, 281). 


Galapageian Province 


Location: Galapagos Islands. 
Ranunculaceae: absent. 


West Indian Province 


Location: Cuba, Jamaica, and other Caribbean 
islands. 

Ranunculaceae: 2 genera, 5 species (e.g., 
Clematis dioica, C. havanensis, Ranunculus cu- 
bensis). 


Central American Province 


Location: Guatemala, Honduras, Nicaragua, 
Costa Rica, and Panama. 

Ranunculaceae: 3 genera, about 20 species. 

Related floras: Mexican Highlands (3/5, e.g., 
Clematis haenkeana, Ranunculus donianus, 
Thalictrum hintonii) and Andean (3/10, e.g., Ra- 
nunculus flagelliformis, R. peruvianus, Thalic- 
irum panamense). 

Observations: 6 endemic species, all Thalictrum 
(e.g., T. guatemalense, T. standleyi). 


Region of the Guayana Highlands 


Location: Guayana Plateau (southern Venezue- 
la, western Guyana, western Colombia, and north- 
ern Brazil). 

Characteristic vegetation: as in the previous 
and next two regions, predominately tropical for- 
ests, savannas, hylaeas, campo limpo, and chaco 
formations. 

Ranunculaceae: 


2 genera, 4 species (e.g., 


Clematis dioica, Ranunculus apiifolius). 


Amazonian Region 


Location: Amazon Basin in Brazil. 
Ranunculaceae: | genus, 2 species (e.g., Clem- 
atis brasiliana). 


Brazilian Region 


Location: main part of Brazil, eastern Paraguay, 
Bolivia, and northern Argentina 
anunculaceae: 2 genera, 


9 species (e.g., 


Clematis hilarii, Ranunculus platensis— 186, 


187, 188). 


Andean Region 


Location: Andean ridges from Venezuela to 
northern Chile. Predominately highlands with sum- 
mits (e.g., Chimboraso and Christóbal Colón) of 
about 5,000-6,000 m 

Characteristic vegetation: predominately high- 
mountain formations of temperate elements. 

Ranunculaceae: 9 genera, 60 species (59, 71, 
83, 186, 

¡A Mexican Highlands (5/20, e.g., 
Clematis sericea, Anemone decapetala, Myosu- 
rus apetalus, Thalictrum podocarpum) and Chile- 
Patagonian (5/10, e.g., Caltha sagittata, Myo- 
surus aristatus, Anemone helleborifolia). 

Observations: 2 endemic genera and about 30 
endemic species. The endemic genera, local and 

monotypic, are represented by Oreithales integ- 
rifolia and Laccopetalum giganteum, which are 
distributed in Peru, Bolivia, and Ecuador at about 
4,000 m. Endemic species belong mainly to Ra- 
nunculus (about 20—e.g., R. pilosus), but some 
endemics include Clematis (e.g., C. albo-rosea), 
Anemone (e.g., A. jamesonii), and Thalictrum (e.g., 


T. longistylum). 


Summing the data of the analysis of the Ra- 
nunculaceae in the floras of the Neotropical King- 
dom, we must keep in mind that North America 
was separated from South America with a relatively 
wide sea space during Cretaceous and Paleogene 
time (250). On the other hand, until the Oligocene 
South America was joined through the subantarctic 
bridge with Antarctica which at that time was cov- 
ered with cool temperate forests (246, 248). The 
Andes were formed in the Pleistocene and after 
that they became an important transtropical bridge 
for the migration of temperate floristic elements 
from North America to South America and vice 
versa (211, 

Species of Thalictrum and Anemone multifida 
migrated through the Andes from the north. In 
fact, most of the species allied to the Andean species 
are distributed in North and Central America (e.g., 
species of Thalictrum, sects. Camptogastrum and 
Pelteria, species of Anemone, series Multifida). 
But some species of Clematis sect. Clematis, sub- 
Riv- 
ularidium, Caltha sect. Psychrophila, and Ra- 


nunculus sect. Trollianthoidea were found almost 


sect. Dioicae and Aristatae, Anemone sect. 


exclusively in South America, richly represented 


1038 


Annals of the 
Missouri Botanical Garden 


in the Andes and with relatives in the extreme 
south of the Southern Hemisphere. 

The Andean high-mountain flora was one of the 
important centers of ranunculaceous differentia- 
tion. There are two distinct monotypic endemic 
genera in this flora, Oreithales and Laccopetalum. 
The former is hexaploid and seems to be derived 
from Anemone (32, 68); the latter differs from 
other Ranunculaceae by possessing a large number 
of nectary glands and possessing large unbroken 
leaves (length over 40 cm) 


HOLANTARCTIC KINGDOM 


Location: territory of South America to latitude 
30°S, some islands in the southern extremity and 
New Zealand. 

Climate: temperate and mainly cold. 

Ranunculaceae: 8 genera, 100 species. 

This kingdom includes four regions: Chile-Pat- 
agonian, Fernandezian, Neozeylandic, and South 
Subantarctic islands. 


Chile-Patagonian Region 


Location: Chile, Uruguay, central and southern 
Argentina, Tierra del Fuego, Falklands and other 
islands. 

Characteristic vegetation: coniferous (Arau- 
caria and other genera), deciduous and evergreen 
(Nothofagus) forests, sclerophyll shrubs, cushion 
heath, dry grassy places, cold boggy sites, and 
alpine meadow ii 

Ranunculaceae: 7 genera, 50 species (186, 

) 


188, 210, 21 


North Chilean and Middle 


Chilean Provinces 


Location: territory of Chile. 

Ranunculaceae: 8 genera, about 30 species. 

Related floras: Andean (5/10), Pampean (3/ 

, €g., Clematis hilarii, Ranunculus platensis) 
and Magellanian (3/5, e.g., Caltha sagittata, Ra- 
nunculus biternatus). 

Observations: 10 endemic species (e.g., Anem- 
one rigida, Ranunculus chilensis). There are some 
species of two endemic genera, Barneoudia (e.g., 
B. chilensis, B. major) and Hamadryas (e.g., H. 

) 


argentea). 


Pampean Province 


Location: southern Brazil, partly Argentina and 
Uruguay. 
Ranunculaceae: 3 genera, about 10 species. 


Related floras: Chilean (3/5). 


Observations: few endemic species (e.g., Anem- 
one decapetala, Clematis bonariensis). 


Patagonian Province 


Location: southern Argentina. 
Ranunculaceae: 3 genera, 5 species (e.g., Ra- 
nunculus patagonicus, Myosurus patagonicus). 


Magellanian Province 


Location: Magellanian and adjacent islands. 

Ranunculaceae: 3 genera, about 15 species. 

Related flora: Chilean (3/5). 

Observations: 10 endemic species (e.g., Ham- 
adryas magellanica, Caltha appendiculata, C. 
dioneifolia). 


Fernandesian Region 


Location: Juan-Fernández and Desventuradas 
islands. 
Ranunculaceae: absent. 


South Subantarctic Islands Region 


Location: Tristan d'Acunha, Kerguelen, and ad- 
jacent islands. 

Ranunculaceae: 1 genus, 5 species (e.g., en- 
demic Ranunculus carolii). 


Neozeylandic Region 


Location: New Zealand and adjacent islands. 

Ranunculaceae: 5 genera, 60 species (5, 45, 
80, 111, 196, 336). 

Related flora: Chile-Patagonian (5/2, e.g., Ra- 
nunculus acaulis, R. biternatus). 

Observations: about 55 endemic species (e.g., 
Caltha obtusa, Clematis australis, Myosurus no- 
vae-zelandiae, Ranunculus gracilipes). Most 
species occur in the South Island only, while some 
(e.g., Ranunculus insignis, Caltha novae-zelan- 
diae) are distributed in North Island and South 
Island. The majority of species are high-mountain 
plants, and a few (e.g., Clematis afoliata, Ra- 
nunculus rivularis) occur in low-mountain zones 
or plains. 


Our analysis showed that within the Holantarctic 
Kingdom the Ranunculaceae are distributed mainly 
in the temperate highlands or in very cold plains. 
Two endemic oligotypic genera, Barneoudia and 
Hamadryas, were formed in the Chile-Patagonian 
E while in the high zones of mountains of 

uthern South America and New Zealand 
some groups of the Ranunculaceae (mainly Ra- 
evidently from common 


nunculus) developed, 


Volume 76, Number 4 


Ziman & Keener 1039 
Geographical Analysis of Ranunculaceae 


ancestors. Fisher (80) noted the predominance of 
polyploids within the New Zealand species of Ra- 
nunculus and listed 10 pairs of alpine species with 
one member in New Zealand and the other member 
a similar South American species (e.g., R. haastii— 
R. semiverticillatus and R. nivicolus- R. capra- 
rum). Consequently, Fisher supposed that high- 
elevation New Zealand Ranunculus was derived 
from the South American one, not from Malesian 
species. 

Mountain ridges in New Zealand formed in the 
Late Pliocene (248), along with the rise of the 
mountains in Australia and New Guinea. According 
to Raven (248) and Wardle (334) the initial species 
of Anemone, Clematis, and Ranunculus arrived 
in New Zealand from South America in the Late 
Pliocene or later, while species of Caltha already 
spread there in the Paleogene. 


AUSTRALIAN KINGDOM 


Location: Australian continent. 

Ranunculaceae: 5 genera, about 50 species. 

This kingdom includes the Southwest, East, and 
Central Australian regions. 


East Australian Region 


Location: Queensland, New South Wales, Vic- 
toria, and Tasmania. 

Characteristic vegetation: high-mountain for- 
est and ericoid formations. 

Ranunculaceae: 5 genera, about 45 species 

(24, 37, ). 
Related pow Neozeylandic (2/5, e.g., Clem- 
atis foetida, Ranunculus rivularis) and Chile-Pa- 
tagonian (2/3, e.g., Ranunculus acaulis, R. bi- 
ternatus). 

Observations: about 30 endemic species (e.g., 
Anemone crassifolia, Caltha introloba, Clematis 
aristata, C. gentianoides, Ranunculus gunni- 
anus), and most of them are endemics of Tasmania 
only (e.g., Ranunculus muelleri, R. scapigeri). 


Southwest Australian Region 


Location: southwestern Australia. 

Characteristic vegetation: dry and rainy scler- 
ophyllous forests of Casuarina and Eucalyptus, 
also scrub. 

Ranunculaceae: 2 genera, 
Clematis microphylla, Ranunculus lappaceus). 


Central Australian Region 


Location: North Territory, Great Sandy Desert, 


Victoria and Gibson deserts, and other areas. 


9 species (e.g., 


Characteristic vegetation: tropical desert and 
dry savanna formations. 
Ranunculaceae: absent. 


The Ranunculaceae in Australia are related to 
those in the Holantarctic floras. Thus, Caltha in- 
troloba, like other species of Caltha in the South- 
ern Hemisphere, belongs to sect. Psychrophila; 
Anemone crassifolia belongs to sect. Rivularid- 
ium; and Australian species of Ranunculus are 
related to South American species. Our conclusion 
is similar to that of Raven (248) who supposed that 
most high-mountain plants, including species of 
Anemone and Ranunculus, migrated to Australia 
and New Zealand from South America in the Late 
Pliocene or later. Raven suggested that a northern 
migratory route for plants into Australia through 

alesia was geologically impossible because the 
alpine flora of New Guinea is younger than t 
Australian one (334). Moreover, Melville (201) 
included the Ranunculaceae with a group of five 
families that invaded in Australia from the north. 


CONCLUSIONS 


The peculiarities of the role of the Ranuncula- 
ceae in the floras of regions and provinces world- 
wide reflect the peculiarities of origin, differentia- 
tion, and distribution of this family, including the 
relationships of tropical, temperate, mountain, and 
lowland floras, and including the relationships of 
the floras of the Northern and Southern hemi- 
spheres. 

According to paleobotanical literature (22, 60— 
63, 93, 253-255), there are no fossil remnants 
of the Ranunculaceae in the Cretaceous. The ear- 
liest such fossils, fruits of some species of Clematis 
and Ranunculus, are Oligocene and come from 
Eurasia (e.g., England, Ukraine, the Urals). Ra- 
nunculaceae are represented in Miocene deposits 
by remnants of few species of Clematis, Ranun- 
culus, and Thalictrum. Pliocene ranunculaceous 
remnants belong to Aquilegia, Anemone, Clema- 

is, Ranunculus, and Thalictrum, about 40 tem- 
perate species altogether. They were found in many 
places within the vast territories of Eurasia, North 
America, and South America. 

These paleobotanical phenomena reflect the 
considerable differentiation of the Ranunculaceae 
and their radiation throughout the world durin 
the Neogene. The sharp increase of the develop- 
ment and dispersion of the Ranunculaceae was 
associated with the Upper Eocene general fall of 
temperature and replacement of tropical and sub- 
tropical floras with temperate ones across Eurasia 


and North America (78, 229, 307). 


1040 


Annals of the 
Missouri Botanical Garden 


There are two different opinions about the origin 
of the Ranunculaceae. Scharfetter (263) placed 
the origin within the Early Tertiary tropical flora, 
while Popov (235) claimed that the Ranunculaceae 
originated within the Neogene warm-temperate flora 
of Eurasia, similar to 50 other derivative angio- 
spermous familie 

e P with both these propositions and 
claim an Early aceous origin and differentiation 
of the Ranunculaceae. This is based partly on the 


peculiarities of the disjunctive areas of such genera 
as Caltha, Anemone, Clematis, Ranunculus, and 
Thalictrum. The origin and primary differentiation 
of the family must have taken place after the 
breakup of the supercontinent Pangaea (or one like 
it). Our data are consistent with the horizontal drift 
of the continents (138, 201, 250, and others). 

Development of the distinctive Holarctic and 
Holantarctic groups of Caltha (sects. Caltha and 
Psychrophila) occurred in temperate parts of 
Gondwana and Eurasia and seems likely to reflect 
the Middle Cretaceous differentiation of the north- 
ern and southern temperate floras. In this point we 
agree with Tolmachev (310) and Moore (211) who 
considered Caltha as one of the most ancient tem- 
perate Angiosperm genera 

The above-mentioned Anemone, Clematis, Ra- 
nunculus, and Thalictrum are temperate genera 
that occur fairly continuously throughout the mon- 
tane Palaeotropics and Neotropics and suggest the 
routes by which their migrations could have taken 
place. Notably, Anemone rivularis and Clematis 
ranunculoides with archaic morphological features 
are tropical-mountain forest elements of the flora 
of Southeastern Asia. Boughey (27) regarded the 
mountain forest flora in Africa and Asia as very 
old and maintained that its survival to the present 
as an insular flora scattered discontinuously over 
huge land masses implies a connection with con- 
tinental drift, land-bridges, and long periods of time. 

Our analysis showed that the majority of species 
of all genera occur in mountains and that, more- 
over, species of more than half of the genera, 
including all endemics, are restricted chiefly to 
mountain locations. Moreover, the most numerous 
groups of species of large genera such as Aconitum, 
Delphinium, Anemone, Clematis, Ranunculus, 
and Thalictrum are distributed mainly in moun- 
tains. Our data confirm Rau (243) who considered 
the Ranunculaceae to be an important member of 
the high-mountain ecosystems and is the basis for 
supposing its origin in mountain locations. 

Within temperate montane floras of the tem- 
perate regions the Ranunculaceae take fourth to 


seventh place, while in tropical montane habitats, 
the Ranunculaceae fall to 15th-17th place (70, 
140) 


Comparison of the ranunculaceous representa- 
tives in the modern floras of the Northern and 
Southern hemispheres (the former contains 51 gen- 
era and about 2,200 species of all 14 tribes, the 
latter is home to 11 genera and about 250 species 
of 5 tribes) and comparison of their endemic genera 
(17 genera of 10 tribes vs. 4 genera of 2 tribes, 
respectively) indicates that primary differentiation 
of the Ranunculaceae took place in the Northern 
Hemisphere. 

From our geographical analysis together with 
data on comparative morphology, we propose that 
the ancestors of the tribes Cimicifugeae, Anemo- 
neae, Clematideae, Aquilegieae, Asteropyreae, and 
Kingdonieae initially developed within the ancient 
floras of eastern Asia, while the tribes Coptideae 
(at least Xanthorhiza) and Hydrastideae were 
formed in North America. It was impossible to 
dide the place of the development of the ances- 

s of the tribes Trollieae, Helleboreae, Delphi- 
nieae, Isopyreae, Ranunculeae, and Thalictreae, 
because some of their genera (e.g., Beesia, Cal- 
athodes, Trollius, Aconitum, Eranthis, Aquilegia, 
Semiaquilegia, Urophysa, and Dichocarpum) 
originated in eastern Asia, while other genera (e.g., 
Helleborus, Nigella, Consolida, Leptopyrum, and 
Aconitella) differentiated in the Ancient Mediter- 
ranean Subkingdom or North America (Enemion 
and Anemonella). Some associated in their origin 
and distribution with the temperate palearctic flora 
Adonis, Trautvet- 
teria, Isopyrum, Paraquilegia, and Paropyrum). 


(Delphinium, Callianthemum, 


In conclusion, we regard the Ranunculaceae as 
an ancient temperate family that originated in Cre- 
taceous time within montane temperate floras of 
the Northern Hemisphere. 


LITERATURE CITED 


. ABRAMS, L. 1951. Illustrated Flora of the Pacific 
eae ie ae ress, d California. 
"red 


mÓTZ 


2. ADA Pr | & R. W. REID. 
1972. nr Pak of nt Univ. West 
Indies, Mona, Jamaica. 

3. ADAMSON, R. S. . SALTER. 1950. Flora 
of the Cape Peninsula. Juta and Co., Capetown, 
Johannesburg. 

4. AICHELE, W. SCHWEGLER. 1957. Die 
— ai der Cattung diapers Feddes Rupe 
Spec. Nov. Regni Veg. 60: 1- 

5: Ms EVA, L. M. 197 » gie E sosudistykh 


rastenyi ostrova Kunashir (Analysis of the flora of 
s as of the Kunashir Island). Pp. 72- 
77 in S. S. Kharkevich (editor), Biology and In- 


Volume 76, Number 4 


Ziman & Keener 1041 
Geographical Analysis of Ranunculaceae 


m 


ee 


e 


mm 
m 


B 


3. BIA AYA, 1974. 


~ 


m 


© 


troduction of Useful Plants of the Sakhalin Region. 
Juzno-Sakhalinsk. [In Russian. 

MOORE & E. Epcar. 1961. 
Flora of New Zealand, Volume 1. Government 


Printer, Wellington, New oe aland. 

ANDRULAJTIS, S. F., e FODORIANOVA M. M. 
IVANOVA, ET AL. wi Sostav ^h, Putorana 
(Composition of the Putorana flora). 40-162 
in L. I. Malyshev (editor), dde b Patecana: 
Nauka, Nereis [In Russian. } 

APALJA, . Lekavicius. 1961. Ranuncu- 


laceae. Pp. rry in A - Mynkevicius (editor), 

Lietuvos TSR Flora, Volume 3. Publ. Polit. Sci., 

Vilna. us Us ] 

AUBR . 1975. La flore australe-papoue. 

Origine et n s Adansonia, Ser. 2, 15 - 
0. 


. —. 1976. Centres tertiaires d'origine, ra- 

ere et migrations des flores Uri A 

tropicales. Adansonia, e 2, 16: 29 

. AXELROD, D. J. € P. H. Raven. pou Late 

b e and Tertiary en history of Af- 
a. Pp. 77-130 in J. A. Werger (editor), Bio- 

iaa and Ecology of Southern Africa. 

unk, The api 

x KER, C. SR. BAKHUIZEN VAN DEN BRINK & 

zt E 1963. Flora of Java, Volume 1. 

. P. Noordhoff, Groningen, the Netherlands. 

Verkhne-Mioceno- 

vaya Flora Juinogo Primorja (The Upper Miocene 

Flora of the South Primorje). Nauka, Leningrad. 


. BALFOUR, B. & W. W. SMITH. 1915. A new 
genus of Ranunculaceae from Burma and Yunnan. 
Not. Roy. Bot. Gard. Edinburgh 9: 63 2 


5: BARBARICH, A. L, D. N. DOBROCHAEV 


DUBOVIK, ETAL. 1986. Khorologiya Flory "krainy 
(The Gborolagy of the Ukrainian Flora). Naukova 
dumka, Kiev. [In Ped 

. BARKLEY, T. M. (editor). 1977. Atlas of the 
Flora of the Great Plains. lowa State Univ. Press, 
Ames, lowa. 

. BATHIE, P. Pad — Biogeographie des Plantes de 


Dz 
t r 
> 
= 
= 


ertiary evolution of the 
Australian flora in the light a latitudinal movements 
of the continents. J. Biogeogr. 4: 11-118. 

BELAYA, G. A., D. P. VOROBJEV, N. N. GURZENKOV, 
ET AL. 1981. alias Pp. 78-89 in S. 
K. Chere pude (editor), Opredelitel rin ae 
Rastenyi E ME. Oblasty (Manual of the V: 
nts 


cular Pl e Kamchatka Region). Nauka, 
Moscow. [In Russin ] 

ELDIE, A. 77. Flora României. Mir A 
vos ae mds Vasculare, Volume 1. Edit. 


Acad. ] r 
i EU T A treatise of the North Amer- 
ican ranunculi. Amer. Midl. Naturalist 40 261. 
. BENTHAM, C. Flora uds iu. Volume 


pend 


2. Berry, E. W. 1924. The Middle and Upper 


Cretaceous Eocene floras of southeastern North 
America. Profess. Pap. U.S. Geol. Surv. 92: 1- 


23. BLATTER, E. 1919. Flora Arabica, Volume 1. 


Calcutta. 


3 


» 


d 


N 
-] 


. Boutros, L. 1977 


[o 


N 


w 
ON 


MaN A. T È 


bo 


Ww 


4 
e 


. BLOMBERY, A. M. 1967. A Guide to Panis Aus- 


tralian Mie Angus & aded 2: dne 

Bonpnov, E. G., V. L. KOMAROV, ae ET 
AL. 7. Ranunculaceae. ps E 539 in V. L. 
Komarov (editor), Flora of USSR, Volume 7. Publ. 
Acad. Sci. U.S.S.R., Moscow & Leningrad. [In 


n. 
. Borin, B. 1966. Énumération des plantes du 
4-596. 


Canada. Provancheria 6: 5 


. BOUGHEY, A. S. 1957. The uc of the African 
Flora. Oxford Univ. Press, Londor 


1965. Comparisons hus ^en the mon- 


tane jos of North America, Africa and Asia. Web 


bia i 507-517. 
: dicar on the flora dE Jordan. 
Candollca 32: 73-80, 98, 99-110, 2 


. BRAMWELL, D. & Z. aed 1974. The wild 


flowers of the Canary Islands. London & Burford. 


. BRAUN, E. L. 1955. The phytogeography of u 


slactated a United States and its inte Ee. 
tion. Bot. . 21: 297-315, 

BRITTON, i L The American species of 
the genus Avene and the genera which have 
been b dE to it. Ann. New York Acad. Sci 


.BRowN. 1943. An lustros Flora 


of the Northern United States, Canada 


British Possessions, Volume 2. Bene New York. 


. BROWN, A. & R. E. Massey. 1929. Flora of the 


Sudan. Marby & Co., London. 


. BRÜHL, P. 1895. Desc riptions of new and rare 


Indian plants. Some new or critical Ranunculaceae 
from India and adjacent regions. Ann. Roy. Bot. 
Gard. Calcutta 51: 71-114 


. Bupantsev, L. J. 1975. Istoriya Arkticheskoi 


Flory Epokhy Rannego Kainofita (The History of 
the Arctic Flora of the E poch of the Early Caino- 
phyta). Synopsis of The E for Doctor of Sci. 


. BURBIDGE, N. T. 1960. The phytoge rari 2 
8: 


the Australian region. Austral. J. Bot 
—— — & M. Gray. 1970. Flora of ihe 6 ye 


tralian Capital tos Austral. Nation. Univ. 


ss, Canberra 

N. OVCHINNIKOV. 1953. 

nunculaceae. Pp. 433-509 in E. P. Korovin (ed- 

je ur Flora of Uzbekistan. Publ. Acad. Sci. Uzbek 
, Tashkent. [In Russian.] 


; i MP, "W. H. 1947. Distribution patterns i in mod- 
l. 


ern plants and i ms of ancient dispersals. Eco 
59-183. 


Monogr. 17: 1 


. CAMPBELL, D. H. 1933. T flora of the Hawaiian 
4-1 


CHANG, CHI. Wu. 1982. Bu i cal study of 
three genera of the family Ranunculaceae in con- 
nection with their systematic position. Acta Phy- 
totax. Sin. 20: 402-409. 

CHANG, CHING WEIT & CHIANG SHUT. 

y on the vertical vegetation belt ol 
the Mt. Jolmo-Lungma (Everest) region and its re- 
lationship with horizontal zones. Acta Bot. Sin. 15: 

9 


. CHATER, A. O., T. G. Turin, B. PAWLOWSKI, ET 
1 


964. Ranunculaceae. Pp. 206-242 in T 
B. "Putin (editor), Flora Europaea, Volume 1. Cam- 
bridge Univ. Press, Cambridge. 


1042 Annals of the 
Missouri Botanical Garden 

45. CHEESEMAN, T. F. 1906. Manual of the New A taxonomic Dm Monogr. Bot. Volume 41. Publ. 
Zealand Flora. NW ligt, New Zealand. Polsk. Bot. Soe 

46. ala eat .K. 19 Sosudistye Rastenija 65. Dosrar, J. 1950. Kvetena CSR. Publ. Prir., 

R (V audae Plants of the USSR). Nauka, Len- Prague. 

iie [In Russian.] 66. DRUMMOND, J. R. & J. J. HurCHINSON. 1920. A 

47. CHEREPNIN, L. L 957. Osobennosty flory juga revision of /sopyrum (Ranunculaceae) and its near- 
Krasnojarskogo is (Pecoliarities of the Kras- er allies. Bull. vere Inf. Kew 15: 145-169. 
nojarsk flora). Ucen. Zap. Krasnojarsk. Gosud. Pe- 67. Du 962. Flora of Panama. uri 
dagog. Inst. 10: 3-11. [In CRM aceae. Ánn. Viam Bot. Gard. 49: 14 

48. CHIANG, SHUT. 1960. The meadows and forests 68. Duncan, T P. PEREZ. 1971. Chromosome 
of Alday mountains in Szechuan Province. Acta of icd >s (Ranunculaceae). Amer. J. Bot 
Bot. 9: 125-136. 989-¢ 

49. us V. E. 1976. Vysokogirna flora Ukrain- 69. DURALD, A H. 10. Sylloge Florae Congolanae. 
skykh Karpat (The high-mountain flora of the Bull. Jard. Bot. Nat Belgique 2: 13-15. 
Ukrainian Carpathians). Naukova dumka, Kiev. [In 70. EICHLER Revelin ied lacie 
Ukrainian. Malesiens. bu Bot. 124 

50. CHOWDHURI, P. K., P. H. Davis & M. boa 71. ENGLER, A. 1906. Ranunc iron dinge Bot. 
1958. Materials foh a flora of Turke Jahrb. 37: 100-408. 
pane 1. Not. Roy. Bot. Gard. Edinburgh 12. —— E Ranunculaceae Africanae. Bot. 

: 403-425. Jahrb. 45: 266-275. 

5l. ME N. E. G. 1979. The sng s of 73. EPLING, C. E H. Lewis. 1952. Increase of the 
Papua New Guinea. Bull. Alpine Gard. . Gr indi range of the genus Delphinium. Evolution 
rit. 47: 1-300. 6: 253-267 

52. Curtis, W. M. 1956. The Students’ Flora of 74. Ewan, J. 1945 As ynopsis of the North American 
Tasmania, Part 1. Government Printer, L. C. Shea, species of De Iphinium. M ^ Colorado Studies, Ser. 
Hobart, began , Phys. Sei. 2: 55- 

53. DARLINGTON, P. J., JR. 1965. Biogeography of 75. ExELL, A. W., J. D MILNE-REDHEAD. 


wa 


c 
~ 


nunc die 


gq 
^c 
CO 


. Davis, P. H. 1960. 


the aie end of the i Harvard Univ. Press, 


Cambridge. 


4. DAUBENMIRE, R. F. 1943. Vegetational zonation 
in the Rocky Mountains. Bot. Rev. 9: 325-393. 
. Davis, K. C. 1900. A taxonomic study of North 


American Ranunculaceae. thesis, Cornell 


Unive me Ithaca, New York. [See also Minn. Bot. 


Stud. 2 343. 1899. (Aquilegia), 345-352. 
1899. m. 431-457. 1900. (Delphin 


ium); 459-507. 1900. (Ranunculus); 509-523 

1900. (Thalictrum).] 

Materials for a flora of Tur- 

key. IV. Ranunculaceae. II. DEAN Not. Roy. 
Bot. Gard. poe 23: 103-16 

———, J. Cutten & M. E. Conor, 1965. Ra- 
|Fp. 95-203 in P. H. is (editor), 

of Turke >y and the East o bleus 9 Vol- 

iv. Press, Edinburgh. 

1901. Die ae von Zentral 

China. Bot. Jahrb. Syst. 29: 16 59. 

————. 1937. Beitrage zur d der Vege- 


Flora 


tation ind Flora von Bude Bibl. Bot. 116: 1 
ol 


60. 


62. 


63. 


~ 
o 
- 


. DOROSZEWSKA, A. E 


DOROFEEV, P. I. 1963.  Tretichnye flory Zapad- 
noy ou oe ruary pee of the Western Sibe ria). 
Publ. . U.S.S.R., Moscow & Leningrad. 


[In nn ] 
———. 1965. O nekotorykh problemakh istoryi 
flory (Coi oncerning some proble ms of Eid. history). 
Bot. Zurn. S scow & Leningrad) 50: 1509 : 
———. 1972. Tretichnye flory Pre sseyna reky 
cea Vien bis of re Omoloj River Basin). 
Pp. 41-112 in I Vasiljchenko (editor), History 
of "à Flora and Vegetation of Eurasia. Nauka, 
Leningrad. i Russian.] 
————. 19 skopaemye tsvetkovye rastenija 

SSR (Fo: M fous ring plants of the USSR). Nauka, 
D [In Russian.] 


974. The genus Trollius L. 


= 


ec 


co 
bo 


o 


7. E EDORON A.N. 1952. 


$ FISHER, 


. FORBE 


. Les espèces Africaines du genre Clematop- 
sis Boj. ex Hutch. Bull. Soc. Roy. Bot. Belgique 
83: 407 e 
—— E-REDHEAD & M. L. GONSALVES. 
Flora de Mie 4. ens ers 
Ultramar, Lisbon. 
Istorija v T fory 
chetvertichnoe vremja kak Į av- 
ios razvitija Tretichnoy flory (The jn 
of the high-mountain Caucasian flora in the Quar- 
ternary as an example of the autochthonous de- 
d daa of the Tertiary floristic base). Pp. 49 
n . Obruchev (editor), Materials on the 


Acad. S 


1972. 
Junta Invest. 


pe rnary in the USSR. Iss sue 3. Publ. 
Moscow. Ln Russ 
957. ne lies apela Kitaja i 


eje : znachenie dlja poz a Ev- 


a of the plant a 
of Eurasia). Pp. 24-50 in E. N. Pawlows B. 
A. Baranov puo Tus riedings. Issue 10 
Publ. Acad. Sci. U.S.S.R., Moscow & Leningrad. 
1971. Flora delle Alpi. A. Martello, 
F. J. F. 1965. The age e of 
New voe New Zealand Dept. Sci. Ind. Res. 
Bull. 165: 192 
H. HemsLeY. 1905. An 
ennmépation of all plants known ge China proper, 
Formosa, Hainan, Korea, the Luchu Archipelago 
and the islands of Hongkong, together with their 
Sind and synonymy, Volume 3. London. 
STER, A. S. & H. J. ARNOTT. 1960. Morphol- 
ogy and dic hetemiaus vasc pea of the leaf of 
se ded uniflora. Amer. J. : 684-698. 
19: 


3. Fos =G A Mos al the ferns 


p ro ming s plants of Bolivia. Contr. Gray Herb. 
184: 1 


Volume 76, Number 4 
1989 


Ziman & Keener 1043 
Geographical Analysis of Ranunculaceae 


eo 


v 


- 


ec 
ec 


© 


me 


ño 


. GOLOSKOKOV, 


B 


n 


E 


No) 


. FRANCHET, M. 1886. Plantae Yunnanensis. 
Ranunculaceae. "s Soc. Bot. France 33: 360- 
3 


. Plantae Davidianae ex Sinarum 
Imperiae. Ranunculaceae. Bull. Soc. Bot. France 


1 FRERE, M. V. 1938. Phytogeographical problems 
of eastern Canada. Amer. Midl. Naturalist 19: 489- 


. FurAK, J., L. BER . HUSAK, ET AL. 


82. 
Flor ke M 3. Veda, Bratislava. [In 


; Caney. V. D., H. G. KuLiEvA & Z. V. VAGABOV. 

1979. Flora i Rastitelinost; Vysokogoryi Talysha 
(The Flora and Vegetation of the High-mountain 
Talysh). Elm, Baku. [In Russian.] 
. GaricH, M., H. DikLicH & M. Jankovich. 1970. 
Ranunculaceae. Pp. 204-317 in M. Josifovich (ed- 
itor), Flora de la Republique Socialiste de Serbie, 
Volume 1. Nauchne Delo, Belgrade. 
. GALUSHKO, A. I. 1978. Flora Severnogo Kavkaza 
(The Flora of the North Caucasus). Publ. Rostov 
Univ., Rostov. [In Russian.] 
. GILI, A. 1954. Neu Ranunculaceae und Papav- 
o aus oe Feddes Repert. Spec. Nov. 

ni Veg. 57: 94-101. 
1964. Beitráge zur Flora Afghanistan. 
4. Ranales Ex Rhoeadales. a ddes Repert. Spec. 
Nov. Regni Veg. 69: 1 
. Gopwin, H. 1956 The T - the British 
Flora. Cambridge Univ. Press, Cambr 
cee L901 cido ete Pp. 
10-132 in N. Ma Pavlov (editor), Flora of Ka- 
zakhstan, Volume 4. Nauka, Alma-Ata. [In Rus- 
sian. 
. Goop, R. 1974. The Geography bs Flowering 
Plants, 4th edition. Vir Lon 
GORCHAKOWSKYI, P. 1960. Endemichnye i 
reliktovye element MN Urala i ikh proiskhoz- 
denye (Endemic x relict elements in the flora of 
the Urals and their origin). Pp. 66-68 in A. I. 
Tolmachev (editor), hao of Botany. Issue 3 
Publ. ru Sci. U.S.S.R., Moscow. [In Russian.] 
h e í Ra stein mir Vysokogornogo 
Urala (Plan word of the High-Mountain Urals). 
auka, Moscow. [In Russian. ] 

. GRAHAM, A. 1972. Outline of the origin and his- 
torical recognition of floristic affinities between Asia 


8 

ham (editor), Floristics and Paleofloristics of Asia 
and North America. Elsevier Publ., Amsterdam. 

. GREENE, E. L. 1897. Ranunculaceae monotypes. 

Pittonia 3: bs 195. 

0 909-1910. Canadian Durs of Tha- 
lictrum. a Naturalist 23: 17-19, 37-40, 24: 
25-3 


. GROSSHEIM, A. 1926. Flora Talysha (Flora of 
the Jalysh). Pu "Tests. Dept. Azerbaidjan, Tiflis. 
.] 


[In Russian 


: 1936. Analyz Flory Kavkaza ge 
of the Flos of the usu) Publ. Acad. Sci. 

U.S.S.R., Tiflis. [In Russ 

. 1950. Flora Kavkaza (Flora of the Cau- 
casus), e 4. Mezniereba, Tbilisi. [In Russian.] 

. GROVES, E. W. Vascular plant collections 


. GRUBOV, V. I. 


j; HARSHBERG ER, J. 1911. 


. HEPPER, P. N. 


from the Tristan da Cunha group | x oe Bull. 

Brit. Mus. (Nat. Hist.) Bot. 8: 

1955. s Flory a ne 

oy Narodnoy Respubliky (Synopsis of wi Flora of 

the Mongolia People's Republic). Publ. Acad. Sci. 
S.R., Moscow & Leningrad. [In usan. ] 

63. Rastenija Tsentraljnoy Azii (Plants 


l of Central Asia), Volume 1. Publ. Acad. Sci. 


U.S.S.R., Moscow & Leningrad. [In Russian.] 
976. Visit heed derivaty vo 
flore Tsentraljnoi Azii (Eastern Asiatic derivates in 


the flora of Central Asia). Bot. Zurn. (Moscow 


un 61: 32- td [In Russian. | 
8. 98 


Sosudistye rastenija Mongolii, s 
atlasom (V xim uis of ei with atlas). 
Nauka, ped [In Russian.] 
& A. N. Feporov. 1964. Flora i ras- 
titeljnostj Kitaja (Flora and vegetation of China). 
3 n V. T. Zaichi = a Pe 
Geography A China. Myslj, Moscow. [In R sian.] 


. GUILLAMIN, A. 194 Flore - ! ae ique 


t Sy- 
Que de la Nouvelle Calédonie os ene 


; Hin J. B. 1966. Biosystematics of the New 


meu r 1945-1964. New Zealand J. Bot. 


2. TER oe. H. 1939-1940. Plantae Si- 
EO 


nensis. Ranunculaceae. Acta Horti Gothob. 13 


3. Hana: H. 1975. New or noteworthy flowering 


a from eastern Himalaya. J. Jap. Bot. 50: 263- 
71. 


An d of the dM 


plants of Pur Publ, Brit. s. (Nat 
AR 


Es. Je Ho ho DSON & T A 
MAN- TERNIEZ. 1980. Contribution to the vascular 
flora of boreal Saskatchewan Canada. Rhodora 82: 
239-280 
Bo dr oe 
survey of North Ns Engelmann & Stecher 
Leipzig & New Yor 


7 


. HEDBERG, O. 1957. Afroalpine vascular epu 
aL B 


A taxonomic revision. Symb. Bot. Upsa 
411. 


1965. Afroalpine flora elements. Web- 
co 


519 O. 
. Heci, G. 1975. Illustrierte Flora von Mittel-Eu- 


opa, Volume 3(4). P. Parey, Berlin & Hamburg. 


r 
20. HEMSLEY, W. B. 1902. hos es 3 Tibet or High 


TA Linn. Soc., Bot. 35: 1-2 
1965. Pu account of the 
phyogeographical affinities of the flora of West 
cal Afric «Mu 19: 593-617 


22. B C. . & A. CRONQUIST. 1973. Flora 


of the Pacific d ue Univ. Washington Press, 
edd bu ton. 


. HOOKER, J. 1872. Ranunculaceae. /n: The 
Part 1. 


Flora of ee India, Volume 1, 


24. Hu, Hsen-Hsu. 1940. Constituents of the flora 


"i m Proc. Sixth Pacif. Sci. Congress 4: 641- 


UN E. 1960. Flora of the Aléutian Islands, 
Volume 1. Academic Press, New Yor 


1968. Flora of je and neighboring 


territories. Stanford, Californ 
27. HUTCHINSON, J. 


1920. lap a primitive 


1044 


Annals of the 
Missouri Botanical Garden 


128. 


pustynj ne Azii (Some results 


130. 


32. Jones, G. W. € € . FULLER. 


33. JoRDANC 


— 
Qu 
~ 


139, 


140. 


. JEN, 


. JUR ,B.A. 1968 


AL. 1979 


T. a sov, B. V. 


genus of Clematideae. Bull. Misc. Inf. Kew 1: 12- 
DI 


idi M. M. 1941. Tretichnye reliktovye ele- 
ienty v taeznol flore Sibiri i ikh vozmoznoe wed 
khoidenie (Tertiary relict elements in the taiga flora 
>p. 257-292 in LV 
aterials on the History of the 
lora and Vegetation of the USSR. Issue 1. Publ. 
S.R., Moscow & Leningrad. [In 
Russian.] 
—————. 1946. Nekotorye itogy mer flory 
of the 


Flora and Vegetation of the USSR 
J.S.S.R., Moscow & Leningrad. [In 


D R. & V. CsAPODY. 1975. Iconographia 
florae partis Austro-Orientalis Europae Centralis. 
Akad. Kiadó, Budapest. 
Hsu. 83. Late Cretaceous and Cenozoic 
vegetation in China. Emphasizing their connections 
with North America. Ann. Missouri Bot. Gar 
490-508. 
1955. Vascular 
Plants id Illinois. Usha, Illinois. - 
NOV, C. ULEV, ET AL. 1970 
Flora of Bulgaria. Publ Acad. Sci. Bulg., Sophia. 
[In ie crt an.] 
Flora Suntar-Khajata (Flora 
of the in -Khajata). Nauka, Leningrad. [In Rus- 


sian.] 
— — — . 1974. Problemy botanicheskoi geografii 
Severo- -Vostoc :hnoi Azii (Problems of botanical ge- 
Vade ih E Northeastern Asia). Nauka, Leningrad. 
[In Russian.] 
V. V. PETROVSKYI, A. A. KOROBKO, ET 
Obzor geograficheskogo rasprostrane- 
nija sosudisty kh i Chukotskoi tundry (Survey 
of tl ascular pi of the Chukotka 
mas Bjull. Moskovsk. Obsc 
14-83. [In Russian. 
1958. Tibet. Fiziko-geograficheskaja 
kharakteristika Physical and geographical char- 
acte ristics of Tibet). Publ. Geogr., Moscow. [In Rus 


:. Isp. Prir. Otd. Biol. 


— 


1947. Ge P and plant distribution. 
137 
( 1962. V m Provintsyi 
Severnogo Kis à; Osnovnye Cherty Flory i Rasti 
teljnosty, Lesorazvedenie (Principal "Pacübstities of 
the Flora and Vegetation, and Forest € esi 
in the Loess Province of C ved Publ. Acad. 
U.S.S.R., Moscow, ue Russian. ] 
KAMELIN, R. V. 1973. iden Analyz 
;stestvennoy Flory Gornoi Srednei Azii (Floroge- 
netic Analysis of the Natural Flora of Mountain 


Middle A Dr Leningrad. [In Russian.] 
l. ——— . 


Kukhistanskyi Okrug Gornoi 
Srednei T: Botako Geograficheskyi Analyz (The 
Kukhistan District of Mountain Middle Asia. The 


Botanical-( EA UM Analysis). Nauka, Lenin- 
grad. [In an.] 
142. Os AEV, M. N. 1958. Konspekt flory Jakutii 
(C /onspectus of Jacutian flora). Publ. Acad. Sci. 
J.S.S.R., Moscow & ni MEL [In Russian. 
143. KARMISHEVA, N. "K. 973. Flora i rastiteljnostj 


— 
on 
w 


-— 
e 
on 


156. 


l 


wn 


. KEENER, C. S. 1967. 


laceae of dis southeastern United States. I. 


. KEM 


tovykh demens flory na severe C 


. KITAMURA, S. 


y nie ae Aksu-Dzabagly (Flora and vegetation 
the Reservation Aksu-Dzabagly). Nauka, Alma- 
Russian.] 
A biosystematic study of 
Clematis subsec tion Mi dL PN eae). 
J. Elisha n itchell S 9! 
—— —.,. 1975 1977 7. Studies in oe Ranuncu- 
Anem 
one L. Castanea 40: 36-46. Il. Thalictrum L. 
Rhodora 18: 457-472. MI. Clematis L. Sida 
33-47. IV. Genera with zygomorphic flowers. Cas as- 
tanea an 12-20. V. Ranunculus L. Sida 6: 266- 
283. VI. Miscellaneous genera. Sida 7: 1-12. 
——— Distribution and biohistory of the 
endemic flora of the Mid-Appalachian shale barrens. 
Bot. Rev. 49: 65-115. 
W 1982. 


DENNIS. The subgeneric 


BAR us of Clematis (Ranunculaceae) i in tem- 


perate North America north of Mexico. Taxon 31: 
Slain 


Hoor. 1987. 


Ranunc ulus sec- 


tion Echine lla (Ranunculaceae) in the southeastern 
7-68. 


es. Sida 12: 5 

ARIA- NATADZE, L. M. 1960. Ranalievye 
na Kav kiss i ikh taksonomija (Ranales in the Cau- 
casus and Det taxonomy). Mezniereba, Tbilisi. [In 
Russian.] 


s State 


1973. Ranunculaceae. Pp. 20-157 in 
a N. Kesldhoveli (editor), Flora of Georgia, Volume 
. Mezniereba, Tbilisi. [In Georgian.] 
& 


: debe: S. S, T. G. Busu. 1976 


Sos 
distye Rastenija Severnoi Korjakii (Vasc ular inn 
of North Korjakia). Bot. Zurn. (Moscow & Ler 
ingrad) 61: 1089 uo [In Russian.] 

— & Osnovnye cherty vy- 


sokogornoi flory ecd Korjakii (Pri incipal fea- 


= 


the high-mountain flora of North Korjakia). 


(editors), Problems of uh Volume 14. Nauka, 


Novosibirsk. [In Russian.] 

. KHOKHRJAKOV, A. 1978. e basseyna 
srednego techenija Reky Omolon (Co E the 
flora of the basin of the middle stream of the River 


Flora and Vegetation of Chukotka. Publ. Acad. Sci. 
S.S.R., V cio [In Russian.] 

Ubezi$éóa mezophyljnykh relik- 
hukotskogo po- 
v basseyne verkhnego techenija Kolymy 
(Refugiun ms of the enr relict floristic ele- 
ments to the north of the Okhotsk Sea coast and 
in the yi o the Kolyma upper part). Bjull. Mos- 
kovsk. *. Isp. Prir. Otd. Biol. 84: 84-96. [In 


en | 


bereza 


1954. New 

collected by Sasuke Nakao, 

Himalayan ae II. 
5: 129-13 

KLEoPov, J. D 


species from Nepal 
: did of the Japan 
a Phytotax. Geobot. 


Osnovnye cherty razvitija 


flory shirokolistvennykh lesov Evropeyskoi chasty 
SSSR (General characters of a A of the 
broad-leaved forests of the European [ of the 
USSR). Pp. 183-256 in omarov (editor), 


Materials on the History of the F lora and Vegetation 
of the USSR. Issue 1. Publ. Acad. Sci. U.S.S.R., 
Moscow. [In Hissin] 


KoLAKOVSKYI, A. A. 1939. Flora Abkhazii (Flora 


Volume 76, Number 4 


Ziman & Keen 1045 
Geographical Analysis of Ranunculaceae 


golii duction to the flor 
y Gla 


163. 


165. 


166. 


72. KUMINOVA, 


. Kom 


: onus J. 197 


2. KOROVIN > 


. KoZEVNIKOv, J. P 


. Knyrov, G. V. 


] Kystirorovit H, A. N. 38. 


. KUMAZAWA, M. 


4. KURENTSOVA, C. e 1965. 


of Abkhaz), Volume 2. Publ. Abkhaz Sci. Research 
Inst., Mor: . [In Rasaan 

i V. L. 1903. Flora Manc p (Flora 
of Manchuria). Trudy Glavn. Bot. Sada 22: 1-196. 
[In E 
——. 1908. Vvedenye k floram Kitaja i Mon- 
as of China and Mon- 
golia). m. Bot. Sada 29: 1-388. 
Flora Poluostrova Vu 


. —— 51. 
(The Flora of the Kamchatka Peninsula), Volum 


. Publ. Acad. Sci. U.S.S.R., Moscow. [In Russian] 
Corresponding taxa and their 
ecological background in the a of temperate 
surasia and North America. Pp. 37- D. H. 
V M vente Taxonomy, Phytogeography and 
Evolut :ademic Press, New York. 
N., A. BAKHTEEV, M. T. TADJITDINOV 
& B. SARYBAEV. 1982. Illustrirovannyi Oprede 
litelj Vysshykh Rastenyi Karakalpaki i liue 
(Illustrated Manual of Higher Plants of Karakal- 
pakia and Kho Pe Fan, Tashke ent. [In Poe 
KOVALEVSKAJA, S. S., M. G. PAKHOMOVA x S: 
SARKISOVA. 1972. Ranunculaceae. Pp. 1 oo 
in O. N. Bondarenko & M Nabiev s s), 
Opredelitelj Rastenyi Srednei Azii (Manual of Plants 
of Middle Asia). Fan, Tashkent. [In Russian.] 
Spok sosudistykh ras- 
tenyi Chukotky (A list of the vascular plants of 
Chukotka). M eR a Vysivkh Rastenyi 18: 
230-247. [In Russi 
Kral, R. 1976 p treatment of A m for 
Alabama and Tennessee. Sida 6 
1 V a Flora 


——— 977. Tretichnye relikty vo n T 
vinskoi ASSR lar de relicts in the flor 
Tuva ASSR). Pp. 4-14 T. Krasnoborov (ed- 
itor), Vegetation al the hon of the Upper Jenisey. 
Nauka, Nov osibirsk. [In Russian.] 

sa Zapadnol Sibiri m 
ests of Western Siberia). Publ. Acad. Sci. U.S 
Moscow. [In Russian.] 

1 


of West Siberia), "Volume 4 


Istorija flory Pa- 
learktiky v techenie Neogena na a alo 
botanicheskykh dannykh (History of the flora of 
Palearctica during the Neogene on the is p the 
paleobotanical data). Pp. 8-9 in A. Kryshto- 
fovich SW apa s Relicts in the Flora of 
the USSR, Volur : _ Scl. USS. Re 
Moscow 5 us [In ES n.] 

19 Pollen i grain d seus 
Lasdésslialaceda and Berber 
daceae. J. Jap. Bot. 8: 19-46 
A. V. 1960. Rastiteh nyi pokrov Al. 
taya (V 2m cover of the Altai). Publ. Acad. 
Sci. U.S.5 » Novosibirsk. [In Russian.] 

Rastitelinyi pokrov Khakassii 


l (V ege tational cover of Khakassia). Nauka, Novo 


Russian. | 
Rastitelinyi pokrov 
Priussuriyskoi Chasty Basseyna Srednego Amura 


(Flora and a of the Ussuri Part of the 


~ 
n 


- 
[oo 
N 


190. 


193. 


5: RU E 
o Gor 


. LAM, 
the island of New Guinea. 
8. ———— 945. 


geography i Celebes. Blumea 
. LAUENER 


. Lr, Hur- 


5. LINCOLN, C. 
Q 
. LOUR 
america oco Darwiniana 9: 397-608 
; Mac BRIDE, 
Tokyo. 

. Maire, H. 


2. MAK 


s ep Middle Amur). Publ. wed EU Acad. 
Vladivostok. [In Rus 

. Vysotnoe goes 
Rastenyi v v Gorakh Putorana (High-Altitude Distri- 
bution of Plants in the Putorana Mountains). Nauka, 
Leningrad. Un Russian. ] 
——. 1985. Kholodnye goljtsevye Pustyni v 
akh Severnogo polusharija {€ zold 
High-Mountain Deserts in the Arctic Mountains of 
e Northern. Hemisphere). Nauka, Moscow. [In 
Russian.] 

1934. Materials saaw a study of 
Blumea 1: 115-159. 
Notes on the preda al phyto- 
5: 600-638. 

. Ae re in eum Not. 
Roy. Bot. Card Edinburgh 2 


New species E records of Ác- 


PA of Nepal Not. Roy. Bot. Gard. Edinburgh 
26: 


—— —— QU M. Tamura. 1978. A Synopsis of 
Ac onitum subgenus ges onitum. Not. Roy. Bot 
nburgh 37: 113-124. 

E ee Lesnye reliktovye (tre 
tichny de Cty mezdu Karpatamy i Altaem (Forest 
relict (Tertiary) centers between the Carpathians 
and Altai). Zurn. Russk. Bot. Obsé. 15: 351 


1885. nce du genre 
Thalictrum Bull. Soc. Bot. Belgique 24: 78-324. 
952. Floristic 
eastern Asi sia and eastern dept America. 
Amer. Philos. Soc. 42: 371-429. 
963. ees al relations in the 
herbaceous flora of the Pacific Coast of North and 
South awe rica. are and historical review. 
uart. Rev. Biol. 09-116 

RTEIG, T Ranunc ulace ‘eas de Sud- 


: relations us between 


Trans. 


a puer de Hope 
tropic ale. a ee ^i. Nat. La Salle 16: 40-42. 
1963. Flora del = ruguay. ae 

. Montevideo. 
J. F. 1936. 
Field Mus. Nat. 


Flora of Peru. Ranun- 
culaceae. Hist., Bot. Ser. 357: 
639-661. 

MaEKAWA, F. 1974. Origin and characteristics 
of Japan's flora. Pp. 33-36 in M. Numata (editor), 
The Flora and Vegetation of Japan. Kodansha Ltd., 


1964. Flore de 
Maroc, Algerie, Tunisie, Tripolitanie, 
et Sahar. Bonon cin: Volume 11. 
ee 


l'Afrique du Nord, 
Cyrenaique 
Lechevalier, 


MHEK, F. 1959. Reljef Zemly. icut re- 
la gnis rima opysanija poverkh- 
nosty Zemly (Relief of the earth. The Method of 
the Re gional Morphological Desc ‘ription of the Sur- 
face of the Earth). Publ. Foreign Liter., Moscow. 
[In Russian. ] 
Martev, V. P. 
Zapadnogo Kavkaza 10 a 

noi istorii ego flory 1 rastiteljnosty (Tertiary 
in 1 the flora of the weste 


1941. Tretichnye re likty vo flore 


Materials on the History of the Flora and Vegetation 


Annals of the 
Missouri Botanical Garden 


of the USSR. Issue l. Publ. Acad. Sci. U.S.S.R., 


. MALYSHEV, L. I. 1965. oo flora Vos- 

tochnogo Sayana (High-mountain flora of the east- 
ern Sayan). Nauka, Moscow. [In Russian 

. ——. 1976. Genesis vysokogornykh flor Sibiri 

(Origin of the high-mountain floras of Siberia). Izv. 

Sibirsk. Otd. / Akad. Nauk SSSR, Ser. Biol. 10: 45- 


. MARK, A.F. & N. M. pe 1973. New Zealand 
Alpine a A. H. & A. W. Reed, Wellington, 


New Zealand. 


7. MARTINENKO, V. A. 1976. Ranunculaceae. Pp. 


40 in A. I. Tolmachev (editor), Flora yide 
vostoka evropeiskoy chasty SSSR (Flora the 
Dig a io Para part of i USSR), E 

. Nauka, Leningrad. [In Russia 
. Monos ICH, C. Flora Tangutica, Vol- 
21. St. Petersbin rg. 

. 890. Plantae oa Dg i 
ceae. Prode Glavn. Bot. Sada 11: 4. [In Rus- 
sian. 

i M EIKLE, R. D. 1954. A survey z ^us flora of 

ipe ie Misc. Inf. Kew 1: 85- 

. MELV ,R. I i EN continental d and plant 

deum Pp. 4 446 in D. H. Tarling (editor), 

i ge of d Drift to the Earth Sci- 

ences, Volume |. Academic Press, London. 


v 


2. vta E.D. 1910. The Malayan, Australasian 


and Polynesian elements in the re flora. 
npe UN "i Buitenzorg, Ser. 2, Suppl. 3, part 
1:2 
3. ——— An Enumeration of Philippine 
Flowering Plants. Vols. 1- anunculaceae, Vol. 
2.] Bureau of Printing Manila. [Reprint, 
A. Achar reer 
———. 1936. Malaysian  phytogeography i in re- 
lation to ‘the Polynesian flora. Pp. 247-261 in T 
N. rro Pu (editor), Essays in Geobotany i in Hon- 
or of W. A. Setchell. Berkeley, California 


o) 
T 


5. MIEHE, C. 1987. An annotated list of cil 


plants collected in the valleys mer of AME Everest. 
Bull. Brit. Mus. (Nat. Hist.) Bot. 05-265. 

. MiNE-REbpHEAD, E. € W. B. oan 1952. 
of Tropical East Africa. Ranunculaceae. Lon- 


Ta MIS. B. A. 1953. Flora Khibinskykh Gor, 


eje analyz i istorija (Flora of the Khibin e 
its analysis and history). Publ. Ac ie Sei. U.S 
Moscow & a ite Russ 

. MIYABE, K. 1943 the species = aad in 
Japan. Acta Payot [m 
——— & M. Tat 1955. p 
to the flora of Norther Tn m Trans. Sapporo Nat. 
Hist M 14: 1-10, 69-86. 


. Moore, D. M. 1968. The js flora of the 


Falken Islands. Brit. Antarct. Su v. Rep. Sci. 60 


._ —. . Connections between cool ten 
perate floras wi pu ular reference to BE E 
South America. Pp. 1 38 in D. H. Valenti 
(editor), Taxonomy, Pus ography and Evolu- 
tion. ii nic Press, New York. 
— 74. Catalogo de las plantas vascu- 
E de Tierra del Fuego. Anales Inst. Patagonia 
: 105-121. 


3. Mu INZ, P. A. 1967-1968. A synopsis of the Asiat- 


. Paczoskyl, J. 1927. 


3. Dou. E. J. 


24. PANFILOV, D. V 


; E M N. 1935. 


. Porozvi, A. V., 


234. POLUNIN, N. 195 


o 


ic species of uo eiie T T Arbor. 48: 
476-545; 49: 166, 23 

Ca a "UE Flora. 

Univ. California Press, Berkeley & Los Angeles. 

AKAI, T. 1930. I flora of Bonin Islands. Bull. 


€ 


Biogeogr. Soc. Japan. 1: 249-278. 
is 


. OHM : M ra of Japan. Smithsonian Institution, 


Washin 


. pis IVER, d 1868. Flora of Tropical Africa, Vol- 


e l. Lor 


e adon. 
; A e P. N. 1971. ee 


Uscelja Varzob (Flora and Vegetation of the Va 


Ravine). auka, Leningrad. [In Russian. ] 
J), | ———— 


JORENKO, I. F. SHIBKOVA, ET > 

Ranunculaceae. Pp. 10-151 in P. N. 
chinnikov (editor), Flora Tadjikistana, TR és 
Nau n Leningrad. In Russian. 


a 
UN 
= 


Ranunculaceae. Pp. 5-69 


in W. eie (editor), Flora Seny Tom 3. Publ. 


Polon. en Cracow. [In Polish. ] 
. Par IAN. 19: 


A survey and collection 
of fores plants in Kansu and Tsing Hai. J. Bot. 
0: 


culaceae (Alexeya vvedenskyi Pachom. - 
nus and species of the Ranunculaceae). Bot. Mater. 


Pide Inst. Bot. Akad. Nauk Uzbek. SSR. [In Rus- 


A. STEYERMARK. 1935. An 
annotated onn of the a. Pec A Mis- 
souri. Ann. Missouri Bot. Gard. 9-54 

1964. oe dus uds Kitsi. Pp. 
593-600 in V. T. Zaychikov (editor), Physical 
oe e China. Myslj, Moscow. [In Russian.] 
PAR 951. Flora d'Iran (La Perse). Tehran. 

lora Zentraljnogo Ka- 

zakhstana (Flora of Central Kazakhstan), vole 
2. Publ. Acad. Sci. U.S.S.R. Moscow. [In Russian. ] 


27. PavLov, V. N. 1974. Osobennosty flory ou 


nogo Tjanj-Shanja p gp oft Y ora of West 
Tien Shan). Probl. : 63 In Russian.] 


. Payson, E. B. a ao Nort American — 
133 


- ee Contr. U.S. Natl. Herb. 


: ARSON, R. Climate and Evolution. Academic 
/ Yor 


P . 
. PESHKOVA, G. A. 1972. Stepnaja flora Bajkaljskoi 


ane iene es of Baikal Siberia). Nauka, Mos- 
w. [In Russ 


979, Ranunculaceae. Pp. -375 


. ——. | 
in L. I. Mal yshev & G. A. de Flora 


Tsentraljnoi Sibiri, Volume 1. Nauka, Novosibirsk. 
[In Russian. ] 
>. MALYSHEVA & V. A. SMIRNOVA. 
1976. Analyz ^n ostrovnykh Prieniseyskykh st 
pei (Analysis of the flora of the steppe Nn of 
Prienisey). Bot. Zurn. (Leningrad & Moscow) 61: 
910-925. [In Russian. ] 
& VJAKINA. 1979. Priledniko- 
a flora Katunskogo F (Periglacial flora of 
he ‘Katun ge). 8-23 in A. I. Tolmachev 
& I. M. rino res Problems of Botany, 
Volume 14. Nau LINE: [In Russian.] 
Circumpolar Arctic Flora. 
C larendon, € Odo. 


. Popov, M.G. 1927. Osnovnye cherty istorii raz- 


Volume 76, Number 4 
1989 


Ziman & Keener 


1047 


Geographical Analysis of Ranunculaceae 


236. 


240. 


299. 


. Posr 


. RADFORD, A. E., 


. RapParcs, R. 


2. E ssEN, H. 1979 


D. continental movements. 
rd. 61: 53 73. 


C alifornia flora. Univ. 
13 


vitija flory Srednei Azii (General features of the 
history of development of the Middle Asiatic flora). 
I redne-Aziatsk. Univ. 15: 239-292. [In Rus- 
sian. | 

—————. 1977. Osobennosty flory Daljnego Vos- 
toka sravniteline s ev ropeiskoy (Peculiarities of the 
Far Eastern flora in comparison with the European). 
Fan, Tashkent. [In Russian.] 

¿ J. E. DINSMORE. o 
Palestine e Sinai, Volume 1. 
QUESEL, 1978. Analysis of he flora of Med- 
e and Saharan Africa. Ann. Missouri Bot. 
Gard. 65: 479-534. 

. E. AHLES & C. R. BELL. 1968. 
Manual of the Vascular Flora of the € oe Univ. 
North Carolina Press, Chapel Hill, North Carolina. 
RAMENSKAJA, M. L. & V. N. ANDREEVA. 
Marron Mer d Rastenyi Murmanskoy Ob- 
lasty i Karelii (Manual of Higher Plants of the 
s District and Karelia). Nauka, Leningrad. 
[In Russian.] 


Flora of Syria, 
u 


1907. A sisak virágnemzetség ren- 
Po eur. Aconiti generis. Nóvényt. Kózlem. 
6: 137-176. [In Hungarian.] 
as genus Knowltonia 
(Ranunculaceae). Opera Bot. 53: 1-44. 
Rau, M. A. 1975. High- RS flowering plants 
India 3: 1 


is West Himalaya. Bull. Bot. Surv. 
34. 
——. 1981. A review of the Indian. zm 
m Bull. Bot. Surv. India 23: 213-216. 
. Raup, H. M. 1941. eeu a in Boreal 
America. Bot. Rev. 7: 1 
Raven, P. H. 


bd relations in 
the foras a North and South America. Quart. Rev. 
Biol. 38: 151-177 


Plant zy 


2 


: in um 

mary. Ann. Missouri Bot. Ga is 59 234-246. 
. ——. 1973. Evolution of o and alpine 
nd. New Zealand J. Bot. 


md groups in New Zeala 
: 177-200. 


. AXELROD. 1972. Plate tectonics 
and Australasian paleobiogeography. Science 176: 
1379-1386 

Angiosperm biogeography and 
Ann. Missouri Bot. 
e Origin and relationships of the 


California Publ. Bot. 72: 1- 


Ru .J. 1978. Clematopsis, genre africano- 
malgache: types biologique et taxonomie. Adansonia 
8: 3-19. 


. E. Rew. 1907. The fossil flora 
of Tegelen Sur- ‘Meuse near Venico, in the province 
of Limburg. Verh. Kon. Ned. Akad. Wetensch., 
Afd. a me Sect. I3: 3-33. 

; A further investigation of the 
Pliocene E of Tegelen. Wersl. Gew. Vergad. 
Natur. Afd. Kon. Akad. Wetensch. 19: 192-200. 

, M. E. CHANDLER & T. GRovEs. 1926. 


. REVERDATTO, B. B 


The Bembridge Flora, Volume 1. Brit. Mus. (Nat. 


Hist.), London. 

1934. Lednikovye relikty vo 
flore Khakasskykh stepey (Glacial relicts in the 
p Pus flora). Trudy Tomsk. Gosu 
Univ. 86: In Russian.] 


257. RIEDL, H. New taxa and combinations in 
line from a and Kashmir. Bull. 
Misc. Inf. Kew 34: 30 

258. Runrsov, N. I. & L. A. Sd ALOVA. 19064. Flora 
Kryma 1 eje geograficheskye svjazi (Flora of the 
Crimea and its geograp d connections). Trudy 
Gosud. Nikitsk. Bot. Sada 3 6. [In Russian.] 

259. RYDBERG, P. A. Flara P E Rocky Moun- 
tains and Adjacent Plains. Hafner Publ. Co., New 
York. 

260. RzazapE, R. Y. & J. J. KARJAGIN. 1953. Flora 
Agor hadjana (Elara of Azerbajdjan), Volume 4. Publ. 

x Sci. Azerb. S.S.R., Baku. [In Russian. ] 

261. SAHNI, R. E. pi Endemic, relict, primitive 
and NUT r tax eastern Himalayan is 
and strategies for ibeir conservation. Ind. J. Fore 
2: 181-190. 

262. SANTAPAU, H. 1973. The flora of the rae "n 
a ior Ind. Nat. Sci. Acad. 45 

203. SCHAR ER 19 ‘Biographien y von E 
spp Springer, Vieni 

264. ScHIFFNER, V. 189 bo mH hellebororum. 
Nova ihe Leop. 56: 198. 

265. ScHOFIELD, W. B. en of 
northwestern North A Bryapby 's and vas 

cular plants. xs o 20: 155-207. 

266. SEDELJNIK P. 1972. ultra i geneti- 
cheskye svjazi b flory Kuznetskogo Ala- 
tau TRAC and genetic relations of the high- 
mountain flora of the Enc Alatau). Bot. Zurn. 
(Moscow & Leningrad) 62: 644-653. [In Russian.] 

267. SERGIEVSKAJA, L. P. 1967. Flora Zabajkalja (Flora 
of Tzansbaical), Volume 2. Publ. Tomsk Univ., 
Tomsk. [In Eu ] 

268. SEWARD, A. C. 1933. Plant Life Through the 
Ages. Cambridge D Press, Londo 

269. SHALYT, M. S. 1972. Ranunc e Pp. 

174 in M. S. Shalyt deu. ), Opredelitelj V a kh 
Rastenyi Kryma (Manua Higher Plants of the 
Crimea). Nauka, Leningrad [In Russian.] 

270. SHIPCHINSK YI, S. 24. O geografiche »skom 
ra sprostranenii vydov me Trollius i o genetiches- 
koi ikh svjaz (€ /oncerning the geographical distri- 
bution of the species of the genus Trollius and their 
genetic oq Izv. Glavn. Bot. Sada R.S.F.S.R 

2: 4. [In Russian.] 

271. SIEBOLD, P. F. & G. G. Zuccarini. 1843. Flora 
Japonic a, Volume 3. Zuccarini, Leiden. 

272. SKALINSKA, M. 1940. pein Aa Proc. 
Linn. Soc. London 152: 328 

213. SMILEY, F. J. 1 d on oe boreal flora 
of ne Ja Nevada of California. Univ. Calif. Publ. 
Bot. 9: 1-423. 

274. SMIT, E C. 1951. The vegetation and flora of 
Fiji. Sci. eren 73: 3-15. 

275. SMITH, P. 73. A revision of Caltha (Ranun- 
ene une 21: 119-150. 

276. SmITH, W. 1913. p alpine and rada 
vegetation a Southeast Sikkim. Rec. Bot. rv. 
India 4: 43 

277. a T. T. “SHIM Mizu, ET AL. 1970. Con- 
tribution to the flora of Southeast ds: South East 
Asian Stud. 8: 171-186. 

278. SOBOLEVSKAJA, K. A. 1953. Konspekt flory Tuvy 
Ud of hue Tuva Flora). Publ. Acad. Sci 

S.R., osibirsk. [In Russian. ] 
279, al v. n Voprosy florogeneza 1 


1048 


Annals of the 
Missouri Botanical Garden 


filotsenogeneza Manchzurskogo smeshannogo lesa 
(Questions of the formation of the flora and vege- 
tation of the Manchurian mixed forests). Pp. 283- 
320 in V. L. Komarov (editor), Materials on the 
History of the Flora and Vegetation of the 


Issue 2. Publ. Acad. Sci. U.S.S.R., Mostów: [In 
ussian.] 

280. Soó, R. 1980. A Magyar Flóra. Kót. 6. Akad. 
Kiado, Budapest. 

281. SrANDLEY, P. C. 1937. Flora of Costa Rica. Ra- 
oo Field Mus. Nat. Hist., Bot. Ser. 18: 

34 

282. STANER, 939. Revision des durar in 
a oe Belge. Bull. Jard. Bot. Etat 15: 307- 

283. E O. 1905. The aconites of India. A Ps 
oe Ann. Roy. Bot. Gard. (Calcutta) 10: 

284. —— ———. 1907. oo gymnandrum. Bot. 
133: oo 114. 

285. STEAR E J. Wittiams. 1978 
PARTE e" fe flowering rn of Nepal. mm 
Brit. iu Nat. Hist. l 

280. STEBBIN 4 AJOR. 1965. i ipn 
and spec ‘tion in nthe C alifornia flora. Ecol. Monog 

35: n 

287. ane NIS, C. C. C. J. VAN. ndo m a bridge 
theory i in botany. Blumea 11: 

288. ————. 1962. The ao ora a the Ma- 
laysia tropics. Endeavour 21: 9: 

289. ———. 1967. es en of p Kinabalu flora. 
Malayan Nat. J. 20: 

290. STEYERMARK, J. A. Ton p of Missouri. Iowa 
State Univ. Press, Ames, lowa. 

29]. SZAFER, W., ULCZYNSKY PAWLOWSKI. 
1924. Rosliny Polskie. Ksiaznica- Atlas, L'vov & 
Warsaw. [In Polish. ] 

292. TACKHOLM, V. 1974. Students’ Flora of Egypt. 
Publ. ene Univ., Beirut 

293. TAKHTAJAN, A. I 1946. 


294, 


2960. 


297. 


208. 


299, 


300. 


2l = 17: 4 


K istorii razvitija ras- 
titeljnosty eu (Concerning the history of the 
development of the vegetation of uar Izv. 
iie Fil. Akad. Nauk SSSR 4: 51-107. [In 
Russ 

of Ar- 
Acad. Sci. Armen. SSR, 


Flora Armenii (Flora 


Publ. 


menia), Volume l. 


Erevan. m Russian.] 
5 is 


7. K voprosu o proiskhozdenyi 
umerennoi flory Evr razii (Con -oncerning the question 
of the origin of the temperate flora of Eurasia). Bot. 
i el ad & Moscow) 42: 1635-1653. [In 
Russian. | 
———. 1970. Proiskhozdenye i Rasselenye 
Tsvetkovykh Rastenyi (Origin and Dispersal of the 
Flowering per N 
—— 978. skye oblasty 
(Floristic "e di of the World). Nauka, Leningrad. 
[In Russian.] 

——— 87 'stema Magnoliofitov oe 
of the us Nauka, Leningrad. [In Rus- 
n.] 
TAM MURA, M. 1966-1968. Morphology, Ei 
and phylogeny i Sai Ranunculaceae. 
Repts. Osaka 14: 27-48; 15: 13- ab Ss 
v 


1968. Ranunculaceae of north western 


Nepal ede by K. Nishioka and O. ra 


in 1958. Acta Phytotax. Geobot. 23: 28-: 
08. 

301. ———. 1970. Archiclematis, a precursory Ad 
nus of Cle matis. Acta Phytotax. Geobot. 24: 146 
152 

302. ———— Ranunculaceae of Thailand. J. 
CRX Taxon. 28: 14-16. 

303. ——— & L. A. LAUENER. 1979. A synopsis of 
Aconitum subgenus L: pease II. Not. Roy. Bot 
Gard. Edinburgh 37: 6. 

304. TANAI, 1972 Tertiary history of vegetation 
in Japan. Pp. 235-278 in M. A. Graham (editor), 
Floristics and Paleofloristics of Asia and Eastern 
North America. Elsevier Publ. Co., Amsterdam 

305. THOMPSON, 977. The vascular flora of 


308. 


309, 


a the flora of the Island Sakhalin). Pp. 


flory 


pi 
ca 


7. ena 


. TOLMACHEV, 


m j "LA A. GRAY. 1838. 


Lost Pari Newton County, Arkansas. Castanea 
42: 61-94. 

— Kuyt. 1976. Montane and su 
alpine plants of the Sweetgrass Hills, Montana, i 
their relation to early postglacial environments o 
AH norther rn Great Plains. Canad. Field-Naturalist 

2 
R. F. 1973. Major disjunctions in the 
ge eographic ranges of seed plants. Quart. Rev. Biol. 
47: 365-416. 


rui p. A. O lesnoi faze v y poslelednikovoi 


. Komarov (editor) 
istry of the Flora oe A ees of the USSR. 


Issue 1. Publ. Acad. S.R., Moscow. [In 
Russian. 

ThE. S. A. 1977. K flore juznogo e 
tinennogo stepnogo rajona Tuvinskoi ASSR (Co 


o de flora. of the south desert-steppe da 
the ^ P 5-32 inl K 


. M. Krasno- 
ix etn, od ie of the Basin of 
Upper ey. Nauka, Nov rsk. [In Russian.] 


A. I. 1958. En pus rue nm os- 
novnykh elementov vysokogornykh flor severnogo 
polusharija (Concerning the origin of the main ele- 
ments of high-mountain floras of the Northern 
Hemisphere). Pp. -300 in V. N. Sukachev 
(editor), Materials on the History of the Flora and 
em » hs USSR. Issue 4. Publ. Acad. Sci. 
w. [In Russian.] 

1959. O flore ostrova Sakhalin (C T 
E P Hw (editor), Komarovs Rea 
ings. oa Acad. U.S.S.R., Moscow & Ler 
ingrad. [In Rican] 
—— ———. 1960. O gesqe si arkticheskoi 
(Qussgons of the or f the arctic flora). 
Vopr. Botany 3: 72-7 
— VV 


Ranunculaceae. 
(editor), Arctic Flora of USSR, Voluts 6. Nauka, 


Leningrad. (In Russian.] 
4. 


——— 74. Vvedenye v Geofragiju Rastenyi 
(Introduction to Plant Geography. Publ. Leningrad 
igrad [In fus ssian. 


= 


Flora of North 
ait Volume 1. Wiley & Putnam, New York. 


Volume 76, Number 4 


1989 


Ziman & Keener 


1049 


Geographical Analysis of Ranunculaceae 


Ww 
= 
kel 


>. Tsur, You-Wan. 
. TURRILL, 
. TWEEDIE, E M. 


. VASILJCHENKO, 


196 
3. Meis O. D. 


5. Mpeg E N., 


. VOROSHILOV, 


Russian.] 
gu ==, 


1958. A phytogeographical sur- 
vey of North-West Sechuan and region of Changtu 
District. Acta Geogr. Sinica 24: 174-188. 

: . The Plant Life of the 
alkan Peninsula A Phytogeographical Study. 
Clarendon Pres Es rd. 

976. Habitats and check-list of 
plants on ihe po side of Mount Elgon. Bull. 
Mis. Inf. Kew 27-25 

T J., M. M. Ivanova & A. K. 
KIsELEVA. 1978. Obzor vydov ip as rastenyi 
Baikaljskogo Zapovednika (Survey « 


4€ hkova (ed 
itor), Flora of Pribaikalj. Nauka, Novosibirsk. [In 


1956. Botaniko-geograficheskoe 
rajonirovanie Vostochnoy Sibiri (Botanical-geo- 
graphical division into districts of eastern Siberia). 
Učen. M eningradsk. Gosud. Pedagog. Inst. 
116: 61-102. [In Russia n.] 

roiskhoždenye flory 1 rasti- 


oo 1958 
teljnosty Daljnego Vostoka i Vostochnoy Sibiri (Or- 


hodky 
the Künashir - Island). Novosty Syst. Vyschykh Ras- 
tenyi 16: -187. [In Russian 1.] 

1953. Ranunculaceae. Pp. 14- 
. K. Zerov dd Flora Ukraine SSR 


152 . 

Pub jm Sci. Ukr. 5 , Kiev. [In Ukrainian.] 
. VopoP JANOVA, N. S., IVANOVA 

KROGULEVICH, ET AL. 1972. Vysoko Baenja Flora 


Stanovogo Nagorja (High-Mountain Flora of the 
Stanovoy Plateau). Nauka, Novosibirsk. [In Rus- 
sian. 
'. N. VorosHILov, E. M. Econo- 
DN Ranunculaceae. Pp. 178-190 
uw ps achev (editor), Manual of the Higher 
Planis al the Sakhalin and Kuril Islands. Nauka, 
Leningrad. [In Russian. ] 
19 Flora o 
POM Vistas (Flora of the Soviet Far E 
Nauka, Moscow. [In Russian. 
—— ——. 1966. Ranunculaceae. Pp. 178-190 
JT. "Tol inache (editor), Manual of Plants of 
Primorje and Priamurje. Nauka, Leningrad. [In 


1982. Opredelitelj rastenyi Sovetskogo 


329. 


Daljnego bodie (Manual of Plants of the Soviet 
Far East). Nauka, Le sningrad. [In Russian.] 
VYKHODTSEV, 1974. Endemizm vo flore 
Tjan- Shan- ides gornogo sooruZenija (Ende 
mism in the flora of the Tier v» yan-Altai Mountain 
Chain). Probl. Bot. 12: 14- 
Wanc, C. W ae of China. M. 
Moors nae agen Publ. Ser. 5: 1-313 
eed The coniferous forests of Tanase, 


— 
al 


on Bull. Dept, Biol Coll. Sci. Tunghai Univ. 34: 
-=58; 


ANG, W. T. 1957. Notulae de Ranunculaceis 
Sinansibus. Acta Phytotax. Sin. 6: 301-391 
— ———. 1962. Critical review of the genus ; Del- 


"ER. a the bes Er cues b of China. 


. WILLDE, H. 1965 


. WILLIS 


, o J. 


. ZIMAN, S. 


u pidtriby i Ta 


l vykh "ead and Phylogeny of 


Er 64-284. 

WARDLE, zu 978. pis of rn po Zealand 

Mountain EE with special reference to trans- 

Tasmanian relationships. New Zealand J. Bot. 16: 
550 


Acta Bot. 


Additional evidence for the 
Africa- Madagascar- India Ceylon land-bridge the- 
ory with special reference to the genera Anisopap- 
pus and Commithora. Webbia 19: 497-505. 

.Scorr. 1918. The sources 
and distribution uf the New Zealand flora, with a 
a to criticism. Ann. Bot. (London) 30: 339- 


Some aspects of plant ge- 
ography ae A Northern Hemisphere during the 
Late Cretaceous ien. Tertiary. Ann. Missouri Bot. 
Gard. 62: 264-27 

^ 19 25: Oc herk istorii "id Vostoch- 


S Gosud. Geogr. Obsc. 71: 1426-1447. 
1975. 
Volume 1. Beijing. 

N. 1977. Obzor ziznennykh form v 
sem. Ranunculaceae. 1. Helleboroideae, Isopyroi- 
deae, Coptidoideae ya y of life forms in the fam- 
ily Ranunculaceae). Pp. 59-96 in Novosty Syst. 
Vyschykh i a zastenyi 1976. Naukova 
dumka, E [In rs an.] 


ializ filogenetichnykh zvjazkiv 


The Alpine Plants of Taiwan, 


of the relative connections in the 

tidinae, Ranunc "eia Ukrajins k. Bot. Zurn. 38: 
4-13. [In 2 raini 
———— " Morfologija 1 “ue ljutiko- 
the Ranun- 
culaceae). Naukova dumka, Kiev. [In Russian.] 


A REVISION OF THE James S. Miller: 
NEW WORLD SPECIES OF 


EHRETIA (BORAGINACEAE) 


ABSTRACT 


Ehretia is a pantropic M genus of about 50 species with centers of diversity in eastern Africa and tropical Asia. 
Although as many as 12 species of Ehretia have been recognized in the New World, field and herbarium studies 
suggest that the patterns of variation in the New World i 
recognized Ehretia anacua and E. tinifolia as distinct ed and have distinguished several species with serrate 
leaf margins. Field studies suggest that characters used to differentiate species with serrate leaf margins often var 


y 
within populations or on a single individual. Thus Ehretia latifolia i is treated in a broad sense, including all segregate 
leaves. 


species with serrate 


The largest genus of the subfamily Ehretioideae, 
Ehretia, contains about 50 species (Airy-Shaw 


» 


1973) distributed tropically and subtropically in 
both hemispheres. The genus is primarily Old World 
in distribution with more species in Africa than in 
Asia; Baker & Wright (1905) recognized 23 Af- 
rican species, mostly in eastern Africa, Johnston 
(1951) recognized 12 species in eastern Asia, and 
perhaps several more occur in the Malay archi- 
pelago and Australia. The genus is poorly repre- 
sented in the New World where it ranges from 
The 
last complete treatment of the New World species 
was that of Miers (1869) who recognized eight 
species. An additional six species were published 
by various authors after his work. Standley (1924) 
recognized seven species in Mexico and one more 
in Costa Rica (Standley, 1938) based on relatively 
few collections of the species with serrate leaf mar- 
gins. With the great increase of collections avail- 


Texas to Panama and the Greater Antilles. 


able since Standley's work, initial studies suggested 
that the characters he used to separate the species 
were of questionable validity, and delimitation of 


the species became problematic. The present re- 
vision combines a study of herbarium specimens 
with extensive field study of the variation of leaf 
and indument characters to resolve these problems. 
The work is supplemented with surveys of leaf 
architecture and pollen morphology throughout the 
Ehretioideae to be published later; only results di- 
rectly relevant to the New World species of Ehretia 
are included 

n addition 1 to o problems with delimitation of 
species of Ehretia in the New World, there have 
been questions regarding definition of the genus. 
An additional aspect of this study was an exami- 
nation of generic limits in the arborescent genera 
of Ehretioideae. Selected species of Bourreria, 
Carmona, Rochefortia, Rotula, and Old World 


species of Ehretia were examined. 


TAXONOMIC HISTORY AND GENERIC 
DELIMITATION 


Patrick Browne (1756) named Ehretia after 
Georg Dionysius Ehret, the illustrator of Browne's 


york was supporte əd in part by National PA ELE grant 3449-86. Fieldwork in Mexico would have been 


! This 
difficult d the gracious hospitality of the sta 


and M. Sou 
Márquez, E. Martinez, J. 


d. J. Nowicke provide 


ff of the Instituto de ag and the Facultad de Ciencias at the 
Universidad Nacional Autónoma de México. I particularly thank A. Del andez, T. i R 

sa for logistical support. G. Campos organized a trip through forse har would hav 
Myers, O. Téllez, R. Torres, 
4 encouragement and collaboration with pollen studies, 
work, C. Jarvis helped with nomenclatural questions, and J. Solomon and M. Thulin provided review 


, H. Hern 


and G. Urquijo provided help and 
erello 


comments. I also thank the curators of the following herbaria for hospitality during visits or for specimens on loan: 
G 


A, B, BH, BM, CAS, CR, DS, F, G, GH, IJ, K, L, LE, LL, 


MA, MEXU, MICH, NY, P, S, TEX, UC, US, WIS. 


* Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. 


ANN. MISSOURI Bor. Garb. 76: 1050-1076. 1989. 


Volume 76, Number 4 
1989 


Miller 1051 
New World Ehretia 


Civil and Natural History of Jamaica. Linnaeus 
(1759) later validated the genus with the binomial 
Ehretia tinifolia. De Candolle (1845) significantly 
enlarged the concept of the genus by treating 58 
species, many of which are now referred to the 
segregate genera Bourreria, Carmona, Rochefor- 
tia, and Rotula. De Candolle recognized four sec- 
tions within Ehretia. All of the species of his sect. 
Ehretia are still referred to the genus but his sect. 
Bourreria was a diverse group of species now as- 
signed to Bourreria and Ehretia. The two species 
that he treated in sect. Carmona have been con- 
sidered conspecific and treated as Carmona retusa 
(Vahl) Masamune (Thulin, 1987). His fourth sec- 
tion, sect. Xeroderma, was another diverse group 
containing species now distributed in Bourreria, 
Ehretia, Rochefortia, and Rotula. Several species 
of sects. Bourreria and Xeroderma have since 
been excluded from Boraginaceae (see excluded 
species). 

John Miers published a series of papers on the 
Boraginaceae shortly after De Candolle's work and 
treated Ehretia (Miers, 1869) in a more restricted 

se. Miers segregated the genus Rotula (as 
Rhabdia) and allied it with Khretia and Cortesia 
(a small genus endemic to saline areas in northern 
Argentina). Miers also sorted the species that most 
recent authors have included in Bourreria into 
Bourreria and Crematomia. Crematomia was 
characterized by having fruits with four pyrenes, 
each retaining an apical attachment to the stigma 
at maturity. Recent authors (Schulz, 1911; No- 
wicke, 1969; Gibson, 1970; Miller, 1988) have 
not accepted this as sufficient for generic recog- 
nition and have included these "Crematomia" 
species in Bourreria. Miers pointed out that Bour- 
reria was distinct in its fleshy calyx with five mar- 
ginally tomentose, valvate lobes; fleshy, rotate co- 
rollas; and fruits that at maturity divide into four 
angular Lied each with a ridged endocarp and 
spongiase mesoc 

Bentham & Hooker (1876) recognized three 
sections e Ehretia, one of which, Carmona, has 
been treated as a segregate genus by most recent 
authors (Johnston, 1951; Nowicke & Miller, in 
press). Johnston (1951) recognized their other two 
sections, Kuehretia and Bourrerioides, in his treat- 
ment of the eastern Asian species. Although John- 
ston devoted most of his research career to Bo- 
raginaceae, he published little on  Ehretia or 
Bourreria. He did recognize Carmona and Rotula 
as distinct from Ehretia (Johnston, 1951). In his 
treatment of the Boraginaceae of the southern West 
Indies (Johnston, 1949) he also considered Bour- 


reria and Rochefortia as distinct from Ehretia. 
Hence, although Johnston never treated Ehretia, 
it appears that his generic concept was similar to 
that of most recent authors 

As currently defined, ilie Ehretioideae consist 
of eleven genera that segregate into four probably 
monophyletic groups, but the relationships among 
them remain obscure. Richardson (1976, 1977) 
separated Tiquilia from Coldenia and these two 
genera of prostrate plants with dry nutlets form a 
distinct lineage within the Ehretioideae. Lepido- 
cordia, including Antrophora (Miller & Nowicke, 
in prep.), is also quite distinct from any of the other 
genera of the Ehretioideae, and Johnston (1950) 
considered the genus intermediate between the Eh- 
The Argentinian 
endemic Cortesia and the Australian endemic Hal- 


retioideae and Heliotropioideae. 


gania comprise a questionable third group of 
shrubby plants of xeric regions with often trifid or 
unevenly toothed sclerophyllous leaves. Whether 
these two genera are closely related or simply share 
a similar appearance because of convergence in 
similar habitats is unclear. Halgania is unique in 
the Ehretioideae in its stamens connate around the 
style. 

The remaining genera of Ehretioideae, Bour- 
reria, Carmona, Ehretia, Pteleocarpa, Rochefor- 
tia, and Rotula, seem to form a rather close group. 
Of these only Pteleocarpa seems to stand apart in 
its unusual winged fruits. Although recent pollen 
studies (Nowicke & Miller, in prep.) supported 
recognition of Bourreria, Carmona, Rochefortia, 
and Rotula as distinct 
appear closely related and form a distinct group 
within the subfamily. (See Table 1 for comparison.) 

Thulin (1987) transferred five African species 
of Ehretia to Bourreria, a genus previously con- 
sidered to be restricted to the Neotropics. These 
species seem to form a monophyletic group distinct 


rom Ehretia, they still 


from Ehretia in their valvate calyces, campanulate 
to urceolate corollas, and ridged endocarps. Eh- 
retia is a more natural genus without these species, 
but they are probably distantly related to the New 
World species of Bourreria, which are character- 
ized by rotate corollas and pollen grains with thick- 
ened ridges on each side of the colpi (Nowicke & 
Miller, unpublished). The characteristics of the 
genera closely allied to Ehretia are compared in 
Table 1 and a key to the genera of the Ehretioideae 
is provided below. Although Johnston (1935) in 

cluded Saccellium in the Ehretioideae, recent pol- 
len studies (Nowicke & Miller, in prep.) indicate 
that it is more closely related to the genera of the 
Cordioideae where Tahktajan (1987) placed it. 


1052 


Annals of the 
Missouri Botanical Garden 


KEY TO THE GENERA OF THE EHRETIOIDEAE 


la. Procumbent shrubs or jen herbs; sid is crowded or few-flowered; fruits dry, of 4 nutlets. 
2a. Ant rld 
a nu 


als with a ititious roots; leave 
lb. Trees or erect « 

but not of 4 nutlet 

3a. Leaves sc lerophyllous low shrubs. 

4a. Anthers connate in a 


asymmetrical; flowers tetramerous, Old Wo 
2b. Perennials E adventitious roots; le eaves symmetrical; flowers 
or Sep shrubs; inflorescences cymose or Sende fruits fleshy at least when young, 


cone surrounding the style; Australia 


RS Coldenia 
entamerous, New World 


T Tiquilia 


Halgania 
Cortesia 


4b. Anthers free; Rae ntina 
3b. Leaves coriaceous to membranace 
Sa. Flowers unisexual, the plants dioecious. 

6a. 


Plants with spines; endocarp dividing into four parts at maturity 
6b. Plants Visas: pues endocarps remaining entire at maturity 


wn 
v 


Flowe "rs all bisex 


eous; erect shrubs or trees. 


ochefortia 


Lepidocordia 


Ta. Style 1, entire or divided only to the middle. 
a 


alyx valvate; fruits with 


8b. Calyx eee. hu with a fles 
not ridged an ver retaining an apica 
9a. "a wen ad bs with flexuous 
Trees or ahaa style usually divided with 2 
7b. Styles 2. f free to the base. 
l Fruits Medien wit 
10b. Fruits drupaceous, not unt leaves toothed at the apex 


THE New WORLD SPECIES OF ÉHRETIA 


Only three of the New World species that De- 
Candolle (1845) included in Ehretia still belong to 
the genus. These are Ehretia anacua (Teran & 
Berl.) I. M. Johnston (as E. elliptica DC.), E. 
latifolia DC., and E. tinifolia L. Miers (1869) 
adopted a very narrow species concept, recognizing 
seven species with entire leaf margins, where only 
two have been recognized by recent authors (Stand- 
ley, 1924; Nash & Moreno, 1981). Various au- 
thors described six New World species of Ehretia 
with serrate leaves between 1891 and 1938. Stand- 
ley (1924) recognized five species with serrate 
leaves in Mexico and an additional one from Costa 
Rica (Standley, 1938). They differed in leaf shape 
and indument, although field studies reveal these 
characters are extremely variable. The present 
study recognizes three species of Ehretia in the 
World. This differs from recent treatm 
by grouping all populations with distinctly serrate 
leaf margins as a single, variable species. This 
variation in leaf morphology of Ehretia latifolia 
is discussed in detail under that species. 


New ents 


The determination of the natural ranges of the 
three New Worl i 
fact that all are cultivated. Ehretia tinifolia is the 
most widely grown, often significantly outside its 
natural range, and the other two species are fre- 


species is complicated by the 


quently planted near houses in areas where they 
occur naturally. These trees seem to be valued 
mostly for shade because of their dense crowns, 
and they are attractive when in flower and fruit. 


1 a spongiose mesocarp, the endocarps with a ridged outer surface 

etaining an apical attachment with the style B 

shy to mucilaginous mesocarp, the endocarps usually 
tyl 


h broad, reticulately winged margins; leaves entire 


ourreria 


al attachment with the style. 
branches; dis simple E obscurely bilobed ... Rotula 
distinct stigmas Vhretia 


mE Pteleocarpa 
Carmona 


Ehretia latifolia is apparently not used for wood 
by local people 

Almost everyone questioned in Mexico had a 
different common name for the local species of 
Ehretia and because of this no common names are 
reported in this study. 


MORPHOLOGY 
HABIT 


All species of Ehretia are trees or shrubs (Fig. 
1). Although New World species are trees, E. an- 
acua is frequently multistemmed. Habit is much 
more variable among the O orld species, which 
range from trees, like E. acuminata fad 
desert shrubs, such as E. rigida (Thunb.) Druce. 
The genus Rotula differs from Ehretia in its arch- 
ing, flexuous branches associated with its rheophy- 
tic habitat. The trunk of Æ. latifolia is usually 

uted, at least on mature plants (Fig. 2 


LEAVES 


All species of Ehretia have simple, alternate, 
nearly lanceolate to nearly round leaves. A few 
Old World species are extreme, for example Eh- 
retia saligna R. Br. of Australia has elongate, 
straplike leaves. Leaf shape is variable within 
species, a single individual often showing a signif- 
icant diversity of leaf shapes on a single branch 
(see E. latifolia). 

Ehretia tinifolia and E. anacua have entire- 


Volume 76, Number 4 
1989 


Miller 1053 
New World Ehretia 


NR. o mm 
AT NT 


Ficures 1, 2. Ha 


rown. e trees are often left as sh 


margined leaves. In some specimens of Ehretia 
anacua, however, a few leaves exhibit short teeth 
along the margin. Ehretia latifolia is unique among 
the New World species in its evenly serrate leaf 
margins. Only rarely are the teeth short and un- 
even. 

Leaf architecture can be valuable in delimitation 
of higher taxa (Hickey & Wolfe, 1975; Lorence, 
1985; Levin, 1986a, b, c) and has been used within 
77). Leaves of 36 
species of 10 genera of i dada d 
for leaf architectural features. Eleven species of 
Ehretia were selected, including the three New 
World species. Leaves were removed from her- 
barium collections, cleared, stained, and mounted 
according to methods of Dilcher (1974), and ar- 
chitectural features were analyzed following the 
system of Hickey (1973, 1977, 1979). A complete 
survey of the genera of the Ehretioideae will be 


genera (Rury & Dickison, 1 


published separately, but results are reported here 
for the New World species of Ehretia and several 
other taxa for comparison. Voucher specimens for 
the leaves examined are cited in the figure captions. 

Leaves of Ehretia appear to be primitively bro- 
chidodromous, this being the condition in nearly 
all of the species, and in the closely related genera 
Bourreria, Rochefortia, and Rotula. Craspedod- 
romous venation is restricted to the few species 
with serrate leaf margins. Of the New World species, 
E. tinifolia has brochidodromous venation (Fig. 4). 
Ehretia latifolia is the only New World species 
consistently with craspedodromous venation (Fig. 
3). Although this might appear to be a conservative 
character, it is variable in E. anacua, which gen- 


bit of Ehretia latifolia. —1. Tree 8 m tall, 
ade trees after other species have been cut.— 


es 
individual. (Both vouchered by Miller & Myers 2802 MO.) 


, Mexico. Note the dense 
hited trunk of the same 


near Guelatao, ud 


erally has brochidodromous venation (Fig. 7) but 
occasional individuals and leaves on juvenile shoots 

ave serrate margins and semicraspedodromous 
venation (Fig. 6). Even in E. acuminata, an 
World species with serrate leaves (Fig. 5), leaves 
are brochidodromous with veins that arise second- 
arily from outer loops beyond the secondaries and 
extend into the teeth. 

The patterns of tertiary and higher-order ve- 
nation vary only slightly among the species of Eh- 
retia. Generally, five or six orders are visible in 
most species. Ehretia anacua (Fig. 8) and E. tin- 
ifolia have randomly reticulate tertiary veins that 
anastomose and form a weak, composite interse- 
condary vein. The higher orders of venation vary 
from random to orthogonal reticulate. The ultimate 
marginal venation is looped in both of these species. 

Ehretia latifolia (Fig. 9) and E. acuminata (Fig. 
10) have randomly reticulate to weakly percurrent 
tertiary veins. Intersecondary veins are weak and 
restricted to the apical portion of the leaf, or are 
completely absent. Ultimate marginal venation is 
incomplete in E. latifolia but looped in E. acu- 
minata 

The New World species of Ehretia are essen- 
tially evergreen. Ehretia anacua is partly decid- 
uous only at the northern extreme of its range in 
Texas. Ehretia latifolia appears to produce most 
of its leaves during the onset of spring rains and 
hold them for about two years. Only in the pop- 
ulation near Monte Alban, Oaxaca, in Mexico did 
ed all of its 
leaves during the dry season. New leaves were 
emerging even as the last few old leaves dropped. 


I observe a plant that seemed to s 


Annals of the 
Missouri Botanical Garden 


Volume 76, Number 4 1055 


Miller 
New World Ehretia 


TABLE 1. Comparison of the major morphological characters of Bourreria (New World), Bourreria (African, 
sensu Thulin, 1987), Carmona, Ehretia, Rochefortia, and Rotula. 


Bourreria 


Bourreria 


African sensu 
Thulin 


Characters New World Carmona Ehretia Rochefortia Rotula 
Habit trees and trees and shrub trees and trees and rheophytic 
: s shrubs shrubs shrubs shrubs with 
arching 
branches 
Leaves entire entire apically re- entire or entire entire 
tuse or serrate 
ri 
Calyx aesti- valvate valvate quincuncial quincuncial imbricate quincuncial 
vation 
Corolla form rotate urceolate to tubular tubular to shortly tu- tubular 
campanu- campanu- bular 
late late 
Corolla tex- leathery thin thin thin thin thin 
ture 
Fruit, num- 4 4 l 1, 2, or 4 4 4 
ber of 
parts 
Endocarp ridged ridged lightly rugu- rugulose to striate reticulate 
lose smooth 
Mesocarp spongiose spongiose fleshy fleshy fleshy fleshy 
INDUMENT without magnification. As the leaf reaches senes- 


Standley (1924) differentiated New World 
species of Ehretia using density and thickness of 
hairs and presence or absence of glandular hairs. 
Field observations indicate that neither of these are 
significant in the sense that Standley used them. 
The widely scattered geographic pattern of plants 
with glandular hairs is not indicative of a mono- 
phyletic group, but rather probably a character 
that is expressed only rarely. 

Collections of Ehretia anacua and E. latifolia 
show considerable variability in leaf indument, much 
of which can be explained as a temporal phenom- 
enon. Young leaves of E. latifolia are densely and 
softly pubescent. Expansion of the leaf blade de- 
creases the density of hairs, and young, fully ex- 
panded leaves are less densely pubescent with long- 
shafted, simple hairs from a multiple-celled, basal 
cystolith. During the life of an individual leaf, the 
majority of the brittle shafts of the hairs are often 
broken off and the leaf may appear nearly glabrous 


cence, the cells of the basal cystolith continue to 
expand and harden. The basal cystoliths with the 
remaining portions of the hair shafts often make 
up what appears to be a scabrous indument. Thus 
an individual leaf may pass through stages from 
velutinous, to sparsely pubescent, to nearly gla- 
brous, and finally to scabrous. e southern 
portion of the range of Ehretia latifolia the dry 
season is less severe. Here leaves are somewhat 
larger, and seldom do herbarium collections contain 
eaves old enough to have reached the last, seem- 
ingly scabrous, stage. These field observations sug- 
gest that indument is not a reliable character for 
the delimitation of segregate species from £. la- 


tifolia. 


INFLORESCENCES 


All species of Ehretia have many-flowered in- 
florescences that are cymose or are paniculate with 
the individual branches of the panicles cymose. 


U 3-7. Clea 
vow (Miller & Télle= 3194 MO, Mexico).—5. E. 
6 


red leaves of Ehretia species.—3. E. latifolia ( 
acuminata (Pételot 6382 MO, 


2690 MO, 
Vietnam). 6 


e —4. E. 
. E. anacua.— 


(Miller et al. 


Juvenile leaf (E. J. Palmer 10809 MO, Texas). — 7. Mature leaf (Duke 3690 MO, Mexico). Scale bars equal 


cm. 


1056 


Annals of the 
Missouri Botanical Garden 


A 


FIGURES 


8 
(Miller et al. 


10. Cleared leaves of Ehretia species. - 
2690 MO, Mexico). — 10. E. acuminata ( 


The inflorescences of Ehretia tinifolia and E. la- 
tifolia have a distinct central rachis and are ba- 
sically paniculate with cymose individual branches. 
Ehretia anacua has a short-pedunculate, cymose 
inflorescence without a central leader. This type 
of cymose inflorescence is widespread in the Eh- 
retioideae and Cordioideae and is probably the 
primitive character state in the two subfamilies. 


FLOWERS 


The floral morphologies of the New World species 
of Ehretia are quite similar. Ehretia anacua has 
hairs on the corolla that are not present in F. 
latifolia or E. tinifolia, but other differences are 
small. Floral features that vary among the New 
2. The Old 
World species are quite different and vary in nu- 
merous floral characters, including features of the 


World species are compared in Table 2. ' 


calyx and size, color, and form of the corolla. 

The diversity of breeding systems in the Borag- 
inaceae is well documented (Bawa & Beach, 1981; 
Ganders, 1979; Vuilleumier, 1967) and second 
only to Rubiaceae in the number of distylous species. 
Ehretia has monomorphic, bisexual flowers with 
the stamens exserted and exceeding the style. This 
is the most common breeding system in the Eh- 
retioideae; the only genera that differ are Lepi- 
docordia (Miller & Nowicke, in prep.) and Ro- 
chefortia (Lefor, 1968), both dioecious. Corollas 
are small and white in the New World species and 
are visited by a wide variety of small generalistic 
Diptera, Hymenoptera, and Lepidoptera. The plants 
| observed of Ehretia latifolia were visited by 
many small insects, mostly dipterans. 


POLLEN 


Pollen is particularly useful in generic delimi- 
tation in the Boraginaceae (Avetissian, 1956; Now- 


CN 


TRU RETIN ORES PERE TOA 


RS: 
AS 
SA 


& 
"ur, 


a. 
D 
č 


^ d 
M. cM 


APA AA ORE. AE 


- 


ees 
x 


: UN 
DEP. m 


8. E. 
Pét 


anacua (Duke 3690 MO, Mexico). 


9. E. latifolia 
elot 60382 MO, Vietnam). Scale bars equal 1 cm. 


icke & Ridgway, 1973; Nowicke & Skvarla, 1974; 
Nowicke & Miller, in press). Pollen of Ehretia is 
more or less prolate and three-colporate (Nowicke 
& Miller, in prep.). Three pseudocolpi or pseu- 
docolpoid depressions alternate with the colpi. The 
apertures have elongate colpi with a bridge of ek- 
texine over the endoaperture. The tectum is more 
or less complete and faintly rugulose, slightly gran- 
ular, or psilate. 

Ehretia is homogeneous and distinct from other 
genera of the Ehretioideae in its pollen morphology 
(Nowicke & Miller, in prep.). Evidence from pollen 
does not support earlier sectional divisions of the 
genus (see Bentham & Hooker, 1876; Johnston, 


FRUITS 

The fruits of Ehretia are drupaceous with a thin 
to leathery exocarp, fleshy to mucilaginous me- 
socarp, and bony endocarp. They vary from white, 
as in E. latifolia, through yellow and orange, as 
in E. anacua, to red, as in E. longiflora Champ. 
(Johnston, 1951). The calyx generally persists ba- 
sally but is not at all accrescent. Although the 
stigma is usually lost, the base of the style often 
persists at the apex of fruits. 

Fruit morphology has been important for generic 
delimitation in the Ehretioideae. Bourreria, Car- 
mona, Ehretia, Rochefortia, and Rotula have es- 
sentially drupaceous fruits, varying in the texture 
of the mesocarp, number of parts into which the 
endocarp divides at maturity, and surface sculp- 
turing of the endocarp. The mesocarp of these 
genera except for Bourreria is fleshy to mucilag- 
inous. That of Bourreria is spongiose, being fleshy 
only when fruits are immature, becoming somewhat 
fibrous with large open air pockets. 

The structure of endocarps was examined by 


Volume 76, Number 4 


Miller 1057 
New World Ehretia 


TABLE 2. Comparison of floral features of the New World species of Ehretia. 
Ehretia anacua E. latifolia E. tinifolia 

Calyx length 2.7-3.5 mm 1.6-2.6 m 1.5-2 mm 
Calyx lobe shape narrowly triangular deltate to triangular ovate 
Calyx indument pubescent to densely strigillose glabrous to strigillose glabrous 
Corolla Sa tubular-campanulate campanulate campanulate 
Corolla length 6- 7.8 mm 3.2-6 mm 4-4.6 mm 

Corolla ig strigillose to puberulent glabrous glabrous 


Du lobe shape obovate 
2.8-3.4 mm 
4-6.3 mm 
2.7-5 mm 


Corolla lobe length 
Staminal filament length 
Style length 


broadly ovate widely oblong to 


1.8-2 mm 2.5-5 mm 
4-4.5 mm 3-4.5 mm 
3-4 mm 2-3.4 mm 


removing mature fruits from herbarium specimens, 
soaking in 5% NaOH for 1-5 days to soften the 
exocarp and mesocarp, washing in running water 
until clear of all outer material, and then drying. 
They were then examined with an Hitachi 5450 
scanning electron microscope. Fruits of 54 species 
of ten genera of Ehretioideae were examined. Col- 
denia and Tiquilia stand apart in having fruits of 
four dry nutlets. The fruits of Halgania, Lepi- 
docordia, and Rotula all differ in having a thin- 
walled endocarp. Bourreria, Carmona, Ehretia, 
and Rochefortia, however, all have somewhat sim- 
ilar fruits. 

Fruits of Ehretia vary in the number of parts 
into which the endocarp divides. Of the species 
examined, Ehretia latifolia is the only species whose 
endocarp remains entire at maturity. Ehretia ac- 
uminata, E. anacua, E. dicksonii, and E. tinifolia 
have fruits with two separate two-seeded pyrenes 
at maturity. Ehretia amoena, E. coerula, E. cy- 
mosa, E. laevis, E. obtusifolia, and E. rigida have 
fruits with four separate single-seeded pyrenes. Of 
related genera, Carmona has an endocarp that 
remains entire. Rotula has a two-parted endocarp 
and Bourreria and Rochefortia have four-parted 
endocarps. 

The surfaces of the endocarps of Khretia species 
have distinct sculptural patterns. Two major types 
occur in Khretia, and several others turn up in 
related genera. Verrucose endocarps are found in 
Ehretia acuminata (Fig. 15), E. dicksonit, E. la- 
tifolia (Fig. 13), and F. tinifolia (Fig. 12). Ehretia 
dicksonii is unusual among these species in having 
an evenly puberulent endocarp with numerous short 
hairs on the outer surface. Areolate endocarps, 
with angular depressed areas, characterize most of 
the Old World species with four-parted endocarps, 
including E. rigida (Fig. 14). Ehretia anacua, of 
the New World (Fig. 11), also has an areolate 


endocarp but differs in being less regularly areolate 
and in ano dividing into two parts at maturity. 
Th of the most closely related genera 


re 
are quite different in sculpturing pattern. Carmona 
retusa has an entire endocarp with few, low, broad 
ridges but is otherwise nearly smooth (Fig. 16). 
Rochefortia has endocarps with broad, thick, 
rounded ridges as exemplified by R. spinosa (Jacq.) 
Urb. (Fig. 19 
varying from erect, as in 
Fig. 18), to ee as in B. nemoralis (Gurke) 
Thulin (Fig. 17). 


Fruit structure suggests that KAretia consists of 


o 


two groups. One has one- or two-parted endocarps 
with a verrucose surface and is found in the New 
World and Asia. The second has four-parted en- 
docarps with an areolate surface and is widespread 
in the Old World. Ehretia anacua is the only New 
World species in the latter group but differs in 
having a two-parted endocarp. 


SYSTEMATIC TREATMENT 


Ehretia P. bae Civ. Nat. Hist., Jamaica 168. 
1756. korg tinifolia L., Syst. Nat. 
ed. 10. pl 159. 


Gaza ii. & Berlandier, joan Comis. Limites 5. 1832. 
zaza anacua d 1 & Berlandier, Mem. 
Dew Limites 5. 


Pubescent or glabrous, trees or shrubs. Leaves 
alternate, petiolate, the margin entire or serrate, 
the venation pinnate, brochidodromous or less com- 
monly semicraspedodromous or craspedodromous. 
Inflorescences terminal, cymose to paniculate. 
Flowers bisexual; sepals 5, imbricate or open in 
bud; corolla white in the New World species (blue 
in Æ. coerula and E. rigida, Africa), tubular with 
5 spreading to reflexed lobes or urceolate; stamens 


1058 Annals of the 
Missouri Botanical Garden 


T 


FIGURES 11-19. Endocarps of species of Ehretioideae. 
MO, Mexico). — 12. E. Mi ea mie r & Myers 2824 MO, Me Eo E. latifolia ( Purpus 5820 MO, Mexico). 
14. E. rigida (Armitage 305/59 MO, S Rhodesia). 15. E. acuminata "m i 7 74 MO, China).— 16. Carmona 
retusa (Merr ill: Species Hlancoana 3 MO, Philippines). 17, 18. Bourreria . B. nemoralis (Gurke) Thulin 
(Frazier 2267 MO, Ke p . B. rubra Lott & Miller (Lott 683 MO, Mexic se -19, Rochefortia spinosa Jacq. 


"ih sner & Gonzalez 12 T Venezuela). Scale bar equals 5 mm. 


meg 11. E. anacua (Hansen & Nee 7489 


Volume 76, Number 4 
1989 


Miller 1059 
New World Ehretia 


9, usually. exserted, the anthers oblong to ellipsoid; 


drupaceous, ovoid to nearly spherical, the endocarp 
separating into 2, 2-seeded pyrenes or into 4, 
l-seeded pyrenes, or rarely remaining entire. 


KEY TO THE SPECIES OF EHRETIA IN 
THE NEW WORLD 


la. Leaf iic dl serrate; endocarp re- 
maining en E. latifolia 
lb. Leaf margin entire or ipei so; endocarp di- 
viding into 2 parts at maturi 
2a. Upper leaf surface duris inflorescence 
paniculate; corolla glabrous ............... E. tinifolia 
. Upper leaf surface scabrous; inflorescence 
cymose; corolla strigillose to puberulent .. 
7. anacua 


N 
c 


Ehretia anacua (Terán & Berlandier) I. M. John- 
ston, Contr. Gray Herb. 70: 24. Gaza 
anacua Terán & Berlandier, Mem. Comis. 
Limites 5. 1832. TYPE: Mexico. Tamaulipas: 
prope Matamoros ubi vulgo adpellatur Ana- 
cua, Berlandier 2369 (lectotype, G, here des- 
ignated; isolectotypes, BM, C, 


Dias os elliptica DC., Prodr. 9: 503. 1845. TYPE: Mex- 
o: Berlandier 2330 (holotype, G-DC, not seen; 
microfiche isotypes, BM, G, GH, K, LE, MO, 
NY sheets, P). 
Ehretia m Kunth & Bouché, Sp. Nov. Hort. Berol. 
12 "Crescit in Mesica,” 
Ehretia us Miers, Ad 
3: 111. 1869. TYPE: U.S.A. Texas: Qum 665 
(holotype, K; pes BM, F, GH —2 sheets, K, 
MO — 2 sheets, NY, P, US). 
Ra exasperata Miers, big & Mag. Nat. Hist. Ser. 
Ji U.S.A. Texas: M. Drum- 
Mad 296 holotype, K, isotypes, BM— 2 sheets, 
GH, — 2 shee 
Ehretia lancifolia mu & “Mociño, Fl. Mex. 51. 1894; 
ed. . M. Johnston, J. Arnold Abt. 
30: 2e irem TYPE: Sessé et al. 5250 
(lectotype, MA, here designated, photo, MO, frag- 
ment, F). 


Tree 3-10(-15) m tall, 
trunks, the twigs puberulent, the older wood with 
thick furrowed bark separating into thin gray or 
reddish scales. Leaves persistent or partially de- 


often with multiple 


ciduous at northern edge of range; petioles 2- 
6(-11) mm long, puberulent, canaliculate adaxi- 
er leaf ta Vide to less commonly ovate, 
1.9-) 2 .2) em long, (1.2-)1.6-3.8(-5) 
cm wide, s apex obtuse or rarely acute to slightly 
acuminate, the base obtuse to rounded, rar 
slightly cordate, the margin entire or sometimes 
unevenly serrate on some of the leaves, often some- 
what revolute, the upper surface scabrous, the 


lower surface scabrous to puberulent, the lateral 
veins 4-7. Inflorescence terminal, cymose, 2.5-6 
cm long, the peduncle puberulent. Flowers bisex- 
ual, fragrant; sepals 5, narrowly triangular, some- 
times slightly connate basally, 2.7-3.: 
0 m wide basally, pubescent to strigillose, 
the hairs 0.3-0.4 mm long; corolla white, 6-7.8 
mm long, strigillose to puberulent on the exterior 
surface, 5-merous, the lobes obovate, 2.8-3.4 mm 
long, 1.8-2.6 mm wide, the tube 2.8-4.2 mm 
long; stamens 5, the filaments 4-6.3 mm long, the 
upper 1.8-3 mm of these free, glabrous, the an- 
thers ellipsoid, 1.2-1.4 mm long; in ovoid- 
1.3-1.8 mm long, 0.9-1.2 mm broad, 
glabrous; disc not evident; style 3 5 mm long, 
the stylar branches 0.7-1.7 mm long, the stigma 


conical, 


lobes capitate to terete. Fruits drupaceous, yellow 
to orange when mature, glabrous, globose, 5-8 
the endocarp dividing into 2, 2-locular 
pyrenes, these randomly reticulate-rugulose on the 


mm diam., 


outer surface. 


Distribution. | Ehretia anacua occurs in 
southern Texas and northeastern Mexico from sea 
level to 1,300(-2,200) m in dry forests (Fig. 20). 
Collections from Guanajuato, Oaxaca, and Que- 
retaro are outside the natural range of the species 
and are not included in the distribution map. 

This common species was known as Ehretia 
elliptica DC. until Johnston (1924) discovered the 
older epithet published in Gaza Terán & Berlan- 
dier. The description of E. scabra indicates that it 
is synonymous although the brief citation of “cres- 
cit in Mexico" does not indicate a type collection. 
The names F. ciliata and E. exasperata are clearly 
synonymous; E. ciliata differs from E. anacua in 
no discernible characters, and the type of E. ex- 
asperata is merely a juvenile shoot with serrate- 
margined leaves. Likewise, the type of E. lancifolia 
does not differ significantly in any character from 
E. anacua. Although most type material for the 
names lacks adequate locality information, it seems 
likely that all were collected in northeastern Mexico 
or Texas, a region in which no other species of 
Ehretia occur. 

n addition to its distinct geographic range, £. 
anacua is easily distinguished from the other New 
World species of Ehretia by its scabrous upper 
leaf surface, cymose inflorescence, strigillose co- 
rola tube, and usually entire leaf margins. It is 
quite distinct in the genus with no apparent close 
relatives. 

Ehretia anacua is valued for its showy flowering 
display and its shade. The specific epithet is derived 


from the common name “anacua” (Nuevo León, 


1060 


Annals of the 
Missouri Botanical Garden 


CARIBBEAN AMERICA 


o 800 kms 


L 9 


" EN ; 
^N 
Ya 
o . d ^ 
ec Ct e : 

e 

^ 

ug i 

N -* Í 


FiGURE 20. Distribution of Ehretia anacua. 


Tamaulipas, Texas). Standley (1924) reported the 
wood to be used for wheel spokes, axes, tool han- 
dles, and yokes. 


Representative specimens examined. U.S.A. TEXAS: 
Esperanzo Ranch, Brownsville, Barber 26 (LL, US); Los 
Borregos, 18 mi. NE of eue e pd 26a (US); 
Odum, cultivated, Bauml et al. O (TEX); Webb 
C 20., d Benavides 81 (TEX); ie Christi, Benke 
9 , F), 5429 (A, F); nar uc res E 
dea 900 (K), 939 (GH); Liv ae 
Th Rivers, Bluhm s.n. (TE ebb e (ra 
Botello & Ayala 51 (TEX); uu (us 152 (TEX, US); 
Hay s Co., back of Speck Hall, Chesser s.n. (FEX); Hidalgo 

»., transplanted from the Grande wooded area, Clo- 
ver 896 (MICH); Las Palmas Da. about 4 mi. SW 
of Brownsville, in palm grove, Correll 14869 (A, GH, 
LL, TEX, UC, US); Gonzalez Co., Palmetto State Par 
Ottine, Correll & Schweinfurth 15533 (A, LL); Hidalgo 
Co., Santa Ana Wildlife Refuge, Correll & Schw einfurth 
15630 (LL); along levee of the Rio Prende, l-2 mi. SE 
of Southmost, Correll & Johnston 17922 
TEX); Hidalgo Co., about West Pond, Santa Ana Wildlife 
Retuge, C ‘orrell E Johnston 18032 (A, LL); Refugio Co., 
in field along route #77, just N of intersection of route 
#13, Correll & Cons ll 1899854 (LL, MO); Calhoun Co., 
nd - M 11626 (GH); Cameron Co., Brownsville, 
Cory 2 1 (GH), 36616 (GH); Aransas Co., Aransas 
Refuge, “Cory 45876 (TEX), 49033 (A, NY), 5 
(MICH, NY); Aransas Co., 6.5 mi. NE 
51230 (GH, MICH, NY); Cameron Co., Rabb Palm Grove, 


ES 


5.5 airline mi. SE of Brownsville, Cory 51436 (GH, 
MICH, NY, UC, US); Goliad Co., 13% mi. N of Goliad, 
Cory 54619 (LL); Victoria Co., cultivated, Cory 55111 
(US); Hidalgo Co., E of Mer ERA Crockett 8026 (LL); 
Austin Co., bank ol Drazo River, Stephen F. Austin State 
Park, Crutc LE 2501 (LL, NY); Cameron Co., ] 


s.n. (TEX); 9 mi. Dod of Brownsville, Davis 46 (TEX); 
eu di ion River at Refugio, Dixon 424 (UC); 
Jacks ' Druskel 9601 (GH); pos son Co., Cordele, 


Druskel p 6 (GH) Aransas Co., 
about 7 
(6); Vic storia Co., hedges in 
Welder Wildlife Refuge, Javelina Bend, A 
Fosberg 44654 (US); Hidalgo Co., Weslac 
(GH); Hidslzo Co., city of Mission, cultivated, Gereau 
940 (MO, NY); We bb Co., Laredo, jus 4 (GH, 
LL); Duval Co., Benavides, Guevara s.n. (US); Bensten- 
Rio Grande V alley State Park, Rio Grande Valley, 10 mi. 
(by air) W of McAllen, 100 ft., Hansen et al. 3766 (LL, 
MICH, US, WIS) Brownsville, Hansen 436 (NY, US); 
Hidalgo Co., in yard beside Motel Matador, in McAllen. 
Harriman & Jansen 12394 (MICH); New Braunfels, 
Harvard s.n. (US — 2 sheets); Neuces Co., Corpus Christi, 
Heller 1502 (A, BH, BM 4 
ae. sheets, P, UC— 2 sheets, US— 3 Sheets Goliad 

, SE side of highway 59, 2.6 mi. SW of the San 
eee River, McNamara Raich, Hill 8234 (NY); Hays 
Co., San Marcos Lake, San Marcos, Innes 816 (GH, 
TEX); roadside 4 mi. E of Choate, dai 1180 (TEX); 
Laredo, Jones 29240 (MO — 2 sheets, UC); without def- 
inite locality, Lindheimer s.n. (NY e sheets, US); Lind- 
heimer 416 (GH); Comanche Springs, New Braunfels, 


Volume 76, Number 4 
1989 


Miller 1061 
New World Ehretia 


Lindheimer 1015 (BM, F, G, GH, LE, MEXU — 2 sheets, 
— 2 sheets, NY, P, TEX, UC, US), 1016 (BM, F, 
G, GH, LE, MEXU, MO — 2 sheets, NY, TEX, UC, US — 
à 1017 a F, GH, MO, NY, P, TEX, UC, 
, levee of San Marcos River, poo 
: f IS); Cameron Co., Brown 
E Lundell & Lundell 87 09 (LL, MICH); TRAE 
, in Palm Grove, S of Brownsville, Lundell & Lundell 
pe (GH, LL, MICH, US); Cameron Co., Brownsville, 
Lundell 10839 (GH, LL, MICH, NY, US); Hidalgo Co., 
in Park, McAllen, Lundell & Lundell 1277 1 (LL); Ken- 
nedy Co., King Ranch, Norias Division, on grounds at 
lodge, headquarters, Lundell & Correll 15221(A, F, LL, 
MO, NY, UC); Comanche Springs, New Braunfels, Mohr 
85 (A); Corpus Christi, Vealy 4 (US); Hidalgo Co., be- 
tween Weslaco and Mercedes, Nogle 9 (F); Neuces Co., 
Corpus Christi Bay, Palmer 870 (A, GH, K, MO, NY, 
US— 2 sheets); Jackson Co., x er: Navidad River, 
Ganado, Palmer 9235 (A, K, US) Jackson Co., 
Vanderbilt, Palmer 9730(A, Mo T S); Wilson Co., river 
banks, Sutherland Springs, Palmer 10809 (A, F, MO); 
Hayes Co., San Marcos, Palmer 11558 (A, F, MO, TEX); 
Uvalde Cc o., Montell, Palmer 12322 (K); Hays Co., San 
Marcos, along rocky bluffs 2 San Marcos river, Pal mer 
33886 (A, MO, NY, P); Texas Co., San Antonio, ina 
RX001 (TEX), 1849 (TEX); pus in the valley o 
Rio Grande, below Donna, Parry et al. s.n. (US); Webb 


Co., Laredo Junior College, Laredo, iin 86 (LL, TEX); 
Hays Co., SC campus, San OS, Paeron 126 
(TEX); Webb Co., Laredo Ramas 32 (WIS); Corpus 


Christi, Reverchon s.n. (A); Mie un tropical nursery, 
Richardson 1408 i DeWitt Co., Wester m, Riedel 
s.n. (GH, TEX); Willacy Co., ay oad 186, 5 
of Raymondville, Rios & Cavazos 274 (LL); Webb Co., 
Laredo, Rodriguez 78 (TEX), MN S c & Rus- 
sell 24278 (NY, US); Hays Co., San s Lake, San 
Marcos, Rose-Innes & Warnock 816 (F). Brownsville, 
Runyon 3226 (TEX), 3227 (TEX), 5175 (TEX), . 
(TEX), 5177 (TEX); throughout the lower Rio a 
Valley, i ideni Runyon 6053 (GH, LL, UC); Cam 
eron vazos tract dove preserve, Cavazos, San- 
derson . 56 (NY San Antonio, Pres nt s.n. D. 2 sheets); 
lorpus Christi, Sargent s.n. (A); Victoria, Sargent s.n. 
Br üinevile, Schiller 101. 3 (MEXU); 
Santa Ana National Wild- 
2089 (MOX Mercertown, Swarz 
an Mae, Tharp s.n. and Gonzalez Co., 
Tharp ue 2 sheets); San Mar Tharp s.n. 
(GH, MO. "TEX. UC); near Braunfels, Thart 1369 (TEX, 
US); Jackson Co., river flood plain, Tharp 8629 (GH, 
MO, NY, TEX, UC, US); poa Saefje Bog, Tharp 
49-1247 (TEX); Karis Co., i S of Karnes City, 
Thompson & Turner 7 m [^ Christi, Tracy 
9209 (BH, BM, F, G, GH, MO, NY, TEX, US WIS); 
Cameron Co., Southmost at edge a Palm Grove, ca. 1.4 
mi. 5 2 end of highway 1419, 20 ft., Traverse 1111 
(F, GH, LL, MO, TEX, US); San Patricio Co., Welder 
NA Foundation, N of Sinton Hackberry di E mi. 
from place where S loop of river road be enc y s 26 
a ie 
, Santa Ana 


N- S road, 80 ft., 
MP enc. ee : 

, Laredo Junior ded Campus, Trevino 48 (TEX); 
rm eron Co., S , Tucker s.n. (US— 2 sheets); 
Gonzalez Co. Palette 25 Park, I td 169 (TEX); 


pen Braunfels, 
ar Southmost, 


Comal Co., 
Warnock 46503 ( 


Comal Spring, Lande Par 
TEX); Cameron b» 


volubilis, et al., raised 
field, Webster & Wilbur 3033 (GH, , 
Lake, Whitehouse s.n. (GH, MICH, MO, NY, TEX); Palm 
Grove, Brownsville, Wiehe s.n. ie Hidalgo Co., 
Santa Ana Wildlife i 
c er, > 

, E of Brownsville, I des 1915 (BH); Goliad Co., 
ie oo 82 d i 


on I. t's 


mountains to Puertecito (via Cuesta de Zozaya), Joh nston 
8890 (GH); Muzquiz, Marsh 1072 (F, GH 
clova, Gloria Mountains, Marsh 1977 (F, TEX). 
GUANAJUATO: jardin du College, ri s.n. (GH); Moin 
definite locality, Duges 153B (GH, US). NUEVO LEÓ 
Highway 60, 18 mi. W of Linares, Ahshapanek 319 
(TEX); C añon de Valle Alto, 2,000 ft., Bowen 42 (TEX); 
Ojo de Agua, Sabinas Hidalgo, 610 ft., Chase 7024 (F, 
GH, MICH); Monterrey-Mexi end 
LL); Monterrey, Dodge 101 (MO, NY, 
Chapinqui, Monterrey, Druskel 9695 (G i. Mere 
Eaton € Edwards 70 (K), Edwards 70 (K, NY); near 
Monterrey, Gregg 769 (MO, NY); 10 mi. b Linares, 
Crum 3271 (GH); Juarez, E la 2 
F); Monterrey, Sintra Madre Mountains, Mueller & 
Mueller 167 (F, GH, e Sierra Madre Oriental, Diente 
Canyon about 12 mi. S of Monterrey, Mueller & Mueller 
1302 (A, F, LE, MEXU, MIC us TEX); Monterrey, Palm- 
er O71 (A, GH, K] P 
SE of Monterrey on M 


~ 
T 


Pequeña et al. 16M 628 (F, TEX); Villa de Santiago, 
dd kins & Hall 356 1 (BH); valley ne ar as rrey, 1,800 
ecu 10171 (BH, BM, F, 2 sheets, 


MO, NY, P, UC, s ull near Monterey, 
Pringle 11640 (BH, GH, L—2 sheets, 
a bs H); Primal TEO MO); a 
Falls, ca. 35 km SSE of Monterrey, 600-700 m, Sanders 
& dd 76077 (TEX); 40 mi. S y Monterrey on 
. Smith & Barkley 17M164 (F—2 sheets, 

LU, MIC H, NY, TEX, WIS); Florida Motel at Mon 
temorelos, Spessard 112 (TEX); Sierra Madre Oriental, 


(G H, MICH, US); Pup cie Vista Hors 35 mi. S of 
Monterrey, 2,200 I 
N of Monterrey on oe 85, iban 10847 (TEX, 


extilán, Ghiesbreght 30 (GH). QUERETARO: "Los Pan 
o a 4 km al S de El Trapiche, Mpio. de Arroyo Seco, 
Tenorio & Hernández 288 (MEXU). SAN LUIS POTOSÍ: 
La Mirall, Mpio. San Antonio, Alcorn 2561 (MEXU, 
TEX); Tzineja, ee Huehuetlán, Alcorn 287 1 (TEX); 
Tanjasnec, San Ene e X rn 3121 (TEX); Rio 
Verde, Pues 46 (F, GH, MO, , US); in the region 
of San Luis Potosí, 6, m 8, Do 4 Parry & Palmer 


1062 


Annals of the 
Missouri Botanical Garden 


591 1/2 (GH, MO, po Tamasopo Canyon, Pringle 
3709 (A, BM, F, G, GH, K, LE, MEXU —2 sheets, 
MICH, MO, NY —2 sheets, P, UC— 2 sheets, o 
sheets); minas de San Rafael, dl 5323'(BM 
MO, NY —2 sheets, US); Rascon, Rio de las D 
Purpus 5323 (F, GH, MEXU, MO, UC, US); i km al 
de El Naranjo, Mpio. de Ciudad de Maíz, 400 m 
PE. ski 7378 (MEXU, MIC D. 7405 (MEXU, MICH, 
TE 4 km al NE de Ciudad del Maiz, 1,300 m, Rze- 
dow ski 10240 (US, WIS); entre Guascama y Minas de 
San Rafael, Mpio. de Villa Juarez, 1, 200 m, Tio 1686 
¿XU). TAMAULIPAS: vicinity of San Jos 000 ft., 
Bartlett 10008 (F, GH, MEXU, MICH); aA del 
municipio de nd Carlos, 500 m, Jimé i 290 (MO); 
lorales, LeSueur 572 (F, GH, TEX, US); S of Mata- 
moros, LeSueur 6. 34 (F, TEX); a 3 km al SE de la Caseta 
Fiscal, en la entrada a Rancho Nuevo y a 1 km al E de 
Rancho El Gachupin, 200 m, Martínez 80 (MEXU —2 
sheets, MO); a 5 km al E de San Rafael (Sierra de San 
José de la Rusias), Mpio. Soto La Marina, 140 m, Mar- 
tinez 131 (MEXU, MO, NY); 2 km al N de San Carlos 
en el camino a Marmolejo, 600 m, Martínez 952 (MO); 
road to Juamave, 6 mi. SW of Ciudad Victoria, McVaugh 
10540 (BM, G, GH, MEXU, MICH, NY, TEX, US); 
Ciudad Guerrero, Montemayer 34 (TEX); San Fernando 
to Jimenez, Mia 001. : 1 ;H, US); bow of Victoria, 
320 m, Palr 7 (F, GH, MO, NY, IS); Hed 
€ i "5 m, Put 23 (BM, bu F, G, 
MO, NY, US); vicinity x Victoria, 320 m, Palmer r 32 
(F, GH, pus NY, UC, US), Palmer 573 (F, GH, MO, 
NY, UC, US); Hacienda Buena Vista, Wooten s.n. (US); 
canon al W de Ciudad Victoria atras de la Sierra Gorda, 
sea level, Yanez 786 (N 
de San Carlos, 700 m, 
Silosuchil, e pee ata Alcorn 1889 (TEX); El Hijo, 
corn 2652B (TEX); en el poblado de Santa Clara, 
Peas zada 4730 (F); El Mirador, Presa 
, Calzada et al. 6162 ur en la 
Ciudad de es an as e cruzar el P lel Rio 
Mund 50 m, Tp dai: et a 6336 (F); camino » paralelo 
a la carretera Pan Tuxpan, a 12 km de Panuco, 20 
m, » Chiang 288 (CH, MEXU `; A road pum Baños de 
izal to Emiliano Zapata (= Carrizal), m SE 
Emilano Zapata, Mpio. Apazapan, 250- 40 JO m, Hansen 
> 7489 (F, MO); vicinity of Panuco, Palmer 365 
(CAS. TF. G, GH, Y, ; Rio Grande, 3.2 
km al S de Santiago Tuxtla; 200 m, E sa 2448 (MEX U); 
plan del Río, Mpio. Dos Ríos, 350 m, Ventura 8022 
(MICH, MO); 250 m, Panda 8552 (MEXU). STATE 
UNKNOWN: without definite locality, Sessé et al. 773 (F), 
774 (E), 775 (F), 5247 (F). 


c 
= 


Ehretia latifolia DC., Prodr. 9: 503. 1845. TYPE: 
erb. amat. (holotype, G-DC, not seen; mi- 
crofiche, 


Ehretia mexicana S. Watson, Proc. Amer. Acad. 26 
14 1. TYPE: Mexico. Jalisco: base of mountains 
near Lake Chapala, C. G. Pringle 3085 (lectot type, 
GH, here designated; isolectotypes, BH, BM, F, 
>H, K, L, LE, MO, NY —2 sheets, UC, US S 2 
d are two collections at GH. neither 
annotated as holotype by ee 

Be pre J. D. Smith, Bot. Ga 
TYPE: Guatemala. Quiche: San Miguel Uspanlán, 


6,000 ft., Apr. 1892, He yde & Lux 3065 (holotype, 
US 94 44452: isotypes, F, G, GH, K, MO, NY, P). 
Ehretia cordifolia Srel Pr Amer. Acad. Arts eh 
319. 1894; png i ard. and Forest 8: 15 
1894. TYPE: Mexic dips o: valley, Zapotlán, P 
May 1893, C. C. Pringle 4382 (holotype, F 106011; 
isotypes, A, BM, G, GH, K, LE, MO, NY, P, UC 
US — 2 sheets 
Ehretia viscosa Fernald, in Un x Trees and doin 
1: 25, pl. 1902. 
risas iim $^ Mo 18 99, C. G. 
(lectotype, F 120287, here vis esign de 
, K, MEXU —2 sheets, MO, , 
UC). In describing this s ipic p^ id cited three 
collections. The order in whic ese are cited and 
his mention of them in his oler indicates that 
he felt that the first one was the 
Phretia te huacana Greenman, Publ. 
39. 1912. TYPE: 
Tehuacán, 2,200 r M 
Dus atti 2220 done F 235156; isotype, CH). 
Ehretia austinsmithii Standley, Publ. Field 
Hist., Chi od Ser., 18: 984 
Costa Rica. 5 aros ero , Austin- 
Smith H. 528 (holotype, F 919653; iu eS F, MO). 


Tree to 15(-25) m tall, the trunk fluted, the 
twigs pubescent or shortly hirsute to nearly gla- 
brous. Leaves persistent; petioles (3-)6-20(-31) 
mm long, puberulent or strigillose to nearly gla- 
brous, narrowly canaliculate adaxially; leaf blades 
ovate to oe or elliptic, (5.5-)7-1 
long, (2-)3-6 cm wide, the apex acute to 
acuminate, nor obtuse, the base obtuse or less 
commonly acute or slightly cordate, the margin 
serrate, rarely minutely so, adaxially scabrous to 
strigillose, abaxially coarsely to minutely puberu- 
lent or strigillose, occasionally nearly glabrous with 
only a few hairs on the major veins, the lateral 
veins 5-7. Inflorescence terminal or subterminal, 
a dense panicle, 4.5-7 cm long, (2-)3-5 cm broad, 
the branches puberulent to nearly glabrous. Flow- 
ers bisexual, monomorphic, sessile or short-pedi- 
cellate; calyx 1.6-2.6 mm long, glabrous to strig- 
illose on the outer surface, sparsely to evenly ciliate 
along the interior margin, the 5 lobes deltate to 
triangular, 0.7-1 mm long; corolla white, 3.2-6 
mm long, tubular with 5 reflexed, broadly ovate 
lobes 1.8-2 mm long and 1.6-2 mm wide, the 
tube 2.5-3 mm long; stamens 5, the filaments 4- 
4.5 mm long, the upper 2.5-3.2 mm free, gla- 
brous, anthers ellipsoid, 1-1.5 mm long; ovary 
ovoid, 1.3-1.5 mm tall, 1 mm broad, disc not 
evident, style 3-4 mm long, faintly bilobed, the 
stigmas capitate. Fruits drupaceous, white, gla- 
brous, broadly ellipsoid, 7-10 mm long, 6-9 mm 
wide, the endocarp remaining entire, 4-locular, only 
lightly ridged on the outer surface. 


Distribution. | Ehretia latifolia occurs from 


Volume 76, Number 4 
1989 


Miller 1063 


New World Ehretia 


CARIBBEAN AMERICA 


o 800 kms 


— 4 ML m 


FIGURE 21. Distribution of Ehretia latifolia. 
Jalisco in western Mexico through Central America 
to Chiriqui Province in Panama in cloud forests 
and dry montane forests at (1,500-)1,800-2,300 
m (Fig. 21). 


Ehretia latifolia exhibits extreme variability in 
leaf shape and indument, which has bred confusion. 
Standley (1924, 1938) recognized six species with 
serrate leaves in the New World. With the excep- 
tion of E. tehuacana, all were based on characters 
of leaf shape and indument. When Watson (1891) 
published E. mexicana, he made no mention of 
how it differed from Æ. latifolia. The types of the 
two are nearly identical, and Standley (1924) re- 
duced E. mexicana to synonymy. Smith (1893) 
published E. luxiana based on a collection from 
Guatemala with broader leaves that was almost 
glabrous abaxially and supposedly had larger flow- 
ers. The populations from the southern portion of 
the range do have slightly broader leaves that more 
often tend toward abaxial glabrousness, but there 
are many exceptions and there is no clear geo- 
graphic discontinuity. At the time that Robinson 
(1894) published E. cordifolia it appeared distinct 
in having cordate leaf bases. However, field studies 
(discussed below) indicate that this is not a valid 


character since it varies on single individuals as 
well as within populations. 

The problem with Ehretia viscosa, a species 
supposedly distinct in having glandular hairs, is 
slightly more complex. The few collections of it 
differ only in the possession of glandular hairs and 
in all other respects are identical to E. latifolia. 
Further, they come from localities scattered 
throughout the Mexican range of FE. latifolia. Glan- 
dular hairs thus appear to be an infrequently ex- 
pressed trait rather than a monophyletic group that 
should be recognized as a species. 

Standley (1924) differentiated Ehretia tehua- 
cana by its corollas about 10 mm long (supposedly 
more than 3 mm longer than the other species he 
recognized). None of the corollas on the type col- 
lection exceed 6 mm (Standley must have based 
his distinction on Greenman's original description) 
and are within the normal range of variation for 
E. latifolia. At the time that Standley (1938) de- 
scribed E. austinsmithii, it appeared to be geo- 
graphically disjunct with the nearest population to 
the north in Guatemala. As currently defined, F. 
latifolia has a relatively continuous distribution in 
Central America, being unknown only in Honduras. 
The Central American populations are quite uni- 


1064 Annals of the 
Missouri Botanical Garden 
TABLE 3. Populations of Ehretia latifolia sampled for variation in leaf shape and indument. 


Numbers of 


individuals 
Population State Voucher numbers Elevations sampled 
Volcan Tacaná Chiapas 2667-2670 2,300 m 4 
Benito Juarez Chiapas 2690-2695 2,100 m 5 
Union Juarez Chiapas 2721 2,010 m l 
San Cristóbal Chiapas 2793-2795 1,800 m 3 
Zinacantán Chiapas 2199-2800 2,190 m 2 
Gueletao Oaxaca 2801-2805 2,200 m 5 
Chicomezuchil Oaxaca 2806-2810 2,200 m 5 
Monte Albán Oaxaca 2817-2823 1,800 m 5 
Acatzingo Puebla 2957 1,960 m l 
Chapulco Puebla 2960-2962 2,000 m 3 
Ciudad Mendoza Veracruz 2971-2975 2,000 m 5 
Ixthuatlancillo Veracruz 2-2993 1,500 m 2 
Tepozt Morelos 2996-299 1,700 m 3 
i DR Morado Guerrero 2888-2892 1,400 m 5 
Querendaro Michoacan 3059-3063 1,870 m 5 
Tiripitio Michoacan 3064-3068 2,040 m 5 


form while most of the significant variation appears 
in Mexico. 


FIELD STUDIES OF VARIATION OF 
LEAF MORPHOLOGY 


The numerous synonyms of Ehretia latifolia 
seem to have arisen via a paucity of collections 
and a failure to understand how leaf shape an 
indument vary on individual plants, within popu- 
lations, and between populations. Herbarium spec- 
imens are only limited samples of individual trees 
and seldom provide an adequate representation of 
variation of leaf morphology. In order to assess 
how leaf morphology varied on a single individual, 
within populations, between populations, and over 
the developmental life span of individual leaves, 
collections were made over the range of F. latifolia 
in Mexico. 

ome variation emerges during the ontogeny of 
an individual leaf (discussed earlier under leaf mor- 
phology) and a small portion correlates with ge- 
ography, but most variation appears random and 
often can be found within a single population. 

Two characters seem to vary geographically. 
Plants from Central America and part of adjacent 
Chiapas tend toward slightly broad, abaxially nearly 
glabrous leaves, while plants from populations fur- 
ther north have narrower, abaxially pubescent 
leaves. other collections from Central 
America have leaves puberulent below, and such 


However, 


leaves are found over a long geographic distance 


Trends of this sort 
occur in other woody Boraginaceae, for example, 
Cordia curassavica (Jacq.) Roemer & Schultes 
(Miller, 1985), and are perhaps a result of a more 


with no clear discontinuity. 


pronounced dry season than occurs to the north 
in parts of Mexico. In Cordia curassavica much 
of the variation disappears in plants raised in the 
greenhouse (Miller, 1985). 
he second geographically correlated variation 
is that plants from the Tehuacán Valley in Puebla, 
Mexico, have leaves with smaller than usual ser- 
rations on the margin. This extreme of variation 
is not even consistent within Puebla and is not 
considered worthy of taxonomic recognition. 
Most variation in leaf morphology appears ran- 
dom, varying in form and indument on individuals 
and within populations. Five leaves from five in- 
dividuals (in some cases five individuals could not 
e found) in each population from various parts of 
the distributional range in Mexico were measured 
for length, width, length / width ratio, broadest point 
(calculated as distance from broadest point of the 
leaf to the base divided by total leaf length), apical 
angle, and basal angle (Tables 3, 4). In addition 
the indument on the adaxial and abaxial leaf sur- 
faces was noted. The leaves selected were at least 
three nodes below the apex of the shoot to avoid 
immature leaves and never adjacent to a branch, 
as these are often quite different in form. 
Field studies reveal considerable variation in leaf 
shape on a single tree, with leaf bases often varying 
from acute to obtuse and sometimes even cordate 


Volume 76, Number 4 
1989 


Miller 1065 
New World Ehretia 


TABLE 4. 
lower figures are standard deviations. 


Variation in leaf shape among Mexican populations of Ehretia latifolia. Upper figures are means and 


Leaf length/ 


Broadest point 
f leaf 


Population width ratio of lea Apical angle Basal angle 
Volcán Tacaná 2.13 0.44 13:19 109.25 
0.33 0.059 18.50 32.14 
Benito Juarez 1.60 0.45 96.80 140.40 
0.24 0.068 20.24 7.20 
Union Juarez 1.95 0.47 83.00 124.00 
0.16 0.049 16.91 13.56 
San Cristóbal 2.37 0.44 68.33 89.33 
0.27 0.049 18.32 18.96 
Zinacantán 2.69 0.50 51.00 66.00 
0.43 0.047 9.95 10.20 
Gueletao 1.81 0.43 70.80 137.20 
0.46 0.043 14.95 36.85 
Chicomezuchil 2:23 0.46 62.00 113.60 
0.32 0.054 13.27 20.91 
Monte Albán 2.10 0.41 61.80 131.60 
0.45 0.044 15.37 30.22 
Acatzingo 2.17 0.45 70.00 95.00 
0.20 0.029 10.49 13.04 
Chapulco 1.88 44 80.00 134.67 
0.22 0.040 17.22 27.41 
Ciudad Mendoza 2.46 0.44 57.80 109.60 
0.46 0.045 18.44 33:92 
Ixthuatlancillo 1.97 0.44 71.50 105.00 
0.25 0.060 14.67 28.28 
Tepoztlán 2.08 0.46 66.67 97.67 
0.22 0.078 8.88 15.37 
Campo Morado 1.97 0.43 75.80 119.40 
0.29 0.054 21.10 28.12 
Querendaro 1.93 0.39 67.40 143.40 
0.25 0.041 15.24 18.91 
Tiripitio 2.07 0.39 56.20 162.00 
0.31 0.055 8.40 33.88 


from one branch to another. Shape of the leaf base 
was the character used to distinguish Ehretia cor- 
difolia, but examination of leaves from different 
branches of a single tree collected near Guelatao, 
Oaxaco, Mexico (Fig. 22) shows this character to 
vary on a single individual. The same type of vari- 
ation in shape of leaf apices often occurs on single 
individuals, as exemplified by leaves from different 
branches of an individual collected near Motozintla, 
Chiapas, Mexico (Fig. 23). 

Most of the Mexican populations examined were 
relatively constant in terms of leaf indument, al- 
though occasional plants were more or less pubes- 
cent than surrounding individuals. There did ap- 
however, 


pear to be, a significant amount of 


variation in leaf shape among individuals from a 
single population. Leaves from the "p wire at 
Monte Alban (Fig. 24) ranged from 1.5 to 3.2 in 
leaf length/width ratio and 45? 
angle. The sampled population near Ciudad Men- 
doza (Fig. 25) had leaf length/ width ratios of 1.7 
to 3.4 and a range of 60? to 240? in basal angle. 
If this degree of variation can be found in a single 


to 100? eN apical 


population, leaf apex, blade, and base shape are 
not reliable characters for delimiting species. 
Mean values and standard deviation are given 
for characters of leaves from Mexican populations 
of Ehretia latifolia in Table 4. Leaf length/ width 
ratio, broadest point, apical angle, and basal angle 
appear to be the best measures of overall leaf shape. 


Annals of the 
Missouri Botanical Garden 


Volume 76, Number 4 
1989 


Miller 1067 


New World Ehretia 


€ 


FIGURES 24, 2 


row consists of five leaves from a single individual. — 24. 


\ 


Variation in leaf shape among five individuals from single populations of Ehretia latifolia. Each 


rom top to bottom, Miller & Myers 2817-2821 (MO). — 


25. From top to bottom, Miller & Torres 2971-2975 (MO). Scale bars equal 5 cm. 


Variation in leaf morphology among populations is 
overlapping in range, and no clear geographic pat- 
terns appear. 


Representative specimens examined. MEXICO, 
CHIAPAS: slope with Quercus and Pinus in the barrio of 
Chihk Ha’, paraje of Yashanal, Mpio. of Tenejapa, 6,000 
ft., Breedlove 9396 (F, NY); slope with Quercus and 
Pinus in the paraje of Kulak'tik, Mpio. of Tenejapa, 6,000 
ft., Breedlove 10947 (DS, F); steep slope along the river 
of Chihk Ha”, barrio of Yashanal, paraje of Matsab, Mpio. 
of Tenejapa, 5,500 ft., Breedlove 11110 (DS); in the 
barrio of Tuk, paraje of Matsab, Mpio. of Tenejapa, 7,500 
ft., Breedlove 12567 (DS, F, MICH); slope with Quercus 
and Pinus, paraje of Kulak' tik, Mpio. of Tenejapa, 5,500 
ft., Breedlove 12856 (DS, F, LL, US); 45-50 km NW 
of Huiztla along road to Motozintla, Mpio. of Motozintla 
de Mendoza, 1,900 m, Breedlove & Thorne 31080 (DS); 
SW side of Cerro Mozotal, 11 km ' of the junction 
of the road to Motozintla along the road to El Porvenir 


— 


and Siltepec, Mpio. de Motozintla de Mendoza, 2,100 m, 
Breedlove 31108 (DS, MO); second growth slopes paraje 
Chikinibalvo, Mpio. of Zinacantán, 1,200 m, Breedlove 
39606 (DS, MEXU); slope with Pinus and Quercus, 6 
km NNE of La Soledad along logging road from Las 
Margaritas to Campo Alegre, Mpio. of La Independencia, 
1,600 m, Breedlove 51268 (CAS, MEXU, MO, TEX); 
a 6 km al N de Oxchuc, sobre el camino a Ococingo, 
Cabrera & Cabrera 6146 (MEXU — 2 sheets, MO); steep 
slopes with Quercus near the paraje of Yal'ichen on the 
road to Chenalo, Mpio. of Chamula, 6,000 ft., Laughlin 
691 (DS, US); slope with Quercus and Pinus in the paraje 
"Apas, Mpio. of Zinacantán, 5,500 ft., Laughlin 1197 
(DS, NY); slope with Quercus and Pinus in the paraje of 
ft., Laughlin 


Sek'emtik, Mpio. of Zinacantán, 6,000 
12224 (DS, LL, US); slope with Quercus on trail from 
Zinacantàn, center to Ixtapa near Paraje Bo' pits, Mpio. 


of Zinacantán, 4,500 ft., Laughlin 2441 (DS, US); 
Ovando, Matuda 495 (F, MICH, US); hills E of Unión 
uárez, lower slopes of Volcán Tacaná 700-2,300 m 


nà, 1,700 : 
Miller et al. 2667 (MEXU, MO), 2668 (MEXU, MO), 


RES 22, 2 


cm. 


Variation in leaf shape on individual trees of Ehretia latifolia. —22. Miller & Myers 2802 
(MO). Note variation in leaf base. — 23. Miller et al. 2690 (MO). Note variation in leaf apex. Scale bars equal 5 


1068 


Annals of the 
Missouri Botanical Garden 


2669 (MEXU, MO), 2670 (MEXU, MO); along the dirt 
road to Ejido Benito Juarez and Siltepec, 4-6 km from 
the iyd aa Mexican highway 190, S of Motozintla, 
ca. 2,10 iller et al. 2690 (MEXU, MO), 
(MEXU Ho 2692 (MEXU, MO), 2693 (MEXU, MO), 
2094 (MEXU, MO), 2695 (MEXU, MO); along the in- 
rt road to Motozintla, hills above Unión Boe 
w slopes m Volcán Pu 
& Myers 2721 (MEXU, MO); along ‘Mexican a 
190 between San Cristóbal de las Casas and Tuxtla Gu 
tierrez » km 52, > se m, pie et al. 2793 (MEXU, 
MO), 2794 (MEXI , MO), 2795 (MEXU, MO); along 
the js road 4 km e of Zinacantán, 2,190 m, Miller 
et al. 2799 (MEXU, MO), 2800 (MEXU, MO); sumidero 
Coyota, Miranda 7752 (US); 3 km NW of 
4 , Thorne & Lath- 
"tik, Mpio. of Tenejapa, 
JERRERO: Campo Morado and 
hills above ca. 1.400 m, Miller & »s 2888 
(MEXU, MO), 2889 (MEXU. MO), 2890 (MEXU, MO), 
2891 (ME! O), 2892 (MEXU, MO). HIDALGO: Cerro 
de Me Isidro, Mpio. i Renesas 2,600 m, Ventura 
414 (MEXU). GUANA 
oom 25 (MEXU Y 
1095 (CAS, MEXU), Arguelles 1223 
ce Guanajuato, Duge s 153 (GH); shore ot us alkaline 
crater lake La Piscina de Yuriria, 1,730 m, Perrott 44a 
(BM): carretera Silao- ica, Cerro Culiacán, Vil- 
lareal de Puga 275 (WIS). JALISCO: vicinity of Guade- 
lajara, A, 500 m, Diquet s.n. (MICH — 2 sheets, NY, P, 
US) E 5 of La Barca, Gregg 841 (MO —2 sheets); Chi- 
quilistlan, Jones 78 (F, MO, US) vicinity of San Juan 
Cosalá, mountainsides above (N of) Lake C 
2,200 m 
Ciudad Ciena, 
4875 (US). MÉXIC 0: dist. e Cumbre Teju- 
pilco, 1,500 m, Hinton 2837 (A, G, GH, K, 
cera ms Nanchititla, Hinton 3419 (A, BM, G, 
GH, NY, US); Mpio. Tezcoco, S icolás ^ 
7 km al E de Tezcoco, 2,600 m, Koc h 821 (CAS 
MIC H, MO, NY, TEX, WIS); Tlatlaya y cercanias, Ma- 
tuda et al. ta (MEXU, MO); Tlapacoyan, Mpio. de 
Chale m, Rzedowski 26798 (CAS, LL, MICH): 
Urbing s.n. e ); San Nicolás Tlamincas, Mpio. Tex 
coco, 2,300 m, Ventura 354 (CAS, NY, WIS); 2,400 
m, Ventura 793 (MEXU). MICHOACAN: Morelia, 2,100 
m, Arséne s.n. (G); Tócuaro, Mpio. Erongaricuaro, 2,000 
m, Bello 86 (MEXU); W of Morelia; Gregg 778 (MO); 
Patzcuaro, 1,850 m, pe 18199 (TEX); 25 mi. W 
Hitchcock & Stanford 
3 Tit, Mpio. Morelia, 1,950 
- Madrigal 2852 (M ; Tócuaro, Mpio. Erongari- 
cuaro, 2,000 m, Vu a 18(MEXU ); along Mexican 
highway between Pe and El Oro, 3 km E of Quer- 
132, 1,870 m, Miller & Telléz 3059 
, 3060 (MEXU, MO), 306 1 (MEXU, MO), 
, 3063 (MEXU, MO); just 5 of Tir- 


, Puga s.n. 


9 
:0, 2,25 


T 
D 


x 


J], MO), vaa: (MEXU, MO), . 
- MO) 3068 (MEX MO); en la desviación a 
C ik 21 km al SW de M carretera a Guadalajara, 
2,140 m, Soto 2382 (C i , MEXU); a 15 km de Ciudad 
Hidalgo, rumbo a Zitcuaro, 1,830 m, Soto 6273 (CAS, 
MEXU, MO). MORELOS: ee Arquelogica dee in 
the Cerro Tepozteca above Tepoztlán, ca. 30 
Cuernavaca, 1,700 m, Miller & Torres 2996 (MEXU : 


— 0 
—- 


MO), 2997 (MEXU, MO), 2998 (MEXU, MO); Chap- 
ultepec Springs near Cuernavaca, 5,000 ft., Pringle 8748 
A, , BM, F, G —sheets, K, L—2 sheets 
MEXU— 2 sheets, MO, NY, P, UC, US), 10261 (BH, 
PM; CAS, F, I í 


(NY, US) « XACA: Oaxaca de Juárez, Bailey 540 (BH, 
MICH); near rione Albán, Camp & Conzatti 27 781 (NY); 
Dist. de Ixtlán, us 175 Tuxtepec a Oaxaca, Guelatao 


de Juárez, 2,200 m, Cedillo & Lorence 643 (MEXU, 
MO); al 500 m : SW de la desviación a La Ascunción, 
carretera a San Agustin Atenango, dist. de Silacayoapán, 
1,980 m, Cedillo & Torres 1449 (MEX U); Rio el Molino, 
4.5 km al N de Ixtlán, camino a Calpulalpán de Méndez 

2,000 m, Cedillo & Lorence 2324 (MEXU); Cedillo & 
Lorence 2330 (MEXU T. a 12 km al SE de Dominguillo, 
por la carretera a Oaxaca, Chiang et al. 157 (MO), 
Monte Albán, “Conz atti 146 (GH); 
San Felipe, 1,800 | m Cons atti 210 (GH, K); Monte 
Albán, 1,700 m, Conzatti 4848 (MIC H); Conzatti 4918 


(CAS); 1,380 m, Conzatti 5245 (N Y); ruins at 
Monte Albán, Croat 935 (MO); Dorantes Plateau , 9,900 
ft., Galeotti 3099 (K), 1 km al SE de - Tamazulapán, 
camino a San Jerónimo, Dist. de Teposcolula Mixteca 


pos 
7 (MEXU); SEDE definite 


2,000 m, García 937 
locality, Kenoyer 4660 (F); Monte Albán, Kenoyer 1541 
(GH); Oaxaca, Liebmann 12722 2); ist. a 2 


Sierra de gr bois de ión a a Xia 

ceda et al. 3 (BM, MEXU, MO); iron rid 

6,000 Messer m 3 (MICH); along Mexican highway 
175 at peA 159 just SW of Guelatao, 2,200 m, Miller 

& Myers 2801 (MEXU, MO), 2802 (MEXU, MO), 2803 

(MEXU, MO), 2804 (MEXU, MO), M MO); 


along the dirt road to C hicomezuchil, l km SE of Mexican 


highway 175, 1 W of Chicomezuchil, ca. 2,200 m, 
Miller Myers 2806 (MEXU, J 


MO), 2808 (MEXU, 2809 (MEXU, MO), 2 
(MEXU, MO); Monte Albán, short deciduous forest sur- 
rounding the ruins and adjacent roadsides, 1,800 m, 
Miller & Myers 2817 (MEXU, MO), 2818 (MEXU, 
MO), 2819 (MEXU, MO), 2820 (MEXU, MO), 2821 
(MEXU, MO), 2822 (MEXU, MO), 2823 (MEXU, MO); 
gi Albán, 5,800 ft., ei ud 6138 (A, BH, BM, C, 

2 sheets, L, MO, NY, P, UC, US —3 sheets), 6 / 36a 
"à BH, BM, G, GH, K, MO, NY, P, UC, US—2 Serie 


UC, US); Dd San Felipe, l, 800 n m, Tine et Sal s.n. 
(MEXU); cerca del Puente de Rio Grande, Mpio. de San 
Juan € emend 1,400 m, Ventura 16098 (MEXU, 

NY). PUEBLA: aprox. 12 km al NW de Tehuacán, rumbo 
a Orizaba, iu et al. F-280 (MEXU, MO); San Nic- 
olás, Tehuac 2,000 m, Conzatti 2221 (F, GH—2 
sheets); TM Liebmann 15166 (F); 5 km al E de 


234, 

Miller & Torres 2957 (M MO); along the road in 
between Tecamachalco a añada Morelos, at Chapulco, 
6 km from Cañada More 2,000 m, Miller Torres 

2960 (MEXU, MO), 2961 (MEX CU, MO), 2962 (MEXU, 
oes serge d San Luis dg ocius Purpus s.n. (BM), 
(BM, , GH, MO, UC sheets, US), 3082 

(BM— 2 eats TF. GH, MO, NY, UC —2 sheets, US); 
Tlaucuislaltepec, Purpus 3976 (L, UC), cerros near San 
Luis Tultitlanapa, Sierra Mixteca, Purpus 4215 (L); near 


Volume 76, Number 4 


Miller 1069 
New World Ehretia 


Oaxaca, in the vicinity of San Luis dere. Purpus 
4249 (DS, F, G, GH, L, P, UC, US), 4251 (BM, F, GH, 
MO, UC, US), 4252 (BM, F, GH, dos P. UC, US); El 
Riego, near Tehuacán, Purpus 5818 (BM, UC); Tehua- 
cán, Purpus 5820 (BM, F, GH, MO, NY, UC, US); road 
Puebla-Tehuacán, Reko 10 (F); Nacozcalco and San 
Antonio Canada, along the Barranca de los Mangos, 1,000- 
1,800 m, Smith 4087 (F, G); Anumbila, Smith & Tejeda 
4530 (US); Chapulco, J. E. Smith F-9 (MICH); 2 
9 km al NE de Tecamachalco por ee carretera a 

Morelos, Villasenor et al. 143 (MEXU). VER pees "Ori. 
zaba, Botteri 1022 (BM, a 1322 (Py Bourgeau s.n. 
(GH, K, P); steep slopes of high valley above route 150, 
4 km W of Acultzingo, 5,500 ft., Long & Burch 3300 
(F); Acultzingo, d 1106 (LL, MEXU, MICH); along 
Mexican highw O at the turnoff to Puente Colorado, 
hills above Ciudad Md ca. 2,000 m, Miller & Torres 
297 | (MEXU, MO), 2972 (MEXU, MO), 2973 (MEXU, 
MO), 2974 (MEXU, MO), 2975 (MEXU, MO); Ixhu- 
atlancillo and vicinity, ca. 10 km N of Orizaba, ca. 1,500 
m, Miller & Torres 2992 (MEXU, MO), 2993 (MEXU, 
MO); Orizaba, Muller s.n. (NY, , 502 (L). STATE 
UNKNOWN: without definite locality, Gregg 778 (MO); 
Sessé et al. 767 (F), 772 (F), 5246 (F), 5248 (F). Gua- 
TEMALA. ALTA VERAPAZ: 3 km de Villa Hermosa 1,400 
m, Molina & Molina d (F, NY). AMATITLAN: San 
Vicente Pacaya, 1,400 Tonduz 472 (G, MO, US). 
BAJA VERAPAZ: Chilasco, EOS 10978 (LL, 5); along 
dirt road 4 mi. NE of Purulhá, 1,500 m, Croat 41360 
(MO); Mpio. Rabinal, at summit of Sierra de Chuacús, at 
border of ‘Mpio: Rabinal and Mpio. El Chol, along road 
pes at summit, 1,800 m, Croat & Hannon 
63635 (MO). 63656 (MO); ^ 8 km S of Rabinal, Harmon 
& Dwyer 3178 (MICH, MO). EL QUICHE: below falls of 
Río de las Violetas, ca. 3 mi. N of Nebaj, 5,300-5,500 
ft., Proctor 25547 (IJ, L Fe pA Gu ATEMALA: with- 
out definite locality, Aguila 7 (F); 
Andrecillo, 1,700 m, Molina & MS 275 57 70 (F, MICH). 
QUEZALTENANGO: region of Las í 
Chile Verde, 2,250 m, Standley 85162 (F); 
Chicharro, on. lower southeast facing slopes of Volcán 
Santa María, 2-4 km S of Santa María de Jesús, 1,400 
1,500 m, Steyermark 34279 (F, NY). SACATEPEQUEZ 
near Antigua, 6,500 ft., Kellerman enin (F, M ICH, NY, 
US) nacimiento del Cuan Cues e 
1,650 m, Molina 1. Pod GH); near “Pastores, 1,560 
LADA m, Standley 5 1 (F); barranca above Du A 
1,590-1,800 m, ond wow B (Ey near Barranca 


Hondo, SE of Alotena O m, Standley 
64959 (F). SAN MARC c bod cas 6 mi. S and W of 
town of Taal, northwestern slopes of Volcan Ta- 


jumulco, 2 2,300 Om, Steyermark 30007 (F); above 
‘inca El Porvenir, between 


El P r, on Potrero Matasano along Rio Cabús, 
Volcán Tajumuleo, 1,000-1,300 m, Steyermark 37631 
(F); Potrero Matasano, Finca El Porvenir, slopes of Volcán 
Tajumulco, 1,200 m, Steyermark 52 124a (F, GH). oy TE 
UNKNOWN: without definite Wim Heyde 572 (US). EL 
SALVADOR. LA LIBERTAD: a Tecla, Calderón OU 
(GH, NY, US); vicinity of uie Tecla, 790-950 m, 
Standley 23081 (GH, NY, US). SAN SALVADOR: Cerro de 
, NY, US — 2 sheets); 
H, US); Santiago de Maria, Calderon 
4 Pe X Cano de San Jacinto, near San Salvador, 


71 m, Standley 20630 (GH, NY, US). 


2 


SONSONATE: highway to Cerro Verde, area S of Lake 
Coatepeque, 3 300 ft., Allen 7072 (F, G, GH, LL, NY — 
2 sheets, US). STATE UNKNOWN: Berlin, Calde pron 2144 
a without definite p. pre 332 (US — 2 sheets); 
Volcán de San Salvador, 1,0 1,800 m, Standley 22884 
oe NICARAGUA. GRANADA: eae Mombacho, 1,100 
ios 24363 (MO). JINOTEGA: vicinity of Jinotega, 
L 03 -1,300 m, Standley 9689 (F); 
rod and Las Mesitas, in Sierra W of Jinotega, about 
1,400 m, Roh 10403 (F). MANAGUA: Sabana 
na Managua, 90-100 m, Suárez 99 (MO). 
A. ALAJUELA: Bella ous de Zarcero, 1,800 m, ausim. 
Smith 139 (F, GH —2 sheets, NY); Zarcero, 5,500 ft., 
Austin-Smith 4384 (F —2 sheets); Zarcero, Alfaro Ruiz, 
1,800 m, Austin-Smith P2500 (GH, UC); Zarcero, Aus- 
tin-Smith 2702 (F); Zarcero, Alfaro Ruiz, 1,900 m, Aus- 
tin-Smith 10048 (F, GH). CARTAGO: La Carpintera, Ech- 
everria 402 (CR, F, UC); cerca del quebrador Ochomogo, 
de eda 459 (CR); Cerro de la Carpintera, 1,500-1, tm 
, Standley 34518 (US); Cerro del Aguacate, La € 
ne. 1,500 m, Williams 11578 (CR, F); on open 
slopes of Cerro Carpintera above Tres Rios, 1,500 
Williams 16156 (BM, F, MO, US); Cerro del Aguac te, 
a Carpintera, 1,500 m, Williams 16578 (F). HEREDIA: 
ane road N of Gethsemini de Heredia, 1,500-1,700 m, 
Hartshorn 1208 (CR, F, MO); en pag al Oeste del 
Monte de la Cruz, San Rafael, 1,700 m, Jiménez 2513 
F). PUNTARENAS: Monteverde, b community, l, 450 
m, Haber 203 (CR, MO); Santa Elena, 5 kr [ 
Elena on road to highway, Haber 1775 (MO); 
Verde, along Río Guacimal below Lechería, 
Hammel & Trainer 13837 EE SAN JOSE: ' 
Córdoba 317 (CR) vicinity of S 
it l, ee m, Standley & Vale orio 44.145 (US). PAN- 
a de Bambito 


— 


a dus llanos de a 6,000 ft., 
MO); at opening to canyon to Bambito, 5,000 ft., 

5870 (MO— 2 sheets); Rio oo Viejo valley, 1 
Bambio, P. White 220 (F, GH, MO). 


& x4 
» 5 


ear 


Ehretia tinifolia L., Syst. Nat. ed. 10. 2: 936. 
1759. TYPE: from Patrick Browne's Jamaican 
herbarium purchased by Linnaeus in 1758 
(lectotype, here designated, LINN (Savage 
Catalog number 254.1); microfiche, MO; pho- 
to, F; isolectotype, BM). 


Ehretia ER ed Miers, a & Mag. Nat. Hist. Ser. 
IV. NTYPES ny ISOSYNTYPES: In 
nerd et vido amaica, in herb. Hook., Lane 


: Meeker (BM, K); Mexico, Oa- 


vins a 3,000 ft., Galeotti 7194 (G, 
Fhreti ala Miers, a & Mag. Nat. Hist. Ser. IV. 
869. SYNTYPES & ISOSYNTYPES: in Antillis, 
herb. Mus. Brit. E identifie , herb. 


Sloan. Gis vii. Fol. 
cult., Jamaica, Houston (BM); i 
maica, Purdie (BM); Cuba, La Sagra (K, LE); C uba, 
Havana, Greene (BM, K). 

Tree to 15(-25) m tall, the twigs essentially 
glabrous. Leaves persistent; petioles (3-)5-10(-14) 
mm long, glabrous canaliculate adaxially; leaf blades 
elliptic, (5-)6.5—12(-16.5) cm long, (2.5-)3-6(-8) 
cm wide, the apex obtuse to rounded, the base 


1070 


Annals of the 
Missouri Botanical Garden 


obtuse to acute, the margin entire, adaxially gla- 
brous and lustrous, abaxially glabrous, the lateral 
veins 6-7. Inflorescence terminal, paniculate, 7— 
15 em long, the branches glabrous. Flowers bisex- 
ual, sessile or short-pedicellate; calyx campanulate, 
1.5-2 mm long, the 5 lobes ovate, 1.5-2 mm long, 
0.7-0.9 mm wide, glabrous externally, ciliate along 
the interior margin; corolla white, tubular-cam- 
panulate with 5 reflexed lobes, 4-4.6 mm long, 
glabrous on the exterior surface, the lobes widely 
oblong to oblong-ovate, 2.5-5 mm long, 1.3-1.7 
mm wide, the tube 1.3-2.1 mm long; stamens 5, 
the filaments flattened, 3-4.5 mm long, the upper 
2-3 mm free, glabrous, the anthers ellipsoid, 1— 
1.3 mm long; ovary ovoid to broadly ovoid, 1-1.2 
mm long, 1 mm broad; style 2-3.4 mm long, 
divided for 0.5-0.7 mm, the stigmas truncate. 
Fruits drupaceous, yellow-orange, glabrous, broad- 
ly ellipsoid, 5-7 mm long, 4-6 mm broad, the 
endocarp dividing into 2, 2-locular pyrenes, these 
ridged-reticulate on the outer surface. 


Distribution. | Ehretia tinifolia occurs from 
southeastern Mexico to Honduras and on Cuba, 
Hispaniola, Jamaica, and the Cayman Islands in 
lowland, moist to wet forests from sea level to 
600(-1,500) m (Fig. 26). Collections cited from 
Guerrero, Michoacán, Nayarit, San Luis Potosi, 
and Sinaloa are not included in the distribution 
map as they are presumed to be out of the natural 
range of the species and are probably from culti- 
vated plants. 


Ehretia tinifolia is distinguished from the other 
New World species by its glabrous, lustrous, entire- 
margined leaves and elongate, paniculate inflores- 
cence. It is widely cultivated, even outside of its 
natural range for its attractive flowering and fruit- 
ing display and as a shade tree. 

The two species described by Miers, Ehretia 
longifolia and E. sulcata, are both clearly syn- 
onymous from Miers's description and their West 
Indian distributions. Unfortunately, Miers cited 
specimens with little information and these collec- 
tions remain difficult to identify. For this reason, 
these names have not been lectotypified, and syn- 
types are cited here. 


Representativ e spec imens examined. MEXICO. CAM- 
PECHE: a 3 km al sur de la carretera Ch ampotón- “Cam. 
peche, sobre el camino a El Zapote, Cabrera & Cabrera 
MO); sobre el camino a Pixoval, a 30 km 


s & Chan 
87 15 (F); environs de Campeche, Linde on 107 (K); Campo 


Experimental, Escarcega, 100 m, Marroquin 294 Pd 
45 km al NE de Campeche, camino a Merida, m, 
Martinez et al. 3011 (BM, CAS, MEXU); ed in 
garden, Seler 4016 (F); near Xujil, 300-400 m, Shep- 
herd 93 (LL, MICH —2 sheets, WIS— 2 sheets); He- 
celchakan, Stewart 52 (GH). CHIAPAS: in the sitios or 
along the streets of Ocozocuautla, Mpio. of Ocozocoautla, 
580 m, Breedlove & McClintock 53514 (DS, MEXU, 
MO); along the streets or in the sitios of Tonala, Mpio. 
Tonala, Breedlove & Thorne 30658 (B, DS, F, LL, MEXU, 
MICH, MO, NY); cultivated, dry open slope with tropical 
deciduous forest, 32 km NNW of Soyalo, along the road 
to Copainala above Chicoasen, Mpio. of Chicoasen, 450 
m, Breedlove 34039 (DS, MEXU, MO); Mpio. of Cin- 
talapa, 23 km W of Las Cruces along the road to La 
Mina microwave station, 870 m, Breedlove & Davidse 


1578 3 a Eeuna cultivated, Matada 16607 (F, 
MEXUJ); Paderón, Tonala, Matuda 1695 1 (F— 2 sheets, 
MEXU); Encanana, arriba Chancona, N Tuxtla Gutierrez, 
Miranda 5263 (MEXU, US); on plaza at Chiapa, Sou- 
viron & Erlanson 103 (US). GUERRERO: Iguala, highway 
95, La Cabana Motel, cultivated, Freeland & Spetzman 
33 (MEXU — 2 sheets); Motel “Costa Azul," cultivated, 
Lorence & WARS 3511 (CAS, MEXU, MO); along 
Mexican highway 95 at Tierra Colorado, ca. 800 m, 
Miller & mid 2954 (MEXU, MO); jadis Botánico, 
Chilpancingo, 1,300 m, cultivated, Toledo & Blanco 50 
(MEXU). MICHOACAN: 1 km S of Nueva Italia along Mex- 
ican highway 37, cultivated at gas company, c 


Miller & Téllez 3079 (MEXU, MO). MORELOS: 


O 
3 


1,050 m, Gutiérrez M. 254 (MEXU); Yautepec, alrede- 
dores del poblado Ignacio Bastidas, 1,520 m, Muskus 40 
MICH); Cuernavaca, carretera a Acapulco, Palacios s.n. 
(DS, MICH, WIS) Villa de Ayala, 1,260 m, Palacios 
EXU); Mpio. Yautepec, Ignacio Bastidas, 1,520 
m, finn 43 (CAS). N e , Acaponeta, Martínez 7 
y Isletas, 5 km E of San Blas on the road to 
M sea level, Miller & od 3194 (MEXU, MO). 
OAXACA: Union Hidalgo, Bailey 572 (BH, MICH); Puerto 
Escondido, Boege 2642 (GH, MEXU); Pinotepa Nacion- 
al, Boege 2660 (MEXU); Cuicatlán, Conzatti & Gon- 


zalez 38 


a 
em 
2 
= E 
S 
5 
fa 
ES 
du 
CERE 
= 
feb} 
s 
un 
ho 


and San Sebas 280 m, Croat 45768 (MO, 
1 Oaxaca, Davis & ene s.n. (TEX); Cuicatlan, 

600 ft., González 38 (GH); El Carrizal, E 2 km al SW 
iE Mono de Mazatlán, o sea a 35 km a / de Salina 
Cruz, sobre la carretera a Pochutla, Mpio. de Tehuan- 
tepec, a ález 473(MEXU, m Extalpec, Mell 2145 
(NY, US); Oaxaca, park in town, 1,600 m, Dd 
Miller & Myers 2824 (MEXU. MOL cerca y a S de 
Cuicatlán, Miranda 4544 ee — 3 sheets); Tlacolula, 

(DS); € ng ide ft. — 

XU, US); Poshutla, m, Re ko 3 3 

(US); Huatulco, 150 m, Reko 6004 co p 
74(NY—2 sheets, US); Cacahuatepec, cultivate an- 
la, Sousa et al. 5216 (CAS, US); Salina Cruz, 0-300 m, 
Williams 9736 (F, GH). QUINTANA ROO: en la brecha de 
Divorciados a la Pantera, por la via corta a Mérida, 
Cabrera & Alvarez 1678 (BM, MEXU—2 sheets); a 1 
km al W de Ciudad Vallarta, sobre el camino a Leona 


sa 14 km al NW de 


rem 
© 
= 
= 
5 
Aa 
~ 
a] 
~ 
oa 
~ 
S 
mn 
mn 
~ 
= 


Vicario, Cabrera et al. 2577 (MO) 


Volume 76, Number 4 
1989 


Miller 1071 
New World Ehretia 


CARIBBEAN AMERICA 


o 800 kms 


TM — 


—— | = 


RENT e 


FIGURE 26. Distribution of Ehretia tinifolia L. 


Calderitas, sobre el camino a Laguna Guerrero, Cabrera 
& Cabrera 3223 (MO); a 5 km " W de La Pantera, 
sobre el camino a Margarita Maza, sobre la carretera via 
corta a Mérida, Cabrera & Cabrera 3320 (MO); a 12 
km E E de San Felipe Bacalar, Cabrera 4527 
(ME — 2 sheets); ee de € hichankanab, Mpio, José 
M. Me 5m, C ; Cobá, 
Lundell & Lundell ; iod (LL, MICH, US); 7e 
(BM, MEXU, MO, NY); Ruinas de Cobá, Téllez 2349 
(MEXU, MO); m & Cabrera 2365a e MO); Bal- 
neario Alvaro Obregón, a 25 km al sur de Mrd Téllez 
2375 (MEXU, MO). san LUIS POTOSÍ: cultivated, Ciudad 
Valles, Hansen & Nee 7357 (F, MO). TAMAU LIPAS: Ejido 
No che Buena, arias su del Hio Soto o La Mene 50 
drano 262 
33 km al E xis Sot La Mai 
ricitas, Mpio. Sot 
2635 (MEXU); al NW de E 
Soto La agri 100 m, Martínez 1 (MEXU); 7 
NW de Cachimba (El Moron), Mpio. de Aldama, 50 m, 
iios 220 (MEXU- 2 sheets, vu pow of Tam- 
1, CAS, F, 

CAS, GH. K, MO, s Chanal, 
Waoren s.n. (US). siNALOA: Culiacán, cultivated Clarke 
et al. 1328-1 (MIC H). vERACRUZ: Tierra Caliente, camino 
Zampoala, Actopan, Aporte et al. 24 a ; Zapotal no. 
1, 5 m, Calderón 1301 (CAS, F—3 sheet ts, MEXU, 
MICH); Sais Andres Tuxtla, parque central, Calzada 833 
F); carretera Tuxpan- Panuco, 5 km antes de Tantoyuca, 
Chiang 3 (CAS, GH, MEXU); a 38 km de Tepetzintla, 
hacia Tantoyuca, Chiang 380 (CAS, F —2 sheets, GH, 


MEXU); Puente Nacional, carretera Xalapa- Veracruz, 
Emiliano Zapata, 250 m, Dorantes 549 (F, MEXU); 
ladera ne del Cerro ue Los M , Dorantes 
; Actopan, 450 m, C 
IEXU, N 

7 km SE of Emiliano Zapata (— Car 
Mpio. in ip m, Hansen & Nee 7492 (F, MO); 
Mirador, Liebmann 15 (US); Ruta 140, ca. 2 km NW 
of San Andres Tuxtla, orilla de la carretera, probabla- 
mente cultivado ice & Ce 'dillo 3136 (MEXU); La 


Bandera, 10 km al SE de Actopán, Márquez & Dorantes 
49 MEXU, MOX Platón Bine hez, Medillín s.n. 
(MEXU 2 sheets); in the city of Veracruz, Nee & Taylor 


19625 (NY); Ln Mpio. Jalapa, 1,308 m, cultivated 
in the city, Nee 26575 (F, G, LL, NY); Baños de Carrizal, 
5 km SE of Emiliano Zapata, eg Puente Nacional, 
220 m, Nee & Taylor 26610 (F, LL, NY); norte de La 
Boca de Laguna de la Mancha (1 km) Actopan, ` level, 
Novelo 402 (MEXU); Playa Paraiso, 2 km al N de La 

La Laguna de La Mancha, PN sea level, 
Novelo 421 (MEXU); km 43 carretera La Concha-Ac- 
topán, Actopán, 190 m, Ortega ndn MEXU, MO, 
NY —2 sheets); La dE Jilotepec, 1,000 m, Or- 
tega 324 (BM, F, G, MEXU, MO); ali of Pueblo 
Viejo, 2 km S of d uo Palmer 431 (BM, CAS 
G, GH, MO, NY, US), 446 (GH, MO, NY, US); Re- 
muladero, Purpus 8753 CH. MO, NY, UC); shore of 
os, near Puente Nacional, Purpus 11168 
2 sheets, UC, US) 
33 (F) camino Atoyac 
Velázquez 285 (F 


Rio los 


f. 
plan del Rio, 
u 


Tepatlaxco, ER 800 m, 


1072 


Annals of the 
Missouri Botanical Garden 


MEXU, 


Ventura 


MO); Plan del Rio, Mpio. Dos Ríos, 265 1 
3738 (CR, DS, MICH, US, WIS); El rex 


Mpio. Dos Rios, 200 m, lentura 8505 (MEXU); Las 
lrancas, Mpio. de Dos Rios, 1,100 m, asada 10264 
(MEXU); La Bocana p Actopan, con la carretera Xala 


apa 
Mpio. de Dos Rios, , bentura 11021 (MEXU); 


Puente Nacional, Mpio. de Puente Nacional, 100 m, 
Ventura 12623 (MEXU); La Ceiba, Mpio. de Puente 
Nacional, 200 m, Fentura 17238 (MEXU — 2 sheets, 


MO), plan del Rio, Mpio. Emiliano Zapata, 300 m, Vil- 
lanueva 116 (NY, WIS), 123 (NY, WIS) La Orduna, 
carretera Xalapa ug Cükténgo, 1,100 m, Zola et 
MEXU — 2 sheets, NY); La Concepción, 1,000 
) 3 ria NY). YUCATÁN: 
Bequaert 52 (F, US); Bruff 1465 
(MEXU); I x Ake 30 km SE us Merida. Butterwick 
141 (F, 3 km al N de Tikal, por la carretera a 
Mérida, aa ra p Cabrera 2376 (MEXU 2 sheets, 
MO); Peto, Castillo s.n. (MEXU — 2 sheets); ruinas de 
Labna, al sur del Mirador, Takax, 80 m, Chan et al. 256 
(MO); 2 km N of the border between Yucatán and Cam- 


80 m, Davidse et al. 20603 (MO); Tunkas, Enríquez 
538 (MEXU); Mérida, 8 m, Flores 8108 (MEXU, MO); 
Izamal, md sn. F, K); without definite locality, 


, DS, F, GH, LE, MICH, MO 


sheets, MON 404 (A, BM, F, GH, MO, NY, US 
WIS); T. de E Gaumer 1721 (A, B, BM, F, GH. 
LE, MO, US); SE Kancabonot, Gaumer et al. 23868 


(DS, F, G, GH, MO, US); Chicher | Itzá, Lundell 7386 
(DS, F, GH, MICH, NY, ; Camino Colonia 
Yucatán- Tizimin, 100 m, Pe pner & Sarukhan 9402 
(A, MEXU, NY); Labna, sur del vs rm o menos a 30 
7 Mpio. Te lérida, Roble 29 
WIS); Schott 139 (BM), 800 (BM: FX Seler 3848 (F. 
6 H); Terul, Seler 3866 (F); Mérida, Souza-Novelo 254 
(US); Chichén Itzá, near Hacienda de Chichén, Steere 
1468 (MICH); without definite locality, Steggerada 32B 
F); Cenote de X-Kikil, Mpio. X-Kikil, m, Ucan 
Burgos 1195 (F); oe ues locality, Valdez 42 (B, 
F, GH, LL shee O, NY —2 sheets, 
IS); Tixcac dom Mpio. Yaxcaba, 25 m, Vargas & 
rd 421 (F); Ruinas de Labná, 28 km al SW de o 
cab, Mpio. Oxkutzcab, Xelhuantzi 5436 (MEXU). B 
LIZE. COROZAL: d definite. locality, Gentle 40 (F. 
4793 (F, GH, Y). ORANGE WALK: Maya ruin, 
Indian Church, Arnason & Lambert 17026 (MO). Gua- 
TEMALA: GUATEMA : Guatemala, Aguilar 530 (F); Par- 
ques de Guatemala, Salas US); cultivated in Gua- 
temala ! ‘TEN: shoreline 


c 


— 


"d 


Harmon & Dwyer 2766 (MO, NY, US). sACATEPEQUEZ: 
near Antigua, 6,500 ft., Kellerman 7426 (NY, US). 
NDURAS. COMAYAGUA: La Libertad, Caballero 155 (NY); 

Valle Comayagua, orillas del Rio Humuya, 700 m, Molina 
0771 (F, G, GH, NY, US); a la salida de Comayagua, 
aped, a Siguatepeque, 400 m, Molina 13695 (F, LL, 
NY, US); El Bar 10 m, Rodríguez 2340 (F); El 
sae :0, 640 m, Rodrigues z 2586 (F —2 sheets, MEXU); 
vicinity of Comayagua, 600 m, Standley & Chacón 5212 
(F, GH, N anco, ds 2580 (GH). EL PARAISO: 

Jamastrán entre Río Los Almendros y 
Molina iL. 393 (BM, F, G, LL, NY 
US); Vegas y matorrales de Río Los Almendros, ‘Valle 
Jamastran, 400 m, Molina 13769 (F, LL, S). 
OLANCHO: orillas de Santa Maria de Real, Valle di Ca- 


, 


tacamas, 500 m, Molina 8416 
ticalpa, 380-480 m, Standley 
n. 400 500 m, 
SWAN ISLANDS: Little S 
Island, Proctor . 32495 (BM, F , GH, J. b. fee ie 
Chandler s.n. (GH). YORO: Ciudad de Yoro y alrededores, 
040 m, Nelson & Martínez 2037 (MO). CUBA. HAVANA: 
roadsides near Havana, Curtiss 724 , BM, E, G, 
GH, MO, NY, P, US); Cabaña, mana 280 (F, G, S, 
US); Havana, below Cabañas Fortress, Jack 4005 (A); 
Cacahual, Alain 4247 (GH, US); near Tapaste, E of 
Habana, Morton 10718 (US); Buena Vista, Schafer 243 
Y); Santiago de las Vegas, Van Hermann 781 
(NY); Fortaleza de la Cabana, Van Heimann 1951 (NY); 
El Morro to Cojimar, Wilson 9131 (NY). ISLA DE PINOS: 


F, MO); BEI: EE Ju- 
y 17604 hs 
V Aid us 


— 


Vivijagua, Britton & Wilson gn (NY). LAS VILLAS 
Farallones de Guajimico, on the t E of Cienfuegos, 
er 10471 (US), 10481 (US) 10491 (US). M: 


NZAS: dry thickets, C in; Britton & Wilson 160 
(NY) Canasi to Boca de Canasi, UR 102. 33 (US); ad 
montes supra Matanzas, Rugel 3: 20 (F, LE, — 2 sheets). 


ORIENTE: vicinity of reas Britton 2277 (NY 
vicinity of Santiago, Britt et al. 914 (MO, NY); 


Ensenada de Mora, Britton et al. 
thickets, Ciudamar, Santiago, Clemente 5331 (GH, U* S); 
coastal thickets, Sardinero, Clemente 5937 (GH — 2 sheets, 
US); Jiguani, Clemente 6064 (GH); coastal thickets, Sar- 
dinero, Santiago, Clemente 6106 (G i 
Ekman 1434 (MICH, S); 
oi on the coral reef, _ 45 
, Monte Oscuro, Ekman 4607 
ay vate, ts del Rey, Ekman 4076 (S); THE in i 
Ekman 6225 S); Banas, Ekman 6572 (S); roadsides 
near La Cienaga, Santiago, Ekman H13235 (LD); ponte 
ros de la Finca "La Nena" Laguna Blanca, cercanias de 
Bayamo, López 648 (Us ; 
Bahia de Santiago de Cuba, López 914 
Marie-Victorin oor (GH); c to Myabe, Shafer 
1415 (F, NY, US); Holguin to Caco Shafer 1526 
(F, NY, US). PINAR DEL RÍO: San Diego de los Baños, 
along stream, Britton et al. 6650 (F, NY); on limestone 
rocks, Mogote de José Maria, Viñales, Alain 2928 (GH 
US); in ER: near El Guama, Palmer & Riley 412 
(US); limestone hills between Rio Cayaguete and Sierra 
Guane, Shafer 10471 (NY); Sierra Caliente. S of Sumi- 
dero, Shafer 13734 (A, US). SANTA CLARA: vicinity o 
Cienfuegos, Britton & Wilson 56009 (NY): district of 
Cienfuegos, Combs 176 (F, GH, MO, NY); Gavilán, How- 
"e 4945 (GH, NY); Dolores pasture, Howard 5410 
(GH, NY); open fields at Gavilán, Howard 6311 (GH, 
NS Soledad, ee MEA 4991 (A); Belmonte, So- 
ledad, Cienfuegos, Jack 5106 (A, US 5); Soledad, Cienfu 


Baracoa, 


fuegos, Jack 6005 (A, G, K, P), 6029 
Belmonte, pou Wu Jack 7153 (A, 
ilán, Jack 78 , BH, F, S); Soledad, Cienfuegos, 25 

STATE UNKNOWN: Vedado, Baleer 


B 24 (BM, L), 37 (BM); Cook s.n. (NY); Ingenio 
Capitolio, a 60 AA e Redondo ad Ronulie, 
Eggers 4663 (A, F, K, P, US); Trinidad Mountains, San 
Blas Buenos Aires, How Hd 527 4 (GH); without definite 
locality, La Sagre s.n. (B, K, Pj Linden 1983 (B 

2 sheets, G, K —2 sheets, P); Rugel 326 (BM, L); Ata- 


laya, Camaguey, Shafer 986 (NY); without definite lo- 
cality, Wright 56 1 (S); prope villam Monte Verde, Wright 
1366 (G, GH, K, MO), 1370 (BM, G, GH— 2 sheets, 


Volume 76, Number 4 
1989 


Miller 1073 
New World Ehretia 


K, LE, MO, NY —3 sheets, P, S, UC, US). CAYMAN 
ISLANDS. GRAND CAYMAN: in coppice, West Bay Village 
area, Correll & Correll 51037 (NY); edge ob coppice 
along roadway near Hell, West Bay Village area, Correll 
& Correll 51047 (F, IJ, MO, NY); N end of Grand Sound 
Road, Proctor 15044 (BM, GH, IJ); NW of Sosdenus m. 
dg 31038 (BM, IJ, LL). JAMAICA. HANOVER: 5 mi. 
Lucea, near sea level, pe 28561 (IJ); i interior 
of fice Purdie s.n. (K). K 
ander 6563 (NY); Kingston, Tower S 
(IJ); corner of Church St. and North St., 
(IJ, NY, US); along East Race Course, Proce 23605 
(LL, MICH, NY, US). MANCHESTER: Spur Tree c near 
Ellington, 1, ane 2,000 ft., Proctor 28751 (BN 


LNY% N elier District, below Spur Tree, T 000- 
1,250 ft., pea 30018 (IJ). ST. ANDREW: Gordon Town, 
1,200 ft., Barry s.n. (IJ); Hope, 600 ft., Barry s.n. (IJ); 


Wolmer's Boys S 
t Marri 6073 (A, B 


Scudamore s.n. (y 


chool, id s.n. (IJ): Green Valley, 
, NY); W slope of Long 
Long Mountain, along 
puras 4987 (A, IJ); 
Webster a s 
8060 (BM, G, IJ, S); in waste area eee to U.C 
campus, 550 ft., Yuncker 17721 (F, MICH, S). ST. ANN: 
vicinity of Green Grotto Caves, 2.5 mi. W of Runaway 
Bay, 5-50 ft., Proctor 36485 (IJ). ST. CATHERINE: Cay- 
manas, . , Adams & Carrington 11513 (BM). sr. 
ELIZABETH: Lititz Savana, 300-900 ft., Harris 11726 
(BM, F, MO, NY, US); limestone hillsides near new build- 
ings, Kaiser mine area S of Gutters, Howard & Proctor 
13858 (A, IJ); Kaiser mine area at Comfort, 500 ft., 
Howard & Proctor 14448 (IJ); 2-4 mi. SW of Lacovia, 
Howard & Proctor 14522 (A, BM, IJ); Parotte, sea level, 
Proctor 28836 (IJ); Giddy Hall, Sangster s.n. 
ST. JAMES: Rose Hall, G. S. Miller 14 19 (US); Greenwood 
estate, 5 mi. ESE of Little River P.O., Stear A, 
BM —2 sheets). sr. THOMAS: Yallahs, : p 1que- 
Molina & Barkley 225438 (BM, IJ —3 ts Yallahs 
Valley -Green Valley, Harris 60838 (F, NY —2 sheets); 
Grant's Pen, near sea. m Proctor 28801 (BM, IJ). 
WESTMORELAND: 0.5 m i i 
Proctor 36729 (IJ). PARISH UNKNOWN: Thicke 
Britton 1077 (NY); near camp, 67 m, Campbell 5690 
, US); asylum grounds, Fawcett s.n. (BM); Malvern 
to Mountain side, Harris 9755 (BM, F, NY, US); road 
from Hall-way Tree to Spanish Town Road, Mantis 10381 
(A, BM, F, NY US); without definite locality, Macfad yen 
s.n (K); Molle s s.n. (GH); Herb. Montinü (S): Swartz s.n. 
(S); Waters s.n. (K); Trelawny, roadside near Duncans, 
Whitefoord 1464 (BM, IJ). Haiti: Fond Parisien, Bailey 
175 (BH, US); Massif du Nord, Port-de-Paix, Bassin Bleu, 
150 m, Piman H3992 (IJ, LL, NY, US); vicinity of Fond 
Parisien, Etang Saumatre, Leonard 4173 (NY, US— 2 
1 l Paix, ridge W of La Coup 
River, Leonard : Leonard 11142(US); vicinity of nen 
Bleu, 630-1,500 m, nel ae 709 (US—2 shee 
DOMINICAN mu BLIC: i, Guayubin, 100 m or 


200 1 , Eggers 
tiago, poda 1990 (BM, G); Santo Domingo, Valle del 
Cibao, Prov. Santiago, Santiago, roadsides, near La Cien- 
ago, Ekman H 13235 (S, US); Barahona, Baum, Fuertes 
296 (BM, F, G, GH —2 sheets, LE, MO, NY, P, S, US); 
Santiago, along banks of the Rio Yaque near Santiago, 
Howard & Howard 9689 (4, B, BM, NY, US); Hotel 


Yaque, near Santiago City, Jiménez 5668 (NY); Peder- 
nales, near Las Mercedes from Cabo Rojo to Aceitillar, 
400 m, Liogier 16688 (NY); Santo Domingo, En Man- 
igua, Pedernales, Liogier 19691 (NY) a orilla de la 


Zanoni 6516 (NY); Azua, Santo Doming 
4005 (NY, US); Santo Domingo, Trujillo City, Schifins 
150 (GH), 162 (GH). 


EXCLUDED SPECIES 


The majority of names published in Ehretia for 
the Neotropics are now considered synonyms of 
Bourreria and Rochefortia. In the following list, 
types have been examined when possible, but many 
of the names can be excluded from Ehretia based 
on the published descriptions. In placing the names 
I have relied heavily on the revisions of Bourreria 
(Miers, 1869; Schulz, 1911) and Rochefortia (Le- 
for, 1968) supplemented with notes from John- 
ston's extensive series of papers on the Boragi- 
naceae. When FEhretia names are assigned to 
species of Bourreria, they should be considered 
cautiously as the best name available from the 
literature since Bourreria is probably the most 
nomenclaturally confused genus of Ehretioideae. 


Ehretia acanthophora DC., Prodr. 9: 510. 1845. 
TYPE: Dominican Republic, Santo Domingo: 
Bertero s.n. (holotype, G-DC, not seen; mi- 
crofiche, MO; isotypes, MO, P). = Rochefor- 
tia iie (DC.) Griseb., Fl. British 
West Indies 482. 1864; O. Schulz in Urban, 
Symb. Antill. 7: dE 1911: León & i Fl. 
Cuba 4: 265. 1957; Lefor, 30. 19 

Ehretia andrieuxii DC., Prodr. 9: pd 1845. 
T exico. Puebla: May, Andrieux 200 
(holotype, G-DC, ). 
= Bourreria andrieuxii (DC.) Hemsley, Biol. 
Cent.-Amer., Bot. 2: : $ 
Bot. 2: 254. 1869; Gibson, Fieldiana Bot. 24, 
pt. 9: 113. 1970; Miller, Fl. Nicaragua (in 

ress). 

Ehretia bogotensis Spreng., Syst. Veg. 1: 648. 
1825. — Aegiphila bogotensis (Sprengel) 
Moldenke, Repert. Spec. Nov. Regni Veg. 
Beih. 33: 114. 1933. 

Ehretia bourreria Desf. (non L.), Ann. Mus. 1: 

; ourreria recurva Miers, Ann. & 
Mag. Nat. Hist. Ser. 4, 3: 203. 1869. 

Ehretia bourreria L., Sp. Pl. 2 ed. 1: 275. 1762. 
— Bourreria d d Jacq., Enum. Syst. 
Pl. Carib. ; O. Schulz in Urban, 
Symb. Antill. 7: s 56, and 62; León & 
Alain, Fl. Cuba 4: 269. 1957. 


v. 
5 
E 


not seen; microfiche, 


1074 


Annals of the 
Missouri Botanical Garden 


Ehretia calophylla A. Rich. in Sagra, Fl. Cuba 2: 
12, t. 61. 1853. = Bourreria rotata (DC.) 

I. M. Taio J. Arnold Arbor. 30: 107. 
1949; León & Alain, Fl. Cuba 4: 268. 1957. 

Ehretia cassinifolia A. Rich. in A. Sagra, Fl. Cuba 
2: 113. 1850. = Bourreria cassinifolia (A. 
Rich.) Griseb., Pl. Wright. Cub. 528; O. Schulz 
in Urban, Symb. Antill. 7: 67. 1911; León 
& Alain, Fl. Cuba 4: 270. 1957. 

Ehretia cirrhosa Lam. = Maripa scandens Au- 
blet; DC., Prodr. 9: 511. 1845. 

Ehretia cumanensis DC., Prodr. 9: 511. 1845. 
= Tournefortia hirsutissima L., Sp. Pl. 140. 
1753. O. Schulz (in Urban, Symb. Antill. 7: 
50. 1911) treated this name as applicable to 
Tournefortia, not Ehretia or Bourreria. The 
name Bourreria cumanensis (Loefl.) O. Schulz 
is based on Rhamnus cumanensis Loefl., 
Hispan. 182. 1758 

Ehretia cuneifolia Sessé & Mocino, Fl. Mexic. 51. 

ed. 2. 47. 1894. = Bourreria spathu- 
lata (Miers) Hemsley, Biol. Cent.-Amer., Bot. 
2: 370. 1882; I. M. Johnston, J. Arnold Ar- 
bor. 30: 107. 1949. 

Ehretia cymosa Willd. ex Roemer & Schultes (non 
Thonn). = d hirsutissima L., Sp. 
Pl. 140. 1753; O. Schulz in Urban, Symb. 
Antill. 7: 50. Tan 

Ehretia divaricata A. Rich. (non DC.) in Sagra, 
Hist. Fis. Cuba, Bot. 11: 113. 1850. = Bour- 
reria succulenta var. revoluta (Kunth in 
Humb., Bonpl. & Kunth) O. Schulz in Urban, 
Symb. Antill. 7: 59. 1911. 

Ehretia divaricata DC., Cat. vo Monspel. 108. 
l Prodr. 9: 506 . = Bourreria 
divaricata (DC.) G. da na Hist. 4: 389. 

1838; O. Schulz in Urban, Symb. Antill. 
69. 1911: León & Alain, Fl. Cuba 4: 271. 
1957. 

Ehretia domingensis DC., Prodr. 9: 508. 1845. 
— Bourreria domingensis (DC.) Griseb., Fl. 
Brit. W.I. 482. 1861; Miers, Ann. & Mag. 
Nat. Hist. Ser. 4, 3: 202. 1869; O. Schulz 
in Urban, Symb. Anull. 7: 64. 1911. 

Ehretia dubia Jacq., Obs. 1, 19. 1764; O. Schulz 
in Urban, Symb. Anull. 7: 71. 1911. 

dia sp. probably C. collococca L. or C. nitida 
l 


Iter 


= Cor- 


Vahl. 
Ehretia exsucca Bert. ex Griseb. (non L.), Fl. Brit. 
Js . 1861. = Bourreria domingensis 
(DC.) Griseb., Fl. Brit. W.I. 482. 1861; O. 
Schulz in Urban, Symb. Antill. 7: 64. 1911. 
Ehretia exsucca L., Sp. Pl. ed. 2. 1: 275. 1762. 
— Bourreria cumanensis (Loefl.) O. Schulz in 


Urban, Symb. Antill. 7: 49. 1911. 


Ehretia fasciculata Kunth in Humb., Bonpl. & 
unth, Nov. Gen. Sp. 3: 66. 1818. TYPE: 
Humboldt s.n. (holotype, P-HBK, not seen; 
microfiche, MO). Rochefortia spinosa 
(Jacq.) Urb. in Fedde, Repert. Spec. Nov. 13: 
472. 1915; Lefor, 1968. 

Ehretia formosa DC., Prodr. 9: 510. 1845. — 
Bourreria huanita (Llave & Lex.) Hemsley, 
Biol. Cent.-Amer., Bot. 2: 369. 1882; O. 
Schulz in Urban, Symb. Antill. 7: 51. 1911; 
Standley, Contr. U.S. Natl. Herb. 28: 1225. 
1924; Gibson, Fieldiana Bot. 24, pt. 9: 114. 
1970. 

Ehretia formosa var. oaxacana DC., Prodr. 9: 
510. 1845. TYPE: Mexico. Oaxaca: Tehuan- 

Andrieux 201 (photo, GH). = Bour- 
reria huanita eis & Lex. a Hemsley, Biol. 
Cent.-Amer., Bot. 369. 1882; O. Schulz 
in Urban, Bab. jdn ri a 191 1; Standley, 
Contr. U.S. Natl. Herb. 28: 1225. 192 

Ehretia grandiflora Poir., Encycl. Bund 2: 3. 

— Bourreria wrightii Alain, Contrib. 
Ocas. Mus. Hist. Nat. Coleg. “De La Salle" 
15: 9. 1956, a new name for Bourreria gran- 
diflora Griseb., non Bertol., O. Schulz in Ur- 
ban Symb. Antill. 7: 52. 1911; León & Alain, 
Fl. Cuba 4: 268. 1957. 

Ehretia grisebachii Maza, Periant. 256. 1890. — 
Bourreria microphylla Griseb., Cat. Pl. Cuba 
210. 1866; O. Schulz in Urban, Symb. Antill. 
7: 69. 1911; León & Alain, Fl. Cuba 4: 271. 
1957. 

Ehretia guatemalensis DC., Prodr. 9: 507. 1845. 
— Bourreria huanita (Llave & Lex.) Hem- 
sley, Biol. Cent.-Amer., Bot. 2: 370. 1882; 
O. Schulz in Urban, Symb. Antill. 7: 51. 1911; 
León & Alain, Fl. Cuba 4: 268. 1957. 

Ehretia havanensis Willd. ex Roemer & Schultes, 
Syst. Veg. 4: 805. 1819. 
culenta var. revoluta (Kunth in Humb., Bonpl. 
& Kunth) O. Schulz in Urban, Symb. Antill. 
7:59. 1911; León & Alain, Fl. Cuba 4: 269. 
1957. 

Ehretia laevis Roxb. var. cymosa Roemer & 
Schultes, Syst. Veg. 4: 805. 1819. — Tour- 
nefortia hirsutissima L., Sp. Pl. 140. 1753. 
Although Ehretia laevis is an Old World 
species, Roemer & Schultes published this va- 
riety from the New World; it was excluded 
y O. Schulz (in Urban, Symb. Antill. 7: 50. 
1911) as a species of Tournefortia. 

Ehretia lanceolata Vell., Fl. Flum. 79. 1829. In 

rodr. 9: 511. 1845, De Candolle listed this 
as unknown in his list of excluded species, 
indicating a distribution in Brazil, where Eh- 


tepec, 


= Bourre ria SUC- 


Volume 76, Number 4 
1989 


Miller 1075 
New World Ehretia 


retia does not occur. This is sufficient to ex- 
clude it from the genus. 

Ehretia microphylla Lam., Tab. Encycl. 1: 425. 
1791. = Carmona retusa (Vahl) Masamune, 
Trans. Nat. Hist. Soc. Formosa 30: 61. 1940; 
Thulin, Nord. J. Bot. 7: 413. 1987. Although 
this species is native in the Old World, it is 
known from cultivated collections from Hon- 
duras and Cuba. 

Ehretia montana Griseb., Pl. Wright. 528. 1862. 
— Bourreria divaricata Don, Dict. 4: 389; 
Griseb., Pl. Wright. 528. 1862; Cat. Pl. Cuba 
210. 1866 

Ehretia montevidensis Sprengel, Syst. 1: 647. 
1825. — Citharexylum montevidense (Spren- 
gel) Moldenke, Phytologia 1: 17. 1933. 

Ehretia radula Chapman (non Poir.), Fl. South. 
U.S. 329. 1872. = Bourreria succulenta var. 
revoluta (Kunth in Humb., Bonpl. & Kunth) 
O. Schulz in Urban, Symb. Antill. 7: 59. 1911. 

Ehretia radula Poir., Encycl. Suppl. 2: 2. 1811; 
DC., Prodr. 9: 506. 1845. = Bourreria to- 
mentosa (Lam.) G. Don, Gen. Hist. 4: 390. 
1838; O. Schulz in Urban, Symb. Antill. 7 
54. 1911 

Ehretia radula Sprengel (non Poir. or Chapman), 
Syst. Veg. 1: 648. 1825. — Bourreria di- 
varicata (DC.) G. Don, Con. Hist. 4: a 
1838; O. Schulz in Urban, Symb. Antill. 
70. 1911. 

Ehretia revoluta DC., Prodr. 9: 507. 1845. = 
Bourreria succulenta var. revoluta (Kunth in 
Humb., Bonpl. & Kunth) O. Schulz in Urban, 
Symb. Antill. 7: 60. 1911. 

Ehretia rupestris Salisb., Prodr. 112. — Bourreria 
succulenta (Jacq.) Urban. 

Ehretia scandens Poir., Encycl. Suppl. 3: 590. 

— Maripa scandens Aublet; DC., Prodr. 
9: 511. 1845). 

Ehretia spinifex DC., Prodr. 9: 506. 1845. — 
Bourreria divaricata (DC.) G. Don, Gen. Hist. 
4: 389. 1838. O. Schulz (1911) treated EA- 
retia spinifex DC., a name De Candolle at- 
tributed to Roemer & Schultes, as a synonym 
of Bourreria divaricata but excluded EKhretia 
spinifex Roemer & Schultes as a synonym of 
Basanacantha spinifex (Roemer & Schultes) 
Urban of the Rubiaceae. León & Alain (1957) 
listed Ehretia spinifex Griseb. as a synonym 
of Bourreria divaricata but made no mention 
of the other names. The proper application of 
De Candolle's and Roemer & Schultes's names 
will require further study, but they certainly 
can be excluded from Ehretia based on their 
descriptions. 


Ehretia spinifex Griseb. (non Roemer & Schultes) 
Pl. Wright. Cuba 528. — Bourreria divari- 
cata Don, Dict. 4: 389; Griseb., Pl. Wright. 
Cuba 528; Cat. Pl. Cuba 210. 

Ehretia spinifex Maza, Periant. 255. 1890. — 
Bourreria cassinifolia (A. Rich.) Griseb., Cat. 
Pl. Cuba 210. 1866; O. Schulz in Urban, 
Symb. Antill. 7: 67. 1911; León & Alain, Fl. 
Cuba 4: 270. 19 

Ehretia spinosa Jacq., Eun. Syst. Pl. 14. 1760. 
— Rochefortia spinosa (Jacq.) Urban in Fedde, 
Repert. Spec. Nov. 13: 472. 1915; O. Schulz 
in Urban, Symb. Antill. 7: 71. 1911; León € 
Alain, Fl. Cuba 4: 266. 1957; Lefor, 1968. 

Ehretia de Spreng. ex DC. (non Jacq.), Prodr. 
9: 510. 1845. — Rochefortia acanthophora 
(DC. Ta Fl. Brit. W.I. 482. 1862; Lefor, 


1968. 
Ehretia ternifolia Kunth in Humb., Bonpl. & 
unth, Nov. Gen. Sp. 3: 66. 1818. TYPE: 
Colombia (holotype, P-HBK; photo, GH). — 
Aegiphila ternifolia (Kunth in Humb., Bonpl. 
& Kunth) Moldenke, Repert. Spec. Nov. Reg- 
ni Veg. Beih. 33: 141. 1933. 

Ehretia tomentosa Lam., Tab. Encycl. 1: 425. 
1793; DC., Prodr. 9: 507. = Bourreria ve- 
lutina (DC.) Gurke in Engl. & Prantl, Nat. 
Pflanzenfam. IV. 3a: 87. 1897; O. Schulz in 
Urban, Symb. Antill. 7: 54. 1911. 

Ehretia tomentosa var. havanensis Gómez, Anal. 
Hist. Nat. Madrid 19: 256. 1890. = Bour- 
reria succulenta var. revoluta (Kunik in 
Humb., Bonpl. & Kunth) O. Schulz in Urban, 
Symb. Antill. 5: 59. 1911. 

Ehretia velutina DC., Prodr. 9: 508. 1845. 
Bourreria velutina (DC.) Gürke in a & 
Prantl, Nat. Pflanzenfam. IV. . 18975 
O. Schulz in Urban, Symb. Antill. pal 1911. 

Ehretia virgata Sw., Prodr. 47. 1788; Flor. Ind. 
Occ. 1: 463; DC., Prodr. 9: 506. 1845. = 
Bourreria virgata (Sw.) G. Don, Gen. Hist. 
4: 389. 1838; O. Schulz in Urban, Symb. 
Antill. 7: 66. 1911; León & Alain, Fl. Cuba 
4: 269. 1957. 


LITERATURE CITED 

AIRY-SHAW, H. K. 1973. A Dictionary of the Flowering 
Plants and Ferns, 8th edition. Cambridge Univ. Press, 
Cambridge. 

AVETISSIAN, E. M. 


956. Morphologie des Microspores 


de Boraginaceae. T. Botan. Inst. Akad. Nauk Arm. 
DO ie ( ~ 60. 
BAKER, J. G. . H. WRIGHT. Boragineae. /n: 
W. T. Ed Dyer (editor), Flora of Tropical Af- 
rica 4(2): 5 


62. 
A, K. S. & J. H. BEACH. 1981. Evolution of sexual 


1076 


Annals of the 
Missouri Botanical Garden 


systems in flowering plants. Ann. Missouri Bot. Gard. 
-2 


8: 2 ; 
BENTHAM, G. & J. D. HOOKER. p Boraginaceae. 
In: E i Plantarum 2: 832-8 
Browne, P. 1756. The Civil and nan History of 
Jamaica in a Three Parts. London 
DE CANDOLLE, A. P. 1845. Boraginaceae. Ehretia. In: 
rodromus Systematis Naturalis Regni Vegetabilis. 
Paris 9: 502-512. 
DILCHER, D. L. 1974. 


Approaches to the identification 
of angiosperia leaf remains. Bot. Re 


ev. (Lancaster) 


GANDER F. R. 197 9. The end of heterostyly. New 

and J. Bot. 17: 607-6 

E T N. 1970. Boraginarcae. E Flora of Gua- 
temala. Fieldiana, Bot. 24(9): 11 67. 

Hickey, L. J. 1973. Classification es de architecture 
of kp caer leaves. Amer. J. Bot. 60: 17-63. 

TT. ratigraphy and P usen of the 

Golden Valley Miu (Early Tertiary) of western 
rth Dakota. Mem. Geol. Soc. Amer. 150: 157- 

163. Erde 2. 

979. A revised classification of the archi- 
mune of dicotyledonous leaves. /n: C. R. Metcalfe 
& L. Chalk ade Anatomy of the Dicotyledons, 
2nd edition ]: 25-39, 

& J. DT 1975. The bases of angio- 
sperm Pres vegetative morphology. Ann. Mis- 
souri Bot. G 538-589. 


mm I. M. 1924. Studies in the Boraginaceae II. 
The Old World genera of the Boraginoideae. 2. 
ves B oe Boraginaceae. Contr. Gray 
Herb. 2-78. 
eae. Studies in the Boraginaceae XI. (3 
New or otherwise noteworthy species. J. Arnold Ar- 
bor. 16: 173-205. 

1949, Studies in the Boraginaceae XVIII. 
Boraginaceae of is. southern West Indies. J. Arnold 
Arbor. en 111-138. 

P de in the Boraginaceae XIX. 
Note 0 species from tropical America. (B) Gar. 
dia section Aena ri A Mexico and Central 
America. J. Arnold Arbor. 31: 172-187. 
1951. Studies in ilie en aceae XX. Rep- 
gni of three subfamilies in eastern Asia. J. 
rnold Arbor. 32: 1-26, 32: 99-122 
i M.S. 1968. A revision of the genus Ras hefortia 
w. (Boraginaceae). M.S. thesis, Univ. of Connecti- 


a 


A) 


T Hanno & H. des AIN. 1957. 
In: Flora de Cuba 4: 252-218. 
Levin, G. idend foliar morphology of 
Phyllanthoideae (Euphorbiaceae). . Conspectus. Ann. 

-85. 


Boraginaceae. 


issouri Bot. Gard. 73: 2 


6b. Systematic foliar morphology of Phyl- 
jd (Euphorbiacea e). Il. Phenetic analysis. 
. Missouri Bot. Gard. 8. 
86c. Systematic foliar morphology of Phyl- 
lanthoideae (Euphorbiaceae). III. Cladistic analysis. 
st. Bot. 11: 515-530. 
LINNAEUS, C. 175 
holm. 


73: 86-9 


9. Systema Naturae, ed. 10. Stock- 


LoRENCE, D. H. A monograph of the Monimi- 
aceae (Laurales) in the Malagasy a (southwest 
Indian Ocean). Ann. Missouri Bot. Gard. 72: 1-165. 

Miers, J. 1869. On the Ehretiaceae. Contr. Bot. 224- 
"n 

MILLER, J. S. 1985. Systematics of the genus Cordia 
Boraginaceae) in Mexico and Central America. Ph.D. 

thesis, St. Louis Univ., St. Louis, Missouri. 

1988. A revised treatment for the Boragi- 

naceae of Panama. Ann. Missouri Bot. Gard. 75: 

21 


Nasu, D. L. & N. P. Moreno. 1981. d 

In: Flora de Veracruz. Xalapa, Veracruz 18: 1-149. 
NOWICKE, J. W. 1 Boraginaceae. /n: Flora of Pan- 
ama. Ann. Missouri Bot. Gard. 56: 33-69 

& MILLER. Pollen morphology of the 
Cordioideae iora 4uxemma, Cordia, and 
Patagonula. Pl. Syst. Evol. (in p 

€ —— —. 


ess). 
bun E ph “Flora of Ceylon 


(in press). 

& J. E. RIDGWAY. Pollen died in the 
genus Cordia (Boraginaceae). Amer. J. Bot. 60: 584 
991. 


SKVARLA. 1974. A palynological in- 
vestigation of ps Agere M (Boraginaceae). 
Amer. J. Bot. 61: 1021-1036 
RICHARDSON, A. T 7 Reinstatement of the genus 

Tiquilia (Boraginaceae: Ehretioideae) p descrip- 
tions of four new species. Sida 6: 235-2 

l Monograph of the genus Ticuilia 
d s lato), Boraginaceae: Ehretioideae. d 
fi 


-1 


572. 
TSAA ES L 1894. Contributions from the Gray 
erbarium of Harvard jsp OR. new series, No. 
oc. Amer. Acad. Arts 29: -330. 
is W. C. DICKISON. D 'af venation 
patterns of the genus iv iai (Dilleniaceae). J. 
dt ee 58: 209-2 
RZE DOWSKI, J. Mure de México. Editorial 


: Beurreria. In: Urban, Symb. 
Antill. 7: 45-71. 
PME J. D. 1893. Undescribed pn m Guatemala 
. Bot. Gaz. (C ev ied cats : 1l- 
STANDLEY, P. C. 1924. es de nw of Mexico. 
ey aaa Contr. US .S. Natl. Herb. 23: 1216- 
1234 


1 doe 7 we m Publ. Field Mus. 
Nat. Hist., Bot. 5 f 
TAHKTAJAN, A. 1987 p bee Magnoliophytorum. 
. & M. Sousa S. 1982. Imagenes de la 
a Quintanarroense, Cioro, México. 
19 Bourre 'ria (Boraginaceae) in tropical 
: 413-417. 


e origin and evolutionary 

development of "asterostyy d in the angiosperms. Evo- 
ution 21: 210-226. 

WarsoN, S. 1891. XI. Contributions to American bot- 
any. Proc. Amer. Acad. Arts 26: 124-163 


NEW AND CRITICAL TAXA 
OF EUPHORBIACEAE 
FROM SOUTH AMERICA! 


Michael J. Huft? 


ABSTRACT 


Study of neotropical Euphorbiaceae for various floristic and taxonomic projects has uncovered several novelties. 


New taxa pr 
Colombia, Tragia ik ue e Peru, and the genus 


the border of Panama and Color 


oposed in this paper are Dalechampia liesneri from souther 
Tacarcuna, including the thr 
bia, 7. amanoifolia from Peru, an 


ela, — S tomentosa from 
ree new speci . gentryi, from 
achirensis from ud. In addition it 


n Venezue 


is shown that Caryode Eon ee) irense Muell. Arg. is the correct name for ihe plant hitherto known as C. grandifolium 


(Muell. Arg.) Pax 


The Euphorbiaceae, one of the largest families 
in the Neotropics, remains one of the least well 
understood. The largest genera, Acalypha, Cro- 
ton, Euphorbia, and Phyllanthus, have not been 
critically revised in this century, and only a handful 
of other, smaller genera have been the subject of 
modern study. Even at the most basic descriptive 
level, a great deal of work remains to be done, and 
the mere discovery of a new species can often lead 
to much insight into the taxonomy of its congeners. 
Such is the case with the collection of taxa proposed 
here, which result from study preparatory to the 
Flora of the Venezuelan Guayana (J. Steyermark 
& collaborators, in prep.), Checklist of the Flora 
of Chocó Province, Colombia (A. Gentry & E. 
Forero, in prep.), or from serendipitous discovery 
in the herbarium. 


Caryodendron janeirense Muell. Arg. in C. 
Mart., Fl. Bras. 11(2): 707. 1874. Centro- 
discus grandifolius Muell. Arg. in C. Mart., 
Fl. Bras. 11(2): 327. 1874, nom. inval. Cary- 
odendron grandifolium (Muell. Arg.) Pax in 
Engler & Prantl, Nat. Pflanzenfam. III. 5: 
52. 1890, nom. illeg. TYPE: Brazil. Rio de 
Janeiro: Riedel 1051 (presumably C). 


Mueller published the name Centrodiscus gran- 
difolius with description and citation of type. Be- 


fore this went to press however, he added as a 
separate item on a different page "Addenda and 
Corrigenda” in which he pointed out that his Cen- 
trodiscus grandifolius belongs to Karsten's genus 
Caryodendron and that it differs from the only 
species, Caryodendron orinocense Karsten, by, 
among other things, having smaller leaves. Ap- 
parently thinking that the epithet grandifolium 
would be inappropriate in Caryodendron, he as- 
signed the new plant the name Caryodendron ja- 
neirense, which is the correct name for the species. 
Centrodiscus grandifolius was not published be- 
fore Caryodendron janeirense, pp. 273-754 of 
Volume 11, Part 2, appearing together on May 1, 
1874 (Stafleu & Cowan, 1981: 333-337). Mueller 
was merely making a last-minute correction to text 
already too far along in press to be changed in- 
ternally. It is therefore clear that the name Cen- 
trodiscus grandifolius Muell. Arg. was not ac- 
cepted by its author when published and therefore 
is invalid according to Article 34.1 of the /nter- 
national Code of Botanical Nomenclature (Greu- 
ter, 1988). The combination of Pax based on that 
name is nomenclaturally superfluous. For the same 
reason, the generic name Caryodendron Muell. 
Arg., published simultaneously with Caryodendron 
grandifolius Muell. Arg. and likewise not acc ee 
by its author when published, is also invali 


I am inde bte d to Dr. 


drawings were 1 


W. Scott Armbruster of the University of Alaska for ae um concerning 


able. Th 


excellent 


2). 
? Missouri Botanical Garden. Mailing address: Department of Botany, Field Museum of Natufal History, Roosevelt 
, U.S.A. 


Road at Lake Shore Drive, Chicago, Illinois 60605 


ANN. Missouni Bor. GARD. 76: 1077-1086. 1989. 


1078 


Annals of the 
Missouri Botanical Garden 


Dalechampia liesneri Huft, sp. nov. TYPE: Ven- 
ezuela. Territorio Federal Amazonas: Depto. 
Atures, stream 0. 5-2 km E of Rio Coro-Coro, 

de Yutaje, 
66°07'30"W. "200 m, 
Liesner & B. Holst 21244 (holotype, MO; 
isotypes, U, VEN not seen). Figure 1. 


(vel nerpa) volubilis; caules et folia glabra vel 
a. Folia simplicia, longipetiolata; lam- 
inae crei basi 5-nerves, apice acutae; basi late 
. Inflorescentiae dissitae, longipedunculatae, flo- 
ribus a Flores masculi ad 8; sepala ovata, acuta, 
sub anthesi reflexa; stamina 8-10, filamentis connatis, 
tubum longum gracilem prats Flores feminei 3, 
subsessilis, pedicello ad fructus maturitatem multo elon- 
gato: sepala 6, lineari-lanceolata; stylus gracilis, elongatus. 
Capsulae glabrae vel brevipuberulae, laeves. Semina ovoi- 
dea vel subglobosa, laevia, brunnea. 


Frutex 


Twining vine; stems terete or slightly angulate, 
glabrous or very sparsely short-puberulent. Leaves 
simple, on petioles to 10 cm long, these pinched 
at apex and at base and puberulent; blades ovate- 
deltate, membranous, glabrous or the principal veins 
minutely puberulent, 5-nerved, the secondaries few 
near apex of blade, ascending, the tertiaries retic- 
ulate, conspicuous, the apex acuminate, the base 
openly cordate, stipellate, the margin minutely and 
remotely denticulate. Inflorescences bisexual, widely 
spaced, compact, the peduncle slender, densely 
puberulent, 1 -4 cm long. Staminate flowers distal, 
up to 8 present, the pedicels 2-5 mm long; bracts 
lanceolate, brown puberulent, acute, 2-4 mm long; 
buds fusiform, acute; sepals 5, ca. 2 mm long, 
membranous, ovate, acute, sparsely hirtellous, re- 
flexed at anthesis; stamens 8-10, the filaments 
united nearly their entire length to form a slender 
tube 4-5 mm long, the anthers free at apex, the 
tube penicillate toward apex. Pistillate flowers 3, 

or 2 sometimes aborting before maturity, prox- 
imal, subsessile at anthesis, the pedicel densely 
puberulous-hirtellous, greatly elongating in fruit (to 
5 cm); sepals 6, linear-lanceolate, 2.8-3.2 mm 
long, acute, sparsely short-puberulent, the base 
thickened, the young fruit densely short-pilose; style 
slender, elongate, to 15 mm long, 0.2-0.3 mm 
thick, densely puberulent toward the base, other- 
wise glabrous. Capsule smooth, 3-lobed, short-pu- 
berulent, not seen entire, apparently 6—7 mm diam.; 
seeds ovoid to subglobose, smooth, ca. 4 mm diam., 
brown. 


The pantropical genus Dalechampia contains 
over 100 principally neotropical species. Numer- 
ous recent discoveries (Armbruster, 1984, 1988, 
1989; Huft, 1984; Webster, 1967, 1988; Web- 
ster & Armbruster, 1979, 1982), many of them 
of taxonomic and phylogenetic importance, attest 
to the need for the critical study this genus is now 
receiving at the hands of Dr. Grady L. Webster 
and Dr. W. Scott Armbruster. 

Dalechampia liesneri did to sect. Rho- 
palostylis Pax & K. , the most primitive 
section of the genus, which oni is restricted 
to the northern and western Amazonian basin. Only 
four species have previously been described from 
this group: D. micrantha Poeppig from western 
Brazil and eastern Peru, D. parvibracteata Lanj. 
from the Guianas, D. olympiana Kuhlm. Ro- 
drigues from northern Brazil, and the recently de- 
scribed D. attenuistylus Armbruster from southern 
Venezuela (Armbruster, 1989). The genus Mega- 
lostylis S. Moore, whose only species, M. poep- 
pigii S. Moore, appears to be based on an isotype 
of D. micrantha, is synonymous with sect. Rho- 
palostylis. 

Section Rhopalostylis is characterized by ab- 
sence of the showy white, pinkish, purplish, or 
green bracts that are so characteristic of the rest 
of the genus; absence of resiniferous glands in the 
inflorescences; and larger, usually basally swollen, 
styles. Dalechampia liesneri is distinctive in its 
few-flowered, long-pedunculate, inflorescences and 
long, thin styles that are not at all swollen toward 
the base. Dr. Armbruster (pers. comm.), who is 
preparing a revision of this group, has discovered 
an additional population of this species near San 

e Cataniapo (ca. of Puerto 
Ayacucho) in southern Venezuela, and he reports 
that it is probably not uncommon in that region. 

It is a pleasure to name this species after Ronald 
Liesner of the Missouri Botanical Garden who has 
collected extensively in Venezuela during the past 
decade and has been responsible for many discov- 
eries and additions to our knowledge of the flora 
of that country. 


Richeria tomentosa Huft, sp. nov. TYPE: Colom- 
bia. Chocó: Municipio de Quibdó, bosque frente 
al barrio Obrero, 24 May 1985, Espina & 
García 1518 (holotype, MO). 


FIGURE 1. Dalechampia liesneri. — eaf.- 
fruits. 4. Seed. (Based on Liesner & Holst 21244. ) 


—b. Inflorescence at anthesis. — c. 


=j 
Inflorescence with immature 


1079 


Huft 
South American Euphorbiaceae 


Volume 76, Number 4 


1989 


1080 


Annals of the 
Missouri Botanical Garden 


Arbor ad 25 m alta, dioecia; cortex crassus exaratus 
Folia alterna brevipetiolata; laminae late obovato-ellipti- 


cae, infra dense brunneo-tomentosae vel glabra 


vel breviacuminatae, basi rotundatae vel 

margine crenato-undulatae. Inflo 

erales sub foliis, ex paniculis s 

longis constans. Fiori masculi in glomerulis sessilibus vel 
il 


subsessilibus. vlores feminei non visi, sessiles vel brevi- 


voideae obscuro ue glabrae; columella non persistens 
alis latis papyrac 
Tree to 25 m high, dioecious; twigs with thick 
bark formed into flattened ridges and narrow fur- 
rows, densely hirsutulous to glabrate toward apex. 
Leaves alternate, short-petiolate; stipules obsolete; 
petioles 0.5-2.5 cm long, stout, densely hirsutulous 
to glabrate; blades chartaceous to subcoriaceous, 
broadly obovate-elliptic, 10-23 cm long, 7-12 cm 
wide, 1.5-2.8 times as long as wide, densely brown 
tomentose to glabrate below, moderately pilose to 
glabrate above, the apex rounded, obtuse, or short- 
acuminate, the base rounded, obtuse, or broadly 
cuneate, the margins entire to obscurely crenate- 
undulate; midrib prominent below, the secondary 
nerves 14-26 on a side, parallel, arching somewhat 
toward the margin of the blade, prominent below, 
light-colored above, neither raised nor impressed, 
the tertiary veins connecting the secondaries at 
right angles, regularly spaced, prominulous below. 
Inflorescences numerous, lateral, below current 
leaves, RE of clustered M 2-4 cm 
long, t ly brown tomentose when young, 
becoming densely puberulent at maturity. Stami- 
nate flowers 1-3 in sessile or subsessile glomerules, 
the buds globose, 0.7-1.1 mm diam. Pistillate 
flowers not seen, sessile or short-pedicellate; calyx 
persistent on young fruits, tomentose outside, the 
lobes 1-1.5 mm long, deltate, ca. 25 the length of 
the calyx; young fruits densely brown tomentose, 
pyriform, the styles 3, ca. 1 mm long, strongly 
ecurved, united at base, glabrous or with a few 
scattered. hairs. Capsules ellipsoid-obovoid, ob- 
scurely trigonous, glabrous, smooth, 11-16 mm 
the styles persistent, minute, 


long, 6-9 mm diam., 
the columella with broad papery wings, not per- 
sistent. 

This very distinctive species is easily distin- 
guished from the four to six other species of Rich- 
eria by the densely tomentose lower leaf surfaces 
on many specimens and by the short, compact, 
highly branched inflorescences. The prominent 
midrib and secondary veins are also distinctive. 
The widespread R. racemosa (Poeppig) Pax & 


Hoffm. has staminate spikes longer than the leaves 


rather than the short, compact, clustered panicles 
that are characteristic of R. tomentosa, and the 
cymules are distinctly pedunculate and two- or 
three-flowered. Two other South American species, 
R. grandis Vahl and R. obovata (Muell. Arg.) Pax 
& K. Hoffm., have longer and less compact inflo- 
rescences than R. tomentosa and are completely 
glabrous; their leaf margins are prominently cre- 
nate-undulate (vs. entire or obscurely crenate-un- 
dulate). The recently described R. dressleri Web- 
ster (Ann. Missouri Bot. Gard. 75: 1094. 1988), 
from Panama and apparently Costa Rica, is dis- 
tinctive in its two-carpellate ovaries and fruits. 


Additional specimens examined. COLOMBIA. VALLE: 
Bajo Calima Concession, ca. 20 km N of Buenaventura, 
behind T Forest Station, 3940'N, 77?0'W, 50 m, 

lay 1987, Faber-Lagoenden et al. 675 (F); Bajo 
Calima, ca. 15 km N of Buenaventura, Cartón de Colom- 
bia Concession, 3%56'N, 77%08'W, ca. 50 m, 14 Feb. 
1983, Gentry et al. 40219 (F); 18 Feb. 1983, Gentry 
& Juncosa 4 ; 1980, Gentry et al. 
53048 (F); bs): Calima, Dindo area, pluvial forest, 3°59'N, 
. 100 m, 20 July 1984, Gentry & Monsalve 
48390 (MO). Bajo Calima, Cartón de Colombia Conces- 
sión, Dindo 4 area, 11 km E of Buenaventura -Río Calima 
road, 3955'N, 77°0’W, 50 100 m, 14 Dec. 1985, Gentry 
O), 27 Feb. 1985, Monsalve 716 (F, 
MO); Bajo Calima, Juanchaco Palmeras area, 3%55'N, 
77°02 50 m, 29 Aug. 1986, Gentry & Cuadros 
55602 (F): a e Concession Pulpapel Buena- 
ventura, 3955'N, ca. LOO m, Monsalve 870 (F). 


Tacarcuna Huft, gen. nov. TYPE: Tacarcuna gen- 
tryl 
Arbor, ut videtur dioecia; foliis cere simplicibus, 
Fon stipulatis; inflorescentiis axillar glomeratis; 
sepalis 3 vel 5, imbricatis; petalis 3 vel 5. ne 9s 
liberis; pistillodio SEM ovariis 3, uniovulatis; stylis 3; 
fructibus capsularibus 


Trees, apparently dioecious. Leaves alternate, 
simple petiolate; stipules small, deciduous. /nflo- 
rescences axillary, sessile or short-petiolate, glom- 
erate. Staminate flowers (seen only in T. gentryi) 
short-pedicellate; calyx lobes 5, imbricate, petals 
5, equaling or exceeding the sepals; stamens 5, 
free; pistillode well developed. Pistillate flowers 
pedicellate, the pedicel elongating in fruit; perianth 
as in the pistillate flowers (sepals and petals only 
3 in 7. tachirensis, in which staminate flowers are 
unknown), persistent on the mature fruit; carpel 
3-locular, the locules uniovulate; styles 3, divided 
at apex. Fruits capsular, the thin outer wall sep- 
arating from the thicker inner wall at dehiscence; 
seeds apparently ovoid-lenticular, smooth or per- 
haps sculptured, often misshapen, rarely seen ma- 


ture. 


This genus of small to medium-sized trees seems 


Volume 76, Number 4 


Huft 1081 
South American Euphorbiaceae 


quite unlike any other Euphorbiaceae in the Neo- 
tropics, and I have been unable to match it with 
any Old World representatives of the family. The 
collections available have mature capsules and 
young fruits that retain the pistillate calyx and 
corolla. Only one specimen (T. gentryi, Gentry et 
al. 16901 MO), however, has staminate flowers, 
and these are partially embedded in mounting glue. 
Additional material of 7. gentryi containing abun- 
dant staminate flowers was collected but has ap- 
parently been lost or misplaced (A. Gentry, pers. 
comm.). Since characters of the staminate flowers 
are of great importance in generic delimitation in 
the Euphorbiaceae, I have been reluctant for some 
time to describe this new genus, pending recovery 
or re-collection of staminate material. However, 
partial dissection of the available staminate flowers 
revealed five free stamens and a well-developed 
staminode; these features along with the glomerate, 
axillary inflorescences, presence of petals, and im- 
bricate calyx mark this as a previously undescribed 
genus. 

Only recently have collections of two additional 
species of this remarkable genus come to light, 
and, even though staminate material is lacking in 
these species, all three species are described to- 
gether in the hopes of stimulating further explo- 
ration. In addition to the characters already men- 
tioned, the three species of Tacarcuna are held 
together by the combination of ovate-lanceolate or 
obovate-lanceolate leaves with very short, stout 
petioles; the sepals and often the petals more or 
less densely puberulous centrally on the outer sur- 
face, and with distinctly hyaline margins; deeply 
lobed capsules somewhat depressed at the apex; 
and a network of low ridges on the mature capsules, 
especially prominent in 7. gentryi, less so in T. 
amanoifolia, and largely obscured by the veluti- 
nous indument in 7. tachirensis. 


KEY TO THE SPECIES OF TACARCUNA 


la. xke ba petioles brown strigillose; sec- 


can E s 5-7, broadly arching; sepals and 
pu ERN T. gentryi 
lb Es Ms and petioles glabrous; secondary veins 


more than 12, arching only at margins; sepals 

and petals 3. 

2a. Capsules glabrous to sparsely bet 

icular; leaves strongly cuspidate 
T. « nro 


sepals nav 
ACUMINATE iii 

2b. Capsules velutinous; se pals more or less 
plane; leaves acute or rarely short-act 
minate T. bo nsis 


Tacarcuna gentryi Huft, sp. nov. TYPE: Panama. 
Darién: lower montane wet forest, S slope of 


westernmost summit of Cerro Tacarcuna Mas- 
sif between Pucuro base camp and Tacarcuna 
summit camp, 1,400-1,600 m, 21 July 1970, 
Gentry, Leon & Forero 16869 (holotype, MO; 
isotypes, COL not seen, DAV not seen, F 
Figure 2. 


r mediocra, dioecia; ramulis junioribus dense pu 


brunneo-puberu 
eer le 8-10 mm altis, 10-12 mm latis, 
rece brunneo-pubescentibus, leviter porcatis. 


Tree 6-10 m high, dioecious; branchlets slender, 
densely puberulent. Leaves alternate; blades lance- 
ovate to elliptic, entire, (8-)10-16 cm long, 3- 
5.5 cm wide, glabrous or brown strigillose below 
toward the base; apex acuminate; base obtuse; mid- 
vein prominent; lateral veins 7-9 per side, prom- 
inent, arching; veinlets forming a prominent retic- 
long, 
glabrous or brown-strigillose; stipules deciduous, 
deltate, 1.6-2.2 mm long, 0.8-1 
lowly cucullate, brown strigillose. Inflorescences 
axillary, glomerate. Staminate flowers apparently 
several in each axil; pedicel ca. 2 mm long, finely 


ulum; petioles short, thickened, 3-5 mm 


mm wide, shal- 


and densely puberulous with short brown hairs; 
sepals 5, valvate, 1.2-1.5 mm long, oblong, round- 
ed at apex, densely brown puberulent outside ex- 
cept the broad hyaline margin, glabrous inside; 
petals similar; stamens 5, free; disk present, the 
features not clearly seen; pistillode well developed. 
Pistillate flowers 2-5 in each axil (seen only in 
fruit); fruiting pedicel 8-14 mm long, stout, finely 
and densely puberulous with short brown hairs; 
perianth as in staminate flowers, persistent in fruit; 
styles 3, strongly recurved, densely brown puber- 
ulent on the outer surface, glabrous on the inner 
surface, bifid at apex for ca. Y4-Y3 their length; 
style branches divergent and recurved. Immature 
capsules brown tomentose; mature capsules prom- 
inently 3-lobed, drying black, 8-10 mm high, 10- 
12 mm diam., pubescent with scattered short brown 
hairs, covered with a network of low ridges, the 
columella unknown; seeds (immature, misshapen) 
apparently reniform, ca. 8 mm long. 


Tacarcuna gentryiis known only from montane 
rainforest and elfin forest on the Cerro Tacarcuna 
massif on the border of Panama and Colombia. It 
is distinctive in its brown strigillose stems and pet- 
ioles, membranous leaf blades with widely spaced 
and broadly arching secondary nerves, and five- 
merous perianth. 

| am pleased to name this species after Dr. 
Alwyn H. Gentry, who in the course of two long 


1082 


Annals of the 
Missouri Botanical Garden 


e 
jè 
"nn 
dq 


IGURE 2. Tacarcuna gentry 
(Same Sn D a and b c. pA MORAN capsule 


expeditions in the mid 1970s was the first botanist 
to collect the rich and biologically significant flora 
on the Cerro Tacarcuna Massif on the boundary 
between Central and South America aa 197 n 


Additional specimens examined. PANAMA. DA 
Cerro Tacarcuna, W ridge Ay toward Río poe s 
below summit camp, 1,50 600 m, Gentry & Mori 


14121 Kis O), 14143 TN m" forest near top of west- 
eak of Cerro Tacarcuna Massif, 1,720 m, Gen- 
i Ph al 16901 (F, MO). 


Tacarcuna amanoifolia Huft, sp. nov. TYPE: 
eru. Maynas: Dtto. Iquitos, Río Momon, 3 


Le ies branch with mature capsules.—b. Bra 
-d. Mature capsule. (Based on a et al. 


nch with Denk capsules. 
6869.) 


km above jet. with Rio Nanay, ca. 105 m, 10 
Jan. 1976, S. McDaniel & M. Rimachi Y. 
20414 (holotype, MO-3622561). 


Arbor mediocra, ut videtur dioecia; ramulis dy 
glabris; folis lanceolate obo vatis, glabris; 
aris 12-16, se alis 3 
ad apicem cucullatis; capsulis fasciculatis, pedicellatis, 5- 
6 mm altis, 9-10 mm latis, parece pubescentibus, leviter 
porcatis 

Tree 7-13 m high, apparently dioecious; 
branchlets slender, glabrous. Leaves alternate, 


Volume 76, Number 4 
1989 


Huft 1083 
South American Euphorbiaceae 


short-petiolate; blades a s to oblanceolate-ob- 
ovate, entire, m long, (1.8-)3-5 cm 
wide, 1.9-2.8 times as is as wide, glabrous, 
somewhat leathery; apex cuspidate-acuminate; base 
acute to acuminate; midvein prominent on both 
sides; lateral veins 12-16 per side, arching near 
the margin of the blade, connected by a fine re- 
ticulum; petioles stout, 3-6 cm long, glabrous; 
stipules fugacious, not seen, leaving a horizontal 
scar 2-2.5 mm long. Inflorescences axillary, glom- 
erate. Staminate flowers not seen. Pistillate flow- 
ers 6-8 in each axil, seen only in fruit; fruiting 
pedicels 3-4 mm long in immature fruits, 6-9 mm 
long in mature fruits, + densely puberulous; sepals 
3, imbricate, strongly navicular, ca. mm long, 
+ densely puberulous except on the broad hyaline 
margins, glabrous inside; petals 3, exceeding the 
sepals, glabrous, cucullate at apex, the margins 
hyaline; styles 3, strongly recurved, ca. 2 mm long, 
+ densely puberulous on the outer surface, es- 
pecially toward the base, glabrous on the inner 
surface, bifid ca. 14-29 their length; style branches 
divergent and recurved. Immature capsules to- 
mentose, conspicuously 3-lobed; mature capsules 
deeply 3-lobed, not drying black, 5-6 mm high, 
9-10 mm diam., 


tered hairs, covered with an inconspicuous network 


sparsely beset with minute scat- 


of low ridges, the columella persistent, 3-3. 

long, with 3 narrow conspicuous wings running the 
length of the axis; seeds (misshapen) apparently 
5-6 mm long. 


This is the only species of Tacarcuna known 
from lowland areas; the two collections were made 
in occasionally inundated mature forest at ca. 0- 
100 m elevation. The leaf blades of T. amanoifolia 
are thicker and more lustrous than in 7. gentryi, 
and are marked by their more or less broadly 
obovate-lanceolate outline with a conspicuous cus- 
pidate-acuminate tip. These features along with a 
sometimes lighter color along the margins impart 
to the leaves an appearance strikingly similar to 
that of 4manoa guianensis Aublet, a widespread 
euphorbiaceous tree of the lowland forests of north- 
ern South America and southern Central America. 
Its short mature pedicels give the fruiting branch 
a more congested appearance than in T. gentryi. 


Additional specimen examined. PERU. MAYNAS: 
Dtto. Iquitos, Rio Nanay, two bends below entrance to 
Mapa Cocha, 14 Jan. 1976, McDaniel & Rimachi 20465 
(MO). 


Tacarcuna tachirensis Huft, sp. nov. TYPE: Ven- 
ezuela. Táchira: Cerro Las Minas, 18 km SE 


of Santa Ana, along steep slopes leading to 


Cerro Azul of Cerro Las Minas, 7?36'N, 72?13' 
W, 1,200-1,380 m, 6 May 1981, R. Liesner 
& M. Guariglai 11874 (holotype, MO; iso- 
types, DAV not seen, F, VEN not seen, and 
to be distributed). 


arva, ut videtur dioecia; ramulis junioribus gla- 
bris; foliis bp ovatis vel lanceolate obovatis, glabris; 
nervis secondariis 17-25; sepalis 3, extus leviter puberulis 
vel glabratis, ad marginem hyalinis, petalis 3, similibus; 
capsulis fasciculatis, pedicellatis, ut videtur 6-8 mm altis, 
12-13 mm latis, velutinis, leviter porcatis. 


Tree to 3 m high, apparently dioecious; branch- 
lets glabrous alternate, short-petiolate; 
blades elliptic, lanceolate-ovate or lanceolate-ob- 


Leaves 


ovate, entire, 7-12 cm long, 3-5 cm wide, 2-2.8 
times as long as wide, glabrous, chartaceous; apex 
acute or scarcely short-acuminate; base obtuse to 
rounded; midvein prominent on both sides; lateral 
veins 17-25 per side, arching near the margin of 
the blade, scarcely prominulous, + inconspicuous, 
connected by a fine reticulum; petioles 2-5 mm 
long, thickened, glabrous; stipules fugacious, lig- 
ulate, ca. 7 mm long, ca. 2.5 mm wide, rounded 
at apex, leaving a conspicuous horizontal scar. 
Inflorescences axilary, glomerate. Staminate 


flowers not seen. Pistillate flowers usually 4 in 


each axil, seen only in post-mature fruit; fruiting 
pedicels 1-1.5 cm long, + puberulous; sepals 3, 
persistent on mature capsules, imbricate, ovate, 
ca. 2.5 mm long, glabrous to sparsely puberulous 
except the hyaline margins, glabrous inside; petals 
3, similar to sepals; styles not evident (no intact 
capsules seen). Mature capsules apparently 6-8 
mm high, 12-13 mm diam., shallowly 3-lobed, 
densely velutinous, the network of low ridges barely 
discernible; columella persistent, ca. 5 mm long, 
thickened at base, with 3 narrow, scarcely dis- 
cernible wings running the length of the axis, these 
rarely larger and membranous; seeds smooth, 
brown, wrinkled, semiglobose with a vaguely tri- 
angular cross section, ca. 6 mm diam. 


This species is known only from a single collec- 
tion on steep forested slopes above 1,200 m. Only 
post-mature (i.e., already dehisced) capsules are 
available. The capsules are distinctive in their 
densely velutinous indument. The leaves are similar 
to those of 7. amanoifolia in their thicker texture 
and lustrous surface, but are more characteristi- 
cally lanceolate-ovate rather than lanceolate-ob- 
ovate, and have more closely spaced and more 
numerous secondary veins; the tips are acute (rare- 
ly short-acuminate) rather than prominently cus- 
pidate-acuminate as in T. amanoifolia. 


1084 Annals of the 
Missouri Botanical Garden 


Volume 76, Number 4 
1989 


Huft 
South American Euphorbiaceae 


1085 


Tragia rubiginosa Huft, sp. nov. TYPE: Peru. 
Loreto: Mishuyacu, near Iquitos, 100 m, Apr. 
1930, G. Klug 1176 (holotype, US-1456205). 
Figure 3. 


Frutex volubilis, monoeicus; caules pilosi. Folia alter- 


axillares, spicatae, ut t videtur unisexuales, longipeduncu- 
latae, plus minusve Pia ae. Flores masculi brevipedicel- 
1.5-1.8 mm longi, elliptici, valvati, 
mina 5; antherae subsessiles, glan- 
dulae discorum 5, intrastaminalos. Flores feminei lobis 
m longis, acutis; stig- 


cristis DOTEN reticulatibus; leviter puberulentes; semina 
ulata. 


globosa, laevia, 4-4.5 mm diametro, ecaruncu 


Vine, monoecious; stem pilose. Leaves alternate, 
on petioles 5-10 mm long, these densely hirsu- 
tulous; stipules narrowly lanceolate, appressed, 3.5— 
5 mm long, sparsely pilose, the base broadly and 
conspicuously hastate, the lobes rounded; blades 
lanceolate to oblanceolate, 7.5-11 cm long, 2.7- 
3.5 cm wide, 2.8-3.7 times as long as wide, gla- 
brous above or sparsely pubescent on midrib, very 
sparsely pilose below, acuminate at the apex, nar- 
rowed to a cordate-hastate base with rounded lobes 
ca. 1 mm long, the margin entire to remotely 
denticulate, 3-nerved from base, the central nerve 
giving rise to another 5-7 pairs of secondaries. 
Inflorescences axillary, spicate, apparently unisex- 
ual, + reflexed, the peduncle densely brown-pu- 
berulent, ca. 3 cm long. Staminate flowers on 
pedicels ca. 1 mm long; calyx lobes 5, drying black, 
1.0-1.3 mm long, 0.7-0.8 mm wide, elliptic, val- 
vate, reflexed at anthesis; stamens 5, the anthers 
subsessile, 2-locular, introrse, disk glands 5, in- 
trastaminal. Pistillate flowers with calyx lobes 5, 
ovate-elliptic, ca. 1.5 mm long, acute, lightly his- 
pidulous; stigmas nearly sessile, 3, thick. Capsules 
(not seen entire), apparently 3-lobed, with low re- 
ticulate ridges, lightly puberulent, apparently 6-7 
mm diam.; seeds globose, cream mottled with red- 
dish brown, smooth, 4-4.5 mm diam., ecaruncu- 
ate. 


This unusual species would key to sect. Bia 
(Klotzsch) Muell. Arg. in the treatment of Pax & 
Hoffmann (1919) on the basis of its five stamens 
and the presence of disk glands in the staminate 


flowers, but it seems hardly related to the other 
species of that section, which are characterized by 
their usually deeply cordate leaves (except 7. les- 
sertiana (Baillon) Muell. Arg.) and especially their 
robust, bipartite inflorescences with one branch 
staminate and the other pistillate. Were it not for 
the presence of the disk glands, however, Tragia 
rubiginosa might seem almost at home in the Cu- 
ban genus Platygyne Mercier, which Pax & Hoff- 
mann distinguished from Tragia by unisexual (vs. 
androgynous) inflorescences, 5-15 (vs. 2-50 usu- 
ally near the lower end of the range) stamens, 
globose hairy receptacles, free filaments, absence 
of a disk, and wide styles emarginate at the apex. 

Liogier (1971: 132, 133) was spurred by his 
study of the Hispaniolan species Tragia biflora 
Urban, which closely resembles Platygyne, to in- 
vestigate the relationship between Tragia and Pla- 
tygyne. He found that the generic characters are 
intermixed in some species referred to Platygyne, 
and particularly in 7. biflora, which has a globose, 
glabrous receptacle with 10-20 stamens and wide, 
but not emarginate, styles. His conclusion that 
Platygyne cannot be kept separate from 7ragia 
is bolstered by the existence of 7. rubiginosa, 
which has the five stamens and monoecious inflo- 
rescences typical of Platygyne but has a disk, flat 
glabrous receptacle, and nonemarginate styles. The 
inflorescences of T. rubiginosa are reminiscent of 
those of T. biflora, which likewise are monoecious, 
axillary, and somewhat reflexed, although consid- 
erably shorter and with a less pronounced peduncle. 
The leaves also seem similar but are longer, nar- 
rower, and less coarsely crenate in 7. rubiginosa. 

The specific epithet refers to the red-brown color 
of the flowers. 


LITERATURE CITED 


ARMBRUSTER, W. S. 1984. Two new species of Dale- 
champia (Euphorbiaceae) from Mesoamerica. Syst. 
Bot. 9: 272-278. 


8 )ecies, section, and synopsis 
of Paleo hampia, (E uphorbiac eae) from Costa Rica. 
Syst. p . 13: 
—_. ; TOR new species of Dalec pem 
(Euphorbiaceae) from Venezuelan Guayana. Brit 
nia =92 


GENTRY, Wi B 1977 7. Botanical exploration of Cerro 
Tacarcuna. Explorers Journal 55: 40-45. 
GREUTER, Chairman Editorial Committee. 1988. 
International Code of Botanical Nomenclature. Reg- 
18. 


GURE 3. iud a —a. Branch with immature inflorescences.—b. Staminate flower.—c. Young 


pistillate flower. — d. 


1086 Annals of the 
Missouri Botanical Garden 
Hurt, M. J. 1984. A new combination in Dalechampia 975. Conspectus of a new classification of 
da e Ann. Missouri Bot. Gard. 71: 341. the M ugue Taxon 24: 593-601. 
LIOGIER, A. 971. Novitates antillanae. IV. Mem. 1988. Three new species of Dalechampia 


New York E. Gard. 21(2): 107-157 
Pax, F. € K. HOFFMANN. 1919. Euphorbiaceae — : 
Das "id 


lypheae- ee In: A. s 
enreich IV. 147. IX (Heft 68): 1-108. 
STAFLEU, F. A. RS. Cowan. 1981. er . Taxonomic 
Literature, Volume 3. ii uod Veg. 105. 
WEBSTER, C. 1967 e. taxa of Dale- 


Tw 
champia. om Missi Bot. Gard. 54: 


(Euphorbiaceae) from Brazil. Brittonia m press). 

& W. S. ARMBRUSTER. 1979. A new Euglos- 
sine-pollinated species of Doral Euphorbi 
aceae) from Mexico. Pins mia 352 

& 1982. An unusual new nod of 
Dalec i Ne (Euphorbiaceae) from Surinam. Syst. 
Bot. 7: 48 


A REVISION OF Carel As Todas 
AMPELOCER A (ULMACEAE) 


ABSTRACT 


Ampelocera, a genus of Ulmaceae with nine species of low- to mid-elevation rainforest trees, occurs from Mexico 
to Brazil. This review deals with the systematics and nomenclature, including the description of three new species, 
A. albertiae Todzia from Colombia, 4. Pun Todzia from Ecua A. macphersonii Todzia from Panama, 
Colombia, and Venezuela. The taxonomic history of ii ion era is nube. followed by a key to the species. Brief 
descriptions and geographic ranges are provided for each speci 


EC 
o 
A 


Ampelocera is a little-known genus of nine is anomalous within the Ulmaceae by virtue of its 
species of neotropical trees. Although a frequent 4-16 stamens. On the basis of a summation of 
component of some low- to mid-elevation neotrop- evidence from various sources (morphology, anat- 
ical forests (A. Gentry, pers. comm.), Ampelocera omy, embryology, palynology, and flavonoid chem- 
is poorly represented in herbaria because it is a istry), Tobe considers Ampelocera best classified 
large rainforest tree and because specimens are in a family of its o 
not readily identified and often are misfiled. Am- In 1847 Klotzsch doaie Ampelocera with 
pelocera is characterized by its large tree habit, one species, Ampelocera ruizii, from a Ruiz 
buttressed trunk, oblique leaf bases, large number Pavón collection. He placed his new genus near 
of stamens, bifid style, and asymmetrical drupa- Celtis. Since then, Ampelocera has generally been 
. The genus ranges from central Mexico aligned within the subfamily Celtidoideae of the 
to Bolivia and southern coastal Brazil, and it occurs Ulmaceae with Celtis L., Trema Lour., Apha- 
in the West Indies. nanthe Planch., and Gironniera Gaudich., al- 

In preparing a treatment for the Flora de Nic- though Planchon (1873) considered its placement 
aragua, the literature dealing with Ampelocera within the Ulmaceae doubtful due to the anomalous 
was reviewed along with all material available from stamen number. 
the following herbaria: BM, CAS, F, K, LL, MO, In 1937 Baehni created the genus Plagioceltis 
NY, TEX, and US. Three new species as well as attributed to Mildbraed on the basis of unpublished 
changes in the currently accepted nomenclature notes on the Ruiz & Pavón type specimen at Ma- 
came to light. Since no summary of the systematics drid (Gentry, 1983). I strongly suspect that the 
and geography exists, a revision of the genus is unnumbered Ruiz € Pavón type specimen of Pla- 
now presented. A short history of Ampelocera is — gioceltis dichotoma is from the same collection as 
outlined below, followed by a diagnostic key for the type of Ampelocera ruizii Klotzsch (Ruiz 
the species and a review of their systematics and Pavón s.n., B!) because both bear leaves and flow- 


nomenclature. ers of exactly the same maturity (very young, not 
Historically included in the Ulmaceae, the genus — fully expanded leaves; flowers at anthesis; no fruits). 
has been the subject of recent studies on embryol- Since Ampelocera was erected in 1847, ten 


ogy (H. Tobe, Kyoto University, Japan, in prep.) species have been added to the original Æ. ruizii. 
and pollen (M. Takahashi, in prep.) that indicate The first addition, Ampelocera cubensis, was de- 
it is not well placed there. Although past authors scribed by Grisebach based on a Wright collection. 
have positioned the genus in the subfamily Celti- It was approximately 50 years before the next 
doideae based on its drupaceous fruit, Ampelocera species of Ampelocera, A. crenulata Urban and 


! [ thank Guy Nesom for providing the Latin diagnosis, Doris Lee Tischler i the drawing, M. Kubala for field 
observations, Robbin Moran for help with references, and L. Albert de Escobar, A. Gentry, M. Nee, G. Rogers, an 
H. van der Werff for their helpful comments on the manuscript. I also wish to ies the dr “of the following 
herbaria for borrowed material and/or help during visits: CAS, F, LL, K, BM, MO, NY, TEX, 

? Dept J: 


. of Botany, University of Texas, Austin, Texas 78712, U.S.A 


ANN. Missouni Bor. Garp. 76: 1087-1102. 1989. 


1088 


Annals of the 
Missouri Botanical Garden 


A. hondurensis J. D. Smith, were described. Those 
species, however, were not members of Ampeloc- 
era, nor even the Ulmaceae, and are presently 
placed into synonymy in genera in the Flacourti- 
aceae and Achatocarpaceae, respectively. In 1925 
Kuhlmann described 4. glabra Kuhlm. and 4. 
verrucosa Kuhlm. followed by 4. edentula Kuhlm. 
(1940) and 4. latifolia Ducke (1943) all from 
Brazil. In the dup uA of the most recently de- 
scribed species, 4. macrocarpa, Forero & Gentry 
(1984) Ed the need for taxonomic study of 
Ampelocera. 


MORPHOLOGY AND TAXONOMIC 
CHARACTERS 


Ampelocera species are small to large trees often 
with slender, channeled buttresses and smooth gray 
bark. The wood is light brown to cream-colored 


and hard. 
LEAVES 


The leaves of all species are pinnately veined 
except for those of Ampelocera hottlei, which are 
trinerved at the base. The texture of most leaves 
is coriaceous, but 4. cubensis and A. macphersonii 
have chartaceous leaves. The leaves of Ampeloc- 
era are glabrous except for those of 4. cubensis, 
which are sometimes softly puberulent beneath. 
Young leaves in most species are often many times 
larger than those on fertile branches. Young leaves 
of A. hottlei and A. macrocarpa are distinctive 
for their bright blue metallic color. 

Nearly all leaves of Ampelocera are oblong to 
elliptic with characteristic oblique bases and atten- 
uate apices. Ampelocera cubensis, A. glabra, and 
A. ruizii often have obscurely and coarsely dentate 
leaves. Petioles are usually thick and have a red 
flaky epidermis. 


INFLORESCENCES 


Ampelocera inflorescences are axillary, some- 
times simple, usually compound dichasia or rarely 
are panicles of compound dichasia. In A. glabra 
the rachises are so short that the inflorescences 
appear fasciculate. The longest and most densely 
flowered inflorescences in the genus with up to 56 
flowers are found in 4. longissima. Inflorescences 
have perfect flowers and functionally staminate 
flowers, the latter being found lower on the inflo- 
rescence axis. 


FLOWERS 


Ampelocera flowers are small, cream-colored, 
yellow, or green and offer a limited number o 


characters. All dimensions provided in the descrip- 
tions are based on dried material. The perianth 
consists of a sepalar cup with four or five usually 
sparsely puberulent lobes. The lobes in 4. cubensis 
are villous. 

Stamen number varies from 4 to 16 with the 
staminate flowers often having fewer stamens (4— 
8) than the perfect flowers. Two kinds of filaments 
are found in the genus: relatively thick filaments 
expanded near the base and drying flat characterize 
A. edentula and 4. macrocarpa, and relatively 
long, thin filaments that twist upon drying char- 
acterize the other seven species. The anther con- 
nective is extended beyond the thecae to form a 
strigose apicule. In the species with the short, broad 
filaments, the connective extension is very pro- 
nounced, while in those with long, thin filaments, 
the extension is smaller. 

The ovary in Ampelocera is puberulent and 
oblong to globose. The two style branches are free 
to the base in most species, but in 4. cubensis and 
A. ruizii they are united in the lower one-fourth. 
Staminate flowers often have cylindrical, ribbed 
pistillodes with the same vestiture as the ovaries 
in the perfect flowers. 

Size and shape of the drupaceous fruits of each 
species of Ampelocera are quite distinctive. Gen- 
erally the fruits of Ampelocera are yellow, asym- 
metrical, and transversely obovoid or ellipsoid; they 
vary from 0.6 to 2.2 cm long. The fruits may be 
glabrous (4. glabra, A. longissima), sparsely pu- 
berulent (4. edentula, 4. macrocarpa), densely 
pubescent (4. macphersonii), or densely pilose (A. 
rulzil). 
usual in the genus, are symmetrically oblong and 
velutinous with longitudinal striations. Ampelocera 


Ampelocera hottlei fruits, the most un- 


macrocarpa typically has pyriform fruits with an 
unusually thick endocarp. 


USES 


The wood of Ampelocera is hard and is used 
for firewood and lumber. In the Pichis Valley of 
Peru, the astringent bark of 4. edentula is used 
by the Campa Indians to tattoo skin. The bark is 
wetted and then bound around the arm, where it 
ulcerates the skin and causes a painful eruption. 
A coloring agent is then applied to the ulceration 
and scar tissue forms (A. Gentry, pers. comm.). 


SYSTEMATIC TREATMENT 


Ampelocera Klotzsch, Linnaea 20: 541. 1847. 
TYPE SPECIES: Ampelocera ruizii Klotzsch. 


ur ag Milbr. ex Baehni, e Field Mus. Nat. 
Hist., Bot. Ser. 13(2): 272. 1937 


Volume 76, Number 4 


dzia 1089 


To 
Revision of Ampelocera (Ulmaceae) 


Small to large trees with slender buttresses. 
Leaves petiolate, entire or coarsely dentate, usually 
glabrous, pinnately veined or palmately veined at 
base, with paired stipules. Inflorescences axillary, 
sparsely branched to much-branched dichasia, or 
the flowers fasciculate, with perfect flowers toward 
the apex and functionally staminate flowers toward 
the base; rachises 0.2-7.8 cm long with 2-57 
flowers. Flowers white, yellow, or green; calyx with 
4—5 lobes, these usually united in the lower half; 
petals absent; stamens 4-16; filaments 1-4 mm 


KEY TO THE SPECIES OF A MPELOCER A 


inflorescences borne on leafless branches; 


long, 0.5-1 mm broad, of two types, either thick 
and expanded near base and drying flat, or long, 
thin, and drying twisted; anther connective ex- 
tended beyond the anther at the apex; ovary su- 
perior, unilocular with a single pendulous ovule, 
glabrous or pubescent; style branches 2. Fruits 
drupaceous, yellow or orange, globose, or oblong, 
or obovoid to transversely obovoid, with persistent 
stamens and style branches; seeds globose, with 


thick cotyledons. 


style branches 


la oie amne ba apn broader than long (not known in 4. albertiae); filaments slender throughout. 
ally 


ntate, occasionally entire; 
3-5 5 mm 
3b. nup d ima not revolute; 


nflorescence rachises 
1.2 cm iun 1.2-1.6 cm broad, 


4b. — qs rachises 0.5-0.8 cm Mni with 4-9 flowers; calyx ca. 1.5 mm ^is fruits 1.5 
brous; southeastern and central coastal Brazil 4. 


, 2-2.3 cm broad, gla 

2b. Leaves is n flowers ya on ind bra 
5a. Inflorescences with : stamer 

anama, 

5b. Inflorescences with (9-)20— 
Hs ids (V is Ecuador. 

ence rachises (2.7-)3.5- 7 


bur 
3a. Leaf margins often revolute; filaments ca. 2 
filaments ca. 3 mm long; $ 
0.9-1.5 7 m iu: with 8-23 flowers; calyx 2-2.5 mm long; fruits 1- 
zl 9. 


northern rure. Rid caido V enezue la i 
flowers; stamens 4-12; leaves subcoriaceous, uniformly colored on 


2. 4. cubensis 


mm long; Cuba and Hispaniola ONCE 
South America. 
A. ruizii 


y pilose; Peru, Bolivia, Amazonian Bra 
D A. glabra 
nches; Vn branches 1-2 mm long. 

es chartaceous, unevenly colored on drying; 
ma icphersonii 


7.8 cm long, elongate, loosely flowered; ovary sparsely 
6. A. 


longissima 


p eden. Ec ene 


6b. Inflorescence rachises 0.6-1.9 cm long, congested, densely flowered; ovary densely pio se; 
Ls 3 


. albertiae 


Colombia 


d 


7a. Leaves trinerved at ase; style branche 


tomentose, with longitudinal striations; Mexico to Nicaragua 
s pinnately veined at the base; style branches 1-2 mm long; stamens (6-)8; fruits abres to 


Fruits obovoid to A not strongly asymmetric a longer than broad; filaments broadened bas a 


fruits densely brown 


4 mm long; stamens ca. 16; 
. hottlei 


sparsely puberulent, yellow; Honduras to Panama, South America. 


"ir densely puberulent; fruit with a thin endocarp; greater Amazon Basin in Surinam, Guyana, 
Colombia, Ecuador, Peru, and 3. 


azil, Venezuela, 


Bolivia . edentula 


8b. inm sparsely puberulent; fruit with a thick endocarp; Honduras to northern Colombia and 


. macrocarpa 


northern Venezuela 


- 


Ampelocera albertiae sp. nov. TYPE. Co- 
lombia. Valle: carretera Cali-Buenaventura, 
km 24, 1,750 m, 15 Oct. 1982, L. Albert 
de Escobar, J. Folsom, J. Brand & D. Sán- 
chez 2788 (holotype, TEX; isotypes, HUA, 
MO). Figure 1. 


Ampelocerae longissimae similis sed inflorescentiis den- 
n et ovarus dense strigosis Mun nflorescentiae 
ad basim ramosissimae, floribus (9-)20-52, sepala 
-2 mm e ongae, stamina 8(- 12) in floribus perfec tis, 
filamenta 1.5-2 mm longae, tenues, fructus ignotus. 
Trees 4 m tall; younger branches very sparsely 
puberulent, reddish brown; older branches gray, 
glabrous; stipules 3-4 mm long, lanceolate, sparse- 
ly to moderately strigose. Leaf blades elliptic, 9— 
12 x 3-6 cm, the apex acute, the base attenuate, 


drying coriaceous, dull green to brown, uniformly 
colored, glabrous and smooth above and beneath, 
the margins entire; lateral veins 5-6, gradually 
arcuate, raised beneath; petioles 0.5-0.6 cm long, 
smooth, drying dark brown. Inflorescences axillary, 
short panicles on leafy branches, often much- 


branched at base, with perfect and functionally 
staminate flowers; rachises 0.6-1.9 cm long, 
densely flowered, with (9-)20-52 flowers, sparsely 
strigose. Flowers white, subtended by ovate, char- 
taceous, acute bracts ca. 1 mm long; calyx 1.5- 
2 mm long, the lobes 4, united at the base, glabrous 
to sparsely strigose, ciliate; stamens 8(-12) in per- 
fect flowers, 4-8 in functionally staminate flowers, 
these with a strigose pistillode; filaments 1.5-2 mm 
long, slender throughout; anthers ca. 1 mm long, 
with the connective extending into a short strigose 


1090 


Annals of the 
Missouri Botanical Garden 


^ d 
NM 
di 


= PIPE AR 38 


Ampelocera albertiae 


Ampelocera albertiae.—a. Fertile branch showing close-up of stipules.— b. Perfect flower 


FIGURE 1. 
Albert de Escobar et al. 2788 TEX.) 


-~ 


From 


Volume 76, Number 4 
1989 


Todzia 
Revision of Ampelocera (Ulmaceae) 


1091 


30 60 50 
| En | 
- + — SE E Za 
20 WN | | 20| 
X | e 
DO 
| we 
E » 
'oj- | z 10 
| 8 
| 
| 
O ^ JR s 
| 2 P 
( S 
| WW d 
e A. macrocarpa mes ) 
; ¿10 
" a A. albertiae e a 
€ A. edentula | El y 
| . / 
0 200 400 600 800 1000km | +f t 2 C — 1 
) 100 200 300 400 500 600 mil | EE Nr Ni | i 
2 [ miles E P m: l ny 
20 | IV ^ ( E XA J LT 
| E Ae PS = i 
90 207 70 : Es * 


FIGURE 2. Distributions of Ampelocera albertiae, 


apicule ca. 0.1 mm long; ovary densely strigose, 
the style branches 1.5-2 mm long, free to base. 
Fruit not seen. 


Distribution, habitat, and phenology. This 
species is known only from the western slope o 
the western cordillera of Colombia (Fig. 2). Flow- 


ering is recorded in October 


Presently known from only one specimen, more 
collections are needed to understand the morpho- 
logical variation and relationships of Ampelocera 
albertiae. Ampelocera albertiae is characterized 
by having entire leaves, densely flowered inflores- 
cences on leafy branches, flowers with relatively 
thin filaments, and style branches 1-2 mm long. 
This species appears to be most similar to Ampe- 
locera longissima but differs by having densely 
flowered inflorescences and densely strigose ova- 
ries. The unusual, new species is named in honor 


A. edentula, and A. macrocarpa. 


of its discoverer, Dra. Linda Albert de Escobar, of 
the Universidad de Antioquia, Colombia (HUA), a 
student of Passifloraceae. 


2. Ampelocera cubensis Griseb., Catalogus 
Plantarum Cubensium. 57. 1866. TYPE. Cuba. 
Pinar del Rio: Loma de Rangel, 28 May 1863, 
Wright 2222 (lectotype [designated by R. A. 
Howard (1988, Fiche 2, B 11)], GOET; iso- 
lectotypes, B, GH, GOET, K, MO). 


P era deo scens e Md den Biol. Soc. Wash. 

: 153.19 T Cuba. Santa Clara: Limones, 
ienfuegos, p? Apr. coon J. G. Jack 6036 
JS; isotypes, A —3 sheets, B, F (frag.), 


ere 
(holotype, | 
, NY). 


Trees 5-30 m tall; younger branches sparsely 
to densely puberulent, light brown; older branches 
glabrous, light gray to brown; flowering branches 


1092 Annals of the 
Missouri Botanical Garden 
90 60 50 
e | 
G | | 
H—É 1 | 
20 | | 
y pu te 
Mi E 
io | 
9 
ol Sl. 
z 
e A. cubensis 
O 
| a A glabra 
e A. ruzi 
9 200 400 600 d TOROK 
20 0 100 200 300 400 500 600 miles 
| 
90 
FIGURE 3. Distributions of Ampelocera cubensis, A. glabra, and A. ruizit. 


with shortened internodes; stipules 2-5 mm long, 
lanceolate, sometimes falcate, acute, pilose. Leaf 
blades narrowly to broadly elliptic, 7.5-17 x 3.5- 
6(-9.5) cm, the apex gradually acuminate with the 
acumen 1-2 cm long, obliquely rounded at base, 
the margins entire, or obscurely dentate with 2-4 
teeth per side, or obscurely crenulate, often rev- 
olute, drying chartaceous, light brown, the lateral 
veins 5-7, arcuate, smooth and glabrous above 
and beneath, the midvein puberulent to glabrous 
beneath; petioles 0.8— cm long, puberulent. 
Inflorescences axillary on leafless branches, fascic- 
ulate or in very short racemes with only perfect 
flowers; rachises 0.2-1 cm long with 2-5 flowers, 
glabrous to puberulent. Flowers (color not known) 
subtended by glabrous to sparsely puberulent bracts, 
the ovate bracteoles 2-3 mm long; calyx ca. 2 mm 
long, the lobes 4—5, united at base, pilose to villous; 


stamens 12-16; filaments ca. 2 mm long, linear, 
slender throughout; anthers 1-1.3 mm long, the 
connective slightly extended into an apicule; ovary 


road, puberulent; endocarp ca. 


0.5 mm thick. 


Common names. hueso de costa, jatia blanca 


Cuba) 


— 


Distribution, habitat, and phenology. 
Uncommon in coastal thickets, limestone cliffs, and 
rocky woods in Cuba and on western Hispaniola 
(Fig. 3) from O to 300 m. Flowering apparently 
occurs February to June when the trees are leafless; 
fruiting specimens have been collected in April, 


May, July, and December. 


Volume 76, Number 4 
1989 


Todzia 1093 
Revision of Ampelocera (Ulmaceae) 


Additional specimens examined. CUBA. HAVANA: 
near s de Mariano, Feb. 1922, Bro. León 10677 
(GH, NY, US); Sierra de Anafe, 8 Aug. 1923, Bro. n 
11486 (GH, NY). LAS VILLAS: on limestone outcrop 
Fanen owns at Gavilán, 1- 20 July 1950, Howard 
et al. 21 (A, NY). ORIE 
1943, 2 Clemente 2984 (GH). rto 
18 Nov. 1915, Te 6629 (F, K, US); WEA p 
Ca suta; 250 m, 2 May 1919, Ekman 9626 (NY); v 
, 21 Feb. 1919, n 9470 (LL, NY, US 
shes). PINAR DEL RÍO: Sierra de Anafe, 18 Dec. 1911, 
P. Wilson 11398 (NY —3 sheets, US). SANTA CLARA: 
Limones, O que us 12 Mar. 1927, E 4911 

NY, US); 3 Sep. 1927, Jack 5396 (US); Eis 
1928, Job Le A — 2 sheets, NY, US); 
Jack 7490 (A, NY, ue va woods, 
eastern Tapata, 1 Aug. 
NY). DoMINICAN REPUBLIC. SANTO DOMINGO: Peninsula de 
peer top of Cerro Jean-José, ca. 300 m, 23 Sep. 
Ekman 7044 (GH). HAITI: L'ARTIBONITE: Massif 

des ad Dessalines, limestone cliffs at Case-à-Roche, 

3 Mar. 1925, ge 3506 (LL); Gonave Island, Anse 
à [ed Aug. ] , Eyerdam 214 (A, GH, NY, US 


ae 
Co 
— 
"ES 
2 

o 

N 
Ot 


This species is characterized by relatively small, 
thin, sparsely dentate leaves; short, fasciculate in- 
florescences that are borne on leafless branches; 
style branches 3-4 mm long; linear filaments; and 
small, puberulent, strongly asymmetrical fruits. 
Ampelocera cubensis is closely related to A. gla- 
bra from central coastal and southeastern Brazil 
and 4. ruizii from Amazonian Brazil, Peru, and 
Bolivia. It differs from 4. glabra by having smaller 
fruits and less prominently dentate leaves and dif- 
fers from 4. ruizii by having shorter inflorescences 
with 2-5 (vs. 8-23) flowers and by having sepals 
1.5 (vs. 2-2.5) mm long. 

Morton (1958) described Ampelocera pubes- 
cens, distinguishing his new species from 4. cu- 
bensis by the vestiture of the lower leaf surface 
and the length of the stipules (4.5-6 mm in 4. 
pubescens vs. 3 mm in A. cubensis). | can find 
no other characters to corroborate the recognition 
of two taxa, and, in fact, vestiture of material 
annotated as 4. pubescens by Morton, including 
the type (Jack 6036), ranges from densely to 
sparsely pilose. 


3. Ampelocera edentula Kuhlm., Annais Prim. 
Reu. Sul. Amer. Bot. 3: 75. 1940. TYPE. Bra- 
zil. Pará: “In silva collina inter locos Piquia- 
tuba et Cipoal," 18 Jan. 1933, Ducke s.n. 
(RB # 24.565) (holotype, RB, n.v.; 
K, US). 


isotypes, 


Mi latifolia Ducke, Trop. Woods 76: 15. 1943. 
TYPE. a Td eni Esperanga, boca do Tox d 
29 Oct Ducke 1039 (holotype, RB, 
isotypes, Me d NY, US) 


Trees 12-30 m tall, up to 90 cm dbh, with 
triangular buttresses 1.5-3 m high; young branch- 
es dark reddish brown, glabrous to puberulent, 
lenticellate, smooth; older branches light grayish 
brown, with exfoliating bark; stipules 2-6 mm long, 
narrowly to broadly lanceolate, acute, strigose. Leaf 
blades elliptic to oblong-elliptic, 8-26(-35) x 3.5- 
12 cm, apex acuminate with tips 0.2-1.2 cm long, 
the base rounded, cuneate to oblique, with margins 
entire, lustrous above, glabrous, drying smooth 
above and beneath, dull green to brown, pinnately 
veined; lateral veins 5-7, arcuate, raised beneath; 
petioles 0.5-1.4 cm long, glabrous or with a red 
flaky epidermis. Inflorescences 1-2 per axil on 
leafy branches, short, much-branched, densely 
flowered panicles, with staminate and perfect flow- 
ers; rachises 0.8-2.8 cm long, branching almost 
to the base, densely flowered, with 6-27(-55) flow- 
ers, puberulent. Flowers white to greenish yellow, 
subtended by bracts 0.5-2 mm long, these reddish, 
ciliate, ovate, acute, chartaceous; pedicels 0.5- 
mm long, narrower than the rachis and flower base; 
calyx 1.5-2 mm long, the lobes 4, united in lower 
half, glabrous to sparsely puberulent; 

-)8 in perfect and staminate flowers; filaments 
3-4 mm long, broadened basally; anthers ca. 1l 


stamens 


mm long, often curved inward, with the connective 
extended into an apicule 0.5-1 mm long; ovary in 
perfect flowers densely puberulent, oblong, 2-3.5 
mm long, with style branches mm long; 
pistillode in staminate flowers oblong, strigose at 
the apex and with 5-6 strigose ridges. Fruits yel- 
low, globose to oblong, only slightly asymmetrical, 
glabrous to sparsely puberulent, scabrous, (0.8-) 
1.6-2.2 cm long, 1-1.7 cm broad; endocarp thin- 
ner than 0.5 mm. 


Common names and uses. | Nina-caspi (Peru: 
San Martin). The fiber of the bark is strong, and 
when bound around the arm it causes painful erup- 
tion; it is toxic and vesicans. The fruit is hypnotic. 
(Data fide Woytkowski 5449.) 


Distribution, habitat, and phenology. This 
species is found throughout the Amazon Basin from 
near sea level to 750 m in Surinam, Guyana, 
Venezuela, Brazil, Colombia, Ecuador, Peru, and 
Bolivia (Fig. 2) in primary tropical moist forest and 
nonflooded and flooded riverine forests. Flowering 
and fruiting apparently occur throughout the year. 


Additional specimens examined. BRAZIL. ACRE: near 
mouth of Rio Macauhan (tributary of Rio Yaco), 9°20’S, 
69*W, 3 Aug. 1933, Krukoff 5266 (K, MO 
NY, US). AMAZONAS: Manaus, Estrada do Aleixo, 27 Aug. 
36 (A, F, NY, US); Rio Purús, E. Ituxi, 
79 (MG) 


2 Sheets, 


1945, Ducke 17 
SE of Lábrea, 29 June 1971, Prance et al. 139; 


1094 


Annals of the 
Missouri Botanical Garden 


near mouth of Rio Embira (tributary of Rio Tarauaca), 
7°30'S, 70°15'W, Krukoff 4877 (A, F, K—2 sheets, 

MO, US): Serra da Neblina, Rio Cauaburi, beyond eek 
of Tucano Igarapé, 125 m, 16 Nov. 1965, Maguire et 
al. 60185 (MO). PARA: near EMBRAPA statal, at km 
23 on road Altamira-Itaituba, 29 Oct. 1977, Berg et 
al. 734 (MO); Tucurui, Cagancho, Rio Tocantins, Jan. 
1981, Lisboa et al. 2114 (MG). RONDÔNIA: Municipio de 
Ouro Preto do Oeste, BR 364, rodovia Cuiabá- Porto 
Velho, km 382, rodovia 470, linha 81, em diregao a 
i 10°1 me Oe ne W, C. A. Cid 
„Arbor no. 


Cerro Neblina, between base camp and * 
along Río rd dun o 180 m, 0°5 ON, ca. 66°8'W, 
26 Apr. ^ & Stein 46979 
COLOMBIA. META: Sierr M la Macarena, 
550 m, 13 Jan. 1950. Philipson et Dur 2107 (l iS). 
VAUPES: Rio Apaporis, entre el Rio Pacoa y el Rio Kan- 
anari, Soratama, 250 m, 28 Sep. 1951, Schultes & 
Cabrera 14147 (ECON — 2 sheets). Ecu ADOR. NAPO: Es- 
tación oo, INIAP-Payamino, 5 km al N de 
Coca, 0%2: 77°00'W, 250 m, 10-15 Se ep. 1986, Neill 
un (TEX); Anangu, Parque Nacional Yasuni, 
0°31 39'S, 76?23'W, 260-350 m, Studies of Ecuador- 
ian Forests 9154 (NY). PERU. HUÁNUCO: vicinity of Tingo 
oan we Rio Huallaga above docks by coffee tanks, 
5 June 1960, Mathias & Taylor 5002 (F). MADRE DE 
DIOS: Tambopata Tourist Camp at in E rios Tam- 
Ay and La Torre, 280 m, 12°49'S 43'W, 24 
July. 1985, doenty et al. 51256 (MO). SAN MARTÍN: Prov. 
, Dtto. Tocache Nuevo, 15 Sep. 1970, 
1395 (F. GH, K, MO, NY, US); Gramalote ad 
o 600 m, 1 Oct. 1959, Woytkow ski 5449 (MO); 
17 x 1962, Woytkowski 7307 (F, GH, MO, NY, 
TEX, US) rer ad Saposoa, 600 m, 14 Apr. 1962, 
Woyikowski 7291 (F, MO— 2 sheets); 20 ei 1962, 
W oytkow ski 7312(MO). BOLIVIA. LA PAZ: Prov. Larecaja, 
Tuiri, near ae (on left bank of Rio Mapin) 490-7 50 
m, 12 Sep. 1939, Tuer 10732 (A, F, K, MO, 
US), qu (A, F, K, MO, 


— 
A 


Ampelocera edentula is characterized by having 
flowers with eight stamens; broad filaments with a 
pronounced terminal extension of the connective; 
and large yellow, globose, symmetrical fruits. This 
species is most closely related to A. macrocarpa 
from which it differs in having generally oblong 
fruits and a thin endocarp (vs. globose fruits and 
a thick endocarp). 

In his description of Ampelocera latifolia, Ducke 
(1943) stated that his new species might be just 
an upper-Amazonian variant of 4. edentula. The 
fruit on the type of 4. latifolia is distinctly oblong 
while that on the type of 4. edentula is smaller 


only a difference in fruit maturity, and since I 
could find no other consistently correlated char- 
acters, A. latifolia is placed into synonymy with 
A. edentula. 


Although Ampelocera edentula occurs through- 
out a broad geographic area in the Amazon Basin, 
it is poorly represented in herbaria, and conse- 
quently, its morphological variation is difficult to 
assess. 
branches, 


eaves are generally shorter on fertile 

12-20 cm long, while sterile branches 
have leaves up to 30 cm long. Some collections 
from Peru and Bolivia (Schunke 4395, Woyt- 
kowski 7307 (Perú: Dept. San Martin); Krukoff 
10727 (Bolivia: La Paz)) have leaves up to 35 cm 
on fertile branches. These populati liffer further 
from typical Brazilian 4. edentula in have shorter 
petioles, more oblique leaf bases, and more pubes- 


cent fruits. A sterile specimen from Peru (Dept. 
Loreto: Prov. Requena, Jenaro Herrera, Rio Uca- 
yali, 73°45'W, 4%55'S, 22 Feb. 1987, Gentry et 
al. 56289 (MO)) is unusual in having sparsely 
pubescent stems, petioles, and the lower surface 
of the primary and secondary veins. As more spec- 
imens become available, the variation presently 
included in 4. edentula may warrant recognition 
of other taxa. 


4. Ampelocera glabra Kuhlm., Arch. Jard. Bot. 
Rio de Janeiro 4: 351, pl. 28. 1925. TYPE. 
Brazil: **Rio de Janeiro ad urbem in selva loco 
Fabrica Allianca," 8 Nov. 1922, Kuhlmann 
6794 (holotype, B; isotypes, K, MO, 


Trees 3-20 m tall, with a spreading crown; 
trunk brown, up to 66 cm diam.; older stems gray, 
glabrous; younger stems brown, glabrous; stipules 
ca. 5 mm long, lanceolate, glabrous or sparsely 
pilose. Leaf blades elliptic, 7.5-14 cm long, 2.7- 
7.2 cm wide, the apex acuminate, the tip 0.3-1.5 
cm long, obliquely rounded at base, the margins 
dentate with 4-6 teeth, seldom entire, smooth, 
glabrous, drying subcoriaceous, the late 
5-8, straight in lower %4, slightly arcuate near 
margin; petioles 0.5-1 cm long, glabrous, drying 
dark brown. Inflorescences fasciculate or very short 
racemes or cymes, axillary on leafless stems, with 


ral veins 


perfect and staminate flowers having rudimentary 
pistils; rachises 0.5-0.8 cm long, sparsely puber- 
ulent, with 4-9 flowers. Flowers green, subtended 

y bracts ca. 1.5 mm long, glabrous to sparsely 
pilose; calyx ca. 1.5 mm long, the lobes 4, united 
at the base, ovate, sparsely pilose; stamens 16 in 
bisexual flowers, 12-14 in functionally staminate 
flowers; filaments ca. 3 mm long, slender through- 
out, with the connective extended into an apicule 
ca. 0.3 mm long; anthers 1-1.5 mm long; ovary 
sparsely puberulent and verrucose, the style 
branches ca. 4 mm long. Fruits yellow, asymmet- 
rical, transversely obovoid, 1.5-2 cm tall, 2-2.3 


Volume 76, Number 4 
1989 


Todzia 1095 
Revision of Ampelocera (Ulmaceae) 


cm broad, glabrous, with persistent style branches; 
endocarp thin. 


Common name. Mentira (Brazil). 

Distribution, habitat, and phenology. This 
species occurs in primary forest in southern coastal 
Brazil (Fig. 3). Flowering August to November usu- 
ally on leafless branches. 


iei ina specimens examined. BRAZIL. ESPIRITO 
SANTO: Reserva Florestal da CVRD, Linhares, Est. Gavea, 
ant. X-2, KM 20,790, 20 Aug. 1979, Foli 98 (MO). 
MINAS GERAIS: Distr. Rio Branco, retiro de Antonio Ave- 
lino, 750 m, 27 Dec. 1930, Mexia 5456 (MO). RIO DE 
JANEIRO: cultivadad no Horto Florestal da Gávea, 7 Jan. 
1932, Fictorio s.n. [RB no. 139428] (B); Huto Florestal, 
. 1928, ilm s.n. [RB no. 139426 
23 Dec. 1926, Kuhlmann s.n. 


“si 


[RB no. 139430] (F, US). 


Ampelocera glabra is distinguished by its small 
fascicles of flowers; large, strongly asymmetrical, 
glabrous fruits; and toothed leaves. It is clearly 
most closely related to and can be easily confused 
with A. ruizii but is geographically disjunct. The 
straight instead of arcuate lateral veins in A. glabra 
are distinctive in the genus. 


5. Ampelocera hottlei (Standley) Standley, 
Trop. Woods 51: 11 


Celtis hottlei pcd Trop. Woods 20: 20. 1929. TYPE 
onduras. Yoro, near Progreso, 4 Apr. 1929, Ba 
tle 32 TU ant F). 


Trees 10-30 m tall; trunk 10-50 cm dbh, with 
narrow buttresses, ca. 2 m tall; crown subglobose 
or spreading; bark smooth, white to gray with dark 
lenticels, hard, with a sweet odor; sapwood creamy 
yellow to light brown; branches light brown-gray, 
lenticellate; stipules ca. 4 mm long, linear-lanceo- 
late, puberulent. Leaf blades oblong to elliptic, 
(7-)8.5-16(-26) x (2.6-)3.3-8(-10.5) cm, the 
apex acuminate with an acumen 0.2-3 cm long, 
obliquely attenuate to rounded at base, the margins 
entire, drying chartaceous to subcoriaceous, dull 
dark green above, dull light green beneath, the 
young leaves blue to purple, glabrous and smooth 
above and beneath, the lateral veins 3-4(-5), ar- 
cuate, only slightly raised beneath, the base pal- 
mately veined with basal pair arising at the petiole; 
petioles 0.6-1.2 cm long, glabrous, minutely ver- 
rucose, usually drying dark brown. Inflorescences 
axillary on leafy branches, 1-2.5 cm long, com- 
pound dichasia with perfect flowers toward the apex 
and functionally staminate flowers toward the base; 
rachises 0.7-1.5 cm long with 8-17 flowers, pu- 
berulent. Flowers purplish to yellowish green, sub- 


tended by puberulent bracteoles 1-2 mm long; 
calyx 1-2 mm long, the lobes 5, united at base, 
puberulent abaxially; stamens ca. 16 in perfect 
flowers, ca. 8 in staminate flowers; filaments ca. 2 
mm long, linear-lanceolate, broadened basally; an- 
thers ca. 1 mm long, anther connective extende 

ca. 1 mm into a puberulent apicule; ovary puber- 
ulent, the style branches ca. 4 mm long, united in 
lower 2-3 mm. Fruits yellow, symmetrical, obo- 
void, 1.2-1.5 cm tall, 1-1.2 cm wide, velutinous, 
with persistent style branches; endocarp ca. 1 mm 
thick 


Common names and uses. Mexico: coquito, 
cautivo, guaya, ojoche blanco, popo mojo. Belize: 
bullhoof, luin. Guatemala: luin, tisón. El Salvador: 
tisón. Nicaragua: cuscano, yayo. The wood is used 
for house construction and railroad ties. 


Distribution, habitat, and phenology. This 
species occurs from central Mexico to Nicaragua 
(Fig. 4) in primary, undisturbed rainforest or trop- 
ical wet forest, sometimes on limestone. Flowering 
reported from February and fruiting from March 
to June. 


Representative Lh imens po MEXICO. CAM- 
PECHE: El Tormento, near Escárega, 17 May 1968, Pen- 
nington & Sarubhn 9639 (A. K NY). CHIAPAS: zona 
comprendida entre los ríos Chancaláh, Chocoljahíto, y 
Tulijá, Selva a 250-450, 29 Nov. 1967, Pen- 
nington & Sarukhán 9363 (NY); road to the ruins, La 

rena, 13 Mar. 1958, Schubert & Gómez Pompa 1731 
(A). oaxaca: Mpio. Sta. María Chimalapa, ca. 7 km al 
oeste de Santa María en la vereda de Chicusaja, 250 m, 
30 June 1984, H. Hernández G. 163 (CAS); Ubero, 30- 
90 m, June 1937, LL. Williams 9497 (A, F, US). TABASCO: 
Villahermosa (Centro), Selva Dos Montes, atrás del aero- 
puerto 3.6 km (2.3 mi.) por camino rural de la entrada 
al aeropuerto, 8 May 1984, Cowan & Zamudio 4672 
(TEX). VERACRUZ: Estación de Biologia Tropical " 
Tuxtlas," Mpio. San Andrés Tuxtla, 26 Mar. 1974, Cal. 
zada 1166 (CAS, MO); Estacion de Donga Los Tuxtlas, 
170-200 m, 1 June 1981, Gentry et al. 32522 (MO); 


1985, Wendt et al. 4803 (TEX). : 
entin, ee July 1936, Lundell 6225 E. NY), 6346 (F, 
NY), 6423 (F). EL TOLEDO: d River, 21 Feb. 1945, 
Gentle 52 12 (F, LL, TEX); near Moffredye Lagoon, 27 
lar. 1945, Gentle 5291 (CH LL, TEX, US); Temash 
River, 200 ft., 4 Mar. 1935, Schipp 1347 (F, K, MO 
NY). GUATEMALA. ALTA VERAPAZ: along Rio Icvolay be- 


and Río Soctelá, 220-210 m, 14 Mar. 
(F); woods SE of Finca Yal- 


tween Rio Apia 
1942, Steyermark 45038 


HUEHUETENANGO: between Ixcan ar 
de los Cuchumatanes, 150-200 m, 23 July 1942, Stey- 
ermark 49219 (F). IZABAL: between Bananera and “La 
Presa" in Montaña del Mico, 40-300 m, e Mar. 1940, 
Steyermark 38175 (F); perd ca. .0 m, 3 May 1939, 
Standley 72886 (A, F), 7295. , NY), Rio Juyamá, 


1096 Annals of the 
Missouri Botanical Garden 
90 80 70 60 50 
EA de 
D | EN d MES i 
{ x " | 
20 | 
^ | F 
eee: | 
es. | 
K f É e | 
os | 
| SSe | 
10 | r 
8 | 
| 
| 
| 
(0) ^ JR ` 
2 o AU 
e 4A.hottlei 
10 m A. macphersonii 
è A. /ongissima 
| 
| 
| 
| 
o 400 600 800 1000km t 7 
20 O 100 200 300 400 500 800 miles | 
90 80 


FIGURE 4. Distributions of Ampelocera hottlet, 


SE of Cheyenne, ca. 15 mi. SW of Bansnera, 50-100 
m, 8 Apr. 1940, Steyermark 39153 (F). PETÉN: Paso 
Caballos, 5 Apr. 1965, Contreras 5387 (LL, TEX). Tikal 
National Park on Remate Road, 4 Mar. 1961, Lundell 
16877 (F, LL; W and NW of NM and 6 mi. W of 
Rio San Román, 50-70 m, 29 Mar. 1942, Steyermark 
45500 (F). RETALHULEU: between Retalhu jeu and Nueva 
Linda, 120-220 m, 18 Feb. 1941, Standley 87284 (F), 
88487 (F), 88520 (A, de SANTA ROSA: Rio de las Pitas, 
E of Taxisco, ca. 225 m, . 1940, pu 78928 
(F). EL SALVADOR. AHUACHAPÁN: Sierra de Apaneca, in 
the region of Finca Colima, 17-19 Jan. 
20149 (G iis HONDURAS. COLÓN: Guarur 
UT 75 O ft., Mar. 1938, Hagen « po 1372 
(F, NY), aie (F, NY). YORO: near Progreso, 11 Apr. 
s A Hottle 108 (F). Nic. > / 
5 km W of Matiguás, 500 m, 5 May 1984, 
ire (MO). ZELAYA: Kurinwacito, ae 100 m, 
Mar. 1984, Moreno 23654 (MO); 
tonio, 2 ae N of Kuikainita, 100 m, 
4157 (MO) Mpio. Siuna 
Ortiz 1938 (MO); El Zapote, 40 km NE of Nueva Guinea, 


— 


between Quebrada El Toro 


A. macphersonii, and A. longissima. 


25-31 Mar. 1984, Sandino 4914 (MO); along Rio lyas 
l , 260-280 m, 13 May 1978, 
Vincelli 367 (MO). 

Ampelocera hottlei is represented by the most 
collections and thus is the best-known species in 
the genus. Unlike other species, description of the 
trunk, bark, and sap is available from collection 
data. It is distinguished from all other species by 
its trinerved leaf base, and obovoid, densely pu- 
bescent, longitudinally lined fruits. As with 4. mac- 
rocarpa, the young leaves are bright metallic blue. 


6. Ampelocera longissima Todzia. TYPE. Ec- 
uador, Napo: Estación Biológica Jatun Sacha, 
Rio Napo, 8 km al E de Misahualli 01?04'S, 
77°36'W, 450 m, 28 Dec. 1987, Neill, Gen- 
try & Manning 8135 (holotype, TEX; iso- 
type, MO). Figure 5. 


Volume 76, Number 4 
1989 


Todzia 1097 
Revision of Ampelocera (Ulmaceae) 


Inflorescentiae pro genere longissimae, (2.7-)3.5- 7.8 
cm longae, flores 33-56. Sepala ala 4 glabra in dimidio 1 in- 


longi. Fructus leviter lutei transverse obovoidei glabri en- 
docarpium tenuissimum. 


Trees up to 20 m tall, with a large, round crown, 
and shallowly channeled buttresses to 1 m high; 
bark light gray; wood cream-colored to light brown; 
young branches very minutely puberulent, dark 
brown; older branches glabrous, gray; stipules 2- 
3 mm long, narrowly triangular. Leaf blades ellip- 
tic, 8.5-17.4 x 3.5-7.1 cm, the apex acute to 
gradually acuminate, the tips up to 
attenuate at the base, rarely oblique, drying sub- 


] cm long, 


coriaceous, green to light brown, uniformly colored, 
smooth and glabrous above and beneath, the mar- 
gins entire; lateral veins 4-6, arcuate; petioles 5— 
14 mm long, glabrous, smooth, and dark brown, 
becoming flaky and gray with age. Inflorescences 
axillary on leafy branches, flat-topped panicles or 
compound dichasia, often curving upward; rachises 
(2.7-)3.5-7.8 cm long, very minutely puberulent, 
loosely flowered, with 33-56 flowers. Flowers bi- 
sexual or staminate with pistillodes, light green; 
sepals 4, glabrous, connate in lower half; stamens 
8-10; filaments ca. mm long, linear, slender 
throughout; anthers ca. 1 mm long; ovary sparsely 
puberulent; style branches 1-2 mm long, papillate 
on inner surface, glabrous to sparsely strigulose on 
outer surface. Fruits light yellow, asymmetrical, 
transversely obovoid, 0.6-1 cm tall, 1-1.4 cm 
broad, glabrous, with persistent style branches; en- 
docarp very thin, thinner than 0.5 mm 


Distribution, habitat, and phenology. This 
species is known only from Napo Province in Am- 
azonian Ecuador (Fig. 4) in primary forest. Flow- 
ering reported in March on leafy branches; fruiting 
reported in January, May, and December 


Additional specimens examined. ECUADOR. "d 
Reserva ou a Pas, Sacha, Rio Napo, 8 km E de 
Misahuallí, 01%04'S, 77?36'W, 450 m, 19-28 Mar. i. 
Cerón 973 (MO, aa 8 Nov. 1987, Cerón 2033 st 
TEX); 18-30 May 1985, Palacios et al. 519 (MO); 2 
Jan. 1986, Palacios & Neill 965 (MO 


Ampelocera longissima is distinctive in having 
the longest inflorescences known in the genus, as 
well as glabrou 
ovoid fruits. Vegetatively, 4. longissima is some- 


s, asymmetrical, transversely ob- 
what similar to 4. edentula, which grows in the 
same area, but can be separated easily by the 
latter's shorter inflorescences and pubescent, sym- 
metrical fruits. 


7. Ampelocera macrocarpa Forero & A. Gen- 
try, Phytologia 55: 365. 1984. TYPE. Colom- 
bia. Chocó: upper Rio Truandó between La 
Teresita and mouth of Rio Ramón, 20 Jan. 
1974, Gentry 9436 (holotype, COL, n.v.; iso- 
types, MO, NY) 


Trees 10-30 m tall; trunk to 50 cm dbh; youn- 
ger branches sparsely puberulent; older branches 
glabrous; stipules ca. 6 mm long, linear-lanceolate, 
sparsely puberulent. Leaf blades oblong to elliptic, 
8-22 3.7-10 cm (sapling leaves 22-38 cm 
long), the apex acute to acuminate with tips 1-3 
cm long, very strongly oblique, base rounded to 
subcordate, the margins entire, drying coriaceous, 
dull green to brown, glabrous and smooth above 
and beneath, pinnately veined; lateral veins 4-11, 
arcuate, raised beneath; petioles 0.6-1 cm long, 
with a flaky red epidermis. Inflorescences axillary 
on leafy branches, compound dichasia with perfect 
flowers toward the apex and functionally staminate 
flowers at the base; rachises (1.8-)2.5-3.7 cm 
long, with 11-34(-57) flowers, glabrous, some- 
times purple. Flowers green, subtended by minute 
floral bracts, ca. 1 mm long, ovate, acute, char- 
taceous; sepals 4-5, 1-2.5 mm long, united at 
base, glabrous; stamens 8 in perfect flowers, 4-6 
filaments 1.5-2 


in staminate flowers; cm long, 


broadened basally; anthers ca. 1 mm long, the 


connective extended into a short apicule less than 


mm long; ovary in perfect flowers sparsely 
puberulent, with style branches 1-2 mm long; pis- 
tillode in staminate flowers cylindrical with longi- 
tudinal strigose ribs. Fruits yellow, globose or drying 
pyriform, 1.8-2.2 cm diam., essentially symmet- 
rical, glabrous or very sparsely strigose, scabrous, 
with persistent style branches ca. 2 mm long; en- 
docarp 1-2 mm this 


Common names. 
Costa Rica: rescaldo. 


Nicaragua: cuscano, yayo. 


This 


species occurs in the Atlantic lowland rainforest of 


Distribution, habitat, and phenology. 


Honduras, Nicaragua, Costa Rica, Panama, north- 
ern Venezuela, and northwestern Colombia (Fig. 
2). It has been reported in second growth evergreen 
forest and on limestone. Flowering specimens have 
been collected in February; fruiting collections are 
known from January, March, May, and August. 


HONDURAS. 
near Tela, 20-600 m, 6 
53125 (F), 54159 
of Rio Santa Cruz, 
tributary of the Rio San "Jum: 42 m, 22 Mar. 1985, 
Moreno 25531 (MO). zELAYA: Comarca de ane Isidro, 


Additional examined. 
pn Lance 
. 1927-20 Mar 


specimens 
tilla Valley, 
. 1928, ewe y 


1098 


Annals of the 
Missouri Botanical Garden 


Ampelocera longissima 


A Z ? Ase SAL be PO £e 
) = 


Volume 76, Number 4 
1989 


Todzia 1099 
Revision of Ampelocera (Ulmaceae) 


above Rio Pajarito, 3 Oct. 1982, Laguna 85 (MO); S of 
Rio Wawa, 60 km NW of Puerto Cabezas, 40 m, 1 
Mar. 1971, Little 25135 (F, MO); 16 Mar. 1971, Little 
25165 (MO); along road from Siuna to El Dos, ca. 1 km 
of Cerro Livico, 400-600 m, 12 Dec. 1980, Stevens 
18687 (MO). Costa RICA. HERE i 0 
m, 25 Mar. 1981, e 9488 (F, LL, MO, TEX); 
Finca La Selva, 30 Ma 974, Hartshorn 1495 (MO, 


968, Correa 738 ( —2 sheets) 
Parque Nacional del Darién, ridge between Rio Topalisa 
1 ucuro, ca. 17 km f Pucuro, 8?03.5'N, 
77°17'W, 850 m, 17 Oct. 1987 “Hammet et al. 16276 
(MO); S of El Real, taplon alba d Altu ique, near 


, 77°40'W, 650-800 m, 21 Aug. 
MO, TEX). PANAMA: Pilota 


se mine, 07°45 

Mc Person i 518 ( 
"à Dem 4 Dec. 1977, Folsom et al. 6776 (TEX). SAN 
BLAS: forests hai Puerto Obaldia, San Blas coast, 0- 
50 m, Aug. 1911, Pittier 4319 (F). VENEZUELA. MIRANDA: 
Cerros del Bachiller, western sector: virgin evergreen for- 


S of Cano Rico and Bachiller, 
Guapo, 10%-7'N, 65?53'W, 20- 
1978, Scr & Davidse 117010 (MO). ZULIA: 6 
km W hes main road os 2 km S of Rio Catatumbo, ca. 
20-100 m, 9°6'N 2'W, 29 Mar. 1982, Liesner & 
Gonz Pie i 3348 (LL, M COLOMBIA. CHOCÓ: trail from 
Rio Tigre base camp a Serrania del Darién W of Unguia, 
300- 600 m, 17 July 1975, Gentry & Aguirre 15286A 
(NY, MO). 


tee 


Ampelocera macrocarpa is characterized by its 
large fruits with a very thick endocarp, relatively 
short petioles with flaky red epidermis, and short 
style branches. The leaves of this species are bright 
metallic blue when young and have strongly oblique 
bases. On saplings and young branches the leaves 
are extremely long and narrow (22-38 cm long). 
Ampelocera macrocarpa appears to be most closely 
related to 4. edentula by virtue of the characters 
given in the key. 


8. Ampelocera macphersonii Todzia, sp. nov. 
TYPE. Panama. Panamá: Canal Area, on hilltop 
W of canal and Panama City, O'N 
79°34'W, 300 m, 3 May 1987, McPherson 
& Stockwell 10901 (holotype, MO; isotypes, 
NY, PMA, TEX). Figure 6. 


» 


Arbores foliis elliptici is chartaceis. pod did es, 
racemi ramosi vel simplices, 0.8-1.2 cm longi, es 3- 
6. Sepala 4, in dimidio distali villosa vel sericea, stamina 
ca. ke filamenta 1.5-2 mm longa. Fructus aurantiaci, 
dense pubescentes, ramis sty li persistentibus, transverse 
doute valde ne endocarpium tenu 


Trees up to 20 m tall; younger branches dark 
brown, puberulent; older branches gray to brown, 


glabrous, sparsely lenticulate; stipules narrowly tri- 
angular to lanceolate, 2-4 mm long, villous. Leaf 
blades elliptic, 7.5-21 x 4-7 cm, the apex acute 
or with acuminate tips up to 1.2 cm long, the base 
rounded to subcordate, sometimes oblique, the mar- 
gins entire, dark green and shiny above, paler 
beneath, drying chartaceous, smooth and glabrous 
above and beneath, light brown to tan, unevenly 
colored on drying, lateral veins 5-10, arcuate; 
petioles 0.5-1 cm long, sparsely puberulent, with 
a smooth epidermis, drying dark brown. Inflores- 
cences axillary, short, simple or compound di- 
chasia, often paired at the node, borne on leafy 
branches; rachises 0.8-1.2 cm long, strigose to 
sparsely villous, with 3-6 flowers; subtending floral 
bracts 1-2 mm long, glabrous. Flowers not seen 
(the following description based on persistent floral 
parts on fruits); calyx ca. 2 mm long, the lobes 4, 
villous to sericeous in distal half; stamens ca. 16; 
filaments 1.5-2 mm long, linear-lanceolate, slender 
throughout; anthers ca. 1-1.5 mm long, connec- 
tive not extended. Fruits yellow to orange, strongly 
asymmetrical, transversely obovoid, 1.2-1.6 cm 
long, 1.6-1.8 cm broad, densely pubescent, sweet, 
with persistent style branches 2 mm long; endocarp 
thin, less than 0.5 mm thick. 


Common name and uses. Panama: Carasu- 


ma. Used as firewood. 


Distribution, habitat, and phenology. This 
species occurs in Panama, northern Colombia, and 
northwestern Venezuela (Fig. 4). It is found in 
undisturbed deciduous forest and in moist forest, 
sometimes on limestone. Fruiting is in May, June, 
and October on leafy stems. 


Re epresentativ e specimens examined. PANAMA. DARIÉN: 
We mi. from Bayano Dam Bridge, A of Canglón, 
ail to S from id to good forest, 50 ft., lay 1980, 
Antonio 4568 (MOY Loma Bini 14 i e Duke 
8108 (MO); Rio Pucro, below village of Puc 23 June 
1967, Duke 13126 (N 10). e BOLÍVAR: t Loma de 
Juan Nepomuceno, ca. 2 km 5 


: Estacion s Primatas de Colos 
75*30"W. 930'N. 17 1981, Gentry et al. 348194 
(MO). VENEZUELA. ZULIA: MA Mara, via entre Embale 
HEN y el Campamento Carichuano, ca. 1-4 km al 
este de Caric SHAH al borde de la vía, en zona de 
100 m, Oct. 1982, Bunting & Smith 
11924 (NY): Dtto. re en e área de reserva carbon 
ifera de Guasare, unos 1.5 km te d | Cosmamentd 


Fic 
Cerón 2633 TEX . Fruit (from Palacios et al. 519 


= 


E 5. O en era longissima. —a. Fertile branch showing inflorescences. — b. Perfect flower. (a & b from 
9 MO.) 


1100 Annals of the 
Missouri Botanical Garden 


Icm 


Ampelocera macphersonii A 


S 
A k T Tse rrÍS 
a 


FIGURE 6. Ampelocera macphe rsonii. —a. Branch with infructescenc —b. Close-up of stipules. — c. Mature 
fruit with pubescence enlarged in inset. (a-c from McPherson & Stockw "ig 10901 TEX.) 
d en bosque seco pero n o O 150 m al. 122921 (MO); Dtto. Mara, undisturbed deciduous 
28 Oct. 1982, Bunting & Smith 1 (NY); Dt tto. forest remnant on low hills, 4 km (by air) N of a 
Mara, $ semievergreen ane ares ee del Norte, ee 'arichuano, 11?2'30"N, 72?216'30"W, 15 
tributary to Río Socuy, 6 km S of Corpozulia, Campa- JO m, 1 > 1980, Steyermark et al. 123074 (MO, 
mento Carichuano, 10%58'N, 72?16'W, Steyermark et d Cerro Cinco de Julio, W slopes to Rio Guasare, 


Volume 76, Number 4 
1989 


Todzia 1101 


Revision of Ampelocera (Ulmaceae) 


8.5-10 km (by air) NW Corpozulia, Campamento Cari- 
d 11?2'30"N, 72*21-22'W, 120-400 m, 4 June 
1980, Steyermark et al. 123297 (MO). 


Ampelocera macphersonii is distinctive in hav- 
ing entire leaves and small, orange, strongly asym- 
metrical, transversely obovoid fruits. By virtue of 
its 16 stamens it is allied to 4. pubescens, A. 
glabra, and A. ruizii. It differs from A. pubescens 
by having glabrous leaves and racemose (not glo- 
merulate) inflorescences, and from both 4. glabra 
and A. ruizii by having entire (not dentate) leaves 
and larger fruits. Vegetatively A. macphersonit 
resembles 4. longissima but can be separated eas- 
ily by the characters provided in the key. The 
occurrence of A. macphersonii in deciduous and 
semievergreen forests is unusual for the genus. 

This species is named in honor of Dr. Gordon 
McPherson whose collecting efforts in Panama have 
yielded many new and interesting plants. 


9. Ampelocera ruizii Klotzsch, Linnaea 20: 
541. 1847. TYPE. Peru: Ruiz & Pavón s.n. 
(holotype, B). 

dics inc verrucosa Kuhlm., Arch. Jard. jus Rio de 
Janeiro 4: 352. 1925. TYPE. Brazil. Pará: Itaituba, 
Rio Tape 20 Oct. 1922, Ducke 19135 nS 


B; , K, US). 
Plagioceltis Hic Kun Mildbr. ex Baehni, Contrib. Mn 
Mu Se 37 


s. Nat. Hist., Bot. 72. 
Peru: Ruiz & Pavón s.n. (holotype, MA, n.v.; i in 
type, F). 


Trees 10-35 m tall; trunk up to 2.5 m dbh, 
with brown, flaky bark, and medium-sized buttress- 
; younger stems brown, glabrous to sparsely pu- 
berulent; older stems gray, glabrous; stipules 3-5 
m long, lanceolate, sparsely puberulent to pilose. 
Leaf blades elliptic to obovate, 6.5-13 x 2.5-6.5 
cm, the acuminate tips 0.5-1 cm long and 2-3 
mm broad, attenuate to rounded at the base, often 
oblique, drying stiffly chartaceous to subcoriaceous, 
glabrous and smooth above and beneath, with 3- 
5 teeth per side, occasionally entire; lateral veins 
5-8, arcuate, raised beneath, puberulent beneath 
when young, becoming glabrous with age; petioles 
0.5-1 cm long, sparsely puberulent, drying dark 
brown. Inflorescences of axillary, compound di- 
chasia, on leafless branches, apparently with bi- 
sexual and staminate flowers (staminate flowers not 
seen); rachises 0.9-1.5 cm long, sparsely puber- 
ulent, with 8-15(-23) flowers. Flowers subtended 
by ovate bracteoles, these 1.5-2 mm long, glabrous 
to sparsely puberulent; calyx 2-2.5 mm long, the 
lobes 4, united at the base, sparsely villous; stamens 
16 in perfect flowers; filaments ca. 3 mm long, 
linear, slender throughout; anthers ca. 1 mm long, 


the connective extended into an apicule ca. 0.5 
mm long; ovary very densely pilose, with style 
branches 3-5 mm long, united in lower 1 mm. 
Fruits yellow, asymmetrical, transversely ellipsoid, 
1-1.2 cm tall, 1.2-1.6 cm broad, slightly flattened, 
densely pilose, with persistent style branches; en- 
docarp thin, ca. 0.3 mm thick 

Distribution, habitat, and phenology. This 
species is known from 0-500 m in lowland rain- 
forest in Brazil, Peru, and Bolivia (Fig. 3). Flow- 
ering time is not known; fruiting is reported from 
August to January. 

Additional specimens examined. BRAZIL. ACRE: near 
mouth of Rio Macauhan (tributary of Rio Yaco), 7 Aug. 
1933, Krukoff 5316 (MO). MATO GROSSO: Serra do Ca- 
chimbo, 12 Nov. 1976, Nascimento : 445 à 


Rio hueso: 28 June 1978, Pires 16046 (MG); 
Rio Xingu, Gleba Bacaja, lote 88, just ae d of 
Rio Bacaja, 27 Nov. 1980, Prance et al. 2 (MO). 
PERU. LORETO: Veradero de Mazán from Ee E. Ama- 
zonas to Rio Napo, 22 Aug. 1972, Croat 19516 (MO). 
MADRE DE DIOS: Parque Nacional del Manu, Río Manu, 
vicinity of Cocha Cashu aranan 18 2 1976, Foster 
& Terborgh 5209 (F, US 


T 
ace 
> 

Ed 

= 

o 
$ 
o 


, 22 Oct. 1979, Gentry et al. 
27135 (MO). SAN MARTÍN: Prov. Mariscal Cáceres, Dtto. 
Uchiza, Caserío Nueva Unión Huicte, se 500 m, 1 
Aug. 1974, Schunke 7964 (MO). BOLIVIA. SANTA ( 
Prov. Sara, Buena Vista, 450 m, J. SEN 1309(G H 
NY, MO). 

Ampelocera ruizii is distinguished from 4. gla- 
bra by having smaller fruits and longer inflores- 
cences. They are similar in having dentate, gla- 
brous leaves and transversely ellipsoid, asymmetrical 


ruits. 
Baehni (1937) separated Ampelocera from Pla- 
gioceltis and the other Peruvian genera of the 
maceae on the basis of the former having twice 
many stamens as the latter. The isotype of 
Plagioceltis dichotoma at F, however, has a great- 
er number of stamens than perianth parts. Since 
stamen number is not mentioned in the Plagioceltis 
description, Baehni’s misplacement of the genus in 
the Flora of Peru key alongside genera possessing 
equal numbers of stamens and perianth segments 
is probably due to Mildbraed's inadequate notes 
(Gentry, 1983). As mentioned under taxonomic 
history, the similarity of the unnumbered Ruiz & 
Pavón type specimen of Plagioceltis dichotoma 
and the type of Ampelocera ruizii is the most likely 
because both originated from the same collection. 


EXCLUDED TAXA 


Ampelocera crenulata Urban, Repert. Spec. Nov. 


Regni Veg. 15: 399. 1919. TYPE. Cuba: Ra- 


1102 


Annals of the 
Missouri Botanical Garden 


mon de la Sagra 413 (B, n.v.). = Casearia 
praecox Griseb. (Flacourtiaceae). 

Ampelocera hondurensis J. D. Smith, Bot. Gaz. 
(Crawfordsville) 54: 244. 1912. TYPE. Hon- 
duras: Dept. Santa Bárbara, San Pedro Sula, 
200 m, May 1890, Thieme 5606 (US—3 


sheets). — Achatocarpus nigricans Triana 
(Achatocarpaceae). 

LITERATURE CITED 

BaEHNI, C. 1937. Ulmaceae. /n: F. Macbride Sa 


Flora of n Contr. Field Mus. Nat. Hist., Bot. S 
13: 272 
DuckE, A. Tv EN forest trees ps climbers of the 
Brazilian Amazon. Trop. Woods 15-32. 
FORERO, E. & A. H. Gentry. 1984. "Ne ew phanerogam 
species from C hoco, Colombia. Phytologia 55: 365- 
371. 


Murs e H. 1983. ori de su- 
ous genus. Taxon 
Bos m. E A. 1988. Charles Wright i in a uba, 1856- 
7. Chadwyck-Healy, Cambridge, England. 
e H, J. F. 1847. Beiträge zu einer Flora der 
A Gegenden der neuen Welt. Linnaea 20: 
337-542 


KUHLMANN, J. 0 Contribuigáo para o conheci- 
mento de algumas pa novas, contendo tambem 
um trabalho de critica e novas combinações. Arch. 
Jard. Bot. Rio de iG 4: 347 7-365. 

1940. Especies novas equatoriais e tropicais- 
orientais brasileiras. Ann. Prim. Reu. Sul. Amer. Bot. 
3(1938): 75-92, pls. 1 

ons is V. 1958. A new v Ampe ^locera from Cuba. 

. Biol. Soc. Wash. 153-154. 

PLANCHON, Er E. 1873. ene In: 

Pro 20517210. 


De Candolle, 


POLLEN MORPHOLOGY OF 
THE SHRUBBY IRIDACEAE 
NIVENIA, KLATTIA, AND 
WITSENIA! 


Peter Goldblatt? and 
John C. Manning? 


ABSTRACT 


Nivenia, Klattia 
have pollen grains ui sculpturing similar to the 
grains have a broa mo 


are unique in Iridace Nivenia species the 


verrucate or micropapillate, gi aos of destinies varyin 
of 


the species examined, and in 
differences between the long- i short-styled morphs. 


, and Witsenta, a monopayietic on 
type for Iridaceae (and for the monocots). The 
and reticulate to briser MAR exine. The supratectal gemmae, found only in Wit 


of 13 species of the southwestern Cape, South Africa, 
nearly spherical 
senia, 


walls of the reticulum are lightly sculptured, sometimes laterally 
g with the species. The pollen grains differ in size for all 
the five heterostylous species of Nivenia there are statistically significant size 


The genera Nivenia (nine species), Klattia (three 
species), and Witsenia (one species) comprise a 
natural alliance of Iridaceae, restricted in distri- 
bution to the southwestern Cape region of southern 
Africa. Uniting the genera are the following spe- 
cialized characters: a shrubby evergreen habit; a 
woody rootstock and aerial stems both having ap- 
preciable secondary growth; nonfugacious flowers 
with a well-developed perianth tube and bearing 
septal nectaries; tangentially compressed seeds; and 
a specialized leaf anatomy with submarginal scle- 
renchyma and girders produced by the extension 
to the epidermis of the sclerenchyma surrounding 
some of the vascular bundles (Goldblatt, in prep.; 
Goldblatt & Bernhardt, in press; Manning & Gold- 
blatt, in prep.; Rudall & Burns, 1989). 

This study of the pollen morphology of the three 
genera was undertaken in conjunction with a broad- 
ly based investigation of their systematics and re- 
productive biology, the latter focusing particularly 
on the heterostylous condition in five species of 
Nivenia. 'These distylous species are made up of 
populations of plants with either long styles and 
short stamens or short styles and long stamens. In 


many plants heterostyly is often associated with 
pollen dimorphism (Ganders, 1979; Richards, 1986) 
and this phenomenon has been considered in our 
examination of exine structure. The homostylous 
Klattia and Witsenia were included in our study 
for the sake of completeness. Klattia and Witsenia 
are probably bird pollinated (Rebelo, 1988; Gold- 
blatt, in prep.) in contrast to species of Nivenia 
which are pollinated by long-tongued insects, either 
anthophorid bees or nemastrinid flies (Goldblatt & 
Bernhardt, in press). Any differences in the pollen 
of the three genera may be significant in light of 
the differences in their pollination biology. 


MATERIALS AND METHODS 


Pollen taken from herbarium specimens of 11 
species (Table 1) was treated in one of two ways. 
For examination of exine sculpturing alone, un- 
treated grains were glued on aluminum stubs an 
sputter coated with gold. For examination of whole 
grains, unacetolyzed pollen was rehydrated by 
warming in 10% KOH for ca. 10 minutes, then 
dehydrated in ethanol, rinsed twice in 50% acetone 


! Support id E study by grant BSR 85-00148 from the U.S. National Science Foundation is gratefully acknowl- 


k Sylvia Feuer, De Paul U niversity, for h 


electron microscopy. We also acknowledge the cooperation o 


extensive help w with the electron microscopy and for her 
y EM Unit, for his assistance with scanning 
frican Department of Environment Affairs 


and the Cape Department of Nature and ir Conservation in providing collecting permits and access to 


field sites where the species used in this study were co 
2B.A 


Krukoff Curator of African Botany, Missouri Botanic al Garden, P.O. Box 299, St. Louis, Missouri 63166, 


U.S.A 


* Compton Herbarium, National Botanic Gardens, Kirstenbosch, P. Bag X7 


, Claremont 7735, South Africa. 


ANN. Missouni Bor. Garb. 76: 1103-1108. 1989. 


1104 Annals of the 
Missouri Botanical Garden 


Species of Vivenia, Klattia, and W itsenia studied with locality and voucher data. All localities are in 
PRE. 


O, NBG, and 


the Duns Province, South Africa. Herbarium vouchers are preserved at M 


Klattia 
K. flava (G. Lewis) Goldbl. 
K. partita Ker ex Baker 
K. stokoei L. Guthrie 


slopes of Landdrost Kop, Goldblatt 8347 
Langeberg, Swellendam, Goldblatt 8698 
Steenbras Ridge, Goldblatt 8342 


Nivenia 
Distylous Species 
N. binata Klatt 
N. capitata (Klatt) Weim. Little Karoo, Rooiberg slop 
M corymbosa (Ker) Baker Bains Kloc weede Tol camp, Goldblatt 7631 
Olifantskloof, Riviersonderend, Goldblatt 8480 
Klein Swartberg, Buffelsrivierpoort, Goldblatt & Manning 8864 


Swartberg Pass, upper id Goldblatt 7959 
, Vlok 1794 


V. dispar Br. 
N. stenosiphon Goldbl. 


Homostylous Species 


N. concinna N. E. Br. Viljoens Pass, Goldblatt 7624 

N. fruticosa (L. f.) Baker Swellendam Langeberg, Esterhuysen 36149 

N. levynsiae Weim. Highlands hiking cu Caledon Div., Goldblatt 8463 
N. stokoei (L. Guthrie) N. E. Br. Palmiet R. Mountains, Levyns 5344 


Witsenia 


W. maura L. Bettys Bay, Goldblatt & Gentry 1600 


tive terminology used follows Walker & Doyle 
(1975). Pollen grains mounted in FLP orcein were 
measured under the light microscope. Measure- 
ments (Table 2) represent the means of 10 grains 
except in the case of the heterostylous species 


and four times in 100% acetone, and critical point 
dried before mounting and coating. Specimens were 
observed at an accelerating voltage of 15-20 kV. 
Klattia partita and K. flava are listed in Table 1 
but were not included in the SEM study. Descrip- 


TABLE 2. Pollen characteristics of Nivenia, Klattia, and Witsenia. Only the equatorial diameter is listed. 
MEM species of Nivenia the diameter of the long-styled morph precedes that of the short-styled morph. The 
asterisk (*) indicates species not examined under the scanning electron microscope (SEM). 


Species Shape Diameter um Sculpturing Muri 

Nivenia 

N. binata spherical 51.26/55.55 reticulate granulate and 

ridged 

N. capitata spherical 59.88 E 2.79 reticulate granulate 

N. concinna spherical 11. microreticulate granulate 

N. corymbosa spherical 44.69. as 69 reticulate granulate 

V. dispar spherical 53.50/52.02 reticulate * 

V. fruticosa spherical- 63.93 microreticulate granulate 

subprolate 

N. levynsiae spherica 65.00 reticulate granulate 

N. stenosiphon spherical 65.00/68.69 reticulate granulate and 

N. stokoei spherical 61.43 reticulate granulate 
Klattia 

K. flava subprolate 63.89 reticulate * 

K. partita subprolate 64.12 reticulate * 

K. stokoei spherical 80.16 reticulate smooth 
IWitsenia 

W. maura oblate- 81.55 microreticulate with granulate 

spheroidal pratectal gemmae 


Volume 76, Number 4 
1989 


Goldblatt & Manning 1105 
Pollen morphology of Nivenia, 


Klattia, and Witsenia 


TaBLE 3. 


Pollen size comparison between long- and short-styled morphs of the five distylous species of Nivenia: 


N = total number of pollen grains measured from three (or four) individuals; X = mean pollen grain diameter in um; 


S.D. = standard deviation; P = probability (from Goldblatt & Bernhardt, in press). 
Pollen morph measurements (equatorial diameters) 
Long-styled Short-styled 
Species N ki S.D. N X S.D. P 
N. binata 39 51.26 3.30 30 55.55 5.12 0.001? 
N. capitata 30 59.88 oui 30 62.79 4.44 0.008? 
N. corymbosa 40 44.69 3.80 40 49.69 2.1 0.000? 
N. dispar 30 53.50 3.36 30 52.02 4.07 0.128? 
N. stenosiphon 30 65.00 3.75 30 68.69 4.07 0.014? 


! Two-tailed t-test. 
? Significant. 
* Not significant. 


(listed in Table 3), where nine or ten grains in three 
or four individuals per style morph were measured 


(Table 3 


OBSERVATIONS 


Grains of all species of all three genera are more 
or less spherical (Figs. 1-6), ranging from near 
prolate to near oblate depending on the species 
(Table 2). The monosulcate grains have a promi- 
nent elliptic to oblong sulcus (Figs. 2, 5, 6); 
are tectate-perforate in Witsenia with a micro- 
reticulate tectum (Figs. 4-6, 9, 10) and semitectate 
with a reticulate tectum in Nivenia and Klattia 
(Figs. 1-3, 7, 8). The floors of the lumina in all 
the species are granulate (e.g., Fig. 7). The muri 
are smooth in Klattia (Fig. 8) but minutely gran- 
ulate in /Vivenia and Witsenia (e.g., Figs. 7, 9, 
10). In N. binata and N. stenisiphon the sides of 
the muri are irregularly ridged (e.g., Fig. 7) (Table 
2). 


In Nivenia the luminae range from about as 
wide as the muri (microreticulate), e.g., /V. 
cinna (Fig. 12), to about twice as wide (reticulate), 
or up to four times as wide in N. capitata (Fig. 
11) 

Supratectal gemmae (ca. 4 um in diameter) are 
restricted to Witsenia (Figs. 5, 6) and are scattered 
more or less evenly across the surface. 

The exine sculpturing ends abruptly at the sulcus 
(Fig. 2) in Nivenia and Klattia, but in Witsenia 
the muri become disrupted into pilae and baculae 
(Fig. 10) along the sulcus margins. 

Pollen grain diameter ranges from 44.69 um in 
the long-styled morph of /Vivenia corymbosa to 
over 80 um in Klattia stokoei and Witsenia maura 
(Table 2). Differences in pollen grain size among 
the species correlate generally with differences in 


style length. Klattia and Witsenia have the longest 
styles and largest pollen in the group, and K. sto- 
koei the longest styles in that genus. Nivenia le- 
vynsiae and N. concinna, however, do not follow 
this trend, having shorter styles than one would 
expect from their comparatively large pollen grains. 

In four of the five heterostylous species of Niv- 
enia the grains of the long-styled morph are sig- 
nificantly smaller (Table 3). Pollen size differences 
between morphs of the four heterostylous species 
late directly with differences in style length 


also 
in the complementary morph (Goldblatt & Bern- 
hardt, in press). Exine sculpturing does not differ 
between morphs within the heterostylous species, 
ut there are notable di the 
species. Nivenia binata exhibits the greatest de- 
gree of sculpturing and V. capitata the least (Figs. 
[maa 


differences between 


» 


DISCUSSION 


The pollen grains of Nivenia and Klattia con- 
form fairly closely to the presumed basic type for 
Iridaceae and for the monocots in general, in which 
reticulate grains appear to be fundamental (Schulze, 
1971; J. Walker, pers. comm.). Similar grains are 
found in species of Aristea (likewise Nivenioideae) 
and in numerous genera of Iridoideae, excluding 
Iris, in which there are multiple unusual special- 
izations, particularly in the form of the aperture 
(Schulze, 1971). A group of New World genera of 
Iridoideae belonging to the tribe Tigridieae stand 
out in having bisulcate grains (Schulze, 1971), 
clearly a secondary condition in the tribe. Members 
of subfamily Ixioideae also differ markedly from 
this basic pollen type, having a tectate-microper- 
ith supratectal spinules (Erdtman, 


Goldblatt & Stein, 1988). 


forate exine w 


1952; Schulze, 1971; 


1106 Annals of the 
Missouri Botanical Garden 


FIGURES 1 76. — 1. Nivenia corymbosa, nonapertural view. — 2. N. corymbosa, oblique apertural view. — 3. Klattia 
Wi 9. W. maura, lateral equatorial view of the 
sulcus. — 6. Oblique apertural view showing differential sculpturing around the aperture. Sca 


stokoel, nonapertural view. L. senta maura, nonapertural view. 


e bar 10 um. 


Volume 76, Number 4 
1989 


Goldblatt & Manning 
Pollen morphology of Nivenia, 
Klattia, and Witsenia 


1107 


FIGURES 7-12.— 7. 
with smooth muri.— 9. Wi 


gemmae. — 1 ( 
exine with Pass granules on the muri. — 12. N. conc 
granulae. Scale bar 1 um for 7-10, 5 um for 11, 12 


Apart from the different exine type in Ixioideae, 
there are no other major specializations in exine 
sculpturing i in Iridaceae. 

minor development evident in Nivenia and 
Witsenia but absent in Klattia is the presence of 
granules on the muri, and the irregular ridges on 
the lateral walls of the muri in N. binata and N. 
stenosiphon appear to be a further development 
in the direction of muri ornamentation. In /Vivenia 


— n binata, reticulate exine with the muri showing lateral ridges. 
senia maura, nonaperturate view showing micror 
W’. maura, exine adjacent to the aperture showing differential sc ulpturi ing. 
‘inna, microreticulate exine wit 


-8. Klattia stokoet, 


supratectal 


eticulate sc ge with 
capitata, reticulate 
1 the muri showing scattered 


the direction of pollen grain evolution appears to 
be from an open reticulate exine with sparse or- 
namentation on the muri, as in the putatively least 
specialized NV. capitata (Goldblatt & Bernhardt, in 
press), to increasing ornamentation and decreasing 
size of the lumina of the reticulum, leading to a 
microreticulate pattern in N. fruticosa and N. con- 
cinna. These two species appear to be indepen- 
dently specialized based on differences in flower 


1108 


Annals of the 
Missouri Botanical Garden 


and inflorescence morphology (Goldblatt, in prep.), 
and their similar pollen sculpturing is presumably 
convergent. The occurrence of ridged muri in N. 
binata and N. stenosiphon, however, appears to 
be an indication of close taxonomic relationship as 
it is correlated with two specialized characters re- 
stricted to these species: blue anthers and pollen, 
and large, dry, brown, crisped bracts (Goldblatt, in 
prep.). 

The grains of Witsenia, with their large supra- 
tectal gemmae, are clearly derived. These distinc- 
tive gemmae, observed by Schulze (1971) under 
light microscopy, are unique in Iridaceae and make 
the pollen grains of this genus immediately rec- 
ognizable. The presence of an unusual exine in 
Witsenia emphasizes the taxonomic isolation of this 
genus, distinctive in its unusual floral and inflores- 
cence morphology. Any significance in the elabo- 
rate exine sculpturing of Witsenia in relation to 
its mode of pollination remains to be demonstrated. 

Pollen sculpturing does not provide any char- 
acter uniting the three shrubby genera of Cape 
Iridaceae, but within the alliance trends of spe- 
cialization are evident in Nivenia, and the supra- 
tectal gemmae in Witsenia constitute an autapo- 
morphy for this genus. 

Mulcahy's (1965) report of a difference in size 
between the grains of the short- and long-styled 
morphs of N. binata is substantiated by our ob- 
servations, although we noted a smaller size dif- 
ference (ratio of grain size between short- vs. long- 
styled morph 1.11 compared with 1.3 reported by 
Mulcahy). We were unable to confirm Mulcahy's 
observation of differences in the size of the lumina 
of the exine reticulum between short- and long- 
styled grains in N. binata, nor did we record mea- 
surable differences in lumina size in the other het- 
erostylous specie 

e close similarity of the exine sculpturing and 
small, though statistically significant, differences in 


pollen grain size between the two style morphs in 
four of the observed heterostylous species of Niv- 
enia are consistent with similar observations in 
many heterostylous taxa (Ganders, 19 Rich- 
ards, 1986). In Nivenia the noolar ea differ- 
ences between the style morphs of the heterostylous 
species appear not to be accompanied by the ex- 
pected self-incompatibility characteristic of most 
heterostylous plants (Goldblatt & Bernhardt, in 
press). 


LITERATURE CITED 


1952. Pollen Morphology and Plant Tax- 
Angiosperms. Almquist & Wiksell, Stock- 


ERDTMAN, G. 
ou 
olm. 
von E R. 1979. The biology of heterostyly. New 
ña Bot. 17: 607-635. 
Systematics E biology of the shrubby 
EROA Nivenia, Klatt Vitsenia (in prep). 
& P. BERNHARDT. 1989. Pollination biology 
of the Cape genus Nivenia Meri dis nd the 
towa rae self comp Mari Israel J. t. (in pre 
. STEIN. 19 E ub of 
Pillansia L. ipa (Iridaceae). Ann. Missouri Bot. 
Minis 
MANNING, J. ee DBLATT. Seed coat structure 
in the shrubby Cape Iridaceae, Nivenia, Klattia and 


m DBLATT, 


~ 


MuLcany, D. L. 1965. Heterostyly within Vivenia (lri- 
aceae). md 17: 349-351. 

REBELO, A. C. 1988. Bird pollination in the Cape flora 
Pp. 83 Em in A. G. Rebelo (edito de A Preliminary 
Synthesis of Pollination Biology ir y Flora. 
5. d Natn. Sei. DRIN Nas Rep. 1 . CSIR, 


> 
RICHARDS, P5 J. 1986. Plant Breeding Systems. George 
Allen & Unwin, London. 
RuparL, P. & P. Burns. 1989. 
woody South African Iridaceae. Kew Bull 
SCHULZE, W. 1971. 
Iridaceae und ihre Bedeutung für die 
Feddes i 82: 101-124. 
WALKER, J. W. € J. A. DOYLE. 1975. Bases of angio- 
nnd pr PSA Ann. Missouri Bot. 
ard. 


Leaf pd of the 
. (in press). 
Beitráge zur Pollenmorphologie der 
Taxonomie. 


SEED MORPHOLOGY 
OF SISYRINCHIUM 
(IRIDACEAE— 
SISYRINCHIEAE) 
AND ITS ALLIES! 


Peter Goldblatt,? James E. Henrich,’ 
and Richard C. Keating? 


ABSTRACT 


Seeds of 25 species of sects. Sisyrinchium, Echthronema, Eriphi ilema, and iai of the large North and South 


— genus Sisyrinchium were examined using the scanning e electron mic 
eeds of representative species of related genera in tribe Sisyrinc ditas. Seeds of sects. Sisyrinchium 
, are s with Vg nd surfaces, and often have a iuis depression, 


compared with s 
and Echh ronema are more or less Pomen 


a large, half-circumferential wing. Seeds o 
probably derived specializations. Based on seed charac 


and light microscope and 


o have brown, more or lessa ular seeds ms 


8, 
to the other genera of Sisyrinchieae, particularly Phones than to sects. Sisyrinchium or Echthronema. 


In the course of a systematic study of the New 
World genus Sisyrinchium for Flora Mesoamer- 
icana (Henrich & Goldblatt, 1987, in press a) and 
Flora de Nicaragua (Henrich & Goldblatt, in press 
b) our attention was drawn to the unusual rounded, 
blackish, umbilicoid seeds of the genus. The seeds 
were novel to us in Iridaceae and distinguish Sisy- 
rinchium from all other genera in the family. 
Seed characters were also useful in establishing 
species limits within the genus. We found no ad- 
equate description or record of Sisyrinchium seeds 
in the literature, and Huber (1969) did not spe- 
cifically describe them in his extensive study of the 
seeds of lilioid monocots. However, Cholewa & 
Henderson (1984) included SEM examination of 
six North American species of sect. Sisyrinchium, 
and Rodriguez (1986) mentioned color and surface 
texture in his revision of the Chilean species of 
Sisyrinchium. 

With this background we undertook the present 
survey of the seeds of Sisyrinchium and its allies 
in tribe Sisyrinchieae of subfamily Iridoideae (clas- 


sification after Goldblatt, 1990). Sisyrinchium as 
presently constituted (Bentham & Hooker, 1883; 
Baker, 2) 

study includes a small but representative sample 
of the total and is sufficient to satisfy our purpose 
in documenting the basic seed types found in the 
genus and to allow a valid comparison with allied 
genera. We included taxa from the two large and 
apparently natural sects. Sisyrinchium (— sect. 
Bermudiana of several authors) and Echthronema 
(Bentham & Hooker, 1883) and the smaller sects. 
Eriphilema and Nuno. Related genera of tribe 
Sisyrinchieae (sensu Goldblatt, 1990) examined for 
comparison were Libertia, Ona, Orthrosanthus, 
Phaiophleps, Solenomelus, and Tapeinia. Seeds 
of Chamelum, which belongs in this alliance, were 
not available. The southern African Bobartia and 
the Australian Diplarrhena, the remaining genera 
of the tribe, appear to be more distantly related to 
the Sisyrinchium complex and neither was in- 
cluded in our survey. Sections Eriphilema and 
Nuno and the other genera examined have seeds 


comprises over 100 species. Our 


! Support for this study by grants DEB 81-19292 and BSR 85-00148 to the first author from the U.S 
thank Jim Solomon for seed samples. 


Science Abb is gratefully acknowledged. We 


. National 


Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, 5t. Louis, Missouri 63166, 


U.S. o 
3 Missouri Botanical Garden, P.O. Box 299, St. 


, Missouri 63166, U.S 


Lou S.A. 
' Department of Biological Sciences, Southern illinois ee Edwardsville, Illinois 62025, U.S.A. 


ANN. Missouni Bor. GARD. 76: 1109-1117. 1989. 


1110 


Annals o 
Missouri Botanical Garden 


fairly typical of Iridaceae in their brownish color 
and more or less angular shapes (Huber, 1969) 
and are very different from the seeds found in 
sects. Sisyrinchium and Echthronema. 


MATERIALS AND METHODS 


Mature seeds were assembled from herbarium 
material, and in a few cases fresh field-collected 
seeds were examined (Table 1). We excluded seeds 
that appeared deformed or showed signs of fungal 
infection. Seeds were examined under the light 
microscope for size and color, and then prepared 
for the scanning electron microscope (SEM) study 
by attachment with water-soluble white glue to 
aluminum stubs and sputter coating with 500—700 
À of gold. They were examined in an ISI Super IIIA 
SEM at 15 kV. 


OBSERVATIONS 
COLOR 


Seeds of all the species of Sisyrinchium sects. 
Sisyrinchium and Echthronema are blackish (Ta- 
ble 2). Surface color intensity ranges from a fairly 
glossy, intense black to matte and dark gray. As 
we used seeds almost exclusively from herbarium 
specimens, we are uncertain how much significance 
to attribute to the color variation as ripening may 
have been affected by the rapid drying, and the 
final seed appearance may thus not be the same 
as in nature. 

Seeds of Sisyrinchium sects. Eriphilema and 
Nuno and of the genera Libertia, Ona, Orthro- 
santhus, Phaiophleps, Solenomelus, and Tapei- 
nia are shades of brown (Table 2), most often red- 
brown, presumably the basic condition in the family 
(Huber, 1969) and an indication of the presence 
of the pigment phlobaphene. Some species, for 
example Orthrosanthus occissapungus, have light 
brown seeds, an apparently significant character 
at the species level. 


SHAPE AND SIZE 


The number of seeds per locule of the three- 
chambered capsules of Sisyrinchieae was not re- 
corded in our study, but all species examined have 
multiseeded loculi. The effect of crowding and al- 
teration of shape by mutual pressure within the 
locules (Huber, 1969; Wagner & Goldblatt, 1984) 
was noted to some degree in most species. Angular 
seeds characterize Libertia and Orthrosanthus 
(Figs. 1-3). Phaiophleps and Sisyrinchium sect. 


Eriphilema have less sharply angular seeds (not 


illustrated), sometimes tending toward rather ob- 
tusely angled and differing consistently in basic 
shape from the rounded seeds of sects. Sisyrin- 
chium and Echthronema. Seeds of two species of 
Solenomelus are cylindric to barrel-shaped with 
distorted angular ends (e.g., S. segethii, Fig. 5). 
In a third, possibly new, species of Solenomelus 
(Fig. 6) the seeds have a broad, membranous, half- 
circumferential wing. The seeds of Orthrosanthus 
occissapungus (Fig. 3) have the proximal and dis- 
tal ends produced into winglike processes, but other 
species of the genus have angular seeds, as illus- 
trated by Takhtajan (1985) for O. chimboracensis. 
In Tapeinia (Fig. 4) the seeds are nearly pyriform 
to elongate-pyriform, narrowest at the chalazal end. 
The raphe is usually distinct and terminates in a 
small depression in which we assume the micropyle 
is located. The least effect of crowding was found 
in sects. Sisyrinchium and Echthronema. There, 


except for the small persistent funicle, the seeds 
(Figs. 7-18) are nearly regularly spherical to 
rounded or sometimes somewhat depressed (e.g., 
S. chiricanum, Fig. 

Seed size varies considerably (Table 2) within 
genera. Thus in Sisyrinchium, S. minus has seeds 

6 x 0.4 mm, whereas those of S. chiricanum 
are ca. 2.5 mm in diameter. The seeds of Soleno- 
melus are even larger, with a maximum length of 
5.5 mm attained by Solenomelus sp. 

There is a regular pattern of small, evenly spaced, 
irregularly rounded depressions oriented in lines 
running along the long axis of the seeds of all three 
species of Solenomelus (Figs. 5, 6). In Solenomelus 
sp. these depressions are limited to the area cov- 
ering the seed body. 

A conspicuous, often large pit, which we here 
call an umbilicus, is an outstanding feature of the 
seeds of most species of sects. Sisyrinchium (Figs. 
9, 10) and Echthronema (Figs. 13-18) (Table 2). 
The umbilicus straddles the raphe, either between 
the hilum and the micropyle (Figs. 17, 18), or the 
umbilicus includes the micropyle (Figs. 13, 15). 


SURFACE MORPHOLOGY AND MICROSCULPTURING 


Outlines of the testa cells often form a pro- 
nounced reticulum owing to the typical condition 
in Iridaceae for the outer periclinal walls to collapse 
while the thickened anticlinal walls persist (e.g., 
Figs. 1-4). The seed surfaces thus have a reticulate 
appearance 
the surfaces exhibit little texture and in S. trinerve 
Fig. 15) are nearly smooth. We did not find ex- 
amples of the somewhat irregularly granular sur- 


. In some species of sect. Echthronema 


— 


faces and diffuse reticulation without cell outlines 


Volume 76, Number 4 


Goldblatt et al. 1111 


1989 Seed morphology of Sisyrinchium 
TABLE l. Voucher data for the species studied (acronyms in parentheses after /ndex Herbarium (Holmgren et 
al., 1981)). 


Species 


Voucher information 


Sisyrinchium sect. Sisyrinchium 
arenarium Poeppig 
campestre Bickn. 
chilense Hook. 
dimorphum Bickn. 
johnstonii Standley 


minus Engelm. & Gray 
ontanum Greene 

nashii Bickn 

pruinosum Bickn. 
Sisyrinchium sect. Echthronema 

chiricanum Woodson 

commutatum Klatt 

convolutum Nocca 

mandonii Baker 

prealtum Kraenzl 

tenuifolium Kunth 

tinctorium Kunth 

trinerve Baker 
Sisyrinchium sect. Eriphilema 

douglasii A. Dietr. 

scirpiforme Poeppig 
Sisyrinchium sect. Nuno 
filifolium Gaudich. 


Libertia 


sp. 
ixioides Spreng. 
Ona 
obscura (Cav.) Ravenna 
Orthrosanthus 
chimboracensis (Kunth) Baker 
occissapungus (Ruiz ex Klatt) Diels 
Phaiophleps 
biflora (Thunb.) R. Foster 
Solenomelus 
pedunculatus (Gillies) Johnst. 
segethii (Philippi) Kuntze 
? sp. nov. 


Tapeinia 


pumila (Forster f.) Baillon 


Venezuela: Dorr et al. 
Bolivia. Murillo, La Paz: Solomon 18360 (MO) 


Chile: Pfister 12279 


Chile: Zollner 11298 (MO 
U.S. 
Chile: Mexia 7994 

Guatemala. Huehuetenango: Breedlove 8730 (F 
Costa Rica. Cartago: Lent 2221 (F) 

U.S.A. Florida: Moldenke 446 (MO 

Costa Rica. San Jos 
U.S.A. Texas: Oliver 417 (MO) 

U.S.A. New Hampshire: Churchill 302 (MO) 
U.S.A. Georgia: Ravenal s.n. 
U.S.A. Texas: Reverchon 2791 (MO) 


) 
A. Nebraska: Tolstead 9715 (MO) 
(MO) 


) 


é: Khan et al. 137 (MO) 


= 
e 


Guatemala: Steyermark 46760 (F) 

Brazil. Paraná: Hatschbach 25417 (MO) 
Honduras: Standley 15493 (MO) 

Costa Rica: Allen 698 (MO) 

Peru. Cuzco: West 7029 (MO) 

Guatemala: Steyermark 50670 (F) 

Belize: Gentle 2225 (MO) 

Costa Rica: Davidse & Herrera 29375 (MO) 


S.A. Oregon: Henderson 478 (MO) 


U. 
Chile. Santiago: Bocher et al. 614 (MO) 
Falkland Islands. Port William: Hooker 98 (K) 


Bolivia. Nor Yungas, La Paz: Solomon 17343 (MO) 
New Zealand. South Island: Anderson 243 (MO) 


Chile. Port Famine: King 113 (K) 


5044 (MO) 


(CONC) 


Chile. Talcar, Camarica: Moreira s.n. (GH) 
Chile. Santiago: Wilkes s.n. (GH) 
Chile/Argentina. Calenfen "Valley: Elwes s.n. (K) 


Ricardi et al. 1156 (CONC) 


reported in some North American species of sect. 
Sisyrinchium (S. idahoense, S. radicatum, 


montanum) by Cholewa & Henderson (1984) ex- 
cept in a possibly fungal-infected sample of S. 


prealatum. 


Although details of the surface of individual cells 
were not our BENE concern, we noted a char- 
acteristic “bumpy” or granular texture (Fig. 19) 
within the outlines of T anticlinal cell walls similar 


to that illustrated by Cholewa & Henderson (1984) 


1112 Annals of the 
Missouri Botanical Garden 


TABLE 2. Seed characteristics in Iridaceae tribe Sisyrinchieae. Presence of a character is indicated as X, absence 


by —. Measurements of seeds are diameter, or if two sizes given, long axis first followed by short axis. 
Size 
Species Shape (mm) Color Umbilicus 


Sisyrinchium sect. 


Sisyrinchium 


arenarium + spherical ca. 1.5 black X 
campestre spherica 0.9 black X 
chilense rounded 1-4 x 1-1.1 black X 
dimorphum + spherical ca. 1.1 black X 
johnstonii + spherical ca. 1.1 black X 
luzula rounded 0.8-1 x 0.8 black X 
miamiense spherical 1-1.1 black — 
micranthum broadly trianguloid 0.8-0.9 black X 
minus rounded 0.6 x 0.4 black X 
montanum + spherical l-1. black X 
nashii spherical 0.8 black X 
pruinosum spherical 0.9 lack X 
rambonis spherical-oblong 0.8 x 0.8-1 black = 
Sisyrinchium sect. Echthronema 
alatum + spherical ca. 1.7 black — 
chiricanum spherical-depressed 2.5 x 2 black X 
commutatum rounded ca. 0.7 black X 
convolutum spherical. depressed 1.2-1.5 black X 
macrocephalum + spher ical ca. 1.8 black — 
mandonii spheri cal 1.2 black X 
prealtum + lis 1.7-2 x 1-1.5 black X 
tenuifolium rounded 0.9 x 0.6 black X 
tinctorium spherica ca. 1.5 black X 
trinerve spherical-depressed 1.1-1.2 black X 
Sisyrinchium sect. Eriphilema 
douglasii + angular 2-2.5 x 1.8-2 red-brown = 
scirpiforme + rounded 1.9-2.1 light brown = 
Sisyrinchium sect. 
filifolium angular 2-2.5 x 1.5-2 red-brown = 
Libertia 
sp. angular 1x1 dark brown = 
ixioides angular 1.8 x 1.5 red-brown = 
Ona 
obscura ovoid ca. 1.9 x 1.5 dark brown — 
Orthrosanthus 
chimboracensis angular 1.1-1.6 x I red-brown — 
occissapungus angular-elongate! 3-3.5 x 0.8-1.1 light brown — 
Phaiophleps 
biflora angular ca. 2.5 x 2 brown 
Solenomelus 
pedunculatus barrel-shaped? 2 x 1.3 ed-brown — 
segethii short-cylindric? 2.3 x ca. 1.4 dark brown 
sp. nov. nearly discoid?? 5-5.5 X ca. 3.5 red-brown 2x 
Tapeinia 
pumila elongate pyriform l.l x 0.5 red-brown 7 


l eles a distal and proximal wing. 


With linear arranged depr 


essions. 


x ^ With a broad half-circumferential win 


8- 
* With a small depression around the micropyle. 


Volume 76, Number 4 
1989 


Goldblatt et al. 
Seed morphology of Sisyrinchium 


FIGURES 1-6. 


occissapungus. — 4. Tapeinia pumila. —5. Solenomelus segethii. —6. Solenomelus sp. Scale bars 


for Sisyrinchium montanum, and 5. pallidum, and 
by Rodriguez (1986) for the Chilean $. azureum, 
all sect. Sisyrinchium. Among the species that we 
examined, this was noted particularly in 5. chtri- 


Seeds of genera of Sisyrinchieae. — 1. 


Libertia sp.—2. Orthrosanthus chimboracencis. 


= 0.1 mm. 


canum, S. convolutum, S. commutatum (Fig. 19), 
and S. tinctorium (all sect. Echthronema). Even 
the apparently smooth seed surface of S. trinerve 


shows this granularity at very high magnification. 


Annals of the 
Missouri Botanical Garden 


7. S. ram 


FIGURES 7-12. Seeds of Sisyrinchium sects. Sisyrinchium (1-10) and Echthronema 1, 12) 
bonis. —8. S. cf. arenarium. S. chil 


hilense. —10. S. commutatum. —11. S. alatum. —12. S. macrocephalum. 
Scale bars 0.1 mm. 


Volume 76, Number 4 Goldblatt et al. 
Seed morphology of Sisyrinchium 


FIGURES d a Seeds of Sisyrinchium sect. Echthronema. —13. S. prealtum. — 14. S. tinctorium. —15. 5. 
trinerve. . S. chiricanum. — 17. S. mandonii. —18. $. ta, Scale bars = 0.1 mm. 


1116 


Annals 
MES ES Garden 


FIGURES 19, 20. 
(20). Scale bars = 50 u 


very high magnification. No granular microsculp- 
turing was noted in species of any other genus 
included in our surve 

In Solenomelus the longitudinal lines of more 
or less circular depressions do not correspond to 
the surface cells. The depressions are larger than 
the individual cells, the outlines of which can be 
clearly seen over the entire seed surface in S. 
segethii (Fig. 5), but are rather obscure in Soleno- 
melus sp. (Fig. 6) and S. pedunculatus. In the 
latter the peripheral cells of the seed are unusual 

being elongate and they have a remarkably 
smooth surface (Fig. 20) 


DISCUSSION 


Most Iridaceae have seeds of a light to middle 
brown, sometimes reddish brown (Huber, 1969; 
Wagner & Goldblatt, 1984), the coloration attrib- 
uted to the presence of phlobaphene (Huber, 1969). 
A few exceptions include the glossy black seeds of 
Belamcanda (tribe Iridoideae) and Babiana and 
Antholyza (tribe Ixioideae), and the bright orange 
seeds of Chasmanthe (also Ixioideae) and Neo- 
marica variegata (Iridoideae tribe Mariceae). 
Whether the black color in the above genera or 
in Sisyrinchium can be attributed to the presence 
of phytomelan, a dense carbon substance that is 
regarded as important in the higher systematics of 
the lilioid monocots (Dahlgren & Rasmussen, 1983; 
Dahlgren et al., 1985), is unknown. 

The most common seed shape in lridaceae is 
more or less regularly angular (Huber, 1969) but 
globose or subglobose seeds are not uncommon. 
Winged seeds occur in Gladiolus and its close allies 
(Ixioideae), and discoid seeds are characteristic of 


ee details of seeds of Sisyrinchium commutatum (19) and Solenomelus pedunculatus 


some species of /ris and Moraea (both Iridoideae- 
Irideae) and Diplarrhena (Sisyrinchieae). The half- 
circumferentially winged seeds of Solenomelus sp. 
are reminiscent of those of Gladiolus (which have 
completely circumferential wings) and represent a 
notable example of convergence. 

Until now the blackish seeds encountered 
Sisyrinchium appear not to have been regarded 
as particularly significant, although the basic dif- 
ferences in seed color have been noted by Rodri- 
guez (1986), black in species attributable to sects. 
Sisyrinchium and Echthronema and brownish in 
sect. Eriphilema. Moreover, the more or less glo- 
bose shape and the frequent presence of a large 
umbilicus along the raphe, either between the hilum 
and micropyle or including the latter, appear to 
have attracted little attention except for occasional 
mention in species descriptions. We believe that 
these features have considerable taxonomic signif- 
icance, particularly in a. genus that is relatively 
unspecialized in the family and has few other dis- 
tinguishing characteristics. 

A conclusion that appears warranted is that the 
black, 


(apomorphic) condition in sects. Sisyrinchium and 


globose, and umbilicoid seeds are a derived 


Echthronema. It is likely, though by no means 
certain, that this seed, including the umbilicus, is 
the basic type in the two sections despite the fact 
North and 


South American species of both sections. 


that an umbilicus is not present in a few 


Seed data indicate that sects. Sisyrinchium and 
Echthronema should be considered sister taxa more 
closely allied to each other than to sects. Eriphi- 
lema and Nuno. The possibility that sects. Eri- 
philema and Nuno may be more closely allied to 
other genera of Sisyrinchieae, Phaiophleps in par- 


Volume 76, Number 4 
1989 


Goldblatt et al. 1117 


Seed morphology of Sisyrinchium 


ticular, than to the rest of Sisyrinchium will be 
explored in a future paper dealing with the phy- 
logeny of Sisyrinchieae. 


LITERATURE CITED 
BAKER, J. G. 1892. Handbook of the Irideae. George 
Bell & Sons, London. 


BENTHAM, G. Hooker. 1883. Irideae. Genera 
o 32) 681-710 
CHOLEW ENDERSON. 1984. Bios 


naie e Sisy rinc jiu section Be déco dii 
of the Rocky Mountains. Brittonia 36: 
63. 


md R. & F. Rasmussen. 1983. Monocotyledon 
evolution: hane ue and phylogenetic states. Evo 
lutionary Biology 16: 255-388. 

. T. cuc ls & P. Yeo. 1985. The Families 
of the Monocotyledons. Springer-Verlag, Berlin. 
GOLDBLATT, P. Phylogeny and classification of 
Iridaceae. Ann. Missouri Bot. Gard. (in press 
HENRICH, J. E. & P. GOLDBLATT. 1987. Mesoamerican 


Sisyrinchium cde new species and records, 
pit cation. Ann. Missouri Bot. Gard. 


and notes on 


74: 903-910 


. Iridaceae. /n: Flora Mesoamericana 


(in press a). 
& . Iridaceae. /n: Flora de Nicaragua 


(in press b). 
HoLmGREN, P. K., W. KEUKEN & E. K. SCHOFIELD. 
981. Index Herbariorum, Part 1. Regnum Veg- 
etabile 106. 
Huser, H. 
hifas der Liliiflorae. . Bot 
samml. München 8: 219-538 
RODRIGUEZ, R. 1986. Die Chilenischen Arten der Gat- 
tung Sisyrinchium L. (Irida iind Mitt. Bot. Staats- 
en 22: 07-20 


Die o ari n Verwandt- 
ot. Staats- 


samml. Münche 


Eon A. (editor). 8 ee Anatomy 
of Seeds. I Monocotyledons. Izdat. Nauka, Lenin- 
grad, 


WAGNER, W. L. & P. GOLDBLATT. 8 A survey of 
seed surface morphology in E "sperantha (Iridaceae). 


Ann. Missouri Bot. Gard. 181-190. 


NEW SPECIES OF Dieter C. Wasshausen? 
MENDONCIA 

(ACANTHACEAE) FROM 

THE VENEZUELAN GUAYANA! 


ABSTRACT 


ecent expeditions in the Venezuelan Guayana have resulted in additional new taxa in the genus d 
ea eae). The following is an effort to place these novelties on record for the convenience of other taxonomists 
prior to the publication of the Flora of the Venezuelan Guayana. Four new Venezuelan Guayana species of Me ndoncia, 
M. steye dea M. neblinensis, M. williamsii, and M. coriacea, are described, Juétratea, por compared with their 
closest relatives 


There are approximately 60 species of Men- 
doncia in Central and South America and in trop- 
ical Africa and Madagascar. The genus, named for 
Cardinal Mendonca, Patriarch of Lisbon, consists 
of twining herbaceous or sometimes suffrutescent 
climbers bearing cylindric red, white, or cream- 
colored axillary flowers with purple streaks in the 
throat. Each flower is subtended by a pair of con- 
spicuous, green, more or less connate bracts. Most 


taxa are limited to wet virgin forests and thickets 
at elevations between 50 and 1,000 m. Due to the 
fact that lianas and vines are generally poorly known 
and collected in the tropics, and given the super- 
ficial similarity of many of the species, the tax- 
onomy of the genus is rather difficult. In an attempt 
to identify recent collections of Mendoncia from 
the Venezuelan Guayana, I have found four un- 
described species of the genus. 


KEv TO THE VENEZUELAN GUAYANA SPECIES OF MENDONCIA 
la. Bracts ovate or ablong- -ovate. 
2a. Flowers 4 or 5 in each axil oe M. cardonae 
2b. Flowers solitary or in pairs in each axil. 
3a. 


ac 
Corolla cream-colored with purple streaks in the throat; bracts 2 cm long and 1 cm broad | 


.M. spruc ei 
3b. Corolla Bein to deep red; bracts 2.3-3 em long, 1.3-1.7 cm broad. 
4a. Bracts yellowish green to brownish green, 2.5-3 cm lone hirsute; leaf blades membranous, 
dn green and lustrous above, glabrous; corolla 3.5 cm long .. M. BER Mu 
4b. Bracts dull green, 2. x 2.5 cm m densely hirsute; leaf blades firmly membranous, dull 
gr bove, sparingly hirsute; corolla 3 cm long 
e oblong, oblong-elliptic, or ns obovate. 
. Bracts narrowly o long or oblong-lanceolat 
6a. Lea cad narrowly to broadly lii rather thin, densely ng, 
S m long, 0.8 cm broad, densely velvety pubescent . hoffmannseggiana 
6b. Leaf “blades. elliptic, ere minutely and sparingly hirtellous; bracts oblong-lanceolate, 2 to 
2.5 cm long, 1 cm broad, de el om rown strigos 
5b. Bracts da: oblong- ellipti ic or dione vate. 
Ta. Bracts oblong or oblong-elliptic, o leaf blades subcoriaceous to thick-coriaceous 
8a. Leaf blades elliptic to oblong, 3-6 cm broad; bracts vinaceous-lavender with green, bling 


E 


stey ante 


pubescent; bracts narrowly oblo 
M. 


M. williamsii 


M. coriacea 
8b. Leaf blades oblong-elliptic, 7.5-9 em broad; bracts green, ded -elliptic d M. phalacra 
Tb. lag oblong-obovate to E -elliptic, not coriaceous; leaf b 
racts a obova 1.3 cm broad, essentially glabrous: stem glabrous; leaf blades 
glabrous, chartaceous, "the veins prominent on ihe lower surfac M. obovata 
9b. B SUE elliptic, 1.5-1.8 cm broad, long-hirsute; stem Penn leaf blades hirsute, 
membranous, the veins inconspicuous on the lower surfac 


es membranous, not coriaceous. 


M. bivalvis 


' My special thanks to Alice "hm who skillfully prepared the line drawings, and the staff of the National 
Museum of Natural History SEM Laboratory for their high-quality pollen ao 
* Department of Botany, Ear aL Washington, D.C. 20560, U.S.A 


ANN. Missouni Bor. Garp. 76: 1118-1124. 1989. 


Volume 76, Number 4 Wasshausen 1119 
1989 Mendoncia 


HN 
m m 
ul IM 


B 


f| i 


Ww 


j 


] 


A 


NI / / ES 
t {hf 
NW 4/4; 
N YG 


Z 
a- 


A Tanger 


C 


FiGURE 1. A, B. Mendoncia neblinensis. — A. Habit (W. W. Thomas 3196). —B. Corolla (M. Nee 30578). C- 
E. Mendoncia steyermarkii (Steyermark & G. C. K. & E. Dunsterville 104353). —C. Habit. — D. Bract. — E. 
Corolla. 


1120 


Annals of the 
Missouri Botanical Garden 


Mendoncia neblinensis Wasshausen, sp. nov. 

PE: Venezuela. Territorio Federal Amazo- 

nas: along Rio Baria just upstream from Base 

Camp on SW side of Cerro de la Neblina, 

0?*49'50"N, 66?09'40"W, 140 m, 27 Jan. 

1985, M. Nee 30578 (holotype, US; isotype, 
NY). Figure 1A, B. 


caulibus ecu cobi) parc 
entibus E de lamina 


Frutex volubilis, 
hirsutis et lepidotis, pilis asce 


vel parce puberula, subtus pallide viridis, glabra vel parce 
hirsuta, costa et venis Mis bus subtus prominentibus; 
petioli subquadrangulares, parce hirsuti; flores (1 vel 2) 
axillares; pedicelli tortiles, aues hirtelli, pilis appressis; 
hisciene d be flavido- vel brunneolo-virides, apice a 


la 
obovatis, leviter émarginalls dap lisse glabra. 


Climbing vine; stem subquadrangular, sparingly 
hirsute and lepidote with stellate scales, the tri- 
chomes ascending. Leaves petiolate, the petioles 
1-2.5 cm long, subquadrangular, sparingly hir- 
sute, the blades elliptic to ovate, thinly membrana- 
ceous, 5-11 cm long, 3-5.5 cm broad, abruptly 
acuminate, the tip aristate, the awn to 6 mm long, 
abruptly acute at the base, the upper surface dark 
green and lustrous, glabrous or bearing a few ap- 
pressed trichomes arising from stellate bases, mi- 
nutely punctate, the lower surface paler and dull 
green, glabrous or sparsely hirsute (especially the 
costa and lateral veins bearing a few appressed 
trichomes), the venation prominent beneath, lateral 
veins 3 or 4 pairs, veinlets coarsely reticulate. 
Flowers axillary, 1 or 2 in each axil, the pedicels 
3.5-5.5 cm long, twisted, gradually enlarged to- 
wards tip, sparingly hirtellous, the trichomes ap- 
pressed upward, straight; bracts ovate, yellowish 
green to brownish green, 2.5-3 cm long, 1.3- 1.7 
cm broad, apiculate, the apicula about 1 mm long, 
the outer surface minutely muricate and hirsute, 
the trichomes straight, rigid, appressed upward, 
the costa rather prominent, the inner surface gla- 
brous, minutely rugose; corolla bright to dark red, 
3.5 em long, the lobes pink within, base of tube 
white, glabrous or sparingly hirsute towards the 
throat, the tube 5 mm broad at base, narrowed to 
2.5 mm at 8 mm above base, the throat 8 mm 
broad, the upper lip about 8 mm long, the lobes 
subequal, elliptic to obovate, 5 mm long and 3.5 
mm broad, shallowly emarginate at apex, the lower 
lip somewhat shorter than the upper lip; pollen 
grains 5-colporate (brevissimicolpate), prolate- 
spheroidal, 34 x 30 um, sexine punctitegillate; 
calyx annular, 0.5 mm long, glabrous; style 2.5 


cm long, glabrous; ovary glabrous. Drupe fleshy, 
black, ellipsoid, 17 mm long, 8 mm wide, glabrous. 


Distribution. Lowland evergreen tropical for- 
est on hummocks along gravelly black-water rivers, 
Territorio Federal Amazonas at elevations between 


100 and 180 m 


Paratypes. | VENEZUELA. TERRITORIO FEDERA 
de AZONAS: Cerro de la Neblina, along Rio Mus 

*50'N, 66*10 W, 140 m, Stein, Gentry & Thomas 1716 
(MO, US); same locality, Davidse & Miller 27452 (MO, 
US); same locality, Lie "DA 15667 (MO, US); same lo- 
i i ), US); same locality, Liesner 
& Funk, 15822 (MO, same locality, Thomas 3318 
(MO, US); same Ep Kral 7 E same loce cality, 
Thomas 33 , US); upper Rio i )*55' 
66?15'W. 100 m, Davide & Miller oe (MO, US); 
between en base camp and "Puerto rq 
warinuma, 0°50'N, ca. 66?98'W, -] 
Stein 46967 (MO, US); 2-6 km I P base c 
66?08'W, 160 m, Thomas 3196 (NY, US). 


'amp, 0°50'N, 


Mendoncia neblinensis is probably related to 
M. sprucei Lindau. It differs markedly from M. 
sprucei in that the latter species has green bracts 
2 cm long and 1 cm broad, sparingly appressed- 
pilose pubescence on the leaf blades, and corollas 
3 cm long. In contrast, M. neblinensis has yel- 
lowish to brownish green bracts cm long 
and 1.3-1.7 cm broad, glabrous leaf blades, and 
corollas 3.5 cm long. Lindau (1897) in his de- 
scription of M. sprucei did not indicate the color 
of the corolla, nor did Turrill (1919) in his revision 
of the genus. Leonard (1951) described the corolla 

f M. sprucei as cream colored with purple streaks 


5: 


in the throat. I believe this to be in error; the 
corolla of M. sprucei is also bright red. Turrill 
(1919) stated in his discussion that M. sprucei 


appears to be very closely related to the red-flow- 
ered M. aspera Nees and that perhaps M. sprucei 
was only a variety of M. aspera. 


Mendoncia steyermarkii Wasshausen, sp. nov. 
TYPE: Venezuela. Estado Bolivar, en el drenaje 
del Rio Cuyuni, al sur de El Dorado, 1,300- 
. 1979, Julian A. Stey- 
L.C. K. & E Dunsterville 104353 
oops. US; isotype, VEN). Figure 1C-E. 


Frutex vokabilis; caulibus subquadrangularibus, sulca- 
pilis ascendentibus. 


integra, supra d parce hirsuta, subtus fulvo-viri- 
a venis lateralibus parce E pue 


dis, glabra vel « 
1 vel 


petioli glabri eee in canalibus parce hirsuti; flore 
2) axillares; pedicelli recti, flavo-virentes, dense "hirtelli; 
bracteae ovatae, obscure virides, apiculatae, externe dense 
use minute muricatae, intus glabrae et minute ru- 


Volume 76, Number 4 
89 


Wasshausen 1121 


Mendoncia 


gosae; corolla atrosanguinea, glabra, lobis rotundatis vel 
leviter emarginatis; drupa subglobosa, nitida, glabra 


Trailing vine; stems subquadrangular, grooved, 
sparingly lepidote (the scales stellate), bearing nu- 
merous ascending, erect, hirsute trichomes. Leaves 
on petioles 1-2.5 cm long, subquadrangular, gla- 
brous or the channels sparingly hirsute with straight 
ascending trichomes, the blades firmly membra- 
nous, ovate, 9-11 cm long and 4-6.5 cm broad, 
apically abruptly acuminate (the tip aristate, the 
awn 6 mm long), the base abruptly acute to obtuse, 
the upper surface dull dark green, sparingly hir- 
sute, the trichomes straight, + ascending, arising 
from stellate bases, the lower surface dull tawny 
green, glabrous or the costa and lateral veins bear- 
ing a few scattered, stiff, straight, ascending tri- 
chomes, lateral veins 3 or 4 pairs, veinlets coarsely 
reticulate, the venation prominent and raised be- 
neath, less so above. Flowers axillary, 1 to 2 in 
each axil, the pedicels yellow-green, 4-6 cm long, 
straight, densely hirtellous, the trichomes ap- 
pressed upward, straight, rigid; bracts dull green 
ovate, 2.3-2.5 cm long, 1.3-1.5 cm broad, with 
an apicula about 1 mm long, the outer surface 
minutely muricate and densely hirsute with straight, 
rigid, trichomes, these appressed upward, the costa 
rather prominent, the inner surface glabrous, mi- 
nutely rugose; corolla deep red, glabrous, 3 cm 
long, the tube 5.5 mm broad at base, narrowed to 
3 mm at 5 mm above base, the throat 8 mm broad, 
the upper lip about 5 mm long, the lobes rounded, 
3.5 mm broad, shallowly emarginate at apex, the 
lower lip somewhat shorter than the upper lip; 
pollen grains 5-colporate (brevissimicolpate), pro- 
late-spheroidal, 42 x 37 um, sexine punctitegil- 
late; calyx annular, 0.5 mm long, glabrous; style 

.8 cm long, the lower portion hirsute; ovary gla- 
brous. Drupe subglobose, black, shining, 15-18 
mm diam., glabrous. 

Distribution. Wooded ridge and tall, moist 
forests with trees on the average 25-30 m tall, 
Estado Bolivar, Venezuela, at 1,100-1,400 m. 


BOLIVAR: Gran. Sabana, El 
1,200 
C. K. hi 


Paratypes. | VENEZU ELA. 


E. Dunstertilla 105454 (US, VEN); El Dorado-Sta. Ele 
de Uairen road, km 127-131, 1,300-1,350 m, Badillo 
& Holmquist 6094 (MY, US); El Dorado-Sta. Elena de 
Uairen road, km 119-122, S of El Dorado, ca. 1,100 
m, Gentry, G. Morillo & B. de Morillo 10460 (MO, US); 
arretera between campamento 125 and km 127, be- 
tween Luepa and Cerro Venamo, 1, t m, Steyermark 
& Nilsson 194 (US, VEN); 134 km S of El Dorado, 
1,300-1,350 m, Buc ark & G. C K. & Dun 
sterville 104450 (US); carretera El Dorado-Gran 5a- 


bana, La Dante, 1,250 m, Ruiz-Terán & López-Palacios 
11460 (MERF, US). 

endoncia steyermarkii superficially resem- 
bles the Colombian lowland forest species M. ca- 
quetensis Leonard. However, in M. caquetensis 
the bracts are green and sparingly and inconspic- 
uously hirsute; and the corollas are white(?) (fide 
Leonard, 1961) with the tube 4 mm broad at the 
base and narrowed to 2.5 mm at | cm above the 
base, and the upper lip is about 1 cm long. In M. 
steyermarkii, the bracts are dull green and densely 
hirsute, the corollas are deep red with the tube 5.5 
mm broad at the base and narrowed to 3 mm at 
0.5 cm above the base, and the upper lip is about 
0.5 cm long. 


Mendoncia williamsii Wasshausen, 
. Territorio Federal Amazo- 


sp. nov. 


30 May 1942, Llewelyn Williams 15697 
(holotype, US; isotype, VEN). Figure 2A-C. 


Frutex asuma caulibus nie qye de stri- 
gosis, pilis brunneis. Foliorum lamina sup ica, bre 
minata, stus a acuta, chartacea, in ie ra, supra 
(in sicco) subnigra, parce et minute hirtella, pilis a dee 
planis stellatis RS TE subtus (in sicco) brunnea, aequal- 
li strigosi; flores 2, axillares; pedicelli 


a aperta; co- 
rolla non visa; drupa ovoidea, leviter compressa, minute 
pubescens. 


Vine; stem subquadrangular, rather densely stri- 
gose upward, the trichomes brownish, to 2.5 mm 
long. Leaves on petioles 1-2 cm long, strigose, the 
blades elliptic, to 15 cm long and 9 cm broad, 
apically short-acuminate with a cusp to 5 mm long, 

basally acute, chartaceous, entire, the upper sur- 
face drying blackish, minutely and sparingly hir- 
tellous with the ascending hairs 0.25-0.50 mm 
long and arising from flat, stellate bases, the costa 
and lateral veins (4 or 5 pairs) rather prominent, 
strigose with brown trichomes about 1 mm long, 
the lower surface drying brown, + evenly strigose, 
both surfaces bearing rather prominent coarsely 
reticulate veinlets. Flowers usually 2 in each axil; 
pedicels 4 cm long, densely brown strigose; bracts 
oblong-lanceolate, 2-2.5 cm long, 1 cm broad just 
above base, gradually narrowed to an acute tip, 
densely brown-strigose, the costa evident, the inner 
surface glabrous; flowers not seen. Drupes 1.5 cm 
long, 6 mm wide, 4 or 5 mm thick, oblique at tip, 
the surface minutely and rather densely pubescent. 

Distribution. 
Territorio Federal Amazonas, Venezuela. 
only from the type locality. 


In dense forest of terra firma, 
Known 


Annals of the 


1122 
Missouri Botanical Garden 


=== 


PA ! 
Ve 
S 

T^ 
| 


WEG 


C. Mendoncia williamsii (L. Williams 15697).—A. Habit. — B. Bracts. —C. Drupe and bract. 
K. & F. Dunsterville 104474). —D. Habit. — E. Bracts. — F. Corolla. 


FIGURE 2. : 
D-F. Mendoncia coriacea (Steyermark & G. C. 


Volume 76, Number 4 


Wasshausen 
Mendoncia 


1123 


FIGURE 3. Scanning electron microscope (SEM) Daraus m of Me a ia pollen. —a. Mendoncia coriacea, 
id view, X 1,200 wire a ark & G. C. unsterville 104474).—b. M. steyermarkii, equa 1atorial 
200 


view, X1,75 de e & C 
(R. Liesner 15667 


Mendoncia williamsii is most closely (but dis- 
tantly) related to M. sprucei Lindau. In M. sprucei 
the bracts are ovate and sparingly appressed-hir- 
tellous without; the flowers are usually solitary; and 
the leaf blades are ovate to oblong-ovate, firmly 
membranous, and about 6 cm broad. In contrast, 
in M. williamsii the bracts are oblong-lanceolate 
and densely brown strigose without; the flowers are 
usually in twos; and the leaf blades are elliptic, 
chartaceous, and considerably wider (about 7.5 cm 


broad). 


K. € E. a 104353). — c. M. neblinensis, equatorial view, 


Mendoncia coriacea Wasshausen, sp. nov. TYPE: 
Venezuela. Bolivar: km 133.5, drenaje del Rio 
Cuyuni, al sur de El piod 1,300-1,380 
m, 2. Dec. 1970, Julian A. Steyermark 
& G. C. K. & E. pi EE 104474 (ho- 
lotype, is isotype, MO). Figure 2D-F. 


Suffrutex volubilis, caulibus subquadrangularibus, gla- 
bris. Foliorum lamina elliptica vel oblonga, apice abrupte 
acuminata et mucronata, basi acuta acea, integra, 
glabra, costa et venis lateralibus prominentibus subtus; 
petioli c Peut. glabri; flores 2, axillares; pedi- 


1124 


Annals of the 
Missouri Botanical Garden 


celli graciles, glabri, olivacei suffuso marronini; bracteae 
e ndulae viridi-vittatae, 


obliquo, lobis suborbicularibus, leviter lisse drupa 
subglobosa, glabra. 

Large suffruticose liana, 25-30 m high; stem 
subquadrangular, glabrous. Leaves on petioles 2— 
3.5 em long, subquadrangular, glabrous, the blades 
subcoriaceous to thick-coriaceous, elliptic to ob- 
long, 8-14 cm long, 3.5-6 cm broad, the tip 
abruptly acuminate (the tip blunt and terminating 
in a mucro 1 mm long), acute at base, entire or 
undulate, both surfaces glabrous, dark green and 
minutely punctate above, dull green below, the 
costa and lateral veins (3-5 pairs) obscure above 
and prominent and raised beneath, the veinlets of 
the middle lower leaves forming a conspicuous re- 
ticulation on the lower surface. Flowers axillary, 
usually 2 in each axil; pedicels 3-4 cm long, slen- 
der, 0.75 mm broad at base, gradually enlarged 
to 2 mm at tip, glabrous, olive green suffuse 
maroon; bracts coriaceous, vinaceous-lavender with 
green, oblong, 2.5-3.5 cm long, 1-1.4 cm broad, 
rounded or obtuse at apex and tipped by a mucro 
l mm long, rounded at base, glabrous and minutely 
rugose without, glabrous and minutely punctate 
within; corolla white with lavender stripes in the 
tube within, glabrous, 4.5 cm long, the tube 4 mm 
broad at base, narrowed to 3 mm at 10 mm above 
base, the throat 8 mm broad, the upper lip about 
l cm long, the lobes unequal, suborbicular, 6 mm 
long and 9 mm broad, shallowly emarginate at 
apex, the lower lip somewhat shorter than the upper 
lip; pollen grains 5-colporate (brevissimicolpate), 
prolate-spheroidal, 58 x 53 um, exine thick, cras- 


sinexious; calyx annular, 0.5 mm long, glabrous; 
style 2.9 cm long, glabrous; ovary glabrous. Drupe 
subglobose, dark wine-purple to black, 18 mm long, 
15 mm in diam., glabrous. 


Distribution. Wooded ridge and tall, moist 
forest with trees on the average 25-30 m tall, 


Estado Bolívar, Venezuela, at 1,200-1,380 m 


Paratypes. 


ls BOLIVAR: between fid 
mento ra 


m, Ruiz- Terán & López oe 11481 (MERE. US ji 


Mendoncia coriacea is perhaps most closely 
related to the widespread lowland Amazonian species 
of Colombia and Peru, M. pedunculata Leonard, 
which differs markedly by having larger (4 cm long, 

1.4-1.8 c road), violet bracts; membranous, 
wider (6-10 cm broad) leaf blades; and longer, 
white corollas marked by reddish brown within and 

to © cm long. In contrast, M. coriacea has 
coriaceous, vinaceous-lavender with green bracts 
2.5-3.5 cm long and 1-1.4 cm broad; subcoria- 
ceous to thick-coriaceous leaf blades 3.5-6 cm 
broad; and shorter (4.5 cm long), white, internally 
lavender-striped corollas. 


LITERATURE CITED 
LEONARD, E x 1951. The Erap of Colombia, 
. Co J.S. Natl. Herb. 31: 11-40. 

"i f a iH novae ve 
criticae 4L Seven new species from Colombia and 
some additional notes. Wrightia 2: 144-146. 

LiNDAU, G. 1897. Acanthaceae Americanae et Asiati- 


cae. Novae vel minus cognitae. Bull. Herb. Boiss. 5: 


647 
TURRILL, WY., B. 


19 A revision sif the genus Men- 
doncia. Kew Bull. 41 


19. 
1919: 417 


NEW SPECIES, NAMES, 
AND COMBINATIONS IN 
AMERICAN COMBRETACEAE 


A. Alwan Al-Mayah and 


Stace! 


A. R. 
C. A. 


ABSTRACT 


Revision of the family Combretaceae for accounts in eight tropical American floras has nese the following 


twelve nov 
subsp. domingensis (Urban) Alwan 
Stace, comb. nov.; 
nov.; T. crispialata d Alwan & Stace, e 
Alwan & Stace, s ; T. uleana Engl. ex Alwa 

Bucida molineti (M. "Cémex) Alwan & Stace, donis nov.; 


mb. no 


nov.; 


elties: Terminalia Vida Anc oe Mer (Bisse) Alwa 


T. ramatuella Alwan & Goce. nom. nov.; 
T, steyermarkii Alwan & 
1 & OR: sp. nov.; 


an & Stace, comb. nov.; 7. chicharronia 
; T. chicharronia subsp. orientensis Monach.) Alwan & 
T. virens (Spruce ex Eichler) Alwan i Stace, comb. 
Stace, sp. nov uhlmanniü 
eichleriana Alwan & Since: nom. nov.; 
and Buc henavia amazonia Alwan & Stace, sp. nov. 


-~ 


l. Terminalia chicharronia Wright ex Sau- 
valle. 

Restricted to Cuba, Haiti, and the Dominican 
Republic, this species is very variable in the size 
and shape of the leaves and in the amount of 
indumentum on the branchlets, leaves, and inflo- 
rescence. These characters reveal four reasonably 
well geographically separated taxa recognized here 
as subspecies. 

Terminalia chicharronia was based on Chi- 
charronia intermedia A. Rich. meg which in 
turn was based on Sagra 185 and 281. The spec- 
imens at G are the best ones, but there is no 
evidence that Richard saw them. In P there is one 
sheet with three shoots—two in flower and one in 
fruit — which Richard did see. The label says “Her- 
barium Richard. No. 185, Combretum sp. nov., 
Guanabo, Marzo 1829, Coll. R. de la Sagra." The 
two flowering shoots are here designated as lec- 
totype because in G there is a note from Sagra 
saying that no. 185 was sent first and was flowering, 
and no. 281 was sent later (both March 1829) and 
fruiting. This typification fixes subsp. chicharronia 
as the subspecies with densely pubescent to rufous 
tomentose leaves with strongly raised venation. It 
occurs most commonly in eastern Cuba (prov. Ori- 
ente), where it is sympatric with the closely related 
T. eriostachya A. Rich., which differs by having 
larger flowers, longer inflorescences, and glabrous 
styles. Terminalia maestrensis Bisse is here con- 
sidered to be a synon 

The other ee i are as follows: 


la. Subsp. neglecta (Bisse) Alwan & Stace, comb. 
nov. BASIONYM: T. neglecta Bisse in Feddes 
Repert. Spec. Nov. Regni Veg. 85: 607. 1974. 


Western and central Cuba (provinces Pinar de 
Río, Habana, and Las Villas, including Isla de Pi- 


nosJ. 


lb. Subsp. domingensis (Urban) Alwan $ Stace, 
comb. nov. BASIONYM: 7. domingensis Urban 
in Symb. Antill. 7: 524. 1913 


Haiti and Dominican Republic. 


1c. Subsp. orientensis (Monach.) Alwan & Stace, 
comb. nov. BASIONYM: T. orientensis Monach. 
in Caribbean Forest. 8: 79. 1974 
Synonyms: T. nipensis Alain, T. aroldoi Bisse, 
T. pachystyla Borh. 


Eastern Cuba (province Oriente, mostly at higher 
altitude). 


2. Ramatuella Kunth 


This former genus was revised by Exell & Stace 
1963), who recognized six species. Subsequent 
collections have shown that this should be reduced 
to three. Moreover, although Ramatuella is still 
recognizable by the combination of characters out- 
lined by Exell & Stace, it seems far better reunited 
with Terminalia, as its distinction is based on only 
one of several combinations of characters found in 
Terminalia. Accordingly, we reduce it to sectional 


— 


status. 


! Department of Botany, University of Leicester, Leicester LEI 7RH, England. 


ANN. Missouni Bor. GARD. 76: 1125-1128. 1989. 


1126 


Annals of the 
Missouri Botanical Garden 


2a. Terminalia sect. Ramatuella (Kunth) AI- 
wan & Stace, comb. . BASIONYM: Rama- 
tuella Kunth, Nov. Gen. Sp. 7: 253. 1825 
(^ Ramatuela," based on misspelling of “Ra- 
matuelle””) 


2b. Terminalia ramatuella Alwan & Stace, 
nom. nov. pro Ramatuella argentea Kunth, 


Nov. Gen. Sp. 7: 254, tab. 656. 1825. 


The epithet argentea cannot be transferred to 
Terminalia as there already exists T. argentea C. 
Martius (1824), a very different species, from Bra- 
zil, Paraguay, and Bolivia. 


2c. T. virens (Spruce ex Eichler) Alwan & Stace, 
BASIONYM: Ramatuella virens 


C. Martius, Fl. Bras. 


comb. nov. 
Spruce ex Eichler in 
14(2): 100, tab. 26, fig. 2. 1867. 

Synonyms: R. maguirei Exell & Stace, R. la- 
tifolia Maguire. 


2d. T. crispialata (Ducke) Alwan & Stace, comb. 
nov. BASIONYM: Ramatuella crispialata 


Ducke, Arq. Inst. Biol. Veg. 2: 65. 193 
Synonym: R. obtusa (Maguire) Exell & bu 


3. Terminalia steyermarkii Alwan & Stace, 
sp. nov. TYPE: dwarf woody plant with branch- 
es spreading from near ground, only 0.5-1 m 
tall; flowers creamy-buff; leaves crowded at 
tip of branch, coriaceous, dull green above, 
gray-sericeous below, the apical youngest 
leaves buff-silvery sericeous; filaments green- 
ish white; ovary and calyx gray-green; peri- 
anth pale green. Venezuela. Bolivar: Cumbre 
de Cerro Guaiquinima, sector nororiental, entre 
las brazos nororiental del Rio Carapo, en una 
meseta arenosa pedregosa y cubierta con veg- 
etación muy esparcida, 5%54/N, 63%25'W, 980 
m, 30 May 1978, Steyermark, Berry, G. & 
E. Dunsterville 117521 (holotype, LTR; iso- 
type, VEN, not seen). 
Frutex 0.5-1 m altus, ramulis dense villosis. Folia 
DAP ordinata ad ramulorum apic es congesta; petiolus 
10 mm longus, dense sericeus; m coriacea, 
sea vel elliptico-obovata, (3 a x 3-6.5 em, 
apice retusa vel rotundata, basi cunea en sa in petiolo 
dec urrente, supra pubescens, subtus ae eo argentea, 
7-9-paribus; venatio brochidodroma in- 


eri; receptaculum inferum 3.5 
mm longum, dense villosum; receptaculum superum 2 x 

n lobi 5 vel 4, 1.5 mm 
a ad 4-5 mm; stylus 5-6 
mm longus, ad medium pon pl Fructus ignotus. 


Terminalia steyermarkii is one of those re- 
markable coriaceous-leaved species of sects. Ra- 
matuella and Pachyphyllum Maguire & Exell that 
are confined to the vicinity of the watershed be- 
tween the Rio Orinoco and the northern tributaries 

of the Amazon. It differs from all but T. ramatuella 
by having a densely sericeous lower leaf surface, 
and from T. ramatuella by having an apparently 
dwarfer habit, broader and thicker leaves, and more 
elongated rachis 
erminalia €—À is known only from 
the type collection. Without fruits its precise af- 
finities are uncertain, but the elongated rachis with 
bisexual flowers borne along its length suggests 
sect. Pachyphyllum rather than sect. Ramatuel- 
a 


A 


. Terminalia kuhlmannii Alwan & Stace, sp. 
nov. TYPE: Brazil. Espirito Santo: Goytacazes, 
Rio Doce. n.v. “Pelada,” arvore frequente no 
mato, da madelta a provei tavel, 16 Dec. 
1943, Kuhlmann 06688 (holotype, RB 
62982; isotypes, NY, R 


. Folia spiraliter ordinata ad ramulorum apices 
conge aie petiolus 1-2 cm longus, glaber vel S arse pu- 
bescens, eglandulosus; lamina chartacea, 5-12 3-6 
cm, obovata, apice subacuta vel aida: dene acuta, 


supra glabra, subtus + glabra; venatio brochidodroma 
conspicua, venis secundariis subtus ra ibus 5-8 
paribu us, venis tertiariis subtus reticulatis, venulis conspic- 


uis vix prominentibus. Pedunculus (in ct tu) 2-3.5 cm 
longus, pubescens; rachis (in fructu) 2-3.5 cm longa, 
dense pubescens. Fructus 2-alatus, transversali oblongus, 
1.2-1.7 x 3.5-4.6 er D. denpe DN corpore 3-4 
mm lato, alis oblongis 1.4- >m lat 


Terminalia kuhlmannii belongs to sect. Dip- 
tera (Eichler) Engl. & Diels, which includes four 
other South American species. Of these, 7. janu- 
ariensis DC. and T. guyanensis Eichler have much 
larger glabrous fruits; T. argentea C. Martius has 
ovate leaves, and leaf lower surface and fruits 
usually tomentose at least when young; and T. 
phaeocarpa Eichler has larger leaves usually acute 
to rounded at apex with eucamptodromous vena- 
tion, and usually glabrous fruits. Terminalia kuhl- 
mannii most closely resembles T. guyanensis, but 
this is geographically remote and differs addition- 
ally in its narrower leaves with less conspicuous 
venation and shorter inflorescence 

A flowering specimen (Silva 356, LTR & CVRD) 
from close to the type locality probably also belongs 
here. It is a tree of 29 m also known as 
The flowers are densely pubescent, cream, and with 
styles pubescent to near the apex. Another flow- 
ering specimen (Almeida & Santos 191, LTR € 


“Pelado.” 


Volume 76, Number 4 
1989 


Alwan Al-Mayah & Stace 1127 


American Combretaceae 


CEPEC), from near Ilheus, Bahia (ca. 550 km 
north of the former) may also be the same species; 
it has peduncles up to 6 cm and rachises up to 7 
cm, but otherwise is very similar. 


en 


. Terminalia uleana Engl. ex Alwan & Stace, 
sp. nov. TYPE: Brazil. St. Catarina: Baum im 
Ufer des Capivary, Tubarào, Jan. 1889, E. 
Ule 1004 (holotype, HBG; isotypes, B (de- 
stroyed, photo F), P, US) 


rbor? Folia alternata, non congesta; petiolus 1 cm 


obovata, apice acuminata vel acuta, basi cuneata, mx 
pubescens vel glabrescens, subtus pubescens; venatio e 
camptodromo- Àrochidodroma: venae secundariae infra 
prominentes, eb. iiie pete fortuito retic- 
ulatae, reola etus Flores ignoti. 
Fructus Pimen 'ersali oblongus, 1.3-1.5 x 
(2.5-)3-4.5 cm, glaber; corpus ovato-ellipticum, 1.5 X 
"m, protuberans vel vy um carinatum; alae latiores 
quam longae, 1.3-1.7 x 1.3-2 cm, deorsum curvatae 


Engler wrote the name “Uleana” on the sheet 
at B but did not publish it. Also written on the 
sheet is the idea that the species is related to T. 
argentea. The new species belongs to sect. Aus- 
trales Engl. & Diels, itself close to sect. Diptera, 
which contains T. argentea. Of the species in sect. 
Australes, T. uleana differs from T. australis Cam- 
bess. and T. reitzii Exell in fruit shape and from 
T. triflora (Griseb.) Lillo by having wider leaves 
and wider fruits with a differently shaped wing. 
The fruit wing of 7: uleana is often curved down- 
ward (retrorsely), and this character is diagnostic; 
a third rudimentary wing is often present. The 
species has not been found since 1889. 


6. Terminalia eichleriana Alwan & Stace, nom. 
nov. pro T. punctata Kichler in C. Martius, 
Fl. Bras. 14(2): 85. 1867, nom. illegit, non 
Roth (1821), nec Spruce ex Eichler (1867) 
(nom. syn. pro Buchenavia punctata Eich- 
ler). TYPE: Brazil. Bahia: Serra d'Acurra, 
Blanchet 2794 (lectotype, BR; isolectotypes, 
BM, BP, BR, F, FI, G, K, LE, P, RB). The 
specimen chosen as lectotype was annotated 

y Eichler, who did not see fruits, although 
the isolectotype at G does bear fruits. The 
isolectotype at FI has the additional data: 
“Certáo do R. S. Francesco, 1838." We know 
of several later collections, all from the state 


of Bahia 


Terminalia eichleriana belongs to sect. Áctino- 


phyllae Engl. & Diels and closely resembles T. 


fagifolia C. Martius in vegetative characters. The 
leaves usually differ in having 3-5 (vs. 5-12) pairs 
of lateral veins with a narrower angle of divergence. 
The fruits are very distinct: in T. eichleriana the 
3(4) wings are 0.5-0.8 x 0.4-0.5 cm, while in 
T. fagifolia the 2 wings are 0.8-1.3 x 0.5-1.3 
cm 


7. Bucida molineti (M. Gómez) Alwan & Stace, 
comb. nov. BASIONYM: Terminalia molineti M. 
Gómez, Anales Soc. Esp. Hist. Nat. 19: 244. 
1890, nom. nov. pro Bucida angustifolia A. 
Rich., Hist. Phys. Cuba, Pl. Vasc. 521. 1846, 
and A. Rich. ex Griseb., Cat. Pl. Cub. 109. 
1866, non DC. (1828). 
(presumably Sagra) specimens connected with 
Richard's 1846 publication, but the specimen 
of Wright 2573 at G, labeled “Bucida an- 
gustifolia Rich. (non DC.) ex Griseb. Cat. Cub. 
p. 109” is the basis for Grisebach's publication 
and can be selected as the lectotype of B 


We have seen no 


molineti. De Candolle's Bucida angustifolia 
is Terminalia amazonia (J. Gmel.) Exell (type: 


French Guiana, C). 


Perrottet s.n., 


Terminale spinosa Northrop, Mem. Torrey Bot. Club 
12: 54, tab. 13. 1902, non Engl. (1895). 

Bucida Hii me ico mcum Ann. Carnegie Mus. 
ll: 201. ; 


B. ophiticola Bisse, Feddes pee Spec. Nov. Regni 
Veg. 85(9-10): 605. 
B. pcs Wilbur, Taxon = 467. 1988, nom. nov. 


o Terminalia spinosa Northrop, non Eng 


Bucida spinosa (Northrop) Jennings is illegiti- 
mate since its basionym (Terminalia spinosa 
Northrop) is a later homonym. Realizing this, Del- 
endick (1984) proposed that B. spinosa Jennings 
be treated as a new species dating from 1917, not 
realizing that the earlier epithet molineti existed 
and should be used. Moreover, recently the com- 

on procedure of legitimizing names in the way 
that Delendick proposed has been called into ques- 
tion (Wilbur, 1988). Wilbur argued that the orig- 
inal intention of the combining author (Jennings) 
should not be ignored, and that names should there- 
fore not be in the way that Delendick 
proposed. Accordingly, Wilbur proposed the new 


ilbur’s argument 


“rescued” 


name B. correlliana. Even if 
is accepted, the name B. ophiticola predates B. 
correlliana. 


8. Buchenavia amazonia Alwan & Stace, sp. 
nov. TYPE: Colombia. Caqueta: wet tropical 
forest of Amazon Basin, 1 km N of Solano, 8 
km southeast of Tres Esquinas, on Rio Ca- 
queta below mouth of Rio Orteguaza, 200 m. 


1128 


Annals of the 
Missouri Botanical Garden 


Tree 8 m high, 20 cm dbh; bark brown, 
smoothish; fruits green, oblong. Wet soil of 
old channel, near river; grass pasture of cleared 
lowland forest and in lowland forest. 7 Mar. 
1945, Little & Little 9626 (holotype, US; 
isotype, COL 


rutex vel arbor parva, 2-8 m alta. Folia spiraliter 
ordinata ad ramulorum apices congesta; petiolus 0.8-2 
cm longus, dense rufo-pubescens, saepe Digiti dorus, 
lamina chartacea, (2-)3-8(-12.5) x 1.5-4(-5.5) cm 
obovata vel anguste obovata vel jen. apice rotundato- 
obtusa vel acuminato-subacuta, b uneata, supra fere 
glabra, subtus fere glabra, venis Sn Bear e riisque 
sparse pubescentibus exc is venatio did x aie vel 


- 


eucamptodromo-brochidodroma; costa i a prom- 
inens; venae secundariae infra prominentes 2d 'aribus; 
venae tertiariae irregulariter percurrente nspicuae; 


areolae parvae, perfecte effectae rescata elon- 
rufo-pubescens; rhachis 1 
4) x 


res 2-3( mm; receptaculum inferum 1-2 mm 


longum, UE Sob ed e culum es gla- 
brum. F lipsoideo-ovoi i 1.3-2 1.2 cm, 
tundato-obtuso vel subacü uto vel ed SES 


basi rotundato-obtusa, sparse puberula; endocarpium in 
sectione transversali circulare, longitudinaliter sulcatum. 


Other specimens have been seen from Colombia 
(Caqueta), Brazil (Amazonas), Peru (Loreto), and 
Ecuador (Napo), and a specimen possibly of this 
species from French Guiana 

his species resembles B. oxycarpa (C. Martius) 
Eichler in leaf characters, but the fruit is pubescent 
and not beaked. The fruit resembles that of B. 
viridiflora Ducke, but the leaves have a much 
closer and more conspicuous vein reticulation, and 
are thinner in texture and usually broader. 


LITERATURE CITED 


DELENDICK, T. J. 1984. Bucida spinosa (Combreta- 
ceae) a new name published by Jennings in 1917. 
Bull. Due Bo t. Club 111: 375-376. 

EXELL, A. W. & C. A. STACE. revision of the 
genera Buc ie vu Ramatuella. Bull. Brit. Mus. 
(Nat. Hist.), Bot. 

WILBUR, R. L. 198 q a assorted extension of 
Article 72. Taxon 37: 464-467 


SOLANUM ALLOPHYLLUM 
(MIERS) STANDL. AND THE 
GENERIC DELIMITATION 
OF CYPHOMANDRA AND 
SOLANUM (SOLANACEAE) 


Lynn Bohs? 


ABSTRACT 


Solanum emo prada has inita been placed in Cyphomandra and in Solanum. This species has a number 
not found in 


thickening. When eac h 
taxa, indicating that the feature 
crossing stu and cytological investigations also supports the 


Solanum pacos aS is d R ible, where 
incompatible, and the chro 


n Cyphomandra, but has 
s they share may not be structurally homologous. New 


omes of Cyphomandra are about 2 


included in the genus because it has a similar 


n of S. allophyllum from Cyphomandra. 


as all but the single domestieatea species of Cyphomandra are self- 


o 5.5 times larger than those of S. a 


llophyllum. 


‘he cae morphology, self. aa and small pa are all consistent with placement of this species 


in Solanun 


INTRODUCTION AND TAXONOMIC HISTORY 


The generic placement of the species here known 
as Solanum allophyllum (Miers) Standl. has long 
been a source of confusion. Various workers have 
placed this species in Solanum, Cyphomandra, 
and Bassovia. It was first described as Pionandra 
allophylla by Miers (1854) in Seemann's Botany 
of the Voyage of the H.M.S. Herald. Pionandra, 
erected by Miers in 1845, is synonymous with 
Sendtner's genus Cyphomandra, created a few 
months earlier (Sendtner, 1845). Accordingly, P. 
allophylla was transferred to Cyphomandra by 
Hemsley in 1882. Standley transferred Cypho- 
mandra allophylla to Solanum in 1927 without 
explanation. 

Georg Bitter, unaware of Miers's species, in- 
dependently described the species as Solanum el- 
lipsoideibaccatum Bitt. in 1913. He noted that 
the tapered anthers with small terminal pores re- 
semble those of Solanum subg. Leptostemonum 
(Dun.) Bitt. but surmised that his species probably 
represented a new section of Solanum. In 1914, 


Bitter received specimens illustrating the lobed 
leaves that often occur on the lower branches in 
this species. He emended his original description 
to include these lobed leaves and described a new 
variety, var. ficilobum Bitt., 
hibiting almost exclusively lobed leaves. 


from a specimen ex- 
Pittier 
(1947) later transferred Solanum ellipsoideibac- 
catum to the genus Bassovia, an error that Hun- 
ziker (1969) later corrected. Solanum ellipsoi- 
deibaccatum and its variety ficilobum are regarded 
here as synonyms of the earlier name Solanum 
allophyllum (Miers) Standley. 

Standley's placement of the species in Solanum 
was followed in the older literature (Morton, 1944; 
1965; Standley, 1928), but 
more recent workers have interpreted it as be- 
longing in Cyphomandra (Bohs, 1986, 1988; Child, 
1984; D'Arcy, 1973). All those who have included 


the species within Cyphomandra have noted its 


Romero-Castaneda, 


atypical anther structure, but placed it in. Cy- 
phomandra on the basis of other similarities. Child 
(1984) called attention to its anomalous features 


! I thank Alan Child for Sending seeds of 5. allophyllum and several C yphomandra species and for his insightful 


discussions on d in 
their lab facilities, and Gre 

Sperry for ud and commenting on the rud ipt. 
? Work done at: Prin 
05405, U.S 


Solanum and Cyphomandra. | 
g Anderson, Dave Barrington, W. G. 


Dave Barrington, John Sperry, and Mel 


D'Arcy, George Rogers, and John 


also thank 


e Herbarium, Marsh Life Science Building, University of Vermont, Burlington, Vermont 
.A. Present qn Department of Biology, University of Utah, Salt Lake City, Utah 84112, U.S. 


ANN. Missouni Bor. Garb. 76: 1129-1140. 1989. 


1130 


Annals of the 
Missouri Botanical Garden 


when he erected Cyphomandra sect. Allophylla 
Child to accommodate this species and the allied 
Cyphomandra phytolaccoides (Rusby) Child (— 
Solanum mapiriense Bitter). 

Why has the generic placement of this species 
been so uncertain? Though $. allophyllum differs 
from Cyphomandra in a number of morphological 
characters, it has been considered as a Cypho- 
mandra because, at least superficially, this species 
shares with Cyphomandra the very features that 
have been used to distinguish Cyphomandra from 
Solanum. Both S. allophyllum and Cyphomandra 
have: 1) a growth habit characterized by a single 
erect trunk and a spreading crown of three main 
lateral branches, 2) three-leaved sympodial units 
with the inflorescences situated mainly in branch 
forks, and 3) tapered anthers that are thickened 
on the abaxial surface. Growth habit and branching 
pattern have been recently proposed as new char- 
acters that distinguish Cyphomandra from Sola- 
num (Bohs, 1986). Anther structure traditionally 
separates the two genera (Sendtner, 1845). Herein 
| reexamine the morphological features of Cy- 
phomandra and S. allophyllum with emphasis on 
these three proposed generic characters. New in- 
formation about the breeding system and mor- 
phology of the chromosomes, pollen, and seeds of 
5. allophyllum and Cyphomandra is also com- 
pared. These investigations help to ascertain the 
generic position of this species and shed new light 
upon the critical characters that separate Solanum 
and Cyphomandra. 


MATERIALS AND METHODS 


Morphological studies were carried out using 
living plants, and dried specimens from the follow- 
ing herbaria: A, BM, C, E, F, GH, K, M, MO, 
NY, P, US, and WIS. Voucher information for 
greenhouse material is given in the Appendix. 

Growth habit is defined by reference to the ar- 
chitectural models of Hallé et al. (1978). This 
system takes into account the dynamic aspects of 
plant growth rather than simply considering the 
shape of a plant at any one time. Terminology 
relating to architecture and branching pattern like- 
wise follows Hallé et al. (1978) 

Compatibility studies were conducted on plants 
growing in pollinator-free greenhouses at Harvard 
University and at the University of Vermont. One 
to several accessions were grown of each species. 
Plants were either selfed or outcrossed, between 
accessions and between individuals of the same 
accession, by tapping pollen onto a clean glass slide 


and rubbing it across the stigma of the female 
parent. 

For observations of meiotic chromosomes, flower 
buds were fixed in Farmer's solution (3: 1 absolute 
ethanol: glacial acetic acid) or Carnoy's fixative (6: 
3:1 absolute ethanol: chloroform:glacial acetic 
acid) shortly after sunrise. Preparations were stained 
with 1-2% acetocarmine and squashed in Hoyer's 
solution. Root tips for mitotic chromosome obser- 
vations were pretreated for up to 24 hours in a 
saturated solution of paradichlorobenzene at ap- 
proximately 4?C, then fixed in Farmer's solution 
for up to 24 hours. Root tips were then stored in 
70% ethanol until use, hydrolyzed in 1 N HCI for 
10 minutes at 60°C, and stained as above 

Histological preparations of anthers were ob- 

tained from flowers that had been fixed in FAA, 
dehydrated in an alcohol series, embedded in Para- 
plast, sectioned, and stained with safranin/ fast 
green. 
Pollen measurements were made from fresh pol- 
len samples obtained from greenhouse plants. The 
grains were shaken into a mixture of 1-2% ace- 
tocarmine and Hoyer's solution and allowed to stand 
for exactly 10 days before measurement. Mea- 
surements of the polar and equatorial axes on 30 
grains per sample were made from a 400 X camera 
lucida projection onto a Zeiss ZIDAS digitizer. Pol- 
len diameter was measured on stained grains only, 
and was taken as the distance between the inner- 
most layers of the pollen grain wall. Pollen volume 
was calculated using the formula for the volume 
of an ellipsoid, V = rPE*/6, where E is the equa- 
torial diameter and P is the polar diameter. An 
analysis of variance revealed no significant differ- 
ence between equatorial diameter as measured in 
polar and equatorial views, so P and E were mea- 
sured in equatorial view only. 

Pollen samples for scanning electron microscope 
SEM) photos were taken from dried herbarium 
material. The grains were mounted on the stub 
with double-stick tape and coated with gold-palla- 
dium. Fresh pollen from greenhouse plants that 
was critical-point dried and coated appeared col. 


lapsed under the SEM 


RESULTS 


DISTRIBUTION, ECOLOGY, AND MORPHOLOGY 


able 1 compares the morphological and eco- 
logical characteristics of 5. allophyllum and Cy- 
phomandra discussed below. 
The ecological distribution, phenology, and gen- 
eration time differ greatly in S. allophyllum and 


Volume 76, Number 4 
1989 


Bohs 
Solanum allophyllum 


TABLE l. Comparison of morphological and other characters in Cyphomandra and Solanum allophyllum (details 
in text). 
Cyphomandra Solanum allophyllum 
Habitat Mesic forest Seasonally dry forest 


Usually > 1 yr. 


Generation time 
i Up to 10 m 


Leaf bases Usually cordate or truncate, rarely de- 
current 
Numer of flowers per 10 
inflorescence 
Pedicel length > 10 mm 


Corolla shape Urceolate, campanulate, or stellate, not 
licat 


Fruit color Red, yellow, orange, purple, or green, 
never white 

Not laterally compressed 

Intermediate between Prevost's and 


Fruit shape 
Architectural model 


Nozeran's 
Sympodial units 3-4-leaved, hete s. leaved 
Anthers Tapered or not, with distinct and en- 


larged connective 


p to 1.5 m 
Subcordate to truncate, decurrent 


4-6 


4-6 mm 
Rotate-stellate and plicate 


White to orange 
Laterally compressed 
“Fragment” of Prevost's or Nozeran's 


models 


-leave 
Tapered, without enlarged connective 


Cyphomandra. Solanum allophyllum has been 
collected from a single site in Honduras and is 
more abundant from Costa Rica and Panama 
through northwestern South America (Fig. 1). 
though the plants are found in a variety of habitats, 
in Costa Rica and Panama the species seems to 
grow in drier sites than Cyphomandra species, 
which are almost always mesic-forest dwellers. No 
herbarium collections of this species from Costa 
ica or Panama were made in the months of Jan- 
h, the dry season in these regions (Coen, 


allophyllum persists by losing its leaves during the 
dry season and perennating via its roots or larger 
shoots, but further field studies of this species are 
needed to ascertain its phenology. In the green- 
house the plants flower and fruit within five months 
from planting. This generation time is much shorter 
than most cultivated cyphomandras, which often 
take several years to reach reproductive age. Sola- j. 
num allophyllum is a weakly woody shrub that 
rarely grows over 1.5 m tall (Fig. 2), in contrast 
to the majority of Cyphomandra species, whic 
develop abundant secondary xylem and may reach 
heights of up to 10 m. 

e subcordate to truncate leaf bases of 5. al- 
lophyllum are decurrent along the petiole (Fig. 3). 
The leaves of Cyphomandra usually have cordate 
or less frequently truncate bases; decurrent leaf 
bases occur only in C. fragilis Bohs. 

The inflorescences of S. allophyllum are un- 


species of Cyphomandra. In the 
presumably also in the field, the fruits fall to the 


branched and bear 4-6 flowers on short pedicels 
4-6 mm long. Those of Cyphomandra can be 
branched or unbranched, typically bear more than 
10 flowers, and have pedicels longer than 10 mm. 
e shape of the corolla of S. allophyllum can 
best be described as rotate-stellate (sensu Correll, 
1962): fairly long corolla lobes are present, but 
these are connected at the base by abundant in- 
terpetalar tissue that is plicate in the bud (Fig. 4 
This type of corolla is often seen in Solanum subg. 
Potatoe (G. Don) D'Arcy. In contrast, the corolla 
f Cyphomandra has various shapes, but never 
rotate-stellate or plicate. 
Solanum allophyllum produces some of the most 
distinctive fruits in the Solanaceae. At maturity, 
they are glabrous and white or occasionally orange 
mottled with green or purplish markings. The fruits 
are ellipsoidal or oblong in outline, and are laterally 
compressed and appear elliptic in cross section (Fig. 


fruits differ from those of all species of Cypho- 
mandra in color and shape, and the seeds are much 
smaller and more numerous than those of any 


ground while still hard and bitter and then ripen 
for several weeks before becoming soft and pal. 
atable and emitting a strong sweet odor. They are 
most likely dispersed by ground-dwelling animals. 
Those of Cyphomandra are held on the tree and 


1131 


greenhouse and 


[T ' 1 1T 1^ —1 
O 100 200 300 400 500 600 miles 


FIGURE 1. 


only fall after they are completely ripe. Their dis- 
persal agents are unknown, but they may be at- 
tractive to birds or bats. 


ARCHITECTURE 


The distinctive growth habit or shape of S. al- 
lophyllum and many species of Cyphomandra is 
a conspicuous feature that allows the plants to be 
recognized easily in the field. In terms of archi- 
tecture, however, Cyphomandra and S. allo- 
phyllum are similar only in the initial phase of 
their growth (Fig. 6). In both, the seedling axis 
produces a single orthotropic, or upright, trunk 
with the leaves spirally arranged in a 2/5 phyl- 
lotaxis. This trunk ends with the production of a 
terminal inflorescence. Usually three plagiotropic, 
or horizontal, shoots then elongate from axillary 
buds located just below the inflorescence to produce 
a spreading crown (Fig. 6A, D). Further branching 
within the crown occurs by sylleptic elongation of 
axillary shoots immediately below the successive 
terminal inflorescences; thus the crown is composed 
of a series of sympodial units where all the flowers 


1132 Annals of the 
Missouri Botanical Garden 
ag 
10 
O 200 400 600 800 1000km 


7 : 
M 
i rl s 
e m 
et SM euh 


Distribution of Solanum allophyllum. (Base map copyright 1979 by the University of Utrecht.) 


and fruits are borne. Beyond this, Cyphomandra 
and S. allophyllum diverge in their architecture. 

In Cyphomandra, after continued sympodial 

rowth and numerous episodes of flowering and 
fruiting, the plagiotropic branches begin to senesce. 
new trunk then arises proleptically from an ax- 
illary bud on the old trunk below the branch tier 
and continues orthotropic growth until another ter- 
minal inflorescence and branch tier is produced. 
In this way, the main axis is composed of successive 
sympodial trunk modules bearing spatially and tem- 
en separated reproductive crowns (Fig. : 
C). I have seen this architecture in at least six 
species of Cyphomandra, and it is probably wide- 
spread in the genus. [n contrast to Cyphomandra, 
5. allophyllum does not produce successive trun 
modules. The whole plant in 5. allophyllum is 
therefore equivalent to a single structural unit of 
the Cyphomandra growth form. 

The architecture of Cyphomandra described 
above occupies an intermediate position between 
Prevost's and Nozeran's models in the scheme of 
Hallé et al. (1978). Both models have an ortho- 
tropic sympodial trunk and tiered plagiotropic 


Volume 76, Number 4 
1989 


Bohs 1133 
Solanum allophyllum 


FIGURES 2-4. 
Scale bar = 2 cm. 


ywers. Scale bar = m 


branches. In Prevost's model both the trunk and 
plagiotropic branches generally have spirally ar- 
ranged leaves, whereas in Nozeran's model the 
orthotropic and plagiotropic branches have highly 
contrasting leaf arrangements, usually with spiral 
phyllotaxis on the trunk and distichous phyllotaxy 
on the branches. Plagiotropy of the branches in 
Nozeran's model is perpetuated if a crown branch 
is independently propagated, whereas in Prevost's 
model the plagiotropy of the crown branches is 
usually lost when they are separated from the trunk. 

he leaf arrangement of the crown branches in 
Cyphomandra is affected by pronounced twisting 


Solanum allophyllum. — 2. Greenhouse-grown plant. Scale bar — 0.25 
Fk cm. 


m. — 3. Leaves from trunk. 


of the axes and differential elongation of branch 
internodes (see section on branching pattern below 
for a more detailed description of leaf arrangement, 
and Danert, 1958, 1967, 
phology of the shoot systems in Solanaceae). There- 


for comparative mor- 


fore, although the leaf arrangement on the plagio- 
tropic shoots is not distichous, the crown branches 
show pronounced dorsiventral symmetry in con- 
trast to the radial symmetry of the spirally arranged 
trunk leaves. Cuttings taken from the crown region 
of C. betacea, C. diploconos, and C. diversifolia 
produce lower, bushier plants than those taken 
from the upright axes (Fletcher, 1979; pers. obs.), 


1134 


Annals of the 
Missouri Botanical Garden 


FIGURE 5. 


so these species apparently exhibit the inherent 
nature of plagiotropy in the crown branches char- 
acteristic of Nozeran's model. At least until an 
accurate interpretation of leaf arrangement on the 
plagiotropic shoots is available, it is probably best 
to consider Cyphomandra as being intermediate 
between Prevost's and Nozeran's model 

The architecture of S. allophyllum does not 
strictly conform to any of the models defined 
Hallé et al. (1978). It most closely resembles Leeu- 
wenberg's model, which is exhibited by several 
n. ies of Solanum and Capsicum (Hallé et al., 
1978). However, in Leeuwenberg's model, all the 
axes are orthotropic and equivalent, whereas in S. 
allophyllum there is evident differentiation be- 
tween the trunk and crown branches. Perhaps the 
yest way to characterize the architectural form of 
5. allophyllum is to consider that it may have been 
derived by "fragmentation" from more woody 
counterparts with multiple trunk modules, such as 
those seen in Cyphomandra. Fragmentation oc- 
curs when only a portion of the original tree model 
is expressed, and is commonly seen in herbaceous 
relatives of woody plants with more strongly de- 
veloped trunk modules. Should it indeed be the 
case that the architectural form of S. allophyllum 
is related in this way to more extensive tree models, 
it may argue that this relatively herbaceous species 
has been derived by reduction and fragmentation 
from woody ancestors. 


Fruits of Solanum allophyllum. Scale bar 


— 2 cm. 


BRANCHING PATTERN 


Three- to four-leaved sympodial units are the 
rule within the crown of nearly all species of Cy- 
phomandra (Fig. 7); the only known exception is 
C. corymbiflora, which frequently has five-leaved 
sympodia. Three-leaved sympodial units are also 
characteristic of S. allophyllum. Commonly, the 
leaf subtending the renewal shoot is carried up to 
a point nearly opposite that of the first leaf pro- 
duced on the axillary shoot so that the seemingly 
opposite leaves are actually members of successive 
sympodial shoot generations. In S. allophyllum 
and in many species of Cyphomandra, two renewal 
shoots grow out from below the terminal inflores- 
cence, thus situating it in a branch fork. This shoot 
structure was previously thought to be peculiar to 
Cyphomandra (Bohs, 1986), and its presence in 
5. allophyllum was evidence supporting the inclu- 
sion of the species within Cyphomandra. Members 
of Solanum may have one- to many-leaved sym- 
podial units, and the inflorescences may be axillary, 
extraaxillary, opposite a leaf or leaf cluster, or 
rarely in branch forks. 

Subtle differences exist in leaf and shoot ar- 
rangement within the crown branches of Cypho- 
mandra and S. allophyllum apart from the leaves 
and branches making up the sympodial shoot struc- 
ture. In S. allophyllum, nearly all the axillary buds 
of the crown expand into short shoots, giving the 


Volume 76, Number 4 
8 


Bohs 1135 
Solanum allophyllum 


ES 
- 


C 


FIGURE 6. Architecture of d and Sola- 

num lb —A, B, ( | stages 

in Cyphomandra, showing the sy mpodial nature of the 

ers and the f the crown branch- 

. Solanum allophyllum due not develop beyond stage 

. Top v of crown, showing three main plagi- 
otropic i he. 


crown a dense leafy appearance. In Cyphoman- 
dra, sylleptic growth is restricted to the axillary 
buds immediately subtending the terminal inflores- 
cences of the sympodial units, the rest of the buds 
remaining dormant unless released by pruning or 
breaking the tip of the branch. In the greenhouse, 
S. allophyllum also undergoes pronounced sea- 
sonal reiteration, with expansion of many buds on 
the trunk by prolepsis and elongation of the short 
shoots of the crown. It is not known whether 5. 
allophyllum reiterates in this way under natural 
conditions; as already mentioned, it may die back 
to the main stem each year and produce a new 
plant conforming to the initial model at the begin- 
ning of the rainy season. 


ANTHER MORPHOLOGY 


Tapered anthers with an abaxial thickening oc- 
cur in Cyphomandra and S. allophyllum (Figs. 
8, 9). Anther shape is not a definitive characteristic 
of either genus, however; tapered anthers occur in 
other species of Solanum (e.g., in subg. Lepto- 
stemonum (Dun.) Bitt., subg. Potatoe (6. Don) 
D'Arcy, and sect. Herposolanum Bitt.), and al- 
though they are common in Cyphomandra, they 
are not exhibited in all of its species. There is no 
present evidence to indicate that tapered anthers 
are homologous within or among these groups. A 
more important criterion for distinguishing Cy- 
phomandra from Solanum is found in the enlarged 
anther connective. In Cyphomandra, the anther 
connective is usually very thickened and prominent 
abaxially and is sharply delimited from the thin- 


Diagram of branching pattern in the crown 


FiGURE 7. 
of Cyphomandra and Solanum allophyllum. Black and 


— 


white shading denotes successive sympodial units. The 
subtending leaf (S) of the axillary shoot has been carried 
up to a level subopposite the first leaf on this shoot (L,). 
The diagram shows a species with four-leaved sympodia; 
in three-leaved sympodia, the leaf L, is absent 


— 


walled anther thecae. Anthers of Solanum can be 
thickened in various ways, but they never have a 
distinct and abaxially prominent connective. 
Solanum allophyllum has a thickened area on 
the abaxial side of the anther that has been inter- 
preted as an enlarged connective like that of Cy- 
phomandra, and has led to the inclusion of S. 
allophyllum within Cyphomandra. Microscopic 
cross sections through the anther region o 
lophyllum and a representative of C mai 
C. diversifolia (Dun.) Bitt., show that anther struc- 
ture is very different in the two taxa. In Cypho- 
mandra (Fig. 8), the thickened portion is expanded 


TABLE 2. Compatibility studies in Cyphomandra and 
Solanum. 
Selfed Outcrossed 
- % Suc 
cessful cessful 
Taxon crosses crosses 
C. betacea 20 55% 36 30% 
C. diversifolia 77 0% 152 60% 
C. diploconos 108 0% 117 bie Ke 
C. hartwegii 38 0% 19 32% 
C. uniloba 33 0% 11 55% 
C. acuminata 60 0% 
C. corymbiflora 32 0% 30 93% 
Solanum 
allophyllum 21 18% 52 31% 


1136 


Annals of the 
Missouri Botanical Garden 


"^9 


FIGURES 8, 9. 
Scale bars = 0.5 1 


abaxially, the thecal walls are free and do not 
contribute to the thickening, and there is a sharp 
demarcation between the swollen connective and 
the thin-walled anther thecae. In contrast, the 
thickened area in S. allophyllum is not expanded 
abaxially and is contiguous with the walls of the 
anther thecae (Fig. 
than abrupt transition for the thickened area on 


There is a gradual rather 


the abaxial surface to the thin-walled anther thecae. 
This type of anther structure conforms to that of 
many other species of Solanum. These differences 
are evident in transverse sections through fully 


UNES sections through anthers. —8. Cyphomandra diversifolia. —9. Solanum allophyllum. 


mature anthers; similar studies on developing stages 
in these species may further emphasize these dif- 
ferences and may also reveal the derivation of the 
tissue involved in the anther thickening in both 
taxa. 


COMPATIBILITY 


Table 2 
linations in the greenhouse of flowers of S. allo- 
phyllum and various species of Cyphomandra. 
The results show that of the seven species of Cy- 


illustrates the results of controlled pol- 


Volume 76, Number 4 
1989 


Bohs 1137 
Solanum allophyllum 


oa ^ 

FIGURE 10. Camera lucida Nr of meiotic chro- 
mosomes a microsporangia. —A. Cyphomandra divers- 
ifolia. —B. Solanum allophyllum. A and B at same mag- 
Action 


phomandra tested, all are self-incompatible with 
the exception of the cultivated tree tomato, C. 
betacea (Cav.) Sendtn. In contrast, S. allophy llum 
appears to be self-compatible. Neither S. allo- 
phyllum nor C. betacea sets fruits in pollinator- 
free greenhouses without deliberate pollination. This 
indicates that these two species may not be autog- 
amous and require a pollinator to transmit self 
pollen, or that they may be apomictic and pseu- 
dogamous. The style in S. allophyllum is not visible 
directly after anthesis, but elongates and emerges 
through the anther cone after the anthers dehisce. 
Thus, protandry may prevent selfing in this species. 


CYTOLOCY 


The most compelling evidence for the exclusion 
of S. allophyllum from Cyphomandra comes from 
chromosome studies. Solanum allophyllum and 
nine species of Cyphomandra that have been ex- 
amined so far chromosomes (Bohs, 
unpublished data; Pringle & Murray, in press). This 


chromosome number is common in Solanum and 


have n — 


A. 


————— 
10^ 


FIGURE 11. Camera lucida drawings of mitotic ER in root tips. — A. C yphomandra acuminata. — 


Solanum allophyllum. A and B at same magnificatior 


the Solanaceae in general. However, chromosomes 
of the two taxa differ markedly in size (Figs. 10, 
11). Investigations of seven species of Cypho- 
mandra have revealed that the chromosomes in 
this genus are very large, averaging about 8 um 
in length with a range of approximately 3 to 14 
um (Bohs, unpublished data). DNA amounts as 
measured by flow cytometry were among the larg- 
est yet known in the Solanaceae (Pringle & Mur- 
ray, in press). In contrast, both the mitotic and 
meiotic chromosomes of 5. allophyllum range be- 
tween | and 2.5 um in length and are thus on the 
order of 2.5 to 5.5 times smaller than those of 
Cyphomandra. 


POLLEN 


Pollen grains of S. allophyllum and Cypho- 
mandra were examined for potential taxonomic 
characters. The grains of 5. allophyllum are tri- 
colporate and have exine sculpturing consisting of 
very small rounded granules best observed with 
SEM (Figs. 12-14). Cyphomandra pollen is also 
tricolporate, and most species have granular exine 
sculpturing like that of 5. allophyllum, although 
several Cyphomandra species (e.g., . pendula 
(R. & P.) Sendtn. and C. pilosa Bohs) lack such 
granules and have a psilate exine. 

Although the aperture type and exine sculptur- 
ing show few taxonomically useful differences, the 
grains of Cyphomandra and S. allophyllum do 
differ somewhat in size and shape. Table 3 com- 
pares the pollen dimensions and volume of S. al- 
lophyllum and seven species of Cyphomandra. In 
the terminology of Erdtman (1952, 1969), all 
species of Cyphomandra have spheroidal or pro- 
late spheroidal grains, whereas those of 5. allo- 
phyllum are more elliptic in equatorial view and 
fall into the subprolate shape class. All taxa ex- 


c 


1138 


Annals of the 
Missouri Botanical Garden 


FicunEs 12-14. 
14. Scale bar = 0.5 


amined have small- to medium-sized grains (again 
using Erdtman's terminology), but those of 5. al- 
lophyllum are much smaller than any species of 
Cyphomandra. Cyphomandra grains have vol- 
umes about 2.5 to more than 10 times larger than 


those of S. allophyllum. 


SEED SIZE 

The seeds of S. allophyllum are substantially 
smaller than those of any of the Cyphomandra 
species examined (Table 3), ranging from about Y, 
to Y, the size of Cyphomandra seeds. 


DISCUSSION 


There are numerous differences in ecology, phe- 
nology, and morphology between 5. allophyllum 
and Cyphomandra, but these alone do not permit 
an unequivocal placement of the species in either 
Solanum or Cyphomandra. Architecture and some 
aspects of the branching pattern further differen- 
tiate S. allophyllum from Cyphomandra, but again 
not enough to place the former definitively in either 
genus. A problem with the use of these characters 
is the wide range in architecture and branching 
pattern seen in Solanum. It looks as if these two 


Pollen grains of Solanum alloph yllum. 


13. Scale bar = 10 um. 


12. Scale bar = l um. 


characters cannot be as useful in distinguishing 
Cyphomandra and Solanum as was previously 
thought. 

In contrast, a consideration of anther morphol- 
ogy shows that S. allophyllum does not have a 
distinct and enlarged anther connective like that 
of Cyphomandra. The anther structure formerly 
interpreted as a similarity between it and Cypho- 
mandra is now revealed to be a difference that 
separates the two taxa. It appears that the presence 
of an enlarged anther connective is the most re- 
liable morphological criterion for distinguishing Cy- 
phomandra from Solanum. 

he fact that 5. allophyllum is self-compatible 
(SC), whereas the majority of Cyphomandra species 
are self-incompatible (SI), is at least an implication 
that 5. allophyllum is not a Cyphomandra. Bot 
SC and SI are known in Solanum, so the inclusion 
of 5. allophyllum in Solanum is consistent on the 
basis of this character. 

Self-compatibility in C. 
indication of its relatively recent origin. SC is con- 
sidered to be a derived character in the Solanaceae 
(de Nettancourt, 1977; Whalen & Anderson, 
1981). If this applies equally well to Cyphoman- 
dra, then the self-compatibility of the cultivated 


betacea could be an 


Volume 76, Number 4 
1989 


Bohs 1139 
Solanum allophyllum 


TABLE 3. 


Pollen size and seed weight in Cyphomandra and Solanum allophyllum. Thirty grains/taxon measured 


for pollen size. Three measurements of 10 seeds averaged for seed weight. Standard deviations given in parentheses 


after measurements. 


Pollen polar Pollen equatorial Seed weight 

iamete diameter Pollen volume (10 seeds) 

Taxon um) (um?) (g x 107?) 
C. diversifolia 33.66 (1.89) 31.92 (1.48) 18,068 (2,628) 3.97 (0.064) 
C. uniloba 31.28 (1.63) 30.58 (1.42) 15,414 (2,182) 3.44 (0.276) 
C. diploconos 29.24 (1.26) 27.73 (1.02) 12,043 (1,801) 1.84 M 

C. acuminata 29.16 (1.09) 26.87 (0.92) 11,057 (1,108) 

C. betacea 22.47 (1.21) 21.93 (0.76) 5,678 (637) 5.10 (0.127) 
C. hartwegii 21.61 (0.96) 20.35 (0.95) 4,706 (587) 24.44 (0.706) 
C. corymbiflora 21.45 (1.16) 19.97 (1.32) 4,520 (780) 4.03 (0.682) 
S. allophyllum 16.54 (0.75) 14.21 (0.75) 1,756 (232) 0.70 (0.025) 


C. betacea would be considered derived, perhaps 
in response to selection pressures relating to its 
domestication. 

e small chromosomes of 5. allophyllum con- 
form to the size reported for species of Solanum 
from several different subgenera (Roe, 1967) and 
are the best indication that this species belongs in 
Solanum. The present study shows that there is a 
tendency toward large chromosomes in Cypho- 
mandra. This appears to be a very useful character 
distinguishing Cyphomandra from Solanum. Fur- 
ther studies of chromosome size in the Solanaceae 
are needed to determine if this character is con- 
sistent in other species. Comparison of genome sizes 
between putatively primitive and derived members 
of s different plant groups demonstrates that 
chang an occur in either direction 
(Ob & Khoshoo, 1986). Additional chromosome 


studies coupled with morphological data may even- 


tually allow us to determine trends in the evolution 
of genome size in Cyphomandra, Solanum, and 
other solanaceous genera. 

The exine sculpturing of 5. allophyllum pollen 
also resembles that of Solanum. Anderson & Gen- 
sel (1976) and Edmonds (1984) reported similar 
granular exine sculpturing in Solanum sects. Ba- 
sarthrum (Bitt.) Bitt. and Solanum, respectively. 

he occurrence of the same type of exine sculp- 
turing in Cyphomandra, however, argues against 
the use of this character to distinguish Cypho- 
mandra from Solanum. This same type of sculp- 
turing also has been reported for other genera 
related to Solanum (Basak, 1967) and is probably 
the general pollen type of the tribe Solaneae 

The pollen grains and seeds of S. allophyllum 
are much smaller than those of any species of 
Cyphomandra. The pollen dimensions of 5. allo- 


phyllum are comparable to some of the smaller 


grains reported for Solanum (Anderson & Gensel, 
1976; Basak, 1967; Murry & Eshbaugh, 1971), 
and the size of the seeds falls within the range for 
Solanum (pers. obs.). Pollen or seed size has not 
been critically examined as a taxonomic character 
separating Cyphomandra and Solanum. 

small pollen and seeds of S. allophyllum 
may be connected with its small chromosome size. 
Bennett (1972) observed that genome size was 
correlated with pollen volume, seed weight, and 
minimum generation time in various species 
herbaceous plants. The association of small chro- 
mosome size with small pollen volume, seed weight, 
and short generation time in S. allophyllum may 
indicate that this correlation also holds in this 
species. Further observations are needed to ascer- 
tain whether these characteristics are correlated in 
other solanaceous species, and whether pollen and 
seed size may be used as indicators of chromosome 
size. 

Affinities of S. allophyllum within Solanum are 
problematic. This species is allied with two others, 
S. mapiriense Bitter from Bolivia and an unde- 
scribed species from Amazonian Peru (Bohs, in 
prep.). At present I know of no existing section of 
Solanum that accommodates these three species. 
They are perhaps best regarded as a new section 
of Solanum, as Bitter (1913) suggested. A taxo- 
nomic treatment of these three species and a con- 
sideration of their placement within Solanum will 
appear in a later paper. 


LITERATURE CITED 


ANDERSON, G. J. . G. GENSEL. 1976. Pollen mor- 
phology and is systematics of uiu sectio n Ba- 
sarthrum. Pollen & Spores 18: 533-552. 

Basak, R. K. 1967. The oni grains of Solanaceae. 
Bull. Bot. Soc. Bengal 21: 49-58. 


Annals of the 


1140 
Missouri Botanical Garden 
BENNETT, M. D. 1972. Nuclear DNA content and min- Ounr, D. & T. N. Knosuoo. 1986. Plant DNA: contents 
imum generation time in herbaceous plants. Proc. and systematics. 1-19 in S. K. Dutta (editor), 
oy. Soc. London, Ser. B, Biol. Sci. 181: 135 A Systematics, Volume II. Plants. CRC Press, 
Birrer, G. 1913. Boca Raton, Florida. 


Solana nova vel minus cognita. VII 
iai e Nov. Regni Veg. 11 4 

. Solana nova vel minus cognita. XV. 
Nov. Regni Veg. 13: 169-173 
The Won and d of Cy- 
Harvard 


Re eper rt. p 
Bons, L. 1986. 
phomandr (Solanaceae). em es thesis, 
ridge, Massachus 

‘he Colombian. species " d 
mandra. Revista Acad. Colomb. Ci. Exa 


— 


1984. Studies in Solanum L. (and related 
rovisional conspectus of the genus 
ex Sendtner. Feddes Repert 


On. x: 
genera) 3. 

D ah pe M. 
: 283- J 


TEN E 1983. Climate. Pp. 3 . H. Jar 
(editor), Lun Rican Natural Boo. a niv. Chicago 
Press, Chicago, Illinois 

CORRELL, D. S. 1962. The Potato and Its is ild ied 
tives. Texas Research Foundation, Renner, Tex 

Croat, T. B. 1978. Flora of Barro Colorado n 
Stanford Univ. Press, Stanford, Califor 

DANERT, S. 1958. Die Verzweigung der Salsa im 
Reproduktiven ood Pos Deutsch. Akad. Wiss. 
Berlin, d Chem. 6: 1 

— 67. Dic eae als Scil 
nn in a Gattung Solanum. Kultu 
pflanze 15: 275- 

iieri W.G. 1973. 

Ann. Missouri Bot. Gard. 60: 57 

Eomonts J. M. 19 Pollen KR EIN of Solanum 

ction Solanum. J. Linn. Soc., Bot. 88: 237 


292. 


Flora of Panama: Solanaceae. 
3 ). 


Pollen Morphology and Plant Tax- 
Almqvist € Wiskell, Stock- 


ED. G. 1952. 
onomy. Angiosperms. 
holm. 

. 1969. 

Copenhagen. 
FLETCHER, W. A. 1979. 

i Ministry of Agric 


Handbook of Palynology. Munksgaard, 


Growing tamarillos. New Zea- 


. & Fisheries Bull. 307: 1- 


m 


TN F., R. A. A. OLDEMAN & P. B. TOMLINSON. 1978 
Tropical Trees and Forests: an Architectural Anal- 
ysis. Springer-Verlag, Berlin. 

Las B. 1882. Biologia Centrali-Americana, 

ota Volume 2. R. H. Porter, London. 

tea a T. 1969, Estudios sobre Solanaceae. V. 
Contribución al conocimiento de Capsicum y generos 
afines (Witheringia, . denistus, e d t, etc.). Pri 
mera Parte. Kurtziana 5: 101-17 

Miers, J. 1845. Contributions to the (m of South 
America. Piomandra. London J. Bot. 4: 353-365. 

1854. Solanaceae. Pp. 172-176 in B. See- 
mann, The Botany of the Voyage of the H.M.S 
Herald. Lovell Reeve, London. 

Morton, C. V 9 Some South American m 
of odd Contr. U.S. Natl. Herb. 29: 41-72. 
Murry, L. E. & W. H. EsupnaUucH.. 1971. ^ prag 
ica al Aa of the Solaninae (Solanaceae). Grana 


HEMSLEY, 


NETUS OURT, D. pr. 1977. Incompatibility in Angio- 
sperms. Springer-Verlag, Berlin. 


Prrrier, H. 1947. mea do la Flora Venezolana, 
Volume 2. Mae a Carac 
PRINGLE, G. J. & . Murray. Karyotype diversity 
and nuclear one variation in Cyphomandra. Third 
eae Symposium on the Biology and System- 

f the Solanaceae (in press). 

Ror, K. ja 1967. rane size in Solanum and 
Cyphomandra: taxonomic -A oe impli- 
cations. Amer ralist | 295-29 

ROMERO-CAST AÑEDA, R 1965. “Flora del A” de Bo- 
livar, M volute 1. Universidad Nacional de Colombia, 


Bogo 
SENDTNER, o. 1845. De Cyphomandra, novo Solana- 
cearum genere tropicae Americae. Flora 28: 16 


Satan, , P. C. 1927. New rape from Central Amer- 
[ 16 


ash. Acad. Sci. 17: 
928. Flora of Hie cdi Canal Zone. So- 
lanaceae. Contr. U.S. . Herb. 27: 327-333. 
WHALEN, M. D. € G. J. ANDERSON. 1981. Distribution 


of gametophytic self. -incompatibility and vide s neric 
classification in Solanum. Taxon 30: : 


APPENDIX. Voucher information. 
md M ed (Miers) Standley 
eed from D'Arcy 9472, in Panama, Prov. 
Blas, in front of Ustupo. Vouchers: Bohs 2339 
T). 


collected 1 


UOTE 


C yphomandra acuminata Rusby 
Seed from Solomon & Escobar 12458, collected in 
Bolivia, Prov. Nor Yungas, 8.7 km below Chuspipata 
on road to Yolosa. Vouchers: Bohs 2338 (GH, V D. 


C LAE RR betacea (Cav.) Sendtn. 
by C. KieI in market, Quito, Ec- 
uador. e Bohs 2274 (GH), 2275 (VT). 


C yphomandra corymbiflora Sendtn. 
n southeastern Brazil, s 


sent by G. ys 
E New Zealand. Vouchers: 3 


gle, D.S.LR Bohs 2 
(GH, VT). 
C dos aaa d onos Mind. ) Sendtn 
d collec razil, Prov. Paraná, city of Curitiba. 
Due ‘Bohs 2335 GH. "b 


C Me diversifolia ( H. & B. ex Dunal) Bitter 
eed from Benitez de Rojas 2744, collected in Ven- 
AER Estado Aragua, Parque Nacional Henri Pittier. 
Vouchers: Bohs 2341 (GH, VT). 


Cyphomandra hartwegii (Miers) Sendtn. ex Walp. 
Seed collected in Colombia, Dept. Huila, Fundación 
Merenberg. Voucher: Bohs 1644 (GH). 


C V era ndo AP by 


eed from Sperli & King 5 »500, collected in Bolivia, 
P rov. Larecaja, between Consata and 
Mapiri. Vouc idiot Bohs 2283 (VT), 2284 (G 


NOVELTIES IN SOUTH 
AMERICAN MALOUETIA 
(APOCYNACEAE) 


Mary E. Endress? 


ABSTRACT 


Three new species of Malouetia (M. bubalina, M. molongo, and M. naias) and one new variety (M. pubescens 
var. pep are described. All new taxa are from northwestern South America. 


The following novelties have accumulated during 
studies of the genus Malouetia, for the most part 
in connection with the Flora of the Venezuelan 


Guayana. 


Malouetia bubalina M. E. Endress, sp. nov. 
Rio Caquetá, 
km en amont de Arara- 
cuara, Sabana de Mosco, 30 Dec. 1976, Sastre 
& Raichel 5062 (holotype, P). Figure 1. 


pi 


Amazonas: 


Arbor lactescens. Folia decussata, lamina sae "pues 
rupte cauda 


coriacea vel su 
illas domatiis conspicuis instructa, nervis lateralibus ca. 

utroque latere; petioli 3-5 mm longi. el 
terminales e fasciculis 12-20. floris compositae. Calyx ex 

tus puberulus. Corolla flavida, hypocraterimorpha, tubo 
6-6.5 mm longo; lobis ovatis acutis mm longis, 
ya 


> 
ta 
| 
mat) 
on 
d 
= 
5 
E 
N 
Uu 
E 
> 
H 
I 
PE 
(= 
c 
OR 
O 
"- 
A 
O 
a 
c 
c 
c 
5 
E 
(E 
£ 
Ss 28 
a 
em] 
p 


faucis tubularis insertae sessiles fere 12 exsertae angie 
lanceolatae dorso pilosae; ovarium dense pubescens. Fol- 
liculi 2 plus minusve crassi fusiformes 8-10 cm i longi, l 
cm diametro versus medium fer 180° usque divergentes; 
semina 2 cm longa l neari iid pilis longissimis retrorsis 
comatum basi obtu 

edium to large trees up to 30 m tall with 
abundant white latex; branches rather slender, an- 
gled and flattened at the nodes, with a glossy black- 
ish sheen in the younger parts, becoming gray and 
conspicuously lenticellate with age. Leaves decus- 
sate, 8-9.5 cm long, 4-4.5 cm wide, the blades 
coriaceous to subcoriaceous, broadly elliptic to ob- 
ovate-elliptic, shortly and bluntly acuminate at the 
apex, narrowly cuneate to attenuate at the base, 
the margins revolute, glabrous on both surfaces, 
the upper surface nitidulous, drying dark brown, 


the lower surface opaque, paler, with conspicuous 
domatia in the axils of the midvein; secondary veins 
ca. 6 on each side, arcuate-ascending, anastomos- 
ing before the margin, impressed on the upper 
surface, prominulent and darkened on the lower 
surface; tertiary veins inconspicuous on both sur- 
faces; petioles 3-5 mm long, canaliculate, gla- 
brous. Inflorescences 12-20-flowered, in terminal 
clusters; peduncle 1-2 mm long, glabrous, black- 
ened, glossy; bracteoles ca. 1 mm long, broadly 
ovate, acute, coriaceous, subglabrous to sparsely 
puberulent outside, glabrous inside. Flowers pedi- 
cellate, pedicels 4-6 mm long, glabrous; calyx 2 
mm diam., puberulent on the outer surface, gla- 
brous on the inner surface with 5 small, flattened 
glands near the base alternating with the lobes, the 
lobes ca. 1.8 mm long, ovate, tapering to a rather 
blunt apex, the margins minutely ciliolate; corolla 
salverform, yellowish, glabrous on both surfaces, 
the tube proper 4.5-5 mm long, 1-1.3 mm diam., 
cylindrical, the upper cupular part 1.5 mm long, 

iam., the orifice constricted by 5 rudi- 
mentary coronal scales, emphasizing the 5 nectar 
channels, the lobes 6-7 mm long, 2.5-3.5 mm 
wide, ovate, acute, densely puberulent on both 
surfaces; anthers inserted at the base of the cupular 
part of the corolla tube, about 12 exserted, sessile, 
narrowly lanceolate, sagittate, 2 mm long, dorsally 
barbellate in the upper half, the side wings glabrous, 
acuminate, incurving at the base; ovary ovoid, 
apocarpous, bilocular, ca. 0.5 mm high, densely 
pubescent, surrounded at the base by a glabrous, 
regularly 5-lobed disc nectary. Fruit apocarpous, 
glabrous, brown-black, composed of 2 stout, thick- 
walled, woody, fusiform follicles, 8-10 cm long, 1 
cm diam. at the broadest point, diverging at ca. 


! | thank Rupert Figur and Peter Endress for help with the Latin descriptions, and the curators of the herbaria 


that specimens for stu 
? [ns 


itut für Sy lo Botanik, Zollikerstrasse 107 


, 8008 Zürich, Switzerland. 


ANN. Missouni Bor. GARD. 76: 1141-1147. 1989. 


Annals of the 


1142 


Missouri Botanical Garden 


JRE 1. 


Fit 


Malouetia bubalina.— A. Habit. — B. Flower. — C. Anther, dorsal view. —D. Anther, ventral view. — 


E. Embryo. — F. Fruit. —C. Seed. —H. Embryo. Based on the holotype. 


Volume 76, Number 4 
1989 


Endress 1143 


South American Malouetia 


180? then curving downward; seeds 2 cm long, 
slender, fusiform, laterally flattened, slightly con- 
vex on one side, the ventral surface longitudinally 
sulcate, the apical end (i.e., toward the fruit apex) 
abruptly converging into a rounded tip 1 mm long, 
proximally tapering to a rather blunt end, the apical 
tip embedded in a dense mat of long tan hairs, 
these attached to the apical half of the seed and 
directed toward the base of the follicle. Beneath 
the hairs the testa very thin, stiffly papery, smooth, 
glossy, yellowish tan, longitudinally striate; the en- 
dosperm forming a leathery case around the em- 
bryo; embryo 1.8 cm long, straight, the cotyledons 
rather fleshly, ca. 314 times as long as the radicle, 
this thick, bent slightly. 


Distribution. Colombia, known only from high 
forest over white sand at the type locality on the 
Rio Caquetá. 


The specific epithet refers to the bifid, stout, 
curving fruit reminiscent of buffalo horns. 

Malouetia bubalina is most closely related to 
M. arborea (Vell.) Miers of eastern coastal Brazil, 
which also has stout fruits and seeds with an in- 
dumentum. But whereas the hairs on the seeds of 
M. arborea (Vell.) Miers are woolly and extend in 
all directions, those of M. bubalina are straight 
and directed toward the base of the follicle. The 
new species also differs by having much shorter 
pedicels and more ovate corolla lobes. 


2. Malouetia molongo M. E. Endress, sp. nov. 
TYPE: Venezuela. T.F. Amazonas: Santa Cruz, 
margen del Rio Atabapo, cerca del boca del 
Rio Atacavi, Foldats 3653 (holotype, NY). 
Figure 

Frutex vel arbuscula lactescens usque 2 m alta. Folia 
4.5-7.5 cm longa, 1.5-2.5 


utroque weak. petioli 2-5 mm longi. 
InPordsoentiae. hosce di axillares e fasciculis 2-3- 
; caly min longus extus glaber; corolla 
xh h ypocraterimorpha t tubo 15.5-17 mm longo, lobis 
ovatis obtusis mm longis n Pu mm P extus glabris 


u 
glabrum; ovula in utroque carpello ca. 12 3-4-seriatim 


affixa. Fructus ignoti. 


Shrubs or small trees up to 2 m tall with white 
latex; branches blackish, glossy and angled in the 
younger parts, becoming terete and grayish with 
age. Leaves opposite, 4.5-7.5 cm long, 1.5-2.5 
cm wide, the blades coriaceous, obovate to elliptic, 
obtuse to rounded at the apex, cuneate at the base, 
the margins revolute, glabrous on both surfaces, 


opaque, slightly paler on the lower surface, domatia 
absent; secondary veins 5-7 on each side, arcuate- 
ascending, anastomosing before the margin, ob- 
scure on both surfaces; tertiary venation inconspic- 
uous on both surfaces; petioles 2-5 mm long, gla- 
brous. Inflorescences 2—3-flowered, in terminal and 
axillary clusters; peduncle 1-2 mm long, glabrous; 
bracteoles broadly triangular, glabrous, ca. 1 mm 
long. Flowers pedicellate, pedicels 5-10 mm long, 
glabrous; calyx 2 mm long, 2-2.5 mm diam., gla- 
brous without, glabrous within with pale, rather 
irregularly shaped toothlike glands alternating with 
the lobes, the lobes 1.5-2 mm long, glabrous, fleshy; 
corolla salverform, white, the tube 13-14 mm long, 
the upper cupular part 2.5-3 mm diam., glabrous 
without, glabrous within except for a short region 
of pilose hairs directly below the anthers and in 
the cupular part near the orifice, the orifice con- 
stricted by 5 weltlike rudimentary coronal scales, 
the lobes 8-9 mm long, 4-5 
obtuse at the apex, the outer surface glabrous, the 
inner surface papillose-puberulent and becoming 
more densely puberulent at the base; anthers in- 


mm wide, ovate, 


serted somewhat above midway in the corolla tube, 
completely included, 3 mm long, oblong, apiculate 
at the apex, the dorsal side glabrous, the side wings 
straight, the bases short and blunt; style-head cy- 
lindrical; ovary ovoid, apocarpous, bilocular, ca. 
0.8 mm high, glabrous, surrounded at the base by 
a low, glabrous, 5-lobed disc nectary, the lobes 
truncate at the apex, basally connate for more than 
V4 their length; ovules ca. 12 per carpel, arranged 
in 3-4 rows. Fruit unknown. 


Par VENEZUELA. TERRITORIO | FEDERAL 
XA cae Cruz, margen del Rio Atabapo, cerca 
del boca del Rio Atacavi, Foldats 36. 54 (NY, VEN). 


Distribution. Known only from the type lo- 
cality in Amazonian Venezuela near the confluence 
of the Rio Atacavi with the Rio Atabapo, on flooded 


river banks. 


The name comes from molongó a frequent local 
name for various Apocynaceae of inundated hab- 
itats. 

This species most closely resembles Malouetia 
gracilis (Benth.) A. DC. from Guyana but differs 
by having five cleft weltlike scales constricting the 
mouth of the corolla tube. In M. gracilis (Benth.) 
A. DC. coronal scales are lacking. 


3. Malouetia naias M. E. Endress, sp. nov. 
TYPE: Colombia. Vaupés: Mitü and vicinity, 
along Rio Vaupés between Hio Yi and Rio 
Kubiyü, inundated margin of river, 9 July 


1144 


Annals of the 
Missouri Botanical Garden 


FIGURE 2. Malouetia molongo.— A. Habit.- 
—— showing coronal scales and positions of anthers. 
. Based on the holotype. 


1976, Zarucchi et al. 1825 (holotype, US; 
isotypes, COL (n.v.), GH (n.v.), HBG, MO, 
USF). Figure 3. 


Frutex vel arbuscula lactescens. Folia lamina lanceolata , 


apice caudata acumine 1-2 cm longa basi rotundata vel 


B. Flow 


D. a E. A 


E F 


-C. Longitudinal section through upper portion of 
-F. A 


nther, dorsal v nther, ventral 


cuneata, utrinque glaberrima, in axillis nervi mediani in- 


terdum domatiis instructa, 5-10 cm longa, 1-3 cm lata 
petioli 1-3 mm ange Due i gos illares et 
(sub?)terminales e fasc s (1-)2-4-floris compositae; 


calyx profunde lobatus intus e glaber, lobis anguste 
bs olatis acuminatis; corolla viridi-alba vel flavida hy- 
pocraterimorpha, tubo 4-7 mm longo, 0.7-1.1 mm dia- 


Volume 76, Number 4 Endress 1145 
1989 South American Malouetia 


FICURE 3. Malouetia naias. — A. Habit. — B. Flower. — C. Longitudinal section through flower showing positions 
Fete, 


of anthers. — D. Anther, dorsal view. — E. Anther, ventral view. —F. Fruit. —G. Seed. Based on the holotype. 


1146 


Annals of the 
Missouri Botanical Garden 


metro Past inflato 1.2-2 mm diametro, lobis lanceolatis 


1, 3 mm diametro; 
semina lineari- S abge indumento dpa d denso 


c 


et pilis dispersis 1-1.5 mm longis obtecta 


Shrubs or small trees 2-4(-8) m tall with sticky 
white latex; branchlets slender, glabrous, glossy 
reddish brown and flattened toward the nodes in 
younger dium. becoming gray and lenticellate 
with age. Leaves opposite, the blades thinly to 
firmly beside 5-)8-10 cm long, (1-)1.5-3 
cm wide, lanceolate, the apex with acumen (1-) 
1.5-2 cm long, rounded to cuneate at the base, 
completely glabrous on both surfaces, the upper 
surface often drying matte dark olive-brown to 
gray-green, the lower surface paler, with or without 
domatia in some of the axils of secondary veins 
with the midvein; secondary veins 8-9(- on 
each side, arcuate-ascending, anastomosing before 
the margin, impressed on the upper surface, prom- 
inulous on the lower surface; petioles 1 -3 mm long, 
glabrous, reddish brown, with reddish brown pec- 
tinate glands at the base on each side. Inflores- 
cences axillary or subterminal clusters of (1-)2-4 
flowers; peduncle 1-2 mm long, glabrous, reddish 

rown; bracteoles ca. 1 mm long, narrowly ovate- 
triangular, glabrous, ciliolate at the margin. Flow- 
ers pedicellate, pedicels (1.3-)2-2.5 cm long, gra- 
cile to filiform, glabrous; calyx split almost to the 
base, glabrous on both surfaces, sometimes mi- 
nutely puberulent near the apex on the inside, with 
5 small flattened glands on the inside, the lobes 
narrowly lanceolate, acuminate, somewhat spread- 
ing, (1.5-)2-2.5 mm long, 0.3-0.8 mm wide, the 
margins densely and minutely ciliolate; corolla sal- 
verform, greenish white to cream or pale yellowish, 
the tube inflated at the base, the mouth occluded 
by 5 rudimentary coronal scales, the upper cupular 
part 1-1.5 mm long, 1-1.5 mm diam., the tube 
proper (3-)4.5-5.5 mm long, 0.7-1 mm diam. at 
the orifice, 1.2-2 mm diam. at the base, glabrous 
without, sparsely pilose within just below the level 
of stamen insertion, otherwise glabrous; lobes lan- 
ceolate (8-)10-15(-23) mm long, (1.8-)2-3.5 
(-5) mm wide, glabrous on the lower surface, gla- 
brous on the upper surface except near the base, 
this granular-pubescent to villosulous; stamens in- 
serted near the orifice of the corolla tube, only the 
very tips exserted, narrowly lanceolate, 2.5-2.8 
mm long, dorsally sparsely pilose, the side wings 
glabrous, acuminate and curving inward at the 
base; style-head cylindrical, 1-1.2 mm long, style 


filiform; ovary ovoid, apocarpous, bilocular, 1 mm 
high, densely puberulent, disc nectary glabrous, 
ca. half as long as the ovary, 5-lobed, the lobes 
connate for ca. % their length; ovules 8-10 per 
carpel, arranged biserially. Fruit apocarpous, gla- 
brous, longitudinally finely striate, medium brown, 
composed of 2 slender, terete follicles 25-28 cm 
long, ca. 3 mm diam.; seeds linear-cylindric, 3 cm 
long, 1 mm diam., the ventral face longitudinally 
sulcate, the entire surface covered by a dense in- 
dument of microscopic hairs interspersed with a 
sparse scattering of thin, whitish to pale gold pilose 
hairs 1-1.5 mm long. 


CDU iS specimens examined. COLOMBIA. VAUPES: 
ú and vicinity, Zarucchi et al. 1810 (COL, GH, US 

USE) 1847 (COL, GH, HBG, MO, US , USF). VENEZUELA. 
AMAZONAS: Cano de Cholo, 15 km NE of San Carlos de 
Rio Negro, 4 km SW of Solano, Lie 2 4059 (MO, Z), 
8560 (MO, VEN, WAG), 8879 WAC), Gentry et 
al. s (HBG). PERU. LORETO: peda Mishana, Río 
, Estación Biológia Callicebus, Vásquez & Jun; 

millo 3280 (MO, Z), 6108 (MO, Z). 


Distribution. Colombia, Amazonian Venezue- 
la, and Peru, in primary, secondary, and seasonally 


inundated lowland rainforest. 


The specific epithet refers to the resemblance 
to the delicate water nymph. 

This new species is most closely related to Ma- 
louetia flavescens (Willd. in R. & S.) Múll.-Arg. 
and M. nitida Spruce ex Müll. Arg. However, M. 
naias is more delicate in all aspects than either of 
the above species. The following key is provided 
as an aid to identification. 


KEY TO THE MALOUETIA FLAVESCENS 

SPECIES GROUP 

la. Leaves 9-18 cm long, 3-7 cm wide; corolla 
ube 6-10 cm long; seeds 3 mm diam. oo 
2a. Anthers exserted 1. flavescens 
2b. Anthers included s Í M. nitida 

lb. Leaves 8-10 cm long, 1.5-3 cm wide; corolla 
tube 5.5-7.5 cm long; seeds 1 mm diam. ..... 

M. naias 


4. Malouetia pubescens Markgraf var. glabra 
M. ndress, var. nov. TYPE: Venezuela. 
Apul: Rómulo Gallegos, Río Capanaparo, near 
Hato Carabali, Davidse & Gonzales 16028 
(holotype, MO; isotypes, NY, VEN). 


A v dni lamina foliorum subtus glabra et 
petiolis libe differt 


Small trees up to 12 m tall with white latex; 
branches glabrous with a glossy blackish varnish. 
Leaves decussate, 8-15 cm long, 1.8-6.5 cm wide, 
the blades membranous, oblong-elliptic, the apex 


Volume 76, Number 4 
1989 


Endress 1147 
South American Malouetia 


acuminate, cuneate to obtuse at the base, glabrous 
and drying intense green on both surfaces; domatia 
usually present in some of the axils of the secondary 
veins with midvein; secondary veins 6-9 on each 
side, slightly impressed on the upper surface, prom- 
inulous on the lower surface; tertiary venation rath- 
er obscure on the upper surface, distinctly retic- 
ulate on the lower surface; petioles 2-4 mm long, 
glabrous. Inflorescences (10-)20-30-flowered, in 
terminal and axillary clusters; peduncle 1-2 mm 
long, glabrous. Flowers pedicellate, pedicels 2.5— 

m long, glabrous; calyx 1-2 mm long, 1-2 
mm diam., minutely puberulent on the outer sur- 
face, the inner surface with 5 small glands near 
the base of the lobes, the lobes 0.5-1 mm long, 
ovate, acute, the margins minutely ciliolate; corolla 
salverform, white to yellowish white, the tube 3- 
3.5 mm long, 0.5-0.8 mm diam., enlarged at the 
base, constricted at the orifice by 5 bilobed rudi- 
mentary coronal scales, glabrous on both surfaces, 
the lobes 4-5.5 mm long, 1.2-2 mm wide, oblong- 
ovate, acute, spreading or slightly reflexed, mi- 
nutely puberulent on both surfaces; anthers in- 
serted near the orifice of the corolla tube, almost 
completely exserted, 1.5 mm long, sagittate, the 
dorsal side barbellate, the side wings glabrous, basal 
appendages fused to the connective, slightly in- 


curved, the ends blunt; style-head conical; ovary 
apocarpous, bilocular, 1 mm high, ovoid, sparsely 
puberulent at the apex, surrounded at the base by 
a 5-lobed disc nectary; ovules 10-12 in each car- 
pel, arranged biserially. Fruit apocarpous, gla- 
brous, dark brown, longitudinally striate, composed 
of 2 slender, thinly woody follicles, 1 1 - 19 cm long, 
4—4.5 mm diam.; seeds linear-fusiform, com- 
pressed, plicate, the ends acuminate, tapering di- 
agonally, 2 cm long, 2.5 mm diam., with a dense 
indument of pale tan to ochre lanate hairs ca. 1.5 
cm long; embryo straight, 1 cm long, the cotyledons 
about 4 times as long as the radicle. 


Additional specimens examined. —— VI- 
CHADA: Parque Nacional Natural *El Tuparro," ca. 7 km 

NE of El Tapón, Zarucchi & Barbosa Jf o (EMB, MO, 
Z), 3766 (FMB, MO, Z). VENEZUELA. BO : Río Par- 
guaza, near Pilon, Wurdack & Monac hino 41116 (F, 
NY, S, US). 


Distribution. Apure and Bolivar, Venezuela, 


and Vichada, Colombia, in gallery forest. 


The new variety differs from var. pubescens in 
having leaves completely glabrous on both surfaces. 

e two varieties, which occur sympatrically in 
Vichada, Colombia, show no habitat differences. 


TICODENDRON: 
A NEW TREE FROM 
CENTRAL AMERICA 


Jorge Gómez-Laurito' and Luis D. Gómez P.? 


ABSTRACT 


Ticodendron incognitum is named as a new genus and species, and the taxonomic position of this tree is discussed. 


Despite many years of floristic research in var- 
ious Central American countries, new findings still 
surprise botanists. Such is the case of a tree, here 
described as a new genus and species, which has 
puzzled many scientists in the field and in herbaria 
and which has produced much correspondence and 
discussion. Based on studies of embryology, paly- 
nology, wood anatomy, and other features, a new 
family of dicots will be published soon for this genus. 


Ticodendron incognitum Gómez-Laurito & 
Gómez P., gen. et sp. nov. TYPE: Costa Rica. 
Alajuela: Reserva Forestal de San Ramón, ca. 
Colonia Palmarena, 900—1,200 m, 10?10'N, 

4?35'W, 30 Mar. 1987, male and female 
tree, G. Herrera, I. Chacón & A. Solís 515 
(holotype, CR; isotypes, F, K, MO). Figures 


Arbor dioicus vel poly gamo: -dioicus, 7-20 m altus, 40- 
80 em diametro ad pectoris. Germir T apom 
Cortex exfolians; interna facies nonnulla aliquanda caeru- 
le Wig alee e Lignum spadiceum, DE a- 
mulis teretis, 3-5 mm crassis, nodosis, sulcatis, griseis, 
1 complanatis, Jo pilosis, pilibus uni- 
cellulatis simplicibus vel raro 


caducae. Petiolis 10-15 mm longis, supra sulcatis, juvenis 
hirsutis (vide supra), demum lenticella hs Foliae alternae, 
lamina elliptico- ovata, 8-13 cm longa, 4-8 cm p sub- 
argine in parte superiore serrata, base cuneata, 
apicem acuminata supra glaberrima, infra ee 
juvenis vide "dre pilibus longioris, rigidis, acutis, 
nervis secundar 3 paribus, regulariter arquatis, sub- 
à eue paralelis, D elevatis, supra canaliculatis, ce- 
n sub > reticulatis, flavislucem transmissa. 
Amenta’ 4 cm longa, 
minea coronatis. 
erosae, 2-4 


=. 
E 
25 


) 
verticillatae dispositae; (verticillum 8-12 mm diametrum) 


latae ovatae vel deltoidea, 5 mm longae, 5 


, mox decidua 


bracteae 3, 
mm latae basoe 


E 'ae pn natae. longitudinaliter dehiscentibus, connec- 

subulato producto. Granula pollinis ca. 2 um crassa, 
ae leviter. Flos feminea solitariae extremitatis 
surculorum dispositae. Pedicelis 3 mm dm a adpreso- 
pilosis. "Bracteae involucratae" 3, ovatae, 2.5-3 mm 
jd adpreso-pilosae, mox deciduae. Perianthium tubus 


l S. 
vula 4 (ut videtur abortu 1), hemitropa. Stigmata 2, 
pilosa, 5-12 mm longa. Fructus asym unilater 


laginosis; in siccitatem on av ellaneum, lignossimum, 
longistrorsum sulcatus. 


Dioecious or polygamodioecious trees, 7-20 m 
tall, 40-80 cm dbh. Germination hypogeal. Inner 
bark reddish or purplish. Wood yellowish, very 
hard. Twigs 3-5 mm thick, flattened toward the 
tips, pilose when young. Stipules subulate, to 15 
mm long, caducous. Petioles 10-15 mm long. 
Leaves alternate, blades elliptic-ovate, 8-13 cm 
long, 4-8 cm wide, subcoriaceous, the margins 
serrate in the upper part. The base cuneate, the 
apex acuminate. Glabrous above, glabrescent be- 
neath, woolly in early stages, with 8-13 secondary 
veins. Male flowers arranged in amentlike inflo- 
rescences 1.5-4 cm long, these simple or branched, 
sometimes crowned by a solitary female flower. 
Stamens many, verticillate, surrounded by 3 bracts, 
these broadly ovate to deltoid, 5 mm long, 5 mm 
wide at base, early deciduous. Filaments 2-3 mm 
long, pilose; anthers oblong, 2 mm long, geminate. 
Female flowers surrounded by bracts, solitary on 
pedicels 3 mm long, these appressed-pilose. Bracts 
ovate, 2.5-3 mm long, appressed-pilose. Ovary 
inferior, bicarpellate, 4-locular, included in the 


l és Dio) Museo Nacional Apartado 749-1000 San n Costa Rica. 
& C. 


Wilson Botanic Garden, 


San Vito, Coto Brus, Costa 


ANN. MISSOURI Bor. GARD. 76: 1148-1151. 1989. 


Volume 76, Number 4 
1989 


Gómez-Laurito & Gómez 1149 


Ticodendron 


FIGURES 1-5. 


Ticodendron incognitum. — 1. Female flowers (G. Herrera et al. 516). 


2. Seed and fruit (Gómez- 


L. et al. 11472). —3. Male m Mflorescence. (G. Herrera et al. 515).—4. Male/female inflorescence (G. Herrera et 
al. 515).—5. Leaves (G. Herrera et al. . 365 Bars = 1 cm. 
perianth tube, with ovule in each locule (3 abort rera vi al. 516 (B, BM, CR, F, MO, NY, USJ); male 


very early); on axile placenta ovules hemitropous; 
stigmas 2, pilose, 5-12 mm long. Fruit asymmet- 
ric, somewhat swollen on one side, round at base, 
acute at the apex, drupelike, to 7 cm long, 4 cm 
thick, greenish, mucilaginous; endocarp very hard, 
longitudinally furrowed. 

A RICA. ALAJUELA: Reserva Fores 

an Ramón, ca. Colonia Palmarena, 900 oe m, 


de 
ee 84°35 W. 30 Mar. 1987, female tree, G. Her- 


Hsia Lp Cost 


tree, Herrera et al. 517 (CR, MO, pus id be dis- 
el Upala, Bijagua, El Pilon, 10?41'N, 00'W, 
1,000- 1, 400 m, female tree, G. Herrera 203210 n MO); 
La Brisa de Zarcero, dustin Smith H 970 
Viejo, Zarcero, 10°15'N, 84?18'W 
mez-L. & A. Carvajal 11100 (CR, USJ). CARTAGO: Ta- 
panti, I. Chacón 1985 (CR). HEREDIA: Sac ramento, i 
del Volcán Barva, 10?06'N, 84°07 female tree, J. 
Gómez-L., L. J. Poveda & M. Grayun 11472 (CR. F, 

USJ); same locality, male tree, J. Gómez-L., L. J. Poveda 
& M. Grayum 11473(CR, F, USI). PUNTARENAS: Reserva 


1150 


Annals of the 
Missouri Botanical Garden 


F 

o 

3 

de URE 6. Ticodendron inc digi — A. Portion of stem of flowering staminate plant. — B. Staminate T 
cence. —C. Stamen. — D. Staminate flower. — E. Bracts E staminate flower. — F. Pistillate Meg 

G, H. Pistillate lower. —I. En do ocarp, front view. —J. Endocarp, cross section. — K. Stipules.—L. Leaf apex (A, K, 


L from McPherson 12493; B-E from Haber 7072; F-H n Haber 6840; J from McPherson 8401. E 


Natural de Monteverde, W. Ha ws gps 2058 (CR, 
MO); same locality, Haber et al. 
7072, 7286 (CR, MO, and to be Fix same lo- 
cality, sterile tree, Hammel et al. 15385 (MO), 
tree, 75396 (MO 
Poveda 864 (CR). : Fortuna 
Dam region, near trail along continental ed 8?45'N, 
82?15'W, 1,250 m, sterile branches with fruits from 
ground, McPherson & Hammel 8401 (MO, C KY 

PMA). CHIRIQUÍ: ao of Fortuna Dam, S of reservoir, 
8?45'N, 82°15'W, 1,100-1,300 m, sterile tree, Mec- 


Eus 11659 (MO); same locality, male and female 

McPherson 12493 (MO, PMA, and to be distrib- 

ated). COCLÉ, N of El Copé, 8%38'N, 80%35'W, 750 m, 

sterile tree, McPherson 12615 s NICARAGUA. ZELAYA: 

erro La Pimienta number 1 range, summit of 

2 peaks, northernmost and central, ca. "13°45! N, 84°59'W, 
900- 1,600 m, sterile tree, Pipoly 5233 (MO). 


Etymology. Tico, a Costa Rican diminutive, 
and dendron, Greek for tree. Its unusual char- 
acters were first noticed by Costa Rican botanists. 


Volume 76, Number 4 
1989 


Gómez-Laurito & Gómez 1151 


Ticodendron 


The first known collection was made in this country 
at the beginning of the century by C. Werckle 
(s.n., US No. 1010451) near San Cristóbal. The 
specific epithet refers to its Hamamelidean rela- 
tionships. 

e new genus and species of Hamamelidean 
affinities (Gómez-Laurito & Gómez, 1989) has 
prompted much discussion: a possible relationship 
to Urticales (e.g., Ulmaceae or Cecropiaceae) has 
been suggested. Dilleniales (Wolfe, 1989) and Ro- 
sales have also been proposed as possible orders 
for Ticodendron. For some time, placement in the 
Ulmaceae was favored and our genus does resemble 
genera of this family (e.g., Aphananthe Planchon), 
but its morphology excludes Ticodendron from 
subfamilies Ulmoideae or Celtidoideae. As in the 
Ulmaceous genus Zelkova Spach, Ticodendron has 
drupaceous fruits and somewhat distichous, pin- 
nately veined leaves, but has four hemitropous 
ovules, three of which abort very early in devel. 
opment. Leaf characteristics suggested to W. Bur- 
ger (pers. comm.) a relationship to the Cecropi- 
aceae, but our observations revealed a bicarpellate 
ovary and two well-differentiated and free stigmas, 
among other differences. The dry, hardened fruits 
bear a faint resemblance to those of Corylus L., 
and, when we first saw a vegetative specimen, we 
thought of Carpinus L., which reaches northern 
Nicaragua. 

Several genera of Hamamelidales (e.g., 
centron Oliver, Sinowilsonia Hemsley, Altingia 
Noronha, and others) show many of the morpho- 
logical characteristics of the new plant and suggest 
that its true affinities lie within this order, although 
its precise placement requires further study. 

he venation and marginal teeth of Ticodendron 
leaves have been compared and found somewhat 
similar to those of fossil Juglandaceae (Hickey, fide 
Raven, pers. comm.). Some botanists initially pro- 
posed Flacourtiaceae as the proper family. 

A possible relationship between Ticodendron and 
the Antillean Picrodendron Planchon, based on 
the similar appearance of the pistillate flowers, has 
been suggested (M. Grayum, pers. comm., 1987) 
but 7icodendron has simple leaves and strongly 
asymmetrical fruits. Further, its pollen grains are 


Tetra- 


unusually small and not of the Olfieldia-type, which 
occurs in the Euphorbiaceae, where most modern 
authors place Picrodendron (Hakki, 1985; Hay- 
den et al., 1984; Webster, 1975). Wood anatomy 
(Carlquist, in press) and reproductive morphology 
(Tobe, in press) point to Fagales. 

Ticodendron incognitum forms sparse stands or 
is found isolated in the montane evergreen forest 
of Central America. In Costa Rica, it usually grows 
from 900 to 2,300 m elevation associated with 
Aiouea costaricensis (Mez) Kostermans, Phoebe 
hammeliana W. Burger, Cedrela salvadorensis 
Standley, Eugenia lepidota Berg, and other species. 


macho and jaül nazareno (male alder 
alder). Both names recall the alderlike habit of 
Ticodendron and the purplish tinge of fresh cuts 
in most of the trees. Also, it is called duraznillo 
and candelillo morado. The wood is used as fire- 
wood. It is fine-grained, dries a pale rosy-brown, 
takes a high polish, and is harder than that of 
Alnus. 


LrrERATURE CITED 


CaRLQUIST, S. Wood and bark anatomy of Ticodendron; 
comments on relationships. Ann. Missouri Bot. Gard. 
(in press). 

Gomez-Laurito, J. & L. D. GÓMEZ. 1989. A new 
hamamelid picem from Central America: a prelim- 
inary report. Mem. of the Evolution, Systematics and 
Fossil History of the Hamamelidae Symposium. Ox 

iv. Press, Oxford (in press). 

Studies on West Indian Plants 3 

On floral morphology, anatomy and relationship of 

Picrodendron baccatum (L.) Krug & T (Eu 

phorbiaceae). Bot. Jahrb. Syst. 107(1-4): 379-394. 

J. 


HAYDEN, W. |. T. GiLLIs, D. E. STONE, C. ET 
& i . WEBSTER. i wie de and paly- 
nology of Picrodendron: furthe ence for rela- 


10) 
dons E the Oldfieldioideae pd J. 
Arnold Arbor. 65: 105-127. 


Tope, H. Reproductive morphology, anatomy, and re- 
ntes: of Ticodendron. Ann. Missouri Bot. Gard. 
(in press). 

WEBSTER, G. L. 1975. Conspectus of a new classifi- 


cation of the Euphorbiaceae. Taxon 24: 593-601. 
1989. Leaf architectural analysis of the 
Hamamelidae. Mem. of the Evolution, Systematics 
and Fossil History of the Hamamelides Symposium. 


Oxford Univ. Press, Oxford (in press). 


A SECOND SPECIES OF 
THE AMPHI-ATLANTIC 
GENUS ALONSOA 
(SCROPHULARIACEAE) 
IN SOUTH AFRICA! 


Kim E. su 


Steiner 


ABSTRACT 


A second species of the predominantly South American genus Alonsoa has been found to occur in South Africa, 


cies in having oil-secreting trichomes in shallow corolla sacs and from the other S 
species in its annual habit, dilated filaments, and acuminate, bilobe 


d stigma. Flowers of Alonsoa unilabiata are self- 


compatible, but are usually out-crossed by Rediviva (Melittidae) oil-collecting bees. Differences between Alonsoa and 


Niascia are discussed. 


Alonsoa Ruiz & Pavón is a small genus of ap- 
proximately 16 species that until recently was be- 
lieved to be restricted to the New World (Edwin, 
1971; Hilliard & Burtt, 1984; Roux, 1986). It is 
concentrated primarily in the Peruvian Andes of 
South America but ranges north through Central 
America to Mexico and south to Chile and Bolivia; 
it also occurs in the Cape floristic region of South 
Africa. 

Alonsoa is one of seven genera that Bentham 
(1876) placed in the Hemimeridae, the others being 
Diascia, Diclis, Hemimeris, Colpias, 
and Angelonia. Since the American genus An- 

elonia is now placed in its own tribe Angelonieae 
(Pennell, 1920; Barringer, 1983, 1985), the Hem- 
imeridae, with the exception of most species of 
Alonsoa, are basically restricted to southern Af- 


Nemesia, 


ca. 

The first Alonsoa (A. peduncularis (Kunze) 
Wettstein) species discovered in Africa was col- 
lected by Zeyher (3485) in the late 1830s from 
woody gorges around Uitenhage in the eastern 
Cape. The original description (Kunze, 1842) was 
prepared from cultivated plants at the Leipzig bo- 
tanical garden. The source of that seed was Leh- 


mann of Hamburg who obtained it from someone 
in South Africa. The original source is uncertain, 
but, due to the rarity of this plant, it may well 
have come from Zeyher’s collection. No subse- 
quent collections of this taxon were made until its 
discovery at a second locality in 1985 (Roux, 1986). 
Since this second site is far removed from areas 
2 past or present cultivation, and since no known 

South American species have corolla sacs, Ben- 
tham’ s (1876) suggestion that the original speci- 
men collected by Zeyher was naturalized from South 
America is highly unlikely. Evidently Bentham’s 
reluctance to accept Alonsoa as indigenous to South 
Africa was due to the unusual amphi-Atlantic dis- 
tribution that this would represent. 

Out of a combined total of about 4,500 genera 
occurring in Africa (including Madagascar) and 
South America, a, 115 (2.6%), in- 
cluding Alonsoa, are restricted to these subconti- 
nents (Thorne, 1973; Hilliard " Burt 1984). O 
these, only seven (0.2% of the full total) occur in 
the Cape flora of South Africa. Among genera 
shared between South Africa and America, Alon- 
soa is the only genus presently centered in the 
Andes. Except for a few widespread genera (e.g., 


! I thank J. P. Rourke for isa ba assistance, and D. Snijman and several 
t. Peter Goldblatt kindly check ked 

Astholm "Mentified Alonsoa meridionalis var. ariete and de cs drew Figure 
Alonsoa pollination has been invaluable. 


one pia sr versions of the manusc 
z S Pavón; 


de of V. B. Sh head in the stud 


commennng 


P ublic a tions 
The 


f 
cion Herbarium, National Botanic Gardens of South Africa, Kirstenbosch, Private Bag X7 Claremont 7735, 


South Africa. 


ió h Associate, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63110, U.S.A. 
ANN. Missouni Bor. Garb. 76: 1152-1159. 1989. 


Steiner 1153 


Volume 76, Number 4 
Alonsoa in South Africa 


FIGURES 1-4. Alonsoa flowers 


front view. —4. 
= 0.5 cm. 


Limosella, Lindernia, and llysanthes) and the 
parasitic genera Melasma and Alectra with tiny 
easily dispersed seeds, Alonsoa is the only con- 
nection between South and African 
Scrophulariaceae. This and other evidence led Ra- 
ven & Axelrod (1974) to suggest that the Scroph- 
ulariaceae probably did not exist when the two 
subcontinents were close together. Thus, Alonsoa 
must have reached Africa subsequently by long 
distance dispersal 

In America, Alonsoa is characterized by race- 
mose inflorescences, resupinate flowers with the 


American 


. A. unilabiata, front view. 


1. peduncu rine 8, 


. A. unilabiata, rear view.— 3. 


A. peduncularis, rear view. Arrowheads indicate sacs containing oil-secreting tric Tb dd Scale line 


corolla tube split to the base between the lower 
(morphologically upper) lobes, and oblong to ellip- 
tic, usually longitudinally furrowed seeds (Edwin, 
971; Hiliard & Burtt, 1984). South African 
members of the genus are additionally character- 
ized by two shallow yellow sacs in the gamopetalous 
portion of the corolla at the base of the lateral 
corolla lobes (Figs. 1-4, 5A, B). These sacs are 
densely packed with oil-secreting trichomes (Hil- 
liard & Burtt, 1984; Roux, 1986; Steiner, pers. 
obs.). 
My current research on the annual Diascia 


cm. 


species from the western Cape has revealed that 
a second species of Alonsoa has been incorrectly 
classified as Diascia unilabiata (Linn. f. th 
The presence of a second Alonsoa species in South 
Africa strengthens the unusual amphi-Atlantic con- 
nection within the genus and warrants an exami- 
nation of the distinction between Alonsoa and 
Diascia in general, and the taxonomic history, 
variation, and reproductive biology of Alonsoa uni- 
labiata (Linn. f.) Steudel in particular. 


HABIT AND HABITAT 


Alonsoa unilabiata is a winter-flowering (July- 
October) annual herb with simple to dissected leaves 
and pink to orangish pink resupinate flowers with 
dark purple-maroon centers. Contrasting with the 
dark center is a yellow translucent window or sac 
at the base of each lateral corolla lobe. 

Alonsoa unilabiata occurs from near the west 
coast of South Africa (Elands Bay and Lamberts 
Bay), where it is found in loose sands of old sta- 
bilized dunes at about 100-200 m, northeast to 
the escarpment of the Bokkeveld Mountains 
(Nieuwoudtville area), and along the Roggeveld 
escarpment from Calvinia south to the northwest- 
ern edge of the Little Karoo (Konstabel and Twee- 


1154 Annals of the 
Missouri Botanical Garden 
FIGURE 5. Alonsoa unilabiata — A. Front view of flow Rear view of flower. — C. Stamens, style, and 
stigmas. — D. Anterior stamens. — E. Posterior stamens. Scale Diu: Pu A and B = 1 cm. " Scale lines for C-E = 0.5 


side) (Fig. 6). On the escarpments of the Bokkeveld 
and Roggeveld, these plants grow primarily in sandy 
to clay loam soils derived from shales, except around 
Nieuwoudtville where they occur on heavy doleritic 
clays. Drége collected 4. unilabiata from the 
southwestern Cape near Cape Town in 1836, 
no recent collections have been made from "m 
area. Alonsoa peduncularis, in contrast, is known 
from only two localities in the southern and eastern 
Cape floristic region (Fig. 

ere appear to be lires fairly recognizable 
ecotypes of A. unilabiata associated primarily with 
different soil types. The differences between the 
ecotypes, involving mainly extent of leaf dissection 
and corolla sac shape, are not, however, great 
enough to warrant taxonomic recognition. 


DISTINCTIONS BETWEEN DIASCIA AND ALONSOA 


Corolla morphology of South American and South 
African species of Alonsoa is remarkably similar 
(Figs. 1-5A, 7). Most obvious is the resupination 
which places the enlarged morphologically lower 
lip in an upright position. Resupination was over- 

i 


and all subsequent botanists except Diels (1897), 
presumably because this character is obscured in 


Volume 76, Number 4 
1989 


Steiner 1155 
Alonsoa in South Africa 


FicunE 6. Distribution of Alonsoa unilabiata (solid 
rica 


pressed material. Diascia species do not have re- 
supinate flowers. Coupled with the rather strange 
appearance of the corolla caused by resupination 
in Alonsoa is the complete split in the corolla 
between the lower (morphologically upper) lobes 
(Fig. 5B), a character that Kunze (1842; in Hilliard 
& Burtt, 1984) erroneously attributed solely to 
Alonsoa peduncularis (as Schistanthe peduncu- 
laris). This condition is also found in A. unilabiata, 
as well as New World species A. meridionalis vd 
Kunze and others (pers. obs.; Edwin, 1971). T 
Diascia corolla always has a short but distinct us 
(i.e., gamopetalous portion) without a split. 
Another distinction between Alonsoa and Dias- 


FIGURE 7. Flower close-up of Alonsoa meridionalis 
subsp. Ds from El Angel, Ecuador 


circles) and Alonsoa peduncularis (star circles) in South 


cia is the form of the androecium. In Diascia (and 
in the related /Vemesia and Diclis) the anterior 
lower) filaments twist around the base of the pos- 


— 


terior (upper) filaments, causing the upper part of 
the anterior stamens to be placed in a posterior 
position and the posterior stamens to appear in the 
anterior position (Hilliard & Burtt, 1984). No such 
twisting of the filaments occurs in Alonsoa, al- 
though a similar result without twisting is achieved 
in A. pe 


(morphologically upper) stamens are situated above 


duncularis, where the anthers of the lower 


the anthers of the upper (morphologically lower) 
stamens. In 4. meridionalis, the stamens are up- 
right; in A. unilabiata (Fig. 5C) and A. pedun- 
cularis they project downward. This downward 
orientation in 4. unilabiata is caused by the turn- 
ing under of the lower (morphologically upper) rim 
of the corolla just above the point of filament at- 
tachment. Alonsoa unilabiata is set apart from all 
other Alonsoa species by its strongly dilated fila- 
ments (Fig. 5C- 

Seed uote has further distinguishes Alonsoa 
from Diascia. As shown in Figures 8-11, 4. uni- 
labiata, A. peduncularis, and other Alonsoa 
species have wingless, oblong to elliptic seeds, usu- 
ally with longitudinal straight (or undulate in A; 
unilabiata) furrows (Edwin, 1971). Diascia species 
have dorsiventrally flattened embryos and usually 
have elaborate wings or cuplike extensions on the 
seed coats. The pollen of 4. unilabiata and A. 
peduncularis is tricolpate in contrast with the hexa- 


1156 


Annals of the 
Missouri Botanical Garden 


Scanning electron micrographs of Alonsoa seeds. 
10. 4. peduncularis. — 


unilabiata from Calvinia District. — 10. 4. 


to octacolpate grains of all examined Diascia 
species. Diascia and Alonsoa are distinguished in 
Ta 


ble 1. 


POLLINATION AND BREEDING SYSTEM 


The major difference between South African and 
South American Alonsoa species is the presence 
in the former of a pair of shallow corolla sacs (Figs. 
1-4, 5A, B) containing a dense patch of oil-se- 
creting trichomes (trichome elaiophores). These tri- 


~ 


FIGURE 12. Trichome elaiophore from the corolla sac 
of Alonsoa unilabiata. Scale line = 15 un 


-8. unilabiata from Elands Bay.— 9. A. 
A. meitduonilis from Peru. Scale lines — 05 mm. 
chomes consist of a two-celled stalk, bearing two 
or three secretory cells (Fig. 12 
The presence of elaiophores has a direct bearing 
on the manner in which these species are pollinated. 
Alonsoa unilabiata is pollinated by female oil- 
collecting bees in the genus Rediviva. Plants grow- 


— 


ABL Characters distinguishing Alonsoa from 
Diascia. 


Alonsoa Diascia 


Flowers resupi- Flowers never resupinate 
nate 
Corolla split to Corolla with a short distinct tube, 
E 


the base be- r split 


=) 


tween the mor- 

phologically 

upper lobes 
Bases of fila- 


ments straight 


Bases of filaments twisted placing 
anterior stamens in a posterior 
position 

Pollen tricolpate en hexa- to octacolpate 

Seeds oblong with Seeds usually dorsiventrally flat- 

longitudinal tened and with wings or with 

furrows and no extensions of the testa fused 


wings into a cup 


Volume 76, Number 4 
1989 


Steiner 
Alonsoa in South Africa 


1157 


ing in the sandveld near Elands Bay are pollinated 
by two small (ca. 10 mm long) undescribed black 
species (Rediviva “ruficornis” and R. 
while plants on the Roggeveld escarpment from 
Grasberg to Sutherland (with the exception of 
Nieuwoudtville) are pollinated by a different un- 
described species R. “alonsoae” and rarely by R. 
“racemosae.” At Nieuwoudtville, plants are visited 
R. “racemosae.” In all 


EET 


“nigra’’), 


by R. “macgregori” and 
cases, bees place their heads at the base of the 
upper (morphologically lower) corolla lobe while 
inserting their forelegs into the oil-containing sacs. 
The dark central portion of the corolla as well as 
the yellow sacs are ultraviolet absorptive and con- 
trast with the UV reflective corolla lobes (Steiner, 
pers. obs.). The presence of an ultraviolet absorp- 
tive region is common in oil-secreting species of 
Diascia and Hemimeris and appears to help orient 
the bee on the flower (Steiner, in press, pers. obs.). 

Specialized scraping and absorptive setae on the 
forelegs of the Rediviva pollinators facilitate ex- 
traction of the oil, which is transferred to the hind 
legs and carried back to the nest (Whitehead & 
Steiner, 1985; Steiner & Whitehead, 1988; Buch- 
mann, 1988). While probing for oil with its fore- 
legs, the bee’s abdomen curls into the cradle formed 
by the filaments, causing the anthers to rub against 
and deposit pollen on the upper tergites. Pollen is 
deposited from the abdomen onto the stigma of the 
next flower as the bee repeats this process. 

The two Rediviva species at Elands Bay do not 
depend solely on 4. unilabiata for oil, but also 
visit at least three other oil-producing species in- 
cluding Hemimeris sabulosa L.f. and two unde- 
scribed Diascia species. On the Roggeveld es- 
carpment, with the exception of the Nieuwoudtville 
area, 4. unilabiata is pollinated almost exclusively 
by Rediviva *alonsoae." Even more unusual, 
however, is the apparent restriction of this bee to 
the flowers of 4. unilabiata for oil, despite the co- 
occurrence of other oil-producing species at each 
site. This bee is the only one of 16 western Cape 
Rediviva species that is restricted to a single oil 
host plant (Steiner & Whitehead, pers. obs.). 

Pollinator observations on other Alonsoa species 
are lacking. Alonsoa peduncularis occurs sym- 
patrically with Diascia patens (Thunb.) Fourcade 
and may share the same Rediviva “lepidos” pol- 
linator that collects oil from and pollinates that 
species, while New World species of Alonsoa, all 
of which lack oil-secreting trichomes, are probably 
pollinated by pollen-collecting bees. Many of the 
American species have large sturdy apically de- 
hiscing anthers that form a cone around the style 
similar to that found in Solanum. These species 


are probably **buzz" pollinated (Buchmann, 1983) 
by large Bombus (Apidae) or Centris (Anthopho- 
ridae) bees. 

The presence of shallow yellow corolla sacs in 
some annual Diascia species (D. capensis (L.) 
Britten, D. diffusa (Thunb.) Benth., D. cardiose- 
pala Hiern, D. elongata Benth. et al.) probably 
accounts for the past confusion between Alonsoa 
unilabiata and Diascia. It is unlikely, however, 
that the presence of these sacs is indicative of a 
close relationship between Diascia and Alonsoa 
(Steiner, unpublished). Instead, it probably indi- 
cates a convergence based on adaptation to polli- 
nation by oil-collecting bees (Whitehead & Steiner, 
1985; Steiner & Whitehead, 1988). Such unre- 
lated parallel evolution of oil secretion has occurred 
numerous times in families as disparate as Orchi- 
daceae, Iridaceae, Malphighiaceae, Krameriaceae, 
Scrophulariaceae, and Primulaceae; in unrelated 
orchid genera such as Oncidium and Disperis; and 
among unrelated Scrophulariaceae, such as Cal- 
ceolaria, Bowkeria, and Diascia (Vogel, 1974, 
1986; Simpson & Neff, 1981, 1983; Steiner, 
1987). 

Alonsoa unilabiata is self-compatible, as are 
other species in the genus, including 4. pedun- 
cularis (Roux, 1986) and A. meridionalis (pers. 
obs.). Both of these last-mentioned species readily 
self-pollinate, and 4. peduncularis can produce 
cleistogamous flowers under certain circumstances 


~ 


pers. obs.). Flowers of Alonsoa unilabiata, how- 
ever, self-pollinate only occasionally because of 
the spatial separation between anthers and stigma. 

Alonsoa unilabiata is most closely allied to 
Alonsoa peduncularis, with which it shares the 
presence of corolla sacs containing trichome elaio- 
phores, characters that must have evolved in Africa 
in conjunction with Rediviva oil-collecting bees. 
The annual habit, curved and dilated filaments, 
and acuminate two-lobed stigma, however, are all 
specialized features that set 4. unilabiata apart 
from A. peduncularis and the South American 
species. 


SYNONYMY AND DESCRIPTION 


Linnaeus f. (1781), Thunberg (1784), and Ben- 
tham (1846, 1876) all failed to realize that Alonsoa 
unilabiata (Linn. f.) Steudel has resupinate flowers. 
As a result, Diels (1897) described a new species 


— 


Diascia nutans Diels) for a collection of A. uni- 
labiata from the Hantam Mountains near Calvinia. 
Discussing the resupinate corolla in his species, 
Diels stated that i 
to the same result as resupination of the Alonsoa 


“evidently this development led 


1158 


Annals of the 
Missouri Botanical Garden 


corolla, in fact the convergence developed so far 
that without the pits in the throat of the corolla, 
our species [D. nutans] would have to be placed 
in Alonsoa.” The type of D. nutans was destroyed 
in Berlin, but the description and recent collections 
from the same locality (Steiner 759) suggest that 
Diel's plant must have been 4. unilabiata (Linn. 
f.) Steudel. 

When Bentham (1836) described D. heteran- 
dra he had not seen the type of Alonsoa unilabiata 
nor did he see it prior to his treatment of Diascia 
in De Candolle's Prodromus (p. 17, 1846). This 
may explain why he described Drége's collection 
as a new species in the first instance and why he 
retained it in his later publication. One of the char- 
acters used by Bentham to MEUS. D. heter- 
andra was the presence of st . Judging 
from the poor preservation of the type as well as 
the relative ease with which anthers become de- 
tached in fresh material, it is likely that the missing 
anthers fell off during preparation and subsequent 
handling, since the specimen matches 4. unila- 
biata in all other respects. 

The type location for Diascia heterandra Benth., 
between Blouberg and Tygerberg, is considerably 
south of the main west coast distribution of 4 
unilabiata. Habitats near the type locality, how- 
ever, are similar to those found around Elands Bay, 
where Alonsoa unilabiata can be locally common. 
The area between Blouberg and Tygerberg has 
been dramatically altered by the spread of urban- 
ization and alien acacias, hence 4. unilabiata is 
probably no longer there. 

Diascia dissecta Hiern is simply a variant of 
A. unilabiata related to growth conditions. Its tall 
and spindly habit and more narrowly dissected leaves 
are typical of plants along the Roggeveld escarp- 
ment that grow up through shrubs. 


Alonsoa unilabiata (Linn. f.) Steudel, Nomencl. 

840. Antirrhinum unilabia- 

tum Linn. f., Suppl. Pl. 279. 1781. Hem- 

imeris unilabiata (Linn. f.) Thunb., Nov. Gen. 

Plant. 4: 78. 1784. Diascia unilabiata (Linn. 

f.) Benth. in DC., Prodromus 10: 257. 1846. 

TYPE: South Africa: e. Cap. b. spei, Thunberg 

s.n. (sheet 14207) (lectotype, here designated, 
UPS, microfiche at BOL). 


iab — Benth. in Hook., Comp. Bot. M ag. 

zd . TYPE: South Africa: Cape Flats, be- 

ween | Blauw Berg and Tyger Berg, below 500 ft., 
7887 (holotype, K). 

TE nutans Diels, Bot. Jahrb. Syst. 23: 472. 1897. 
TYPE: South Africa: Hantam Mountains, Meyer s.n. 
(holotype, B, destroyed), ex char. only. 

Diascia dissecta Hiern, in Fl. Capensis 4: 153. 1904. 


TYPE: South Africa: Gerrit Snyman's farm in the 
Roggeveld Karoo, Burchell 1289 (holotype, K). 


Annual herb, erect, simple or branching; main 
stem 30-400 mm high, up to 2 x 2 mm in cross 
section, quadrangular, green to reddish purple, gla- 
brous to glandular-villous or hirsute, more densely 
so towards base. Leaves simple, opposite, subop- 
posite or upper stem leaves alternate, in up to 5 
pairs per branch, 4-55(-60) x 3-25 mm, lan- 
ceolate to ovate, dentate to pinnatifid, upper sur- 
face glabrous, lower surface glabrous to glandular- 
villous-hirsute with translucent hairs, apex acute, 
base obtuse or attenuate; lobes 1.5-9 x 1.0-5.0 
mm, linear to ovate, occasionally secondarily lobed 
or divided, the apex acute; petioles 2-20 mm long, 
glabrous to glandular-villous-hirsute. Flowers 1—4 
in a loose terminal raceme, alternate; bracts 4-31 
X 1-6 mm, becoming smaller upwards, linear to 
ovate, entire to pinnatifid, glabrous to glandular- 
villous-hirsute; pedicels 5- 18 mm long, ascending, 
resupinate, slightly longer in fruit, glabrous to glan- 
dular-villous-hirsute. Calyx segments 2.5-3.0 x 
1.0 mm, narrowly elliptic, entire, lowermost (mor- 
phologically uppermost) slightly narrower and 
shorter, ca. 2 mm long, obtuse, entire, hirsute with 
minutely glandular translucent hairs. Corolla bila- 
biate, 12-24 x 13-26 mm; upper lip (actually 
lower lip in upper position due to resupination) of 
3 lobes, upper lobe (morphologically lower lobe) 
ca. 6-11 x 7-13 mm, very widely ovate, sparsely 
glandular-villous-hirsute outside, glabrous inside; 2 
lateral lobes 5-10 x mm, depressed ovate, 
bases oblique, pink to orangish pink; gamopetalous 
portion below the 3 lobes 3-5 x 5-9 mm, very 
dark purple, bisaccate; sacs 2.2-3.3 mm long, 
openings 1.5-2.0 x 1.3-2.1 mm, cordate to trans- 
versely elliptic in outline, sacs projecting back and 
downward, yellow, inner surface densely covered 
with oil-secreting trichomes; lower lip completely 
split to the base; lower lobes (morphological upper 
lobes) ca. 2-4 x 4-7 mm, transversely elliptic 
and partly folded under, bases oblique; pink to 
orangish pink. Stamens didynamous, projecting 
downward and involute to form 2 concentric cor- 
date rings, with outer ring 4.0-4.5 mm diam. and 
inner ring ca. 2.5 mm diam.; posticous filament 
6-8 mm long, broadly dilated in the lower half into 
a kneelike projection and in upper half involute; 
anticous filament 4-5 mm long, lacking a broad 
dilation, involute; both deep purple, glandular-pu- 
berulent; anthers ca. 1 mm long, bearded with light 
blue hairs; thecae yellowish blue, opening outward; 
pollen tricolpate. Ovary 1.2-1.5 x 1.2-1.5 mm, 
widely ovate in outline, obtuse, glandular-puberu- 
lent; ovules 20-30; style 5.5-8 mm, involute, su- 


Volume 76, Number 4 


Steiner 
Alonsoa in South Africa 


1159 


bulate, persistent; stigma minutely bifid, with acute 
lobes; capsule 6.0-8.5 x 4-6 mm, pyriform, apex 
emarginate; seeds 1.4-1.9 mm, oblong to elliptic 
in outline, with shallow undulating longitudinal fur- 
rows, dark reddish brown. 


Additional specimens examined. South Africa: 
31.19 (CALVINIA): ca. 10 mi. from Nieuwoudtville on 
road to Kareebooms, Lewis 5868 (NBG); near Grasberg, 
N of Nieuwoudtville (AC), Barker 9348 (NBG); Van Rhyns 
Pass, Bond 1156 (NBG), Esterhuysen 6142 (BOL); top 
of Van Rhyns Pass, Lewis 5461 (NBG); Van Rhyns Pass 
on S-facing side of W-facing slope, Steiner 901 (NBG); 
Klip Koppies, Nieuwoudtville, Lewis 5854 (NBG); be- 
tween Oorlogs Kloof and Papkuilsfontein, Leipoldt 3454 
(NBG); Farm Glen Lyon, S of Nieuwoudtville, Oliver 46 
(PRE); Matjesfontein, Schlechter 10919 (SAM); road to 
Theunissedrift, 0.5 km NE of junction with Grasberg road, 
Steiner 1355 (NBG); Farm Klipbok (AD), Lewis 5882 
(NBG); Farm Toren, 2 km E of Klipwerf road, 16.4 km 
N of Calvinia- Loeriesfontein road (BC), Steiner 759 (NBG); 
near Augustfontein Mtn., S of Calvinia (CB), Barker 9301 
De 31.20 (WILLISTON): Blomfontein, E of Middelpos 
C) Barker 10781 (NBG, STE); near Elandsfontein 


Calvinia ‘oul a Rooiwal), Steiner 817 (NBG); Farm 
Bloomfontein/Hartbeesfontein, NW of Middelpos, Stei- 
ner 822 (NBG). qe a LANWILLIAM): Zuurfontein, near 
Lambert's Bay (AB Schlechter 8556 (BOL); Ven- 
sterklip, N of elsa e (AD), Boucher 2832 s 
ay, 7 km S of 


Un 


s Ba mNo 
Redlinghuys road (R366), Steiner 7 
tween Lamberts Bay and Elands Bas: 5kmS of junction 
with Road 365, Steiner 1543 (NBG); between Clanwilliam 
and Graafwater (BB), L. Bolus 23172 (NBG, BOL); in 
arenosis near Alexander’s Hoek (BC), R. Se hlechter 5124 
(BOL); Het Kruis (DA), Stephens & Glover 8 (BOL). 
32.20 (SUTHERLAND): d SW of Sne land es- 
(AD), 4402 (STE, PRE); Houthoek, 
Sutherland (CA), Hanekom 1068 (STE, PRE); Klipbanks- 


(N1) (DC), Steiner 1061 (NBG); below road to Sutherland 

(R354), 45.1 km N of junction with National Road (N1) 

at Matjiesfontein, Steiner 1051 (NBG). 33.20 (MONTAGU): 

; Compton 3014 (BOL), Konstabel (AD), 

y 20 mi. N of Matjiesfontein 

(BA) "Hall 3262 (NBG). WITHOUT PRECISE LOCALITY: Cape 

Flats, between Blauw Berg and Tyger Berg, below 500 

ft., Drége 7887 (K); a Elandskloof and Clanwil- 
liam, Leipoldt s.n. (BOL 21270). 


LITERATURE CITED 


BARRINGER, K. Monopera, a new genus of 
ey amass from South America. Brittonia 35: 
111-1 


D Revision of the genus Basistemon 
(5 crophulariaceae). Syst. Bot. 10: 125-133. 
BENTHAM, C. Scrophulariaceae. /n: Hook. Comp. 


—. 1846. Scrophulariaceae. In: A. De Candolle, 
Prodiomus 10: 186-5860. 
7 Scrophulariaceae. i 
. Hooker, Genera Plantaru 
Buc ae S.L. 1983. Buzz olution in in ms. 
Pp. 73-113 in C. E. Jones & R. J. Little cora 
Hendboo k of Bo eal Pollination Biology. 
Nostrand Rheinhold, New 
—— ——. 1988. The ecology of oil pod and their 
bees. Annual Rev. Ecol. Syst. 18: 34 9 
Diets, L. 1897. Beiträge zur Kenntnis p Scrophu- 
lariaceen Afrikas. Bot. Jahrb. Syst. 23: 471-496. 
EDWIN, G. a aoe In: Flora of Peru: 
Publ. Field Mus. Nat. Hist., Bot. Ser. 13(5-b) : 
HILLIARD, O. M. & B. L. Burtr. 1984. A revision of 
ee section Racemosae. J. S. Afr. Bot. 50: 269 
340. 


a pnm & 


Kunze, O 842. 2 ae peduncularis Kunze. 
Linnaea 16: 109- 
LiNNAEUS F. 1781. Supplementum Plantarum. Bruns- 


vigae (Impensis orphanotr ophei). 

PENNELL, F. W. 1 Scrophulariaceae of Colombia. 
Proc. Acad. Nat. Sci. ne hia 72: 136-188. 

Raven, P. H. & D. I. AXELROD. 1974. Angiosperm 
biogeography and past pee movements. Ann. 
Missouri Bot. Gard. 61: 539-673 

Rou e " E 1986. Alonsoa rediscovered. S. African J. 

: 7-9, 

e 3 B. & J. L. Nerr. 1981. Floral rewards: 
alternatives to pollen and nectar. Ann. Missouri Bot. 
Gard. 68: 301-322 
& 1983. Evolution and diversity of 

floral rewards. Pp. 142-159 in C. E. Jones J. 

Little (editors), Handbook of Experimental oo 
Biology. Van Nostrand Rheinhold, New Yor 

STEINER, K. E. Oil-producing orchids t oil- 
collecting bees in southern Africa. Abstr. S. African 
Assoc. Bot., Durban. 

The Diascia (Scrophulariaceae) window: an 
orientation cue for oil-collecting bees. J. Linn. Soc. 
Bot. on aa 
. B. WHITEHEAD. 1988. The association 
ao al: producing flowers and oil- collecting bees 

a. In de zold- 


~ 


Botany. Monogr. Syst.  Misscun 
211. 
Nomenclator Botanicus, 2nd 
. I.G. Conas Stuttgart. 
P n F. 73. Floristic relationships between 
tropical e gie e gie America. Pp. 27-47 in 
. J. Meggers, E. S. Ayensu & W. D. Duckworth 
(editors) Mum iden ied D in Africa and 
So : a Comparative Review. Smithsonian 
p 


nst. 
THUNBERG, C. P. 1784. Nov. . Plantarum 4: 78. 
1974. Olblumen ul da Bienen. 
ou ye turwiss. Kl., Trop. Sub- 
trop. Pflanzenwelt 7 
198 VR is olsammelnde Bienen, 
unite Folge. Akad. Wiss. Abh. Math.-Naturwiss. 
Kl., Trop. Subtrop. Pflanzenwelt 54: 1-168 
WHITEHEAD, V. B. & K. E. STEINER Oil- col- 
PE a in South Africa. African Wildlife 39: 
144- 


NOTES 


A NEW SPECIES OF 
ISOETES FROM TERRITORIO 
FEDERAL AMAZONAS, 
VENEZUELA 


While preparing a treatment of the genus /soetes 
for the Flora of the Venezuelan Guayana, several 
specimens from Territorio Federal Amazonas in 
Venezuela were found to represent an undescribed 
and distinctive species. This species is described 
and compared with other species of the northern 
Andes. 

The various terms for organs and their character 
states follow Hickey (1985, 1986a); terminology 
for spore morphology follows Hickey (1986b 
dimensional shape characterizations for the ligule, 


labium and sporangium follow Radford et al. (1972). 


WoO- 


Isoëtes vermiculata Hickey, sp. nov. TYPE: Ven- 
ezuela. Territorio Federal Amazonas: Depto. 
Atabapo, Cerro Marahuaca, submerged in 
small rivulet, 03°37'N, 65°23'W, 2,520- 
2,650 m, 26-27 Feb. 1985, Steyermark & 
Holst 130758 cia MO; isotypes, MU, 


Ven n.v.). Figures 1-3 


alis speciebus neotropicis differt subula triquetra, 
velo fere completo, ligula habenti pulvino a -ovato, 
auriculato, et macrosporis rugulatis vel subreta 


Corm globose, 20-35 mm diam., bilobed; roots 
arising synchronously from the circumbasal fossa. 
Leaves numerous, stiff, erect to strongly recurved, 
to 85 mm long, 8-19 mm wide at the base, 1.3- 
2.0 mm wide at mid-length; alae chartaceous to 
submembranaceous, hyaline proximally, casta- 
neous to blackish brown or dark green distally, 
1.5-2.5 mm wide at the sporangium, 10-20 mm 
long (24-29% of the total leaf length), the apices 
on outer leaves acute to free, those on inner leaves 
attenuate; subula triquetrous, dark green, the apex 
acute to attenuate, corneous; fibrous bundles ab- 
sent; scales and phyllopodia not seen. Sporangia 
obovate to widely depressed-obovate in surface view, 
hyaline, unspotted, 4.0-4.5 mm long, 2.5-4.0 mm 
wide, basal. Velum complete to incomplete, cov- 
ering 66-100% of the sporangium. Ligule margins 
delicate, ephemeral; the cushion dark, persistent, 


ANN. MISSOURI Bor. 


GARD. 76: 1160-1162. 


depressed-ovate, 0.23-0.40 mm high, 1.1-1.4 mm 
wide, auriculate. Labium absent or only pen 
developed. Megaspores white, slightly shiny, ru- 
gulate to subretate, often with a a weakly developed, 
smooth girdle, 880-990 43) um diam.; 
proximal ridges straight, brandly rounded, the 
equatorial ridges straight or scalloped, broadly 
rounded. Microspores pale reddish brown, scab- 
rate, 35.5—45.8 (x = 40.1) um long, 25.2-34.4 
(x — 29.5) um wide. 


Paratypes. VENEZUELA. TERRITORIO FEDERAL AMAZO- 
as: Depto. Atabapo, summit, Cerro Marahuaca, 03°50'N, 
65°28'W, 2,5 2,600 m, 30 Mar.-1 Apr. 1983, Stey 


e & Delascio 129259 (VEN); submerged in Anu. 
attached to m pi in shallow water, 03?40'30"N, 
Es 20"W. 2,5 0 12 Oct. 1983, Steye d 
129586 (MO, l 


Isoetes vermiculata is known only from stream- 
banks on the summit of Cerro Marahuaca of the 
Territorio Federal Amazonas in southern Venezue- 
la. The species is named for the distinctive wormlike 
markings on the proximal and distal faces of the 
megaspores. 

Isoetes vermiculata is a distinctive element of 
the /soetes flora of northern South America. When 
fertile, it is most easily identified by its large, ru- 
gulate to subretate megaspores and depressed-ob- 
ovate sporangia with nearly complete vela. It can 
be identified vegetatively by a combination of the 
triquetrous (three-sided, with rounded angles) subu- 
la, depressed-ovate, auriculate ligular cushion, and 
absence of a well-developed labium. The only other 
species of /soetes known from this region of Ven- 
ezuela is /. triangula, a lowland, seasonal species 
having a trigonal (three-sided with sharp angles) 
subula, large labium, and pustulate megaspores, 
and not having a velum. 

Not enough is known about /. vermiculata to 
judge affinities with other species in the genus. The 
triquetrous subulae and rugulate to subretate mega- 
spores of /. vermiculata are, however, similar to 


1989. 


Volume 76, Number 4 Notes 1161 
1989 


= 


FIGURE 1. Leaf base morphology of Isoëtes vermiculata. — A. Microsporophyll showing nearly complete velum 
and partially eroded ligule, Steyermark & Holst 130758 (MU).—B. Megasporophyll with a nearly complete velum 
and an entire ligule, Steyermark 129586 (UC).—C. Microsporophyll on which only the ligular cushion remains, 
Steyermark & Holst 130758 (MU). D, E. Variation in ligule form, Steyermark & Delascio 129259 (VEN).—D 
Ligule cushion with all of the ligular margin eroded. — E. Same as D but with portions of the upper ligular margin 
and marginal auriculate base remaining (black — foveola; stippled — cushion). 


FIGURES 2, 3. Spores of Isoëtes vermiculata. — 2. Distal and equatorial views of rugulate megaspores showing 
a nearly smooth girdle distal to the equatorial ridge, Steyermark & Holst 130758 (MU). Bar = 1 mm.— 3. Dis 
and equatorial views of scabrate microspores, Steyermark & Holst 130758 (MO). Bars = 10 um. 


1162 


Annals of the 
Missouri Botanical Garden 


those of I. novogranadensis, I. andina, and I. 
killipii Both /. novogranadensis and 1. killipii 
are distinctive in having levigate microspores, 
whereas those of /. andina have the plesiomorphic 
echinate condition. The large spores of I. vermic- 
ulata suggest a hybrid of high ploidy level, perhaps 
10x or 12x. Whether this hybridization event took 
place when other taxa inhabited the area (perhaps 
during the Pleistocene) or whether it represents a 
morphologically and geographically distinct seg- 
regate population of a more widespread hybrid is 
open to question. 


This research was supported by NSF Grant BSR 
860672 


LITERATURE CITED 


HICKEY, R. J. 1985. Revisionary studies of go e 
. Ph.D. dissertation. The University of Con 
necticut, Storrs, Connecticut. 
1986a. The early evolutionary and morpho- 
logical diversity of d with descriptions of two 
new Er um species. Syst. Bot. 11: 309-321. 

. 86b. qu megaspore ‘surface morphol- 
ogy: DE. variation, and systematic impor- 
tance. Amer. 13. 76: 1-16. 

RADFORD, A. E., W. C. Dickison & C. R. BELL. 1972. 
Vascular Plant Sip Univ. North Carolina, 
Chapel Hill, North Carolin 


—R. James Hickey, Botany Department, Miami 
University, Oxford, Ohio 45056, U.S.A 


A NEW VENEZUELAN 
VIROLA (MYRISTICACEAE) 


This note describes and illustrates a new species Virola is a genus endemic to the American con- 
of Virola from Cerro Sarisarinama in the Vene- tinent with its center of diversity and greatest species 
zuelan Guayana. richness in the western Amazon region. With this 


p ; P 7 
4f y 


C 


fh 
] 
2mm Imm 


FIGURE l. Virola steyermarkii (from Steyermark et al. 109178 
from male flower. 


. — A. Habit. — B. Male flower. — C. Androecium 


— 


ANN. Missouni Bor. Garp. 76: 1163-1164. 1989. 


1164 


Annals of the 
Missouri Botanical Garden 


new species of Virola, there are now 13 Venezuelan 
species in this genus of 48 species. 


Virola steyermarkii W. Rodrigues, sp. nov. TYPE: 
enezuela. Bolivar: Meseta del Jaua, Cerro 
Sarisarinama, cumbre, porción nor-este, 
04?41'40"N, 64°13'20"W, quebrada baja en 
selva más alta de promedio de 15-20 cm de 
alto, 1,400 m, 16-18 Feb. 1974 (fl), J. A. 
Steyermark, | varreno-Espinhoza & C. 
Brewer-Carias 109178 (holotype, VEN; iso- 
type, INPA). Figure 1 


officinali Warb. maxime similis nervatio et inflo- 
rescentiae habitu, sed praecipue androe reviore et 
ds dcn apicem versus angustato statim tes enda. 


Tree 20-25 m tall, branchlets robust, subterete, 
darkly ferruginous tomentose when young (hairs 
usually sessile or short-stalked, stellate, few- 
branched, up to 0.5 mm long), soon glabrescent; 
petioles canaliculate, stout, 3-6 mm long, 1.5-2 
mm diam., darkly ferruginous tomentose; leaf blades 
chartaceous, oblong-lanceolate, 5-12 cm long, 1.8- 
4.0 em broad, obtuse at base, acute at apex, gla- 
brescent or sparsely and minutely puberulent be- 
neath (hairs sessile or subsessile, stellate, 4—5- 
branched); costa shallowly grooved and glabrous 
above, prominent and tomentellous beneath, sec- 
ondary nerves 22-26 per side, straight, impressed 
above, raised-tomentellous beneath, tertiary nerves 
subparallel, impressed above, obscure beneath. Male 
inflorescences narrow, once-branched, 6.5-7.5 cm 
long, peduncle conspicuously flattened, 2.5-3.5 
cm long, rachis 5-7 branched, each branch 5-8 
mm long; branchlets few, short, with flower clusters 


sessile toward the apex; vestiture of branchlets and 
flowers darkly ferruginous tomentose; bracts not 
een; flowers arranged in clusters of about 25, 
pedicels slender, up to 3 mm long. Perianth of male 
flowers thin, 1.5-3 mm long, 3-lobed ree to 
base, the lobes obtuse; androecium 1.2-1 
long, androphore 0.7—0.8 mm long, carnose, eh 


mm 


narrowed towards the apex, the anthers 

0.5 mm long, obtuse at apex, rounded and rd 
cuneate at base. Female inflorescences and fruits 
not seen. 


Virola steyermarkii has a close affinity with the 
officinalis, particularly in the nu- 
merous, close secondary nerves and in the narrow, 
once-branched male inflorescence, but differs mainly 
by having a shorter androecium and androphore 


allopatric V. 


narrowed from the bottom toward the apex. Fur- 
ther, V. steyermarkii is known only from the type 
locality in Venezuela, very far from the restricted 
distribution of V. officinalis in the Brazilian states 
of Bahia and Minas Gerais. 

The new species is named in honor of the late 
Julian A. Steyermark for his valuable contributions 
to the study of the neotropical flora. 


I am grateful to the John Simon Guggenheim 
Memorial Foundation for the fellowship awarded 
to me in 1982 for continuation of my taxonomic 
studies of Myristicaceae, to William Leite for draw- 
ing the plate, and to Bruce Walker Nelson for 
reviewing the English translation. 


— William A. Rodrigues, Instituto Nacional de 
Pesquisas da Amazónia, Caixa Postal 478, 69083 


Manaus, Amazonas, Brasil. 


A NEW SPECIES OF 
CLITORIA (LEGUMINOSAE) 
FROM VENEZUELAN 
GUAYANA 


Two recent collections of a distinctive species 
of Clitoria L. (Leguminosae) were discovered while 
preparing a treatment of the genus for the Flora 
of the Venezuelan Guayana. This species is as- 
signed to subg. Bractearia (Mart. ex Benth.) Fantz 
sect. Brachycalyx Fantz. 

Clitoria comprises 60 species distributed pri- 
marily within the pantropical-subtropical belt; 49 
species are native to the Neotropics. The genus is 
characterized by showy, resupinate, papilionaceous 
flowers; infundibular calyx with persistent brac- 
teoles at the base; persistent stipules and stipels; 
and stalked ovaries with a geniculate, bearded style. 
Members commonly bear a pubescence of micro- 
scopic uncinate (hooked) trichomes best viewed at 
20-30%. 

Fantz (1979) circumscribed sect. Brachycalyx 
and in 1980 added three new species. Members 
are characterized by short tubular calyces with 
minute lobes, inflorescences cauliflorous emerging 
from denuded nodes preceding the leaves, erect 
habits (trees or shrubs), and fruits that become 
weakly convex around the seeds and that are con- 
spicuously depressed transversely between the seeds. 
Species of this section are distributed from Guyana 
to central Panama to Ecuador with isolated col- 
lections of Clitoria glaberrima Pitt. collected as 
far north as southern Mexico. 


Clitoria steyermarkii Fantz, sp. nov. TYPE. Ven- 
ezuela. Bolivar: cumbre de Cerro Guaiquini- 
ma, Salto Rio Szczerbanari (Rio Carapo), 1— 
2 rio arriba del Salto Szczerbanari, 
5944'4"N, 63?41'8"W, 750 m 25 Jan 
1977, Steyermark et al. 113464 (holotype, 
F 1878216; isotypes, F 1878213, 
2769908, US 2821976, VEN). 

ex ad 2 m altus. Folia trifoliata, foliola coriacea, 


margine purpurea. C 
ves. Stylus ovario duplo 


F arvi, resupinati, albi, 
breviter infundibularis, lobi e 
longior. Legumen incognitu 


Slender shrub to 2 m tall. Branches 2-5 mm 
diam., microscopically uncinate and sparsely stri- 
gose, becoming glabrous with age; pith hollow. 
Leaves 3-foliate, 
green to bluish green and glabrous, abaxial surface 


coriaceous, adaxial surface dull 


pale green to gray green, ceriferous, glabrous to 
sparsely strigose on veins; leaflets oblong-elliptic to 
lance-elliptic, apex with short acumen 0.5-1 cm 
long, mucronate, base broadly cuneate to nearly 
rotund, primary lateral veins 7-9 pairs, lamina 5— 
11 em long, 2.5-6 cm wide. Petioles 5-8 cm long, 
subquadrangular, striate to slightly canaliculate at 
apex, microuncinate and sparsely strigose; rachis 
1.5-3 cm long; petiolules subquadrangular, rugose, 
dark colored, sparsely pubescent, 4-6 mm long. 
Stipules persistent, linear, acute, striate, strigose 
apically, 3-4 mm long, 1 mm wide; stipels persis- 
tent, shorter than petiolules, linear-subulate, striate, 
3-4 mm long, ca. 0.3-0.5 mm wide. /nflores- 
cences cauliferous (1—3-fascicled) to 5 mm long, 
pseudoracemose, nodose, with 4-8 flowers, these 
paired at nodes, axes microuncinate; pedicels 3— 
4 mm long. Bracts in 3 series, striate, microun- 
cinate, strigose-pilose apically; inner and middle 
ones inconspicuous, ovate, acute, ca. 0.4-0.6 mm 
long, 0.2-0.4 mm wide, outer ones linear, 2-3 
mm long, ca. 0.3-0.5 mm wide. Bracteoles linear, 
acute, striate, microuncinate, 3-5 mm long, ca. 

mm wide, inserted 1-2 mm below calyx 
base. Flowers resupinate, small, 2.5-3.5 cm long, 
white with purplish margin and purple venation. 
Calyx infundibular, 1 0-nerved, uncinate (trichomes 
conspicuous at 10-12 x), tube 9-11 mm long, 2- 
3 mm wide at base expanding to 4-6 mm wide at 
throat, lobes acuminate-deltoid, 2-3 mm long, 1.5- 
2 mm wide basally, ventral lobe 3-4 mm long. 
Vexillum microuncinate toward apex and margins, 
blade complicate, obovate, emarginate, 1.5-3 cm 
wide, claw 4-6 mm long. Alae extended beyond 
carina 3-4 mm, blade 9-10 mm long, 4-5 mm 
wide, claw 12-14 mm long. Carina falcate, blade 
6-8 mm long, 3 mm wide, claw 15-18 mm long. 
Staminal tube falcate apically, 18-21 mm long, 
ree filaments 1-2 mm long; anthers linear, ca. 


ANN. Missouni Bor. GARD. 76: 1165-1166. 1989. 


1166 


Annals of the 
Missouri Botanical Garden 


1.5 mm long, 0.4-0.5 mm wide. Gynophore ca. 
2-3 mm long; ovary linear, 7-9 mm long, seri- 
ceous with tawny trichomes ca. 0.6-0.8 mm long 
masking dense uncinate microtrichomes beneath; 
style dark colored, ca. twice the ovary length, 14— 
17 mm long, bearded, geniculate 7-8 mm from 
distal end. Legume unknown 


Paratypes. | VENEZUELA. BOLÍVAR: cumbre de Cerro 
Guaiquinima, Salto Rio Szezerbanari (Río Carapo), 1-2 


KEY TO SPECIES OF SECTION BRACHYCALYX 


km río arriba del Salto Ure 95?44'4"N, 
63%41'8"W, 750 m, 20-25 Jan. 1977, Steyermark et 
al. 113282 (MO 2769907, US 2821975, VEN). 


This species is named in honor of Julian A. 
Steyermark who collected the species and who 
established the Flora of the Venezuelan Guayana 
project. 


la. Flowers 4-6 cm long; legume 10-15 mm wide; bracteoles 2-3 mm lon 


2a. Inflorescence 5-22 c 
or treelet, 2-8 m ken (Ecuador) 


m long; staminal tube 27-33 mm long, sir di leaflets glabrous above; shr 


ub 
brachystegia Benth. 


bo 
c 


. Inflorescence to 
shrub to 2 m bes (Colomb; a). 


m long; staminal tube 23-26 mm long, microuncinate; leaflets hispidulous abov 
oa 


ei 


S 


Flowers 2.5-4 c 
3 


a. as to 0.5 cm long; leaves glabrou 


ong; legume 16-30 mm wide; bracteoles 3-5 m 


us or glabrate below; pean 1-1.5 mm long. 
4a. Leaves membranous, concolorous, nonceriferous; style 11-13 m 


ermannii Fantz 


mm long, dual kaai length; 
e 


stipules & stipels caducous, 4-7 mm long; tree (Central America to northern Ven 


4b. Leaves coriaceous, pale green to gray green and ceriferous below; style 14- 17 n n 
the ovary length; stipules & stipels persistent, both 3-4 n 


Venezuela) 


glaberrima Pitt. 
m lc ca. twice 
mm long; small shrub (southeaste ern 


w 
c 


Liana; petiolules 5-7 
elow (western Brazi 


. iris 1-7 em long; leaves conspicuously pubescent below; anthers 1.5-3 mm long. 
: m long; bracteoles 3-4 mm long; anthers 2.5-3 mm long; leaflets pilose 


». steyermarkii Fantz 


9b. Tree or shrub; petiolules On mm long; bracteoles 1.5-3 mm long; anthers 1.5- 


leaflets strigose or velutinou 
6a. Calyx cup-shaped, 8- 12 mm wide at 


roesit ae 
n 


throat; flowers white; gynophore 6-7 mm long; style 
geniculate 6-8 mm from distal end (western Guyana and adjacent Brazil) 


c 
c 


tube 7-11 mm long, lobes 


C. brachycalyx i 


. Calyx narrowly infundibular, 5-7 mm wide at throat; flowers dull purple; gynophore 1-3 m 
long; style geniculate 3-6 mm from distal end. 
7a. Ce : 


2-3 m 


m long; staminal tube 16-22 mm long; keel 


e E 9 mm long, 2-3.5 mm wide; leaflets velutinous below; tree (Venezuela, aces np 


Tb. Calyx tube 13-15 mm long, lobes 
blade 


3-4 mm long; staminal tube 24 
14-16 mm long, 4-5 mm wide; leaflets strigose below; shrub enezuela) 


C. dendrina Pitt. 


4-28 mm long; keel 


I am grateful to the curators of the herbaria (F, 
MO, US, VEN) for the loan of specimens and to 
Julian A. Steyermark and Bruce Holst for assis- 
tance on the Flora of the Venezuelan Guayana 
project. Paper No. 11981 of the Journal Series 
of the North Carolina Agricultural Research Ser- 
vice, Raleigh, North Carolina 27695-7643, U.S.A. 


LITERATURE CITED 


FANTZ, P. R. 1979. Taxonomic notes & new sections 
of Clitoria subgenus Bractearia (Leguminosae). Sida 
8: 90-94. 


anescens Pitt. ex Fantz 


80. New species of Clitoria subgenus Brac- 
tearia section Brachycalyx (Leguminosae) from Co- 
lombia, Venezuela, and Brazil. SIDA 8: 304-311. 

— Paul R. Fantz, North Carolina State Univer- 

sity. Mailing address: Dept. of Horticultural Sci- 

ence, Box 7609, North Carolina State Univer- 

sity, Raleigh, North Carolina 27695-7609, 

U.S.A. 


A NEW SPECIES OF CABOMBA 
(CABOMBACEAE) FROM 
TROPICAL AMERICA 


The genus Cabomba is restricted to the Western 
Hemisphere, ranging from the eastern United States 
through Central America and the Caribbean to 
Argentina. Species of this genus are quite similar 
in habit, and few morphological characters are 
useful for distinguishing among them. The only 
complete taxonomic treatment of Cabomba, by 
Norman Fassett (1953), separated species primar- 
ily using characteristics of the perianth and leaf 
segments. From my fieldwork in tropical America 
and an examination of much of the material that 
has been collected from this region, some taxo- 
nomically significant characters not employed by 
Fassett have been discovered. Such characters are 
the number of leaves per node, stamen number, 
and seed size and shape. Other differences among 
taxa for the characters used by Fassett have also 
been noted. 

In applying this enhanced scrutiny to the ma- 
terial included by Fassett in C. piauhyensis Gard- 
ner (= C. furcata'), I have become convinced of 
the existence of two distinct taxa within this entity. 
Typical C. piauhyensis always has whorled leaves 
at most or all nodes, commonly six (occasionally 
four or five) stamens, and subglobose seeds. This 
has been verified using an isotype of Gardner's 
original collection (Gardner 2478 US) and is also 
readily apparent from the original illustration 
(Gardner, 1844). The distribution of C. piauhyen- 
sis (C. furcata) includes Brazil, northern Bolivia, 
Venezuela, eastern Colombia, the Guianas, Trini- 
dad, and outlying populations are known from Cuba 
and Costa Rica as well. A second taxon, which 
Fassett failed to recognize, has opposite leaves, 
often three stamens (sometimes four or five or 
rarely six), and slightly larger subglobose to oval 
seeds. This taxon occurs from Honduras to western 
Ecuador and northern Venezuela and in the Great- 
er Antilles. The principal area of sympatry for the 
two species is northern Venezuela. The second 
taxon is here described, and a key to all members 
of Cabomba that occur within its range is provided. 


Cabomba haynesii Wiersema, sp. nov. TYPE: 
Panama. Canal Zone: Gamboa, Chagres Riv- 
er, ca. % mi. up river from hyacinth control 
building near dock, 8 Feb. 1973, H. Kennedy, 
C. von Chong & J. Steiner 2323 (holotype, 
MO; isotypes, UNA, 


Folia natantia larninis simplicibus linearibus usque 1.5 
cm longis et 1 mm latis; folia submersa opposita dissecta 
(divisionibus isa ute dichotomis et trichotomis) seg- 
mentis terminalibus 0.1-0.4 mm latis; flores plerumque 
6-8 mm longi ubi Kum stamina saepe 3, interdum 4 
vel 5, raro 6; re xs 3, plerumque 2; semina 1-3, 
1.8-2.5 mm longa, -1.9 mm lata, 1.1 
quam lata 


-2x longiora 


Aquatic herbs with creeping rhizomes rooted in 
substrate and with elongate, distally floating, leafy 
stems. Floating leaves alternate, opposite or borne 
opposite submerged leaves, with petioles to 3.5 cm 
long, the laminae simple, linear, to 1.5 cm long, 
to 1 mm wide, peltate, entire. Submersed leaves 
opposite (rarely whorled at some nodes), with pet- 
ioles to 2.5 cm long, palmately dissected into usu- 
ally 5-7 primary segments, these with 1—3 addi- 
tional and mostly dichotomous divisions, except the 
central 2-3 segments with secondary and often 
tertiary divisions trichotomous, terminal segments 

m wide. Flowers solitary, axillary from 
the upper nodes, bisexual, radially symmetrical, 
mostly 6-8 mm long when closed, borne at or 
above the water surface on peduncles to 5 cm long; 
sepals 3, white to pink-purple; petals 3, coloration 
as in sepals, with protruding auricles near base; 
stamens often 3, sometimes 4 or 5 or rarely 6; 
carpels 1-3, mostly 2, each tapering to a short 
style and a spherical terminal stigma, the ovaries 
superior, the ovules -3. Fruits indehiscent, 
coriaceous, achenelike, or folliclelike; seeds 1-3, 
1.8-2.5 mm long, 1.2-1.9 mm broad, 1.1-2 times 
as long as broad, with longitudinal rows of papillae 
at maturity. 


! Communication with Niels Jacobsen and Mrs Ber of the Royal Veterinary and Agricultural University, 


Frederiksberg mark, who have examined the 


of Cabomba furcata Schultes & Schulte s f. (Roemer & 


Schultes, Syst. Veg. 7(2): 1379. 1830.), indicates i C. furcata is the correct name for the species previously 


known as C. piauhyensis. 


ANN. Missouni Bor. GARD. 76: 1167-1168. 1989. 


1168 


Annals of the 
Missouri Botanical Garden 


Habitat and distribution. 


northern Venezuela, and the Greater Antilles. 
Additional specimens examined. 
28633 (WIS), Standley 20955 (US). He 


(US) Lake Yojoa, Cienaga section, Howard et al. 
(US). NICARAGUA. CHONTALES: 12.5 
más, Haynes 8635 (MO, UNA); 1 km NW 
Diaz, Haynes 8324 (MO, UNA). JINOTEGA: 
Marin. Haynes 8274 

an del Norte, Seymour 5301 (MO, UNA), Stevens 
20849 (MO. UNA). z 


— 


& Schery 951 (US). DARIÉN: Rio Pirre, Duke 
nuo 8309 (US). COLOMBIA. CAUCA: across Rio Naya 


KEY TO CABOMBA OF CENTRAL AMERICA, WEST INDIES, 


la. Floating leaves broadly elliptic, flowers yellow 
lb. Floating leave 
28. Submersed leaves mostly who 


Submersed in stag- 
nant or gently flowing water to 1 m deep in ditches, 
ponds, swamps, marshes, rivers, or streams to 1,000 
m of altitude. Honduras to western Ecuador and 


EL SALVADOR. LA 
UNIÓN: Laguna Maquigua, 18 km W of La Unión, Fassett 
)NDURAS. CORTÉS: 
Lake Yojoa, near Agua Azul, Williams & Molina 17872a 
662 
km SE of Santo To- 
of Puerto 
Presa El 
MO, UNA). RÍO SAN JUAN: San 


ZELAYA: Santa Marta, Haynes 6417 
lau UNA). PANAMA. CANAL ZONE: Madden Lake, Wood- 
& 


from E] Pastico, Gentry & iod 40602 (MO, UNA). 
ECUADOR. GUAYAS 18 km S f Balzar, Davenport et al. 
2090 (UNA, USF). ros níos: Juaneche forest, Canton 
Vinces, Gentry et al. 30760 (SEL, UNA). VENEZUELA. 
ARAGUA: 1.6 km down road to Emo. Camatagua, Hayne ^s 
7645 (UNA); La Victoria, dec ios 79 ( 
Embalse between Upata an 


:alabozo, Davidse 
3799 (MO, US). | PORTUGUESA: Bs e Lucre near 
Acarigua, Salazar 24 (VEN). ZULIA: Caño La Maro 
near Santa Barbara, Trujillo 12044 (MY). Cu IBA: Her- 
redura, Cook 135 (US); unspecified locality, Wright 1861 
(US). Jamaica: lake at Dromilly, Proctor 24494 (NY 
PUERTO Rico. Añasco, Sintenis 5601 (Z); Aguadilla, Sin- 
tenis 5766 (Z). 


Named in honor of Robert R. Haynes, who 


played an important role in the collection and rec- 
ognition of this species. 


AND NORTHERN SOUTH AMERICA 


. C. aquatica Aublet 


s linear to narrowly elliptic, flowers white to pink- purple. 
rlec 


urcata Schultes & Schultes f. 


(including C. warmingii apa and C. piauhyensis Gardner) 


2b. Subrcend leaves opposit 


Ww 
oP 


d )2-4, blades of petals 


c 


4b. Stamens usually 3, sometimes 4 or 5, 


arpel 1, blades of pe ul nir ds narrowed to base (rarely auricled) 
auricled toward 
4a. Stamens usually 6, floating ed more than ] mm yae 


plas C. palaeformis Fassett 
base. 
'aroliniana A. Gray 
(in luding C. “pulec herina ne Fassett) 
rarely 6; floating iu up to lr wi e 
`. hayne esiiWiersema 


I thank Steven R. Hill for the Latin description, 
Lauritz B. Holm-Nielsen for supplying information 
on the type of C. warmingii, Niels Jacobsen and 
Marian Orgaard for providing information on the 
nomenclature of C. piauhyensis and the curatorial 
staff at MO, NY, UNA, US, WIS, and Z for 


providing access to their specimens. 
LITERATURE CITED 


Fasserr, N. C. Vb 
tanea 18: 116- 


"à e of Cabomba. Cas- 


GARDNER, C. Cabomba piauhyensis Gardn. Icon. 
id 


1844. 
Pl. 7 641 
—John H. Wiersema, Systematic Botany and 
Mycology Laboratory, USDA/ ARS, Bldg. 265, 
BARC-East, Beltsville, Maryland 20705, U.S.A. 


A NEW SPECIES OF 
HYPTIS (LAMIACEAE) 
FROM THE VENEZUELAN 
GUAYANA 


While examining collections of Lamiaceae for 
the Flora of the Venezuelan Guayana, | encoun- 
tered a single gathering of a Hyptis species that 
was unfamiliar to me. It was surprising to find a 
new Hyptis species at an altitude of 120-150 m 
in northern South America. 

Although clearly belonging to sect. Hyptis, it 
differs from all other known species in its elongate, 
interrupted, pseudospicate inflorescences com- 
posed of small sessile capitula and situated in the 
axils of much-reduced bracts. The new species is 
also remarkable for its almost sessile leaves, the 
petiole apparently never more than 2 mm long. In 
this respect, it recalls H. vilis Kunth & Bouche, 
from Central America and northern Venezuela but 
differs in floral characters and in leaf shape, and 
by having sessile rather than long-pedunculate ca- 
pitula. The lack of a ring of hairs in the calyx tube 
places H. guanchezii in subsect. Marrubiastrae 
Benth., along with H. parkeri Benth., which is also 
found in the Territorio Federal Amazonas. 

Hyptis guanchezii is named for the collector 
and discoverer Francisco Guánchez. 


Hyptis guanchezii R. Harley, sp. nov. TYPE: 
Venezuela. Territorio Federal Amazonas: Depto. 
Atures, bosque-laja en Cerro Uchonhua (len- 
gua Piaroa), a unos 5 km N del casério San 
Pedro de Cataniapo (en lugar quemado du- 
rante la sequia de 1979), a unos 60 km al 
SE de Puerto Ayacucho, 05?41'N, 67?1 I'W, 
120-150 m, 9 Nov. 1980, Guánchez 365 
(holotype, VEN; isotypes, K, TFAV). Fig- 
ure |. 


Ab aliis speciebus sectionis Hyptidis foliis subsessilibus 
vel petiolis usque 2 mm longis, lamina basim truncata vel 
leniter subcordata, inflorescentia spiciformi interrupta 
capitulis sessilibus in axillis bracteis deminutis subtentis, 
differt. 

Erect herb to 80 cm high, with slender purple- 
tinged stems branching from a central rootstock. 
Stems quadrangular with coarse, spreading villous 


hairs below, these sparser and more slender above, 
the nodes densely tufted with whitish, antrorse hairs. 
Leaves 3.5-7.5 X 

ing in size above and often shorter than the inter- 


.0-3.2 cm, markedly decreas- 


nodes, ovate-lanceolate to narrowly ovate with acute 
apex and narrowing to a truncate or weakly cordate 
base; upper surface of lamina appearing dark green, 
villous with scattered rather broad-based hairs, the 
veins weakly impressed; lower surface paler, villous 
with weak hairs restricted to the rather prominent 
veins and margin, glabrous between the veins and 
with sunken sessile glands; margin coarsely and 
bluntly serrate; petioles 1-2 mm, the upper leaves 
often appearing almost sessile. Inflorescence spi- 
ciform, 10-15 em long, composed of opposite ses- 
sile capitula in the axils of inconspicuous bracts, 
the lower internodes usually 5-6 cm long. Bracts 
up to 4-6 mm long (the upper bracts shorter), 
lanceolate to ovate-acuminate, with broadened base 
and somewhat recurved margin, midrib prominent 
abaxially. Capitula 10-12 mm diam., with an in- 
volucre of narrowly lanceolate, curved bracteoles 
up to 5 X 1 mm, these glabrous with the midrib 
prominent beneath. Flowers subsessile, surrounded 
at base by dense tuft of long, silky, white (pale 
brown in herbarium material) hairs. Calyx tubular, 
at anthesis ca. 4 mm long, the tube 2-2.5 mm, 
glabrous throughout; calyx teeth 1.75-2 mm long, 
narrowly deltoid-lanceolate, ciliate with long-acu- 
minate apex. Corolla 5-6 mm long, white, the tube 
ca. 4.5 mm long; stylopodium absent. Ripe nutlets 
not seen. 


I acknowledge Julian Steyermark's vision in con- 
ceiving the idea of a Flora of the Venezuelan 
Guayana, which helped turn my attention to this 
floristically rich area with its particularly interesting 
phytogeographical patterns in the Lamiaceae. 


— Raymond M. Harley, Royal Botanic Gardens, 
Kew, Richmond, Surrey TW9 3AB, England. 


ANN. Missouri Bor. Garb. 76: 1169-1170. 1989. 


» 
(^ 
A, 


son 
<= SAS 
Ze, 
m~ 


Y 


SS 
EZ 
Ro e. 


We 
Na / 
GTA 
qn 


Ag 


C. Leaf detail: upper surface.—D. Leaf 
(Drawn by 


abit.—B. Leaf, lower surface. 
side view. —F. Flower, side view. —G. Calyx, outer surface. 


FIGURE 1. Hyptis guanchezii. —A. Hab 
detail: lower surface. — E. Sessile capitulum, s 
Milton Andrews from Guanchez 365. 


NEW COMBINATIONS 
IN SORGHASTRUM 
(POACEAE: 
ANDROPOGONEAE) 


Nash (1901:71) separated Sorghastrum from 
Sorghum Moench on the basis of the reduction of 
the pedicellate spikelets to simple pedicels in Sor- 
ghastrum. Morphological, anatomical, and cyto- 
logical data (Davila, in prep.) support Nash's con- 
clusion and reinforce the generic recognition of 
Sorghastrum. Davila (1988) recognized 15 Amer- 
ican and two African species of Sorghastrum. Two 
new combinations in this genus are presented be- 
low. 


Sorghastrum balansae (Hackel) Davila, comb. 
nov. Sorghum balansae Hackel, Fl. Bras. 2(3): 
277. 1883. TYPE: Paraguay: Balansa 209a 
(holotype, P). 


Sorghastrum incompletum (Presl) Nash var. 
bipennatum (Hackel) Davila, comb. nov. 


Andropogon bipennatus Hackel, Androp. 
Nov. Fl. 68(8): 142. 1885. TYPE: Central 
African Republic: Djur and Scriba ghattas, 
Schweinfurth 2486 (holotype, US). 


LITERATURE CITED 

DaviLA, P. 1988. Systematic revision of the ae 
Sorghastrum (Poaceae: Andropogoneae). Ph.D 
sertation. Iowa State Univ., Ames, lowa. 

Nasu, G. V. 1901. Poaceae (P ars.). Page 71 in N. L 
Britton (editor), Manual of the Flora of the Northern 
States and Canada. Henry Hold & Co., New Yor 


— Patricia D. Davila, De; to de Botá- 
nica, Instituto de Biología, U nnd Nacion- 
al Autonoma de Mexico, Apartado Postal 70- 
233, 04510 México, D.F., México. 


ANN. Missouni Bor. GARD. 76: 1171. 1989. 


ANN. Missour! Bor. 


PASSIFLORA MALLETII, 
A NEW SPECIES IN 
SECTION DECALOBA 
(PASSIFLORACEAE) 
FROM MESOAMERICA 


A number of undescribed species of passion- 
flower have been found during a revision of the 
Passifloraceae for the Flora Mesoamericana proj- 
ect (in press); most of them are in the small-flowered 
subgenus Plectostemma Masters. Newly described 
here is one such species; it is restricted to Panama 
and known from three recent collections. 


Passiflora malletii J. MacDougal, sp. nov. TYPE: 
Panama. Panamá: near Cerro Jefe, 3 mi. along 
road past Alto Pacora toward Cerro Brewster, 

?15'N, 79?25'W, 700-750 m, 19 June 
1988, G. McPherson 126124 (holotype, MO; 
isotype, PMA). Figure 1. 


Passiflora pubescens scandens; petioli eglandulosi; folia 
eglandulosa integra acuta vel acuminata, vel r 


3) mm; flores parvi subvirides vel pallido- undi. 
flavi, perianthiis ad basim purpurascentibus; coronae fil- 
amenta biseriata, fi ioribus filiformibus 3.8- 
5 mm longis; operculum plicatum purpureum setulosum; 
androgynophorum 2.5(-3) mm longum; ovarium dense 
pubescens pilosum; bacca atropurpurea; semina 5-6(-7)- 
sulcata. 

Vine 3-5 m, densely short-pubescent to pilos- 
ulous throughout. Stems subangulate or subflat- 
tened, substriate. Stipules (3-)5-8 x (0.3-)0.6- 
| mm, linear to linear-lanceolate, sometimes slight- 
ly falcate; petioles eglandular; laminas (1.5-)4-10 
X (0.8-)1.5-6 cm, entire, not variegated, ovate 
(rarely ovate-oblong), unlobed or occasionally 
3 lobed (if lobed then usually unlobed leaves also 
present on the branch), the leaves of determinate 
axillary branches usually unlobed, the lateral lobes 
if present acute to acuminate, the central lobe 
acuminate (to acute); laminar nectaries absent. 
florescence often present, of determinate, short, 
axillary shoots with reduced or no leaves. Peduncles 

(-2) per node, 0.6-1.4 cm long, uniflorous; bracts 
(3.5-)4-7 x (0.3-)0.7-2(-3) mm, linear, fusi- 
form, oblanceolate, or obtrullate, entire or deeply 
3-toothed to 3-cleft. Flowers pale to light yellow- 
green, slightly flushed purplish; floral stipe up to 


GARD. 


0.5 mm long; sepals 8-10 x 4.2-5 mm, with no 
apical projection; petals 1-2 mm long (or obso- 
lete?); coronal filaments in 2 series, the outer series 
3.8-5 mm long, filiform, pale greenish yellow or 
greenish white, basally purplish?, the inner series 
ca. 3.5 mm long, capitellate, whitish with purplish 
apices; operculum 1.5 mm long, membranous, pli- 
cate, sparsely setose below the apex, purplish; 
staminal filaments connate 2.5(-3) mm along an- 
drogynophore, the free portions ca. 4 mm long; 
ovary densely pilose, dark green. Fruit 1.3-1.8 
cm diam., subglobose, (dark green to) purplish black, 
estipitate; arils unknown; seeds 2.8-3.1(-3.3) x 

9-2.1 mm, transversely sulcate with 5-6(-7) 
sulci, the ridges slightly rugulose. 


Habitat and distribution. The one field ob- 
servation by Mallet and the three collections are 
from 300—750 m at edges of tropical wet forest 
and premontane wet forest, within 25 km of each 
other in central Panama. The label on the type 
describes the locality as “Pacific slope near divide, 
rich wet forest.” 


Name. The specific epithet honors the ecol- 
ogist Dr. James Mallet, whose specialty is the Heli- 
coniine butterflies that feed on Passifloraceae. He 
observed this species along the Llano—Carti road 
in March 1982, made field notes about the plant, 
sent living collections to the University of Texas 
at Austin for study (which died), and pointed out 
(in litt.) that it was apparently undescribed. He has 
made numerous and detailed observations on the 
Passifloraceae and realized that a number of species 
he encountered were new. By kindly allowing free 
access to his field books, his detailed notes have 
enhanced the accuracy of the revision for Flora 

esoamericana and contributed much to our un- 
derstanding of Passiflora. 


Additional CE ns examined. PANAMA. PANAMA: 
El se no to Carti road, 9.3 km N of Panamerican Hwy., 
1977, Folsom 3601 (MO); El Llano- Carti pu 
9 d from Panamerican Hwy., 9%16'N, 78%57'W, 
: 1983, J. S. Miller et al. 851 (BM. 
MEXU, MO, PMA). 


76: 1172-1174. 1989, 


Volume 76, Number 4 Notes 1173 
989 


FIGURE 1. Passi, 


B. Bracts. —C. Stipule 


flora malletii, composite drawing from J. S. Miller et al. 851 and Folsom 3601. — ^. Habit. — 
le. — D. Seed. 


D. Seed 


Passiflora malletii is a member of subg. Plec- — row stipules, and transversely sulcate seeds. Within 
tostemma Masters sect. Decaloba DC. as revealed sect. Decaloba it belongs with the species group 
by the small flowers with biseriate filamentous co- that includes P. sexflora A. L. Juss., P. rugosis- 
rona and plicate operculum, small free bracts, nar- — sima Killip, P. porphyretica Masters, and two new 


1174 


Annals of the 
Missouri Botanical Garden 


species from Mexico. From these it is remarkably 
distinguished by its mostly unlobed, sharply pointed 
leaves, an evidently derived condition. Mallet's field 
notes describe the stem as strongly flattened, 
suggestive of an unusual phyllotaxy for the family, 

ut the dried specimens of the other collectors 
appear to be subangulate to partially flattened. The 
phyllotaxy is not clear because of deformation of 
the stems in drying; both 7; and /j are known in 
the related species of this group. 

The flowers of the new species are notable for 
their short androgynophore and very reduced pet- 
als. The sparsely setulose operculum is also dis- 
tinctive since the floral parts are only rarely pu- 
bescent in this genus. The peduncles are uniflorous 
and the flowers are generally crowded on deter- 
minate axillary branches, much like its close rel- 
ative P. porphyretica. 

The seeds of nearly all of the species of sect. 
Decaloba are probably bird dispersed, but obser- 


vations are few. The label on J. 5. Miller et al. 
851 notes "fruit green and being eaten by tou- 
canettes." I believe the mature fruits to be purplish 
black because in this collection the pericarps turned 
black during the process of preparing the specimen, 
a phenomenon commonly seen with nearly mature 
fruits in this section. In most species of Passiflora 
the seeds and arils mature before the pericarp 
changes color (pers. obs.). It thus appears that the 

irds were eating submature fruit, probably with 
sweet mature arils. 


The Missouri Botanical Garden, with a grant 
from the Jessie Smith Noyes Foundation, gener- 
ously funded my postdoctoral research. John Myers 
prepared the illustration. 

—John M. MacDougal, Missouri Botanical Gar- 
den, P.O. Box 299, St. Missouri 63166- 
0299, U.S.A. 


Louis, 


A NEW COMBINATION IN 
OCOTEA (LAURACEAE) 


Nees (1836) described Nectandra arnottiana 
in his Systema Laurinarum and cited only a spec- 
imen from the Arnott Herbarium which is now 
incorporated in the Herbarium of the Royal Botanic 
Garden, Edinburgh (E). This specimen has not been 
seen by later botanists working on Lauraceae, who 
treated the name incorrectly. Meissner (1864) 
treated N. arnottiana as a variety of Nectandra 
laurel Nees and stated that he did so only guided 
by Nees's description. Mez (1889) accepted N. 
arnottiana as a valid species and treated Pleuro- 
thyrium chrysothyrsus Meissner as a synonym of 
it. He cited only Mathews 3031 (K, BM), the type 
of P. chrysothyrsus, under N. arnottiana and 
obviously had not seen the specimen from the Ar- 
nott Herbarium. I have now seen the Edinburgh 
specimen annotated by Nees as N. arnottiana. This 
specimen is Mathews 1429. Meissner (1864) rec- 
ognized a duplicate of Mathews 1429 in Kew as 
an undescribed species and made it the type of his 
Pleurothyrium ferrugineum, a species up to now 
known as Ocotea ferruginea (Meissner) Mez. But 
because the Nees epithet has priority, I transfer it 
to Ocotea and present the synonymy for the re- 
sulting combination, with the Meissner name in 
synonymy. 


Ocotea arnottiana (Nees) van der Werff, comb. 
nov. BASIONYM: Nectandra arnottiana Nees, 
Syst. Laurin. 289. 1836; Nectandra laurel 
Nees var. arnottiana (Nees) Meissner, DC. 
Prodr. 15(1) 147. 1864. TYPE: Mathews 
1429 (holotype, E). 


Pleurothyrium ferrugineum Meissner, DC. Prodr. 15(1): 
169. 18 PLNRA name for N. arnottiana; 
typified by differ ecimen from same collection 
Har) Ocotea pes (Meissner) Mez, Jahrb. 

. Garten Berlin 5: 265, 1889. TYPE: Mathews 

1439 (holotype, K). 


Duplicates of Mathews 1429 have also been 
reported from BM, G, OXF, and P; these are all 


isotypes of Ocotea arnottiana (Nees) van der Werff. 


Excluded from the synonymy is Nectandra ar- 
nottiana sensu Mez, non Nees, Jahrb. Bot. Garten 
Berlin 5: 402, 1889. Mez based his description on 
Mathews 3031, the only collection cited, and his 
description of leaves and flowers does not apply to 
Ocotea arnottiana. 

Ocotea arnottiana is known to me from Andean 
cloud forests at 2,000-3,000 m in northern Peru 
Bernardi (1975) placed 
Ocotea steyermarkiana Allen in synonymy under 
O. ferruginea and identified several collections from 


and southern Ecuador. 


Venezuelan Guayana as O. ferruginea. However, 
I consider these Venezuelan collections distinct from 
O. arnottiana; 
monodaphne steyermarkiana (Allen) van der 


their correct name is Rhodoste- 


Werff. Similar coriaceous leaves with raised retic- 
ulation and a dense tomentum on the lower leaf 
surface, which misled Bernardi, occur in several 
unrelated, montane species of Lauraceae and are 
probably a habitat adaptation. Meissner (1864) 
already noted the strong similarity between **Pleu- 
rothyrium ferrugineum 
and Persea ferruginea HBK; and an as yet un- 
described species of Ocotea (not related to O. ar- 


EE) 


(= Ocotea arnottiana) 


nottiana) from high elevation cloud forests in Ec- 
uador and Colombia also has these characteristics. 

I thank the curator of E for the loan of neo- 
tropical Lauraceae. 


LrTERATURE CITED 


BERNARDI, L. 1975. 
additis notulis ad Lauraceas americanas nonnulas. 
eee 30: 

F. 


Ocotea budowskiana species nova, 


255-261. 


MEISS 2G. 1864. Lauraceae. In: A. de Candolle, 
Prodromus Systematis Naturalis Regnae Vegetabilis 
15(1): 1-260. 

Mez, ( lagn, 


en Americanae. Jahrb. Kónigl. 
. Berlin 5: 1-556. 
183 36. ‘ane Laurinarum, Berlin. 


NEES, C. G. 


— Henk van der Werff, Missouri Botanical Gar- 
P.O. Box 299, St. Missouri 63166- 
U.S. A. 


Louis, 


den, 
0299, 


ANN. Missouni Bor. GARD. 76: 1175. 1989. 


SISYMBRIUM AREQUIPANUM 
(BRASSICACEAE), A NEW 
SPECIES FROM PERU 


The Brassicaceae (Cruciferae) of Peru were 
treated by Macbride (1938) more than half a cen- 
tury ago. 
changes have since been proposed (see for example, 
Al-Shehbaz, 1989a, b; Khanna & Rollins, 1965). 
The family as represented in South America needs 
to be revised, and many undescribed taxa, partic- 


any new taxa and nomenclatural 


ularly in the genera Draba L., Lepidium L., and 
Sisymbrium L., should be accounted for. The pres- 
ent writer is presently working on Draba in South 
America as a first step toward achieving that goal. 

Sisymbrium arequipanum Al- Shehbaz, named 


endemic, is somewhat isolated morpho- 
logically from all of the Peruvian species of Sisym- 
brium. lt is easily distinguished by having narrowly 
linear-lanceolate, remotely dentate basal leaves, 
auriculate linear-lanceolate cauline leaves, a few 
small flowers, and straight, terete fruits on short, 
ascending, straight pedicels. The species resembles 
superficially members of Arabis L., but this genus 
does not occur anywhere in Central and South 
America, and it always has flattened fruits. 


digi WO Mus Al-Shehbaz, sp. nov. 
TYPE: Peru. Depto. Arequipa: Pichu Pichu, 
des hillside, 5 Jan. 1937, 13,500 ft. [ca. 
4,423 m], Dora Stafford 674 (holotype, F; 
photo, GH). Figure 1 


erba perenna, caulibus erectis glabris, 7-21 cm lon- 
gis; folia basalia petiolata, a lineari- ince hs re- 
moti dentata, ciliata, attenuata, 3.5-14 cm longa, 2- 
mm lata; folia caulina dope. lineari-lanceolata, in- 
tegra; racemi ebracteati, pedicelli fructiferibus recti, cras- 
ibus, 2-6 mm longi; sepala erecta, oblonga, 
sparse hirsuta, 3-3.5 mm longa; petala alba, spathulata, 
3.5 mm lata; siliquae teretae, 
linea gae, ca mm latae; stylus 0.4 
0.7 mm Pis semina immatura uniseriata. 


Perrenial herbs with a slender to thick caudex 
covered with petiolar remains of previous years. 


Stems simple, erect, glabrous, 1 or few from the 
caudex, 7-21 cm high. Basal leaves petiolate, nar- 
rowly linear-lanceolate, 3.5-14 cm long, 2-8 mm 
wide, remotely dentate, ciliate, acute to acuminate 
at apex, attenuate at base, sparsely to densely 
hirsute on both surfaces, the trichomes simple, 
straight, 0.3-0.6 mm long; petioles slender, ciliate, 
flattened and broadened at base. Cauline leaves 
few, sessile, auriculate, linear-lanceolate, 1.5— 
long, 2-2.5 mm wide, entire, ciliate. Inflorescence 
an ebracteate, few-flowered, corymbose raceme, 
elongated considerably in fruit; fruiting pedicels 
thick, ascending, straight, sparsely pubescent, 2- 
6 mm long. Sepals erect, oblong, 3-3.5 mm long, 
1-1.3 mm wide, 
margin, sparsely hirsute. Petals white, spatulate, 
mm long, 1.2-1.4 


nonsaccate at base, scarious at 


mm wide, rounded at 
apex, attenuate at base to a claw ca. 2 mm long. 
Filaments erect, white, ca. 3 mm long; anthers 
oblong, ca. 0.8 mm long. Nectar glands confluent, 
subtending the bases of filaments. Fruits ascending, 
linear, terete, obscurely torulose, 1.5-4 cm long, 
ca. 1.5 mm wide; valves glabrous, with a conspic- 
uous midvein; septum complete; style 0.4-0.7 mm 
long; stigma slightly 2-lobed. Immature seeds uni- 
ca. 2.4 X m 


seriate, 

Vd d ro imen examined. PERU. DEPTO. ARE- 
QUIPA: SW slope of Nevado Chac dager rocky quebrada 
with steep opos, 3,950 m, 3 Nov. 1963, Richard M. 
Straw 2326 (US). 


The holotype of Sisymbrium arequipanum dif- 
fers from the paratype in being sparsely instead of 
densely hirsute. The two collections are otherwise 
very similar. 


I am most grateful to the directors and curators 
of the Field Museum and the Smithsonian Insti- 
tution for the loan of specimens. I thank Neil A. 
Harriman for his critical review of the manuscript 
and Barbara Nimblett for typing the manuscript. 


Sisymbrium are raays =a. 
a and f kam Stafford 674; b-e from Straw 


Plant. —b. 
2320. Scales: 


Basal leaf. 
a,b, f = 1 en 


-c. Flower.—d. Sepal.— e. Petal. —f. Fri 


1; c-e = | mm. Illustrations by the eien 


ANN. Missouni Bor. GARD. 76: 1176-1178. 1989. 


Notes 1177 


Volume 76, Number 4 


1989 


1178 


Annals of the 
Missouri Botanical Garden 


LITERATURE CITED 

AL-SHEHBAZ, I. A. 1989a. The South American genera 
Brayopsis and E fiu rocharis (Brassicaceae). Nordic 
J. m 8: 619-6 

989b. ee elos ardamum (Brassicaceae), a 

en new genus from Peru. J. Arnold Arbor. 


KHANNA, K. R. & R. C. RoLLins. 1965. A taxonomic 
revision of Cremolobus (Cruciferae). Contrib. Gray 
. 195: 97 


Cruciferae. (Fl. Peru). Publ. 
3(2): 937-983. 


MacBRIDE, J. F. 1938. 

Field Mus. Nat. Hist. Bot. 1 
— Ihsan A. Al-Shehbaz, Arnold Arboretum, Har- 
vard University, 22 Divinity Avenue, Cambridge, 
Massachusetts 02138, U.S.A 


INGA NEBLINENSIS 
(LEGUMINOSAE- 
MIMOSOIDEAE), UNA 
NUEVA ESPECIE DEL 
SUR DE VENEZUELA 


ENGLISH SUMMARY 


Inga neblinensis (Leguminosae) is named and described from Venezuela. 


Una serie de colecciones recientes de la base y 
las laderas de Cerro de la Neblina en Territorio 
Federal Amazonas, Venezuela, ha confirmado la 
presencia de una nueva especie de /nga, la cual 
se describe a continuación. 


Inga neblinensis Cárdenas & De Martino, sp. 
nov. TIPO: Venezuela. Territorio Federal Ama- 


zonas: Departamento Rio Negro; gravelly 
banks of large island in Rio Mawarinuma just 
above Neblina Base Camp, ca. 160 m, 


00*50'N, 66?10'W, small tree, flowers yellow, 
3 Dec. 1984, R. Kral 7 1954 (holotipo, MY). 
Figura 1. 
Arbor ramis glabris, striatis vel costatis, fuscis. Folio- 
umque 3-juga, elliptica, gla- 
bra vel subglabra, nitida, 2.6-20 cm longa, 1-6. 
lata, apice acuta, basi cuneato- rotundata. Flores magni, 
mm longo; corolla tub- 


compressum, nitidum, fu , nervis transversis prae- 
ditum, basi rotundatum, apice cuspidatu 


Arbusto o árbol pequeno 3-8 m alto. Ramas 
flexuosas, glabras, estriadas o costilladas, castano- 
ennegrecidas; lenticelas dispuestas longitudinal- 
mente, generalmente de color más claro que e 
resto de la superficie de la rama. Hojas con 2-4 
pares de folíolos, más comúnmente 3 pares, el distal 
siempre más grande; estipulas subuladas o lanceo- 
ladas, estriadas, 3-6 mm largo, 1-1.5 mm ancho, 
glabras o las márgenes pubescentes; peciolo des- 
nudo o alado, pubescente, 4-17 mm largo (incluye 
el pulvínulo); raquis alado, 2- 7.8 cm largo, ala 2- 
10 mm ancho (incluye el raquis), ancha y cordada 
en el ápice, angostada hacia la base; peciolulo an- 
trorso-pubérulo, ennegrecido como el pulvinulo, 2- 
2.5 mm largo, 1-2 mm ancho, articulado con e 


ANN. 


raquis; foliolos elípticos, 2.6-20 cm largo, 1-6.5 
cm ancho, glabros o subglabros, nitidos por ambas 
caras, arriba castano-ennegrecidos, abajo pardos, 
nervadura pinnada sobresaliente, el envés conspi- 
cuamente reticulado, margen entero, ondulado, 
glabro o antrorso-aplicado, base cuneado-redon- 
deada, ápice agudo o algo obtuso, a veces acu- 
minado, la punta mucronulada; glándula entre cada 
par de folíolos, subglobular u ovoide, cupuliforme, 
levantada sobre la superficie, castaño-ennegrecida, 
adentro casi negra, 1.3-2.8 mm diámetro. Flores 
en racimos acortados corimbiformes; pedúnculos 
antrorso-pubérulos, 1-2 en las axilas de las hojas, 
2-4.2 cm largo, 1 mm diámetro; brácteas caedizas, 
2-6 mm largo, 0.1-1.5 mm ancho, pedicelos pu- 
bérulos, 1-2 mm largo, 0.5-1.5 mm diámetro; 
cáliz tubular, pubérulo, nervado, 5-dentado, 5-11 
mm largo, 1.5-3 mm diámetro, dientes irregulares; 
corola tubulosa, tomentosa a escaso pubérula, ner- 
vada, 5-lobulada, 1.3-2.2 cm largo, 1-3 mm dia- 
metro, abajo unida al tubo estaminal en una dis- 
tancia de 2-4 mm, lóbulos 2- 
márgenes involutos; estambres numerosos (más de 
50), 4-5.4 cm largo, tubo estaminal exerto o in- 
cluso, 2-2.6 cm largo; granos de polen en poliades; 
pistilo 3.5-5.2 cm largo, ovario sésil, glabro o corto 
pubérulo, 2-2.5 mm largo, 0.2-0.9 mm diámetro. 
Legumbre aplanada, 10-13.3 cm largo, 2.1-2.5 
cm ancho, 2-3 mm grueso, base redondeada, ápice 
cuspidado, superficie brillante, castario-ennegreci- 
da, con numerosos nervios transversales. 


mm, profundos, 


Paratipos. VENEZUELA. TERRITORIO FEDERAL AMAZO- 


10'W, 21 Feb. 5 (fr), Boom & 
Vaid p (MY); Neblina hus Camp on the Rio 
Mawarinuma, 00%50'N, 66?10'W, m, 17 July 


1984 (fr di, dein & n. ms (MY); 
Missouni Bor. GARD. 76: 1179-1181. 1989. 


1180 


Annals of the 
Missouri Botanical Garden 


Y 
E 
x (0) 3 cm 
<= 


FIGURA 1. 


margins of Rio Mawarinuma below Cerro Neblina Base 
Camp, 140 m, 00%50'N, 66°11'W, frequently inundated 
bamboo swamp forest, 4 May 1984 (fl), Gentry & Stein 
Ve Y); near Cerro de la Neblina, Base Camp, Rio 
Mawarinuma, 140 m, 00%50'N, 66%55'W, 5 Feb. 1984, 
(fr inmaduro), Liesner 15664 (MY); Cerro de la Neblina 
Camp IV, 15 km NNE of Pico Phelps, N branch of river 
in canyon, 780 m, 00%51'N, 65%57'W, 15-18 Mar. 1984 


Inga neblinensis (Kral 71954)—a. Rama florifera.— b. Flor, corte longitudinal. 


pm 


fr inmaduro, restos de fl), Liesner 16710 (MY); along 
Cañon Grande E of Cumbre Camp, Cerro de la Neblina, 
Rio Yatua, 1,100 m, 24 Nov. 1957 (fl), Maguire, Wur- 


Mawarinuma) just upstream from Base 
Camp, SW side of Cerro de la Neblina, 140 m, 00?49'50"N, 
66*09'40"W, 16 Feb. 1985 (fr inmaduro), Nee 30876 


Volume 76, Number 4 
1989 


Notes 1181 


(MY) ca. 2 km S of Base td alluvial we Ad 
forests along Mawarinuma River, 140 m, 00%50'N, 
66?10'W, 11 Feb. 1984 (fl, Thomas & Plowman 3008 
(MY); along Río Mawarinuma, 2-6 km of base camp, 
between base camp and “Puerto Chimo," 160 m 00%50'N, 
66°08'W, 26 Apr. 1984 (fl), Thomas 3212 (MY); along 
Rio jer at "Puerto Chimo" camp, ca. 6 

of bas mp, 170 m, 00%50'N, 66°07'W, 23 April 1984 
(fl), Thomas 3224 (MY). 


T 


Inga neblinensis pertenece a la sección Pseu- 
dinga Benth. Se caracteriza por sus hojas glabras, 


nitidas, raquis alado, flores grandes amarillentas y 
por el color castaño-ennegrecido de la planta (en 
muestras de herbario). Solamente se ha localizado 
en el sur del Territorio Federal Amazonas, en el 


Cerro de la Neblina. 


— Lourdes Cárdenas & Giovanna De Martino, 
Facultad de 
Agronomia, Universidad Central de Venezuela, 
Apartado 4579, Maracay 2101, Venezuela. 


Instituto de Botánica Agrícola, 


REDISCOVERY OF 
TETRAMOLOPIUM 
ARENARIUM SUBSP. 
ARENARIUM 

VAR. ARENARIUM 
(ASTERACEAE: ASTEREAE) 
ON THE POHAKULOA 
TRAINING AREA, HAWAII 


The initial step in the U.S. Army's Land Con- 
dition Trend Analysis Program (LCTA) is to con- 
duct a comprehensive floristic and vegetation in- 
ventory of each installation. One of the first group 
of installations to be inventoried was the Pohakuloa 
Training Area (PTA) on the Island of Hawaii, Ha- 
waii (Fig. 1). Pohakuloa Training Area is a 44,100- 
ha area located in the saddle between Mauna Loa 
and Mauna Kea. Approximately half of the in- 
stallation is ordnance impact area. Because of the 
rough terrain and inaccessibility of much of the 
area, only about 4,100 ha are used for infantry, 
artillery, and tracked-vehicle maneuvers. 

Three endangered plant species have been iden- 
tified previously from PTA (Herbst & Fay, 1979): 
Haplostachys haplostachya (A. Gray) St. John 
var. angustifolia (Sherff) St. John (Lamiaceae), 
Stenogyne angustifolia A. Gray var. angustifolia 
(Lamiaceae), and Lipochaeta venosa Sherff (As- 
teraceae). The species of Lamiaceae were thought 
to be extinct until their rediscovery on PTA (Ay- 
ensu & DeFilipps, 1978; Herbst & Fay, 1979). 
All three of these species occur in a designated 
endangered plants habitat (approximately 1,600 
ha) on the northwest side of PTA within Kipuka 
Kalawamauna (Fig. 1). This kipuka is a vegetated 
older lava flow isolated by newer, sparsely vege- 
tated, or barren lava flows. The elevation of the 
area is approximately 1,545 m. The kipuka re- 
ceives about 380 mm of precipitation annually. 

During the LCTA survey of the vegetation at 
PTA, Tetramolopium arenarium (A. Gray) Hil- 
lebr. subsp. arenarium var. arenarium (Astera- 
ceae: Astereae) was collected by P. Douglas (Doug- 
las 3191) (Fig. 2). Ayensu & DeFilipps (1978) 
and Lowery (1986) reported this taxon as presum- 
ably extinct because no collections had been made 
since the late 1800s. The only previously known 
localities for this taxon were Puu Hualalai and 
Nohonao Hae (near Waimea) on Hawaii and Kula, 


East Maui (Lowery, 1986). A botanical survey 
conducted in 1977 as supporting documentation 
for an environmental impact statement for PTA 
listed T. arenarium (Lawrence Hirai, pers. comm.). 
We have been unable to locate any voucher spec- 
imens for this reported collection. 

Douglas 3191 was collected in the endangered 
plants habitat, near the three endangered plant 
species. The population of Tetramolopium covers 
an area about 200 m by 60 m and is bisected by 
an unpaved road. One part of the population grows 
about 20 m east of the road. When last observed, 
it consisted of 33 robust individuals. The other 
portion, with 81 adult plants and 14 juvenile or 
seedling plants, occurs approximately 80 m west 
of the road. Consequently, the two portions are 
about 10 m apart. Fifteen dead individuals were 
found in the larger portion of the population. The 
cause of death of these plants is unknown. The 
occurrence of juvenile or seedling plants indicates 
that viable seed is being produced and that envi- 
ronmental conditions necessary for seed germina- 
tion and seedling establishment occur periodically. 
Mature plants were observed flowering in January, 
April, and August. 

The collection was made in a shrubland domi- 
nated by Dodonaea viscosa Jacq., Dubautia lin- 
earis (Gaud.) D. Keck, and Sophora chrysophylla 
(Salisb.) Seem. A weedy, introduced grass (Pen- 
nisetum setaceum (Forssk.) Chiov.) is aggressively 
expanding into the area. The Tetramolopium pop- 
ulation occurs on a mesic ridge that runs west- 
northwest. The soil type at this site is rockland 
(rRO) (USDA, 1973). 

Tetramolopium arenarium is a member of sect. 
Sandwicense Lowery, which is composed of gy- 
nomonoecious, upright shrubs with corymbose- 
paniculate heads, narrowly funnelform disk corol- 
las, and fewer than 15 disk florets per head (Lowery 
& Crawford, 1985; Lowery, 1986). This section 


ANN. Missouni Bor. Garb. 76: 1182-1185. 1989. 


Volume 76, Number 4 
1989 


Notes 


1183 


Pohakuloa Training Area 


Kipuka Kalawamauna 
Endangered Plants 
Habitat 


[ICM | 
um pact 
L Area | 


Critical 


High Impact Area 


Scale 
1000 O 1000 3000 5000 Meters 


Miles 


Hawaii 


FIGURE l. Map of Pohakuloa Training Area, Island of Hawaii, Hawaii, showing the extent of the ordnance impact 
a, and locations of the Kipuka Kalawamauna Endangered Plants Habitat and Critical Bird Habitat (Palila, Loxioides 


bailleui Oustalet). 


also includes T. consanguineum (A. Gray) Hillebr., 
T. conyzoides (A. Gray) Hillebr., and 7. lepidotum 
(Less.) Sherff. Of the four species in the sect. Sand- 
wicense, two have been considered extinct: T. are- 
narium and T. conyzoides (Ayensu & DeFilipps, 
1978; Lowery, 1986). The other two species have 
extinct infraspecific taxa (Lowery, 1986). 


Tetramolopium consanguineum never has hir- 
sute or pilose leaves (Douglas 3191 has densely 
and evenly hirsute leaves). Tetramolopium cony- 
zoides has 1-3 disk florets per head (Douglas 
3191 has 5-9 disk florets per head). Tetramolo- 
pium lepidotum has glandular punctate leaves 
(Douglas 3191 has glandular but not punctate 


1184 


Annals of the 
Missouri Botanical Garden 


FIGURE 2. General habit of atan arenar- 
ium subsp. arenarium var. arenarium within the Kipuka 


ne 


Kalawamauna Endangered Plants Habitat, Pohakuloa 
Training Area, Island of Hawaii, Hawaii. (Scale in deci- 
meters.) 


leaves). The only other member of sect. Sandwi- 
cense that occurs on PTA is Tetramolopium con- 
sanguineum. 

Tetramolopium arenarium is composed of two 
subspecies, both presumably extinct: subsp. are- 
narium on Hawaii and Maui and subsp. laxum on 
Maui (Lowery, 1986). Tetramolopium arenarium 
subsp. laxum has open, lax inflorescences, flexuous 
pedicels, granuliferous-glandular achenes, and fo- 
liage that retains its yellow-green color when dried. 
In contrast, T. arenarium subsp. arenarium has 
compact inflorescences, densely strigose achenes, 
nonflexuous pedicels, and brown foliage upon drying. 
Douglas 3191 has compact capitula, densely stri- 
gose achenes, nonflexuous pedicels, and brown dried 
foliage. 

Tetramolopium arenarium subsp. arenarium is 
separated into two varieties: var. arenarium and 
var. confertum. Tetramolopium arenarium subsp. 
arenarium var. confertum has dentate leaf mar- 


gins, and involucres 4-6 mm wide and 2-3 mm 
long. Tetramolopium arenarium subsp. arenar- 
ium var. arenarium has entire leaf margins, and 
involucres 6-9 mm wide and 3-5 mm long. Doug- 
las 3191 has entire leaves and involucres that are 
6—7 mm wide and 4-5 mm long. 

We verified our collection by comparing it with 
two specimens of 7. arenarium subsp. arenarium 
var. arenarium at the U.S. National Herbarium 
(US): Mann & Brigham 519, collected at Puu 
Hualalai; and Hillebrand 319, which has no lo- 
cality information. Both of these specimens were 
annotated by Lowery (1986) as T. arenarium subsp. 
arenarium var. arenarium. Lowery (1986) stated 
the lectotype for this taxon was “Hawaiian Islands: 
Hawaii: 1840, U.S. Exploring Expedition (US!, 
mixed collection with isotype of T. arenarium subsp. 
laxum).” The staff at US could not locate nor has 
any record of this specimen; thus, we were unable 
to compare our specimen with the lectotype. 

Tetramolopium arenarium subsp. arenarium 
var. arenarium has no federal or state protection, 
because it has been listed as extinct by the U.S. 
Fish and Wildlife Service, Office of Endangered 
dp pin l (Andy Robinson, pers. comm.). 
The U.S. Army, however, has begun a protection 
program for this taxon at PTA. 

Herbst & Fay (1979) reported that grazing and 
browsing by feral animals, cutting for camouflage 
by military units, trampling by foot traffic or ve- 
hicles, and accidental fires set by hunters and/or 
military ordnances may adversely affect the three 
endangered plant species found at PTA. Our ob- 
servations indicate that the most likely threats to 
the endangered plant species and the Tetramolo- 
pium arenarium population are from feral animals, 
fire, and competition from invading, introduced 
plants. We have observed numerous feral sheep, 
goats, and hogs in the endangered plants habitat. 
Also, the area has been burned (date unknown). 
We have not observed off-road vehicular traffic, 
foot traffic, or cutting of vegetation for camouflage. 
Fencing, removal of all feral animals, reduced use 
of the area for training, and continued efforts by 
the Army to control fires may help protect these 
plants. 

We are researching the ecology and population 
biology of the endangered species and Tetramo- 
lopium arenarium with funding from the U.S. 
Army. Propagation of the species is being inves- 
tigated as a means of ensuring its survival. 

This research was funded by the U.S. Army 
Corps of Engineers, Engineering & Housing Sup- 
port Center (CEHSC-FN), Donald M. Bandel, Chief 


of Cultural and Natural Resources. We appreciate 


Volume 76, Number 4 
1989 


Notes 1185 


the assistance and encouragement given us by Col. 
E. T. Teixeira, Commanding Officer, PTA, and the 
men and women under his command. We also 
thank K. A. Schulz, E. D. O' Regan, and E. Weg- 
linski for field assistance, the Colorado State Uni- 
versity Herbarium (CS) for handling specimens, 
and D. H. Wilken, B. Berg, and E. L. Painter for 


assistance with this manuscript. 


LITERATURE CITED 

AYENSU, E. S. & R. A. DeFiLiPPS. 1978. Endangered 
and Threatened Plants of the United States. Smith- 
sonian Institute an nd the World Wildlife Fund, Inc. 

Hrnpsr, D. R. & J. J. Fav. 1979. Endangered and 


threatened wildlife and plants; determination that three 


Hawaiian plants are endangered species. Fed. Reg. 
44: 62468-62469. 
Lowery, T. 
Hawaiian Teramolopium (Compositae: 
Allertonia 4: 365. 
& D. 


biosystematic revision of 
Astereae). 


es FORD. 1985. Allozyme diver- 
gence and ev a in Tetramolopium (Compositae: 
ini ae) on the Hawaiian Islands. Syst. O: 


TM 197 3. Soil Survey of Island of Hawaii, State of 
Hawaii, U. S. Government Printing Office, Washing- 
ton, 

— Patricia P. Douglas & Robert B. Shaw, De- 
partment of Range Science, Colorado State Uni- 
versity, Fort Collins, Colorado 80523, U.S.A.; 
and Victor E. Diersing, U.S. Army Corps of En- 

ineers, Construction Engineering Research 


Laboratory, Champaign, Illinois 61820, U.S.A. 


MISCELLANEOUS 
CHROMOSOME COUNTS 
IN ASTERACEAE, 
BIGNONIACEAE, 
PROTEACEAE, AND 
FABACEAE 


Since the publication of my review of the chro- 
mosome cytology of the Fabaceae (Goldblatt, 
1981a) and the documentation of the original chro- 
mosome counts cited therein (Goldblatt, 1981b), I 
have made several more counts in the family. These 
are reported here with associated collection data 
(Table 1). Most of the counts are particularly note- 
worthy as they are either the first records for 
genera or the first for species. 

Counts are also reported for species of Astera- 
ceae, Bignoniaceae, and Proteaceae. The signifi- 
cance of the counts in relation to the existing cy- 
tological record is briefly discussed for each species. 


MATERIAL AND METHODS 


All counts were made from root tips of germi- 
nating seeds, grown in the greenhouse. All seeds 
were collected in the wild for this study. Tips were 
harvested in midmorning and pretreated in either 
0.001 M aqueous hydroxyquinoline for 6-8 hr. at 
refrigerator temperature or in saturated aqueous 
M-bromonaphthalene for 4 hr. at room tempera- 
ture. After rinsing in tap water, the tips were fixed 
for 2-3 min. in 3: 1 absolute ethanol: glacial acetic 
acid, rinsed again in tap water, and then hydrolyzed 
in 10% HCI for 6 minutes at 60°C. Root apices 
were squashed in FLP orcein (Jackson, 1975). The 
species studied and collection data are listed in 
Table 1, in which the diploid numbers are pre- 
sented. 


DISCUSSION 


ASTERACEAE 


Of the estimated 48 species of the Old World 
and predominantly African Dicoma (tribe Muti- 
sieae), only an Indian species, D. tomentosa Cass., 
has been counted, Bhandari & Singh (1977) having 
reported 2n = 22. The report here for the southern 
African D. schinzii provides additional support for 


assuming a base number of x — 11 for the genus, 
although x — 9 is almost certainly basic for the 
tribe Mutisieae as well as for the entire family 


(Raven, 1975; Goldblatt, 1987). 


PROTEACEAE 


Already counted as Aulax cneorum, the record 
here of 2n = 22 in A. umbellata, confirms de 
Vos's (1943) report of n = 11 in Aulax. Other 
African Proteoideae have x — 12, except Leuca- 
dendron, which has x = 13 (de Vos, 1943; Wil- 
liams, 1972). 4ulax and Leucadendron are the 
only dioecious African Proteoideae, and the kary- 
ological data support the contention, based on other 
criteria (Rourke, 1987), that dioecism evolved in- 
dependently in the two genera and that they are 
not closely allied. 


BIGNONIACEAE 


The count here of 2n — 40 for Parmentiera 
valerii, the first record for this species, Is consistent 
with the number reported for three other members 

f this neotropical genus of tribe Crescentieae 


10) 
(Goldblatt & Gentry, 1979). 


FABACEAE 


The count for Zenia insignis, a taxonomically 
isolated and apparently primitive member of Caes- 
alpinieae—Cassieae, accords with the putatively ba- 
sic chromosome number for the tribe, x = 14 
(Goldblatt, 198 1a). 

In Papilionoideae, 2n = 20 in the monotypic 
Graziodendron is consistent with other numbers 
recorded for Dalbergieae, including the closely re- 
lated genus Andira (Goldblatt, 1981a). Pterocar- 
pus, also considered closely related (de Lima, 1983) 
probably has x = 11, but at least two species also 
have n = 10 (Goldblatt, 1981a). The diploid num- 
bers for the previously uncounted Chadsia and for 
Lonchocarpus pycnophyllus, 2n = 22, accord 


ANN. Missouni Bor. GARD. 76: 1186-1188. 1989. 


Volume 76, Number 4 
1989 


Notes 1187 


ABL r 
collection data. 


Chromosome counts reported here for Asteraceae, Bignoniaceae, Proteaceae, and Fabaceae, with 


iploid 
chromosome 
m 
Species n Collection data 
ASTERACEAE 
MUTISIEAE 
Dicoma schinzii O. Hoffmg. 22 Namibia, Kupferberg Pass, Goldblatt & Manning 8849 (MO) 
PROTEACEAE 
Aulax umbellata (Thunb.) R. Br. 22 South Africa, Cape Prov., Vogelgat, Hermanus, Williams s.n. (no 
voucher) 
BIGNONIACEAE 
Parmentiera valerii Standl. 40 Costa Rica, Volcán Miravalles, Poveda 2478 (MO) 


FABACEAE- CAESALPINIOIDEAE 


CASSIEAE 
Zenia insignis Chun 28 China, Kwangsi, Yu s.n. (PE) 
PAPILIONOIDEAE 
DALBERGIEAE 
Graziodendron rio-docensis 20 Brazil, Espirito Santo, Linhares, de Lima 1928 (RB) 
Lima 
TEPHROSIEAE 
Chadsia grevei Drake 22 Madagascar, Fort Dauphin-Tulear, Dorr et al. 4075 (MO) 
Lonchocarpus pycnophyllus 22 Dominican Republic, Peninsula de Barahana, Zanoni & Pimentel 
Urban 
ROBINIEAE 
Poissonia hypoleuca (Speng.) ca. 32 Argentina, Jujuy, Río San Lorenzo, Vervoorst s.n. (no voucher) 
Lillo 
DESMODIEAE 
Brya buxifolia (Murr.) Urban 22 Dominican Republic, Prov. San Juan, Zanoni & Mejía 19785 


PHASEOLEAE-ERYTHRININAE 


Puerto Rico, Vivaldi s.n. (no voucher) 


PHASEOLEAE-PHASEOLINAE 


Neorudolphia volubilis (Willd.) 22 
Britton 
Neorautanenia brachypus 22 


(Harms) C. A. Smith 


Zimbabwe, Wedza, Corby 727 (SRGH) 


AESCHYNOMENEAE 


Pachecoa prismatica (Sesse & 20 
Moc.) Standley & Schubert 


Mexico, near Vera Cruz, Norman 801 (F, GH) 


with the presumed base number for Tephrosieae, 
= 11 (Goldblatt, 19812). In the previously un- 
counted genus Poissonia (tribe Robinieae), P. hy- 
poleuca has 2n — 32, and this species appears to 
be oe Base number in Robinieae is either 
= nies or 10, but dysploidy is evident in the tribe, 
and 6 recorded in some genera 


— 9, 8, 
C nie: 1981a). A base number of x = 8 seems 


likely for Poissonia, a hypothesis consistent with 
the patterns of karyological variation in the tribe. 

Basic chromosome number for Desmodieae is 
ipee x = 11, but at least Pycnospora and the 
one species of Brya previously counted have n — 
10 (Goldblatt, 19812). This first count for B. buxi- 
folia is consistent with the earlier record for the 
genus. 


1188 


Annals of the 
Missouri Botanical Garden 


Basic number for Phaseoleae is x — 11, 

number also probably basic for all eight subtribes 
(Goldblatt, 1981a). The counts here for Neoru- 
dolphia (Erythrininae), a monotypic genus previ- 
ously uncounted, and Neorautanenia brachypus 
(Phaseolinae) are consistent with this pattern. The 
count for Neorudolphia is of interest in view of 
its possible relationship with the large genus Ery- 
thrina, whic 
er in Neorudolphia confirms data from other 


as x — 21. The chromosome num- 


sources that it is probably not closely allied to 
Erythrina 
My count of 2n — 20 in Pachecoa prismatica 
(Aeschynomeneae-Stylosanthinae), cited by Nor- 
man & Gunn (1985), accords with the base number 
of x — 
the nnus four genera of the subtribe. Arthro- 
m is uncounted while Chapmannia has x — 
1 “Goldblatt. 1981a, b). The last-mentioned two 
genera are most closely allied to Pachecoa (Nor- 


man & Gunn, 1985). 


10 in Arachis and Stylosanthes, two of 


I extend my thanks to all those who provided 
the living material used in this study. 


LITERATURE CITED 


1977. In IOPB 


Taxon 26: 107 


iiie M. M. & D. M. SINGH. 
romosome number reports LV. 
DE LIMA, H. C. 1983. 


Novos taxa de Leguminosae 


~ 


Papilionoideae (tribo Dalbergieae) do Brasil. Bradea 
3): 399-406. 
DE Vos, M. P. 1943. pcc d studies in the genera 
of the Proteaceae. S. African J. Sci. 194 l; 
99 


GOLDBLATT, P. 198la. iip d as the phylogeny of 
Leguminosae. Pp. 427-463 in R. M. Polhill & P. 
H. Raven (editors), Advances in une Systematics, 
Part 2. Royal Botanic Gardens, Kew 
1981b. C Vicar numbers i in legumes Il. 
en Bot. Gard. 68: 551-557. 
87. C bo es cytology of Olde ee 


Ann. 


m Mone eae). Ann. Missouri Bot. 
= Pe 
& / H GENTRY. 197 x oo of Bigno- 
niaceae. js Not. 132: 475 


Jackson, R. 1975. ia nus in Haplo- 
pappus gracila: a centric transposition race. Evo 
lution 27: 243 

Norman, E. M. & C. R. GUN NN. 1985. Pachecoa pris- 
matica Wood taxonomy and phylogeny. Brit- 
tonia 37: 

Raven, P. H. 1975. The bases of ao ia 
eny: cytology. Ann. Missouri Bot. Gard. 

76 


ROURKE, J. P. 1987. The inflorescence morphology and 
PA Aulax (Proteaceae). S. African J. Bot. 
53: 480 

Vibes, 1 J. M. 1972. A revision of the genus 
Leucadendron. (Proteaceae). Contr. Bolus Herb. 3. 


— Peter Goldblatt, B. A. Krukoff Curator of Af: 


rican Botany, Missouri Botanical Garden, P.O. 
Box 299, St. Louis, Missouri 63166, U.S.A. 


LEPIDIUM BOELCKET AND 
L. JUJUYANUM 
(BRASSICACEAE), 

NEW SPECIES FROM 
JUJUY, ARGENTINA 


Several new taxa and nomenclatural changes 
have been published (Al-Shehbaz, 1986; Boelcke, 
1964, 1984, 1986) following the treatments of 
Thellung (1906) and Hitchcock (1945) on the South 
American species of Lepidium. The genus, which 
is represented in South America by probably more 
than 50 species, is in need of a comprehensive 
study. Species boundaries within certain complexes 
are artificial, and the variation of many taxa is 
poorly understoo 

he two novelties described below were initially 
sent to Professor Reed C. Rollins for identification. 
I am grateful to him for making them available for 
my study. Each of these is known only from the 
holotype. 


Lepidium boelckei Al-Shehbaz, sp. nov. TYPE: 
Argentina. Jujuy: Depto. Humahuaca, Cerro 
La Soledad, 3,500 m; 23 Jan. 1929, S. Ven- 
turi 8859 (holotype, US 1549309). Figure 
a-c. 
erba perennis caudicibus lignosis simplicibus, 1 -2 cm 
in dud: caulibus villosis subdecumbentibus, e 
cm longis; folia basalia spathulata integra vel dent 
racemi floriferi subcorymbosi, fructiferi bres ong 
sepala oblonga — pies albo-marginata; petala 
alba; stamina 2; siliculae anguste ellipticae vel apdor 
oblongae; glabrae, obtusae vel retusae, 6-7 mm longae, 
.6 mm latae; styli exserti, 0.6-0. 8 mm longi; semina 
ilios: nigella, compressa, exalata, ca. 3 x 1.5 mm. 


Perennial herbs, with a woody, unbranched cau- 
dex 1-2 cm diam. Stems few to several from the 
caudex, ascending to subdecumbent, villous, 6-15 
cm long. Basal leaves spatulate, petiolate, obtuse 
to rounded at apex, attenuate at base, earlier ones 
entire, later ones dentate to subincised, 1.6-4 cm 
long, 3-8 mm wide, the blades glabrous or hirsute 
along the midrib; petioles flattened, densely hirsute, 
1-2.5 cm long. Cauline leaves few, spatulate to 
oblanceolate, entire, attenuate and subsessile at 
base, 5-11 mm long, 1-3 mm wide. Inflorescences 
densely flowered, ebracteate, corymbose racemes, 
considerably elongated in fruit; fruiting pedicels 


ascending, densely puberulous to subhirsute, 5-7 
mm long. Sepals oblong, persistent, glabrous, some- 
what leathery, 2-2.5 mm long, 1.2- 


with a conspicuous white margin 0.2-0.5 mm wide, 


1.5 mm wide, 


repand to suberose in the distal half. Petals spat- 
ulate, white, rounded at apex, attenuate at base, 
3-3.2 mm long, 0.7-0.9 mm wide. Stamens 2; 
filaments white, 2-2.5 mm long; anthers oblong, 
0.7-0.8 mm long. Fruits narrowly elliptic to ellip- 
tic-oblong, conspicuously flattened, glabrous, wing- 
less, obtuse to obscurely notched at apex, 6-7 mm 
long, 3-3.6 mm wide; style exserted, 0.6-0.8 mm 
long; stigma entire, wider than the style. Seeds 
oblong, blackish, wingless, flattened, ca. 3 x 1.5 
mm. 


Lepidium boelckei, which is named after Dr. 
Osvaldo Boelcke, authority on the Argentinian 
Brassicaceae and author of several species of Le- 
pidium, is easily distinguished from all of the pe- 
rennial South American species in having narrowly 
elliptic fruits 6-7 mm long, persistent sepals with 
a broad white margin, conspicuous styles well- 
exserted beyond the apical fruit notch, and sub- 
decumbent stems from a thick woody caudex. The 
Brazilian L. grandifructum C. L. Hitchc. is the 
only other South American species with fruits as 
large as those of L. boelckei. However, the former 
is a robust, erect plant with caducous sepals, six 
stamens, and included styles. Evidently, the two 
species are unrelated, and L. boelckei has no close 
relatives. 


Lepidium jujuyanum Al-Shehbaz, sp. nov. TYPE: 
Argentina. Jujuy: Depto. Humahuaca, Tres 
Cruces; 3,750 m; 6 Apr. 1973, Barbara 
Ruthsatz 506/7 (holotype, GH). Figures ld- 


h, 2 


erba perennis caudicibus tenuibus simplicibus; folia 
Wes longe petiolata, oblonga vel oblongo-oblanceolata, 
laminis 13-23 mm longis, 6-10 m Diu serrato-cren- 
atis, superne villosis, petiolis persisteniibus 3-4 cm longis, 
in dimidio proximale stramineis, valde alatis, glabris, in 


ANN. Missouni Bor. GARD. 76: 1189-1192. 1989. 


Annals of the 


1190 


Missouri Botanical Garden 


Volume 76, Number 4 
1989 


Notes 1191 


j 
AÑ 


Ñ És 


FIGURE 2 


cid distale exalatis, villosis; sepala papyracea, persis- 
entia, 2-3 mm longa; petala alba; stamina 2, filamentis 
a ee dilatatis; siliculae rhomboideae, glabrae, emar- 
ginatae, 3.5-4.5 mm longae, 3-4 mm latae; septum in 
dimidio apicale ee urs exserti, 0.6- E 8 mm lon- 
gi; semina ovata e m longa, 1.4-1.6 mm 


lata. 


;:474. 


Cespitose perennial with an unbranched, slender 
caudex. Stems few, glabrous, 1.5-6 cm long, much 
shorter than the basal leaves. Basal leaves long- 
petiolate; blades oblong to oblong-oblanceolate, 13- 
23 mm long, 6-10 mm wide, serrate-crenate, ob- 
tuse at apex, cuneate at base, glabrescent below, 
villous above with crisped, subappressed, simple 
trichomes, the midrib prominent in the proximal 
half; petioles persistent, 3-4 cm long, the distal 


" fu N 


Leaf trichomes of Lepidium jujuyanum. Scales: a — 100 um; b= 


10 um. 


half green, villous, slender, wingless, the proximal 
straw-colored, glabrous, with 2 conspicuous lateral 
wings to 1.2 mm wide. Cauline leaves few to sev- 
eral, similar to the basal ones but gradually reduced 
in size upward. Inflorescences ebracteate, corym- 
bose racemes, slightly elongated in fruit; fruiting 
pedicels divaricate, usually straight, densely villous, 
4-7 mm long. Sepals oblong to slightly obovate, 
papery, persistent, spreading, membranaceous and 
entire to repand at margin, a pubescent i in 
the middle, 2-3 mm long, 1.5-1.8 mm wide. Petals 
spatulate, white, rounded at apex, 2-2.6 mm long, 
ca. ] mm wide. Stamens 2; filaments white, 1.4— 
1.6 mm long, considerably dilated at base; anthers 
oblong, ca. 0.6 mm long. Fruits rhomboid, 3.5- 
4.5 mm long, 3-4 mm wide, with an apical notch 


FIGURE 1. aoe boelckei and L. poo a-c. L. boelckei. —a. Plant. —b. Fruit. —c. Petal. 
leaf. —f. S 


Jupuyanum. — d. Plant. — e. Basal le 
e ; others — =1 mm. Illustrations by n par. 


—g. Fru 


d-h. L. 
uit.—h. Replum with perforated septum. Scale: a, d, and 


1192 


Annals of the 
Missouri Botanical Garden 


0.3-0.4 mm deep; valves glabrous, thin, obscurely 
veined; septum membranaceous, perforated at apex; 
replum 1-1.2 mm wide; style exserted from the 
apical notch, 0.6-0.8 mm long. Seeds ovate, flat- 
tened, dark brown, wingless, 2-2.4 mm long, 1.4— 
1.6 mm wide; cotyledons incumbent. 


Lepidium jujuyanum, which is named after 
Province Jujuy, Argentina, belongs to the L. mey- 
enii Walp. complex, of which the component species 
are perennials with exserted styles. The new species 
is easily distinguished from all the South American 
lepidiums in having long-petiolate leaves with vil- 
lous, crenate-serrate blades and straw-colored, con- 
spicuously winged, persistent petioles, strongly di- 
lated filament bases, perforated septa, and rhomboid 
fruits with exserted styles. 


LITERATURE CITED 


AL-SHEHBAZ, I. A. 1986. Lepidium solomonii (Crucif- 
erae), a new bg: ies from Bolivia. Ann. Missouri Bot. 
Gard. 73 -831. 


ERRATUM 


BoELCKE, O. 1964. Notas sobre especies x E CRDI 

e la Er ee Darwiniana 13: 506-5 
1984. Notas sobre Cruciferas Argentinas I I. 

SENSUS en el género Lepidium. Parodiana 3: 2 


2 . 


986. Notas sobre Cruciferas Argentinas II. 
Dos nuevas especies Chaqueñas del género L epidium 
y P de sus especies en el NE Argentino y paises 
nos. Parodiana 4: 35-61. 

dne HCOCK, C. 945. The South American species 
of Lepidium. Lilloa 11: 34 

19 Die oe Lepidium (L.) R. B 
A cem Allg. 


iis 
Eine ee es Studie. di 
Schweiz. Naturf. Ges. 41(1): 1 
— Ihsan A. Al-Shehbaz, Arnold Arboretum, Har- 
vard University, 22 Divinity Avenue, Cambridge, 
Massachusetts 02138, U.S.A. 


This is an erratum for the R. J. Rodin volume, The Ethnobotany of the eel Ovambos, published i in di 
np 


by the Missouri Bot 


nical Garden. APONOGETONACEAE; 


Aponogeton (pp. 50, 147): the description give 


50 is for a junceus TER subsp. junceus, not A. rehmannii Oliv. See H. van Bases (1973) Bull. Jard. a p 


Belg. 43: 191-233 


Instructions for Authors, Annals of the Missouri Botanical Garden 


The Annals publishes original articles in systematic 
botany ie related fields. Text may be in English or 
Spanish. When in Spanish, an English abstract in addition 
to a Spanish abstract is required. Short pes nel 
nomenclatural matters and new taxa, r 
“Notes,” which have their own format as described ios 
Authors are urged to contact the editor prior to debui. 
manuscripts. Manuscripts sent already reviewed an 
vised are generally subject to the full normal review pro- 
cess initiated he . Manuscripts not proper prepared 
may be returned for revision prior to re . This includes 
pens n with improper cS E or with diacrit- 

cal marks not indicated. 

O per page, although charges may 
be reduced nader certain circumstances. Corrections in 
proof will be billed nonnegotiably to the author at $2.50 
per line. 

Manuscripts must be typed (output from dot matrix 
printers must be entirely clear) on one side of nongos 
8⁄2 x 11 inch paper. Leave at least one ir in 
all around, except on the 
e left blank at iu top. Th 

lcd riple-spaced throughout, in- 
cluding abstract, notes, np o tables, ont lists, 
Literature Cited, and footnotes. De if 
right-hand margin, and do | there. 
Italics, boldface, or other special palcos should not be 
used —they will appear later as communicated to the 
printer by the editor. W ords to be set in italics should be 
underlined. Accents, umlauts, and other Mee 
marks are t ir Onde of the author. 

Footnotes should be typed double-spaced as ade 


on the first page. first footnote contains acknowledg- 


ments, including information on granting agencies, and 
herbaria that loaned s ‘imens. The second footnote is 
the author's addr ddresses for additional authors are 
each separate ee footnotes. No other footnotes are 
permitted, except footnotes in tables, and these are best 
avoided. In papers formatted as "Notes," there 


p are 
footnotes: the acki yodedgmenis make up the iced 


graph e 
rticle esses of hale 
authors are all placed in the same paragraph. 

ong, complex tables Tables not neat 
and easily er will be returned for improvement. 
Leave out vertical lines, and avoid horizontal lines. Lines 
are added as needed by the editor and/or printer. Type 


re discouraged. 


table captions double-spaced as paragraphs at the tops of 
the tables. 

A concise abstract should accompany each paper other 
than “Notes.” 

eriods are used after all abbreviations (which should 

be minimized) except metric measures, compass direc- 
tions, and herbarium i tions. States are not abbre- 
viated, and cities are usually spelled out, although “st.” 
is acceptable. Abbrevi 


Abbreviations Compiled at ie Herbarium 
otanic 


a however, that book 
abbreviations are to be capitalized, and that books liste 
n the Literature Cited section uld have their titles 
spelle when feasible. Note further that names o 
taxonomic authorities may S out in major taxo- 


nomic citations such as the formal citation at the head of 
a taxonomic treatment. Except by prior arrangement with 


authors not having access to crucial references, the edito 


does not provide the service of looking up abbreviations. 


ar 

Do not give masc part numbers after volume 

numbers pa ss each part is po separately, which 
hor, follow this style: 


is unusual. more "han one aut 


author's e name, initial(s), second author" s initial(s), last 
ird author's initials), last name. Citations of 
books should appear as follows: au uthor's last name, ini- 
i r. Full Title (edited by Editor), ard edition, 
>. Publisher, Place. C itations of work rep. 


— 


urces are aant 


2 x 
EE 
N 


sary, the ot perform library research to fill 
in missing biblio uris information. 
A brief Latin diagnosis for e: 


to a complete Latin description. 
U 


ch new taxon is preferred 
Ilse one paragraph per basionym as follows: Taxon 
author, literature citation, ype n e.g., Bauhinia 
dipetala Hemsl., Diag. Pl. Nov. 1 En Da 
tala (Hemsl. uen & Rose, N. Jonas E^ 23:2 93( 

TYPE: Mexic eracruz: Valle de enge te 
17 13 (holotype, K; isotypes, F, GH, P, US). Exclamation 

use 


points are not ed for specimens Hbri Indicate 


unseen types as "n.v. 

Kevs should be dichotomous, indented, and consistent 
with Rapid du Couplets should be numbered as 
lb, 2a, and so forth. 

Cite specimens as follows: 

Additional specimens examined (or AEA speci- 
mens examined or Paratypes). ae O. OAXACA: Sierra 
San Pedro Nolesco, Talea, 3 Feb. (fl), j^ rgensen 
865 (BM, G, K, US). Dates and deb status are 
optiona 

Send. copies of illustrations; do not send originals until 
accepted. Illustrations will 


All illu 


requested if the manuscript isa 


be returned after ublication. strative materia ls 


mm) or full page width (5/2 inches or ca. l 
Figures are numbered in Arabic numerals in the order of 
their citation in the text. Parts of figures are labeled with 
capital letters. Poetic should be black-and-white 
prints. Transparencies and ne gatives are not epted. 
nto composite po when possible and 


a 

Map should ir perm to » iste ‘and longitude. 
e bars a: appear on dec on photc graphs, 

i on maps. Magnifications/ reductions indicated in cap- 

tions are unwelcome to the point of usually not being 

accepted. Bs electron ip ees er" be free 

of sunset uous ioo eid Axes on graphs labeled. 


tis to appear in figures. ond , prov ide 


following the style in current issue ; 
tyle, cons edi a recent issue of the 


editor. Communication with the editor 
at all points of the publication process is encouragec 

rr nee (three copies) and ~ e PRET 
be addressed to the Editor, Annals of the Missouri Bo- 
tanical PE P.O. Box 299, St. Louis, Missouri 63166, 
U.S.A. 


ANNALS OF THE 
MISSOURI BOTANICAL GARDEN 


VOLUME 76 
1989 


Colophon 


This volume of the ANNALS of the Missouri Botanical Garden has been set in APS Bodoni. The text is 
set in 9 point type while the figure legends and literature cited sections are set in 8 point type. 


The volume has been printed on 604 Warrenflo (No. 1) and 70st Vintage Gloss (Nos. 2, 3, 4). These 
cid-free papers designed to have a shelf-life of over 100 years. Vintage Gloss is manufactured by 
"i Potlatch Paper Company. Warrenflo is manufactured by S.D. Warren Paper C 


Photographs used in the ANNALS are reproduced using 300 line screen halftones. The binding used in 
the production of the ANNALS is a proprietary method known as Permanent Binding. 


The ANNALS is printed and distributed by Allen Press, Inc. of Lawrence, Kansas 66044, U.S.A. 


© Missouri Botanical Garden 1989 


ISSN 0026-6493 


VOLUME 76 


ALMEDA, FRANK. Tessmannianthus, an Arborescent Genus on Melastoma- 
taceae New to Panama 


AL-SHEHBAZ, IHSAN A. Sisymbrium arequipanum (Brassicaceae), a New 
Species from Peru 


AL-SHEHBAZ, IHSAN A. Lepidium boelckei and L. jujuyanum (Brassicaceae), 
New Species from Jujuy, Argentina . 
ALWAN AL-MaYaH A. R. A., & C. A. SraceE. New Species, Names, and 


Combinations in American Combretaceae 


BARRINGTON, Davip S. New Species and Combinations in Tropical American 
Polystichum (Dryopteridaceae) . 


Berry, PAUL E. A Systematic Revision of Fuchsia Section Quelusia (On- 
AgTACEAE) MINI LE 
Berry, PauL E. (See Takuji Hoshino & Paul E. Berry) 


Bons, Lynn. Solanum allophyllum (Miers) Standl. and the Generic Delim- 
itation of Cyphomandra and Solanum (Solanaceae) .. 


Bremer, BIRGITTA. The Genus Argostemma (Rubiaceae-Argostemmateae) 
in Borneo 


BUNTING, GEORGE S. Notes on Araceae 


CÁRDENAS, LOURDES & Giovanna DE MARTINO. Inga neblinensis (Legu- 
minosae-Mimosoideae), Una Nueva Especie del Sur de Venezuela ...... 
CARNEVALI, GERMÁN & Ivón RAMÍREZ. New or Noteworthy Orchids for the 
Venezuelan Flora. VII. Additions in Maxillaria from the Venezuelan 
Guayana 


CARNEVALI, GERMÁN & Ernesto FoLDAaTSs. A New Aspidogyne (Orchida- 
ceae) from Venezuelan Guayana 


CARNEVALI, GERMAN. (See Gustavo A. Romero & Germán Carnevali) ........ 
CARR, GERALD D. (See Harold Robinson, A. Michael Powell, Gerald D. Carr, 


Robert M. King & James F. Weedin) aia l 


CuuNc, Myonc Gi. Hosta jonesii (Liliaceae/Funkiaceae), a New Species 
from Korea 


CONTRERAS, DoMINGO. (See Carlos Ramirez, Enrique Hauenstein, José San 
Martin & Domingo Contreras) 


CRUDEN, RoBERT WILLIAM. A New Echeandia (Liliaceae) from Venezuelan 
Guayana 


D'Ancv, WiLLIAM G. & ARMAND RAKOTOZAFY. Solanum toliaraea, a New 
Species from Madagascar 


Davita, PATRICIA D. New Combinations in Sorghastrum (Poaceae: Andro- 
pogoneae) . 
De Luca, PAoro. (See Aldo Moretti, Paolo De Luca, Jean Pierre Sclavo & 
Dennis W m. Stevenson) M 


Dr MARTINO, GIOVANNA. (See Lourdes Cárdenas & Giovanna De Martino 


1989 


934 
1179 


DIERSING, VICTOR E. (See Patricia P. Douglas, Robert B. Shaw & Victor 


A aio l 


DoucLas, Patricia P., RoBERT B. SHAw & VICTOR E. DIERSING. Rediscovery 
of Tetramolopium arenarium Subsp. arenarium var. arenarium (As- 


teraceae: Astereae) on the Pohakuloa Training Area, Hawaii |... 1182 
Dunn, Davip B. (See Ana Maria Planchuelo & David B. Dub ar. 303 
EnprEss, Mary E. Novelties in South American Malouetia (Apocynaceae) 1141 
ENGLEMAN, M. (See J. Márquez-Guzmán, M. Engleman, A. Martínez-Mena, 

E. Martinez & C. Ramos) s LLL 124 
ERICKSON, SALLY. The Publications of Edgar Anderson: Additions ...... 942 
ESCOBAR, LINDA K. A New Subgenus and Five New Species in Passiflora 

(Passifloraceae) from South America LLL 877 
Fantz, PAUL R. A New Species of Clitoria (Leguminosae) from Venezuelan 

soda 1165 
FoLDATS, ERNESTO. (See Germán Carnevali & Ernesto Foldats) 00 596 
FORERO, ENRIQUE. Book Review 1L 363 
FORTUNATO, RENEE H. Contribución al Genero Mimosa (Mimosaceae) ..... 381 
GENTRY, ALWYN H. A New Species of Allomarkgrafia (Apocynaceae) and 

Notes AIN 923 
GEREAU, Roy E. Three New Species of Bomarea (Alstroemeriaceae) from 

Mesoamerica o . 998 
GOLDBLATT, PETER. Miscellaneous Chromosome Counts in Asteraceae, Big- 

noniaceae, Proteaceae, and Fabaceae |... LL. 1186 
GOLDBLATT, PETER & Jonn C. MANNING. Chromosome Number in Walleria 

(Tecophilaeaceae) LLL 925 
GOLDBLATT, PETER & Jonn C. MANNING. Pollen Morphology of the Shrubby 

Iridaceae Nivenia, Klattia, and Witsenia 0. 1103 
GOLDBLATT, PETER, JAMES E. HENRICH & RICHARD C. KEATING. Seed Mor- 

phology of Sisyrinchium (Iridaceae-Sisyrinchieae) and Its Allies .......... 1109 
GOLDBLATT, PETER. (See John C. Manning & Peter Goldblatt) |... 932 
GÓMEZ-LauRITO, JORGE & Luis D. Gómez P. Ticodendron: A New Tree 

from Central America 1. LLL 1148 
Gómez P., Luis D. (See Jorge Gómez-Laurito & Luis D. Gómez RI. 1148 
GRAHAM, ALAN. Studies in Neotropical Paleobotany. VII. The Lower Mio- 

cene Communities of Panama— The La Boca Formation 0 50 
Hate, W. H. G. Note Concerning Festuca henriquesii (Gramineae) in 

Northern Portugal . = 393 
HAMILTON, CLEMENT W. A Revision of Mesoamerican Psychotria Subgenus 

Psychotria (Rubiaceae), Part I: Introduction and Species 1-16 |... 67 
HAMILTON, CLEMENT W. A Revision of Mesoamerican Psychotria Subgenus 

Psychotria (Rubiaceae), Part II: Species 17-47 386 


HAMILTON, CLEMENT W. A Revision of Mesoamerican Psychotria Subgenus 
Psychotria (Rubiaceae), Part III: Species 48-61 and Appendices ..... 


HAMMEL, Barry. New Combinations and Taxonomies in Clusiaceae .......... 


HAMMEL, Barry E. & GEORGE J. WILDER. Dianthoveus: A New Genus of 
Cyclanthaceae 


HanLEY, RayMonp M. A New Species of Hyptis (Lamiaceae) from the 
Venezuelan Guayana 


HAUENSTEIN, ENRIQUE. (See Carlos Ramirez, Enrique Hauenstein, José San 
Martin € Domingo Contreras) ... 
HENRICH, James E. (See Peter Goldblatt, James E. Henrich & Richard C. 
Keating) 


HERNANDEZ, HéÉcroR M. Systematics of Zapoteca (Leguminosae) .................. 
Hickey, R. James. A New Species of /soétes from Territorio Federal Ama- 

zonas, Venezuela .. 
Horsr, Bruce K. (See Julian A. Steyermark, Bruce K. Holst & Collabo- 


rators) 


Hosuino, Takuji & PauL E. BERRY. Observations on Polyploidy in Fuchsia 
ects. Quelusia and Kierschlegeria (Onagraceae) 


Hurr, MicuakL J. New and Critical Taxa of Euphorbiaceae from South 
America . 


Jones, SAMUEL B., Jk. Hosta yingeri (Liliaceae/Funkiaceae): a New Species 
rom Korea 


Jupp, WALTER S. Taxonomic Studies in the Miconieae (Melastomataceae). 
III. Cladistic Analysis of Axillary-flowered Taxa 


Kawano, SHoICHI. (See Masamichi Takahashi & Shoichi Kawano) ................ 
Keatinc, RICHARD C. (See Peter Goldblatt, James E. Henrich & Richard 
C. Keating) 
KEENER, CARL S. (See Svetlana N. Ziman & Carl S. Keener) «0 
Kinc, RoBERT M. (See Harold Robinson, A. Michael Powell, Gerald D. Carr, 
Robert M. King & James F. Weedin) 
Kramer, K. U. Lindsaea mesarum, a New Fern Species from the Roraima 
Sandstone 


LANDRUM, LESLIE R. A New Species of Calycolpus (Myrtaceae) from the 
Venezuelan Guayana ..... 


LANDRUM, L. R. Book Review ... 


MacDoucar, Jonn M. Passiflora citrina, a New Species in Section Xero- 
gona (Passifloraceae), from Mesoamerica .... 


MacDoucaL, John M. Two New Species of Passiflora Section Decaloba 
(Passifloraceae) from Costa Rica ........ 


MacDoucar, Jonn M. Passiflora mayarum, a New Species Related to P. 
prolata in Subgenus Passiflora (Passifloraceae) 


1160 


MacDoucaL, Jonn M. Passiflora malletii, a New Species in Section De- 
caloba (Passifloraceae) from Mesoamerica |... 
MANNING, JOHN C. & Peter GorpBLATT. Chromosome Number in Phena- 
ospermum and Strelitzia and the Basic Chromosome Number in Stre- 


litziaceae (Zingiberales) 555 
MANNING, JOHN C. (See Peter Goldblatt & John C. Manning) A 
MANNING, JOHN C. (See Peter Goldblatt & John C. Manning) ss. 


MÁRQUEZ-GUZMÁN, J., M. ENGLEMAN, A. MARTÍNEZ-MENA, E. MARTÍNEZ & 
C. Ramos. Anatomia Reproductiva de Lacandonia schismatica (La- 
Candona AA Eu RR E 

MaRTÍNEZ, E. (See J. Márquez-Guzmán, M. Engleman, A. Martínez-Mena, 
E. Martinez & C. Ramos) s LLL 

MARTÍNEZ, ESTEBAN & CLARA HiLDa Ramos. Lacandoniaceae (Triuridales): 
Una Nueva Familia de México LLL 

MARTÍNEZ-MENA, A. (See J. Márquez-Guzmán, M. Engleman, A. Martínez- 
Mena, E. Martinez & C. Ramos) 1. LLL 

MEEROW, ALAN W. Systematics of the Amazon Lilies, Eucharis and Cali- 
phruria (Amaryllidaceae) |... LLL 

MILLER, JAMES S. Notes on the Cordia panamensis Complex (Boraginaceae) 
and a New Species from Colombia |... LLL 

MILLER, James S. Two New Species of Tournefortia (Boraginaceae) from 
Colombia SNP NERONE 

MILLER, James S. A Revision of the New World Species of Ehretia (Bo- 
ETTO risa 

Moretti, ALDO, Paoro De Luca, JEAN PIERRE ScLavo & DENNIS WM. STE- 
VENSON. Encephalartos voiensis (Zamiaceae), a New East Central 
African Species in the E. hildebrandtii Complex 0... oc 

MonLEY, Thomas. New Species and Other Taxonomic Matters in the New 
World Memecyleae (Melastomataceae) |... LLL 

NEILL, DaviD A. & ELENA MARIA DE LAMARE OccHIONI. A New Species of 
Stryphnodendron (Fabaceae: Mimosoideae) from Amazonian Ecuador 

OccHIONI, ELENA MARIA DE LAMARE. (See David A. Neill € Eléna Maria 
de Lamare Occhioni) 1. LLL 

OYEWOLE, S. O. A New Species of Urginea (Liliaceae) in Nigeria 0 


PENG, CHING-I. The Systematics and Evolution of Ludwigia Sect. Micro- 
carpium (Onagraceae) ........... 


PLANCHUELO, ANA Maria & Davip B. Dunn. Two New Species of the 
Lupinus lanatus Complex Ls 
POwELL, A. MICHAEL. (See Harold Robinson, A. Michael Powell, Gerald D. 
Carr, Robert M. King & James F. Weedin) 
PRUSKI, Jonn F. Compositae of the Guayana Highland—II. Novelties in 
Gongylolepis and Stenopadus (Mutisieae) ooo 


1112 


1103 


1004 


RAKOTOZAFY, ARMAND. (See William G. D'Arcy € Armand Rakotozafy) .. 351 
Ramírez, CARLOS, ENRIQUE HAUENSTEIN, JosÉ SAN MARTÍN & DOMINGO 
CONTRERAS. Study of the Flora of Rucamanque, Cautín Province, Chile 444 
RAMÍREZ, Ivón. (See Germán Carnevali & Ivón Ramirez) ... 374 
Ramos, C. (See J. Márquez-Guzmán, M. Engleman, A. Martinez-Mena, E. 
Martinez & C. Ramos) iconos . 124 
Ramos, CraRA HiLDA. (See Esteban Martínez & Clara Hilda Ramos) .......... 128 
RENNER, SUSANNE S. A Survey of Reproductive Biology in Neotropical 
Melastomataceae and Memecylaceae ............—— 496 
RoniNsoN, HAROLD, A. MICHAEL POWELL, GERALD D. CARR, ROBERT M. KING 
& James F. WkEDIN. Chromosome Numbers in Compositae, XVI: 
Eupatorieae II n 1004 
Ropricues, WitLIAM A. A New Venezuelan Virola (Myristicaceae) .............. 1163 
ROGERS, GEORGE (EDITOR). Steyermark Recollections „t 627 
Romero, Gustavo A. & GERMAN CARNEVALI. Novelties in the Orchid Flora 
of Southern: Venezuela ii ee 454 
San Martín, José. (See Carlos Ramirez, Enrique Hauenstein, José San 
Martin & Domingo Contreras) a A nn 444 
ScLavo, JEAN PIERRE. (See Aldo Moretti, Paolo De Luca, Jean Pierre Sclavo 
emus Wm. Stevenson) ii o assi ann 934 
Shaw, Ropert B. (See Patricia P. Douglas, Robert B. Shaw & Victor E. 
Diersing) t 182 
SMITH, ALAN R. € CaroL A. Topzia. Augustus Fendler's Venezuelan Col- 
lections of Ferns and Fern Allies cc anet 330 
Stace, C. A. (See A. R. A. Alwan Al-Mayah & C. A. Stace) „u 1125 
STEIN, Bruce A. & HirosHi Tose. Floral Nectaries in Melastomataceae and 
Their Systematic and Evolutionary Implications ......... 519 
STEINER, KiM E. A Second Species of the Amphi-Atlantic Genus Alonsoa 
(Scrophulariaceae) in South Africa „uinn 11542 
STEVENSON, DENNIS WM. (See Aldo Moretti, Paolo De Luca, Jean Pierre 
Sclavo & Dennis Wm. Stevenson) ar ida 934 
STEYERMARK, JULIAN A., BRUCE K. Horsr & COLLABORATORS. Flora of the 
Venezuelan Guayana— VII. Contributions to the Flora of the Cerro 
Aracamuni,. Venezuela 2 cese reos ee 945 
TAKAHASHI, MASAMICHI & SHOICHI Kawano. Pollen Morphology of the Me- 
lanthiaceae and Its Systematic Implications „i 863 
TayLor, Mary Susan. Plant Taxa Described by Julian A. Steyermark ..... 652 
Tose, HirosHI. (See Bruce A. Stein & Hiroshi Tobe) eee 519 
Tobzia, CAROL A. Augustus Fendler’s Venezuelan Plant Collections ............ 310 
Topzia, CAROL A. A New Species of Hybanthus (Violaceae) from Panama 300 
Topzia, CAROL A. A Revision of Ampelocera (Ulmaceae) cian 1087 


Topzia, CAROL A. (See Alan R. Smith & Carol A. Todzia) ssh hl 330 
VAN DER WERFF, HenK. Novelties in the Lauraceae from Venezuelan Guay- 

A ———————— ÉÉ"-——— AMT 462 
VAN DER WERFF, HENK. A New Species of Persea (Lauraceae) from Suri- 

nam, With a Discussion of Its Position Within the Genus 22222 939 
VAN DER WERFF, HENK. A New Combination in Ocotea (Lauraceae) ...... 1175 
WASSHAUSEN, DIETER C. New Species of Mendoncia (Acanthaceae) from 

the Venezuelan Guayana 1. 1118 
WEEDIN, JAMES F. (See Harold Robinson, A. Michael Powell, Gerald D. 

Carr, Robert M. King & James F. Weedin) o 1004 
WiERSEMA, JouN H. A New Species of Cabomba (Cabombaceae) from 

Tropical ol "M 1167 
WILDER, GEORGE J. (See Barry E. Hammel € George J. Wilder) ................. 112 
ZIMAN, SVETLANA N. & CARL S. KEENER. A Geographical Analysis of the 

Family Ranunculaceae s LLL 1012 


Volume 76, Number 3, pp. 627-944 of the ANNALs OF THE MISSOURI BOTANICAL GARDEN was 
89. 


published on August 4, 19 


Rediscovery of Tetramolopium arenarium Subsp. arenarium var. arenarium (Asteraceae: 


Astereae) on the Pohakuloa Training Area, Hawaii Patricia P. Douglas, Robert 
Shaw & Victor E. Diersing 1182 

Miscellaneous Chromosome Counts in Asteraceae, Bignoniaceae, Proteaceae, and 
Fabaceae Peter Goldblatt 1186 


Lepidium boelckei and L. jujuyanum (Brassicaceae), New Species from Jujuy, 
Argentina Ihsan A. Al-Shehbaz 


Cover ilesustieo: Solanum toliaraea D'Arcy & Rakotozafy, by John Myers. 


CONTENTS 
Flora of the Venezuelan Guayana — VII. Contributions to the Flora of the Cerro Aracamuni, 
Venezuela Julian A. Steyermark, Bruce K. Holst & Collaborators -_. 945 


Compositae of the Guayana Highland— II. Novelties in Gongylolepis and PEE 
(Mutisieae) John F. Pruski 
Chromosome Numbers in Compositae, XVI: Eupatorieae II Harold Robinson, A. Mi- 


chael Powell, Gerald D. Carr, Robert M. King & James F. Weedin 1004 
A Geographical Analysis of the Family Ranunculaceae Svetlana N. Ziman & Carl S. 

Keener 1012 
A Revision of the New World Species of Ehretia (Boraginaceae) James S. Miller ....... 1050 
New and Critical Taxa of Euphorbiaceae from South America Michael J. Huft ............ 1077 
A Revision of Ampelocera (Ulmaceae) Carol A. Todzia 1087 
Pollen Morphology of the Shrubby lridaceae Nivenia, Klattia, and Witsenia Peter 

Goldblatt & John C. Manning ... .. 1103 


Seed Morphology of Sisyrinchium (Iridaceae-Sisyrinchieae) and Its Allies Peter Gold- 
- blatt, James E. Henrich & Richard C. Keating 1109 
New Species of Mendoncia race from the Venezuelan Guayana Dieter ae 
Wasshausen . : 
New Species, Names, and Combinagpus.] in baila Combretaceae A. R. A. Alwan 
Al- Mayah & e A. Stace .. 1125 
Solenum allophyllum (Miers) Standl. and the Generic Dio of Cyphomandra and 
; Solanum (Solanaceae) Lynn Bohs E 


_ Novelties in South American Malouetia (Apocynaceae) Mary E. Ehdress ........ 
Ticodendron: A New Tree from Contest America ae dae Gómez-Laurito & Luis D. 
E Góme. mez 1148 


i — d 


A Second Species of the Amphi-Atlantic Genus Alonsoa (Scrophulariaceae) in South 


| Africa. Kim E. Steiner s x odere n V 
E New Species of Isoëtes from Territorio Federal Amazonas, Venezuela E James : ie 
Hich Esc eee 


— 22 


A New. Coda: Virola: (Myristicaceae) William A. Rodrigues - uie 
i A New Species of Clitoria (Leguminosae) from Vepepielas Guayana * - Paul R. Faniz. fi 1165 P 
A New Species of Cabomba (Cabombaceae) f ll John H. Wiersema 1167 D + 


1169 23 


ams | 


e d 


o 


E ae = of shea (Lamiaceae) from e Venezuelan t Guayana Raymond M 
New Conkinatcis in | Sorghastrum (Poaceae: Anieopogonese) - : Panici a D. Davila — 18 
$ ; eodd malletii, a New Species in Seetion Decaloba  Possilraceae) from 
ee . Mesoamerica - John M. MacDougal .. anat Y 
a ^ New Combination i in Ocotea (Lauraceae) | Henk van de "v 
‘Sis ES erequipuiin. risiceceno) a New. a from Peru = Ihsan P AL 


ae