Annals
of the

Missouri Botanical

Garden

CE

Volume XLV
1958

Published quarterly at Galesburg, Illinois, by the Board of Trustees of
the Missouri Botanical Garden, St. Louis, Mo.

Entered as second-class matter at the post-office at Galesburg, Illinois,

under the Act of March 3, 1879.

MISSOURI BOTANICAL
GARDEN LIBRARY

Annals
of the

Missouri Botanical Garden

A Quarterly Journal containing Scientific Contributions from the
Missouri Botanical Garden and the Henry Shaw School of Botany of
Washington University in affiliation with the Missouri Botanical

Information
The ANNALS OF THE MISSOURI BOTANICAL GARDEN appears four times
during the calendar year: February, May, September, and November. Four
numbers constitute a volume.
Beginning with Volume 45, 1958:
Subscription Price ___ $12.00 per volume
Single Numbers |... — 3.00 each

Contents of previous issues of the ANNALS OF THE MISSOURI BOTANICAL
GARDEN are listed in the Are Index, published by the H. W. Wilson
Company.

TABLE OF CONTENTS
Flora of Panama. Part VII, Fascicle 1 (Passifloraceae to Cactaceae) ._.._.
— Robert E. Woodson, Jr., Robert W. Schery and collaborators

Flora of Panama. Part VII, Fascicle 2 (Thymelaeaceae to Myrtaceae) _..

— Robert E. Woodson, Jr., Robert W. Schery and collaborators

Flora of Panama. Part VII, Fascicle 3 (Melastomataceae) ----------------------

Robert E. Woodson, Jr., Robert W. Schery and collaborators

Syntbyris li F. W. Went

The New World Species of Cynometra

z John D. Dwyer

General Index to Volume XLV

1-91

93-201

203-304

305-312

313-345

347-354

MISSOURI BOTANICAL GARDEN

EDGAR ANDERSON,
Curator of Useful Plants

HENRY N. ANDREWS,
Paleobotanist

HucH C. CUTLER,
Executive Director

CARROLL SX DoDGE,
Mycologi

STAFF

Director
Frits W. WENT
Oscar E. GLAESSNER,
Controller

Norton H. NICKERSON,
Morphologist
TRIFON VON SCHRE

Eas ome Curator greta of
onomic Plants

JULIAN A. STEYERMARK,
Honorary Research Associate

RGE B. VAN SCHA

ROBERT L. DRESSLER,
GEO
Cum ator of Grasses = aa

Taxonomist and dito?
of the ANNALS

Joun D. Dw

Research idm

ROBERT E. Woopson, JR.,

Curator of Herbarium

BOARD OF TRUSTEES

Presiden
ROBERT BROOKINGS SMITH
Vice-President
LEICESTER B. FAUST
Second Vice-President
HENRY B. PFLAGER
DANIEL K. CATLIN JOHN S. LEHMANN
DuDLEY FRENCH CHARD
HENRY HITCHCOCK
WARREN McK. SHAPLEIGH

EX-OFFICIO MEMBERS
STRATFORD LEE Piscina
President of the Academy of
Science of St. Louis

ETHAN A. H. SHEPLEY,

ARTHUR C. LICHTENBERGER,
Bishop of the Diocese of
Missouri

JAMES J. MCCAFFERY,
m si z E Board ai Chancellor of Washington
Educa of St. Louis University

RAYMOND R. TUCKER,
Mayor of the City of St. Louis

Oscar E. GLAESSNER, Secretary

Volume XLV Number 1

Annals
of the

Missouri Botanical

Garden

| FEBRUARY, 1958 —

Flora of Peak. Part. VII, Fascicle 1 (Passifloraceae to Cactaceae) ge
R Sob E. docente = Robert W. Schery and Collaborators 1-91

„PUBLISHED de AT CALESEURO, ILLINOIS e
BY THE RARE TEEN cen gut som | BOTANICAL | GARDEN

Vol. XLV

FLORA OF PANAMA
BY

ROBERT E. WOODSON, JR.
AND
ROBERT W. SCHERY

AND COLLABORATORS

PART VII

Fascicle 1

PASSIFLORACEAE

CARICACEAE

LOASACEAE

BEGONIACEAE (L. B. Smith and B. G. Schubert)
CACTACEAE

ANNALS
OF THE
Missouri BOTANICAL GARDEN

FEBRUARY, 1958

FLORA OF PANAMA

Part VII. Fascicle 1*

PASSIFLORACEAE
1. PASSIFLORA L.
PASSIFLORA L. Sp. Pl. 955. 1753; Killip, in Field Mus. Publ. Bot. 19:1-613. 1938.

Granadilla [ Tourn. ex Rupp.] ue Fam. 2:408. 1763.
Cieca Medik. Malvenfam. 97. 1

Abe A. Bory i Ann Gén. Sci. Phys. 2:138.

Antbactinia Bory, loc. cit. 135. 1819.

Polyantbea DC. in Mém. Soc. Phys. Genéve 1:435. 1822.
Disemma Labill. Sert. Austro-caled. 78. 1824.
Astropbea Rchb. Consp. 132. 1828.

epocha Raf. Fl. Tellur. 4:103. 1836.

Ma or 836

Ceratosebalum Oerst. Rech. Fl. La Centr. 18. 1863.

Herbaceous or woody lianas, usually climbing by tendrils, rarely erect herbs,
shrubs, or small trees. Leaves alternate, petiolate, basifixed to peltate, simple or
very rarely compound, entire to deeply lobed, occasionally with large irregular
glandular areas (ocellate) upon the blade; petiole frequently with few to several
more or less conspicuous glands. Tendrils volubile, solitary, simple, axillary or

* Assisted by a grant from The National Science Foundation.

Issued March 20, 1958.

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[Vol, 45
2 ANNALS OF THE MISSOURI BOTANICAL GARDEN

produced from the inflorescence. Inflorescence axillary, usually 1-flowered, less
frequently cymose and few-flowered, the peduncle jointed and there producing
usually 3 inconspicuous and setaceous to large and petaloid involucrate bracts.
Flowers hermaphrodite, actinomorphic, frequently large and showy, perigynous.
Hypanthium shallow to relatively deep, the sepals 5, valvate, usually with a cornic-
ulate process toward the tip in the large-flowered species; petals 5, rarely absent,
inserted at the margin of the hypanthium, green to highly colored; corona of
numerous petaloid filaments in 1 to several centripetally decreasing series inserted
on the margin of the hypanthium at the base of the corolla. Pistil and stamens
borne upon a more or less elongate common androgynophore; stamens 5, borne
immediately beneath the pistil, the anthers 2-celled, versatile; pistil 3- or very
rarely 4-carpellate, the ovary 1-loculate with 3 or very rarely 4 parietal placentas
bearing numerous ovules, the 3—4 stigmas usually quite sessile, rather elongate,
radial. Fruit an inflated berry, frequently large, globose to fusiform, containing
numerous small hard lenticular seeds immersed in abundant mucilaginous pulp
derived from the testa. |

Although the Passifloraceae include a few small genera of the Old World as
well, the large genus Passiflora is confined to the Americas in its indigenous distri-
bution, although numerous species with showy flowers and edible fruits have been
introduced into the tropics of the Eastern Hemisphere and are frequently culti-
vated in greenhouses in the colder latitudes. In America native species are found
from the south-central United States to Argentina.

Their extremely complicated and frequently large and brightly colored flowers
have attracted several botanists to the special study of the Passifloras, the most
recent of whom is E. P. Killip. In his study of Passiflora, Killip recognizes 353
species which are distributed amongst no less than 22 subgenera, chiefly based upon
rather intricate differences in flower structure. Twenty-six species have been
discovered in Panama to which at least a few additions undoubtedly will be made
in the future. The present account has been drawn very largely from that of
Killip’s monumental work, although the keys are original and the species concept
has been changed in a few minor instances. The authors of this Flora are grateful
to Mr. Killip for the generous cooperation which he has given them in the naming
of their study material over many years.

Early Spanish missionaries to the New World saw in the radiant flowers of
Passiflora a symbol of the Crucifixion, and from this inspiration the Spanish popular
name pasionaria and the English ‘“Passionflower” have had their origin. Less rev-
erent Spanish names of frequent application to various species are calzoncillo and
ala de murciélago. 'The large pulpy berries of various species, particularly the
frequently cultivated P. edulis and P. quadrangularis are known as granadillas and
are eaten raw or more often cooked with milk to make a kind of dessert.

a. Flowers typically rather — mostly less 5 in diameter
usually paired or several at the nodes Cassidy soley in xk punctate);
sepals not corniculate t a the tip; bracts — and setose, very

inconspicuous; petiolar Puede usually 2 when presen

(2)

1958]
FLORA OF PANAMA (Passifloraceae)

b. Flowers 2 upon a common elongate peduncle, the associated tendril
terminal; plants Por and very glaucous; leaves small, sub-
orbicular, e wył or nearly so; petiole oh 2 foveate glands at the
A: p + blade [ceci NL pU:

. Flow or ral upon paired (occasionally solitary) peduncles,
the UM ki basal.

c. Petioles eglandular

d a owers

c
ia

y or li upon each peduncle; plants densely
dd. Flowers solitary upon each peduncle; plants glabrous or very
inconspicuously pubescent.
e. Leaves basifived, stipules minute and se
f. Peduncles jointed at about the middle. or somewhat below;
flowers usually paired at the nodes.
ff. Potencia jointed far above the middle, almost directly
beneath the fl

ower.

g. za about twice as long as broad or nearly so, very
broadly 3-lobed "ide dir central lobe st asc occasion-
ally nearly entire; flowers usually paired at the nodes.......

88. Leaves about as broad as long; flowers usually eid at

h. Leaves w ith 2 strongly pra eteri lobes, rarely
with a weak central lobe;

eb sai equally 3-lo sk La s unb c
g- Leaves uch broader than long, with 2 ipe transverse
obes; flowers usually solitary at the nodes...............
ee. Leaves eccentrically peltate, entire or if 3-lobed the central
lobe far stronger; e large and bezie amplexicaul,
ciliate-serrate, deciduo

cc. Petioles glandular
es cccentrically peltate, deeply 3-lobed, the lateral lobes

nearly transverse and as strong as the «a to much stronger;
stipules pochód and subsetaceous
dd. Leaves basifixed.
e. Flowers typically 2 or more upon a short common peduncle;
leaves broadly 3-lobed, the lobes rounded, i mu-
cron „ape rd central = stronger; petiolat landed inconspic-

uou oadly poculifor
ee. "ne “solitary upon Qu peduncle; leaf lobes acute to
acumi j ele

pe ic
f. Leaves entire or subentire with the lateral hub reduced to
inconspicuous angles; petiolar piede Pind large and auric-
ular, borne toward the base of the
ff. Leaves pera ely 3-lobed; seria) sa Eme inconspic-
us, b out tdo or higher upon the petiole.
M Pa glande b about midway upon „gore petiole;
, linea

Bg. Petiolar ‘glande poppa almost directly beneath the leaf
tals a

ade; peta
aa. — typically rather ag, torii more than 5 cm. in diameter, kę
h es (except occasionally in P. a); sepals us
cicle corniculate toward the E IA in P. oe Li j
membranacea) ; bracts conspicuous an ucrate, foliaceous or peta
ceous; petiolar glands usually more chan 2 i» present.
b. Involucral bracts entire, serrulate, giam ane
c. Sepals not corniculate; pata pot sland dular
Leaves cuneately 2-lobed, occasionally w ich a small — uen
tipules inconspicuous and inlata taceous; involucral bra

vate, obtuse or acute at the base, 1.0—1.5 cm. ct ni asia
4-6 cm. in diameter

E
3
8

uborbicular, irita i 3-lobed; stipules conspicuously
E involuc nb racts suborbicular, „dee ei A ordate, about
about 7.

; flow
c. Sepals brine irs i inia siii. t.

(3)

. P. TRYPHOSTEMMATOIDES
2. P. SEXFLORA
. P. BIFLORA
4. P. LANCEARIA
APETALA
6. P. PANAMENSIS

. P. PUNCTATA

. P. HAHNII

. P. CORIACEA

. P. HOLOSERICEA

. P. AURICULATA

. P. PEDICULATA

. P. suBEROSA

14. P. PULCHELLA

5. P. MEMBRANACEA

[Vol. 45
4 ANNALS OF THE MISSOURI BOTANICAL GARDEN

d. Leaves palmately m to ree to below the middle, the central
ard the base.

e. de olucral bes semis shorter sper the penis free and never
ca” = da bud, s e or lacer

f£ 7-lobed pats E lobed) paz glands

ver n yá ad prominently stipitate, bor most directly

beneath gm leaf “blade; ias Gace re ine

urple and whit 16. P. ADENOPODA
ff. Leaves 3-lobed; petiolar glands inconspicuous and sessile,
borne toward s base of the petiole; involucral bracts
yellow

glandular-serrulate; aa scarlet an 17. P. VITIFOLIA
ee. Involucral bracts almost as long as the sepals, united toward
the > „and completely enveloping the flower bud, entire;
flowers violet and white 18. P. WILLIAMSII
dd. Leaves p or palmately 3-lobed above or about s middle
and with the central lobe not constricted toward the bas
e. Toni 3-lobed.
f. Plants more or less on leaves lobed above the faery the

central lobe much stronger; petioles with 2 to several pairs
of glands; da: mt greoi rather narrowly Ak p foli-
aceou 9. P. MENISPERMIFOLIA

ff. Plants genes leaves lobed to about the middle, the 3 lobes
subeq with 1 pair o „kij ds; involucral bracts

o ova E somewhat petalaceor . P. SUBPELTATA
. Leaves entire, deeply cordate i in some speci
co n ules very z pn foliaceo

eous.
Sti pules 2 ilateral, deciduous; involucral bracts about as
ong as the sepals

h. Stems not winged; leaves deeply cordate; petioles with
filiform glan 21. P. LIGULARIS
hh. Sun » prominently wi inged; leaves rounded at the base
to obscurely cordate; petioles with ovoid glands............. 22. P. QUADRANGULARIS
gg. Stipules "ut dada I aisi 3 taca bracts
about half as long as the P. OERSTEDII
ff. Stipules i kawa: and s eous or barely foliaccous,

g. Leaves obtuse or very Cs ci cordate at the base; in-
volucral bracts free
. Leaves enun petioles glandular at the middle or
below; flowers maroon, pur rple and white 24. P. AMBIGUA

middle; flowers white sk. purp 25. P. NITIDA
gg. Leaves deeply cordate at the pe petioles glandular
a ectly beneath the leaf blade; involucral bracts united
t the base; eons pute and white. 26. P. SEEMANNII
bb. d ika complexly ri -pinnatifid, each filiform division
gl ; plants i petioles with PENSO. gland-tipped
hairs; ie greenish white more or less tinged with purple............... 27 P. FOETIDA

Other species, at present known from Costa Rica and Colombia, are to be
expected in Panama as well.

1. PASSIFLORA TRYPHOSTEMMATOIDES Harms, in Engl. Bot. Jahrb. 18: Beibl.
46:6. 1894; Killip, in Field Mus. Publ. Bot. 19:81. 1938.

Passiflora gracilimma Killip, in Journ. Wash. Acad. Sci. 14:112. fig. 2b. 1924.

Plants slender, climbing, glabrous and very glaucous throughout. Leaves long-
petiolate, simple, entire, suborbicular, about 1.5—4 cm. long and 1.5—3.0 cm. broad,
very pale beneath, darker above; petioles very slender, about 1-2 cm. long, bearing
2 rather inconspicuous foveate glands at the juncture with the leaf blade; stipules
setose, very inconspicuous. Inflorescence of 2 greenish white flowers upon an

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1958]
FLORA OF PANAMA (Passifloraceae) 5

elongate common peduncle, the associated tendril terminal; peduncle 0.5-4.5 cm.
long; pedicels 1.5-4.0 cm. long, jointed somewhat above the middle, the bracts
minutely setose, deciduous. Flowers 1.5-4.0 cm. in diameter; sepals oblong, about
0.7-1.5 cm. long, not corniculate; petals narrowly oblong, 0.5-1.0 cm. long; fila-
ments of the corona in a single series, filiform, 3-5 mm. long. Berry subglobose to
ovoid, 3-6 cm. long, greenish yellow.

Panama and Colombia, in wet forests of middle elevations.

BOCAS DEL TORO: Fish Creek Hills, Von Wedel 2421, 380. cHIRIQUI: exact locality
uncertain, Hart 104.

There seems to be little room for doubt that P. trypbostemmatoides and P.
gracilimma are conspecific (cf. Killip, in Field Mus. Publ. Bot. 19:82. 1838),
since they are not closely related to other species, their characters overlap or at
least intergrade, and both have been reported from closely adjacent stations both
in Panama and Colombia. No intermediate stations between Bocas del Toro
(probably Hart's "Chiriqui") and the Atrato valley in Colombia have been dis-
covered as yet, but they are to be expected since the Caribbean coast of Panama
still is known so poorly.

2. PASSIFLORA SEXFLORA Juss. in Ann. Mus. Hist. Nat. 6:110. pl. 37. fig. 2.
1805; Killip, in Field Mus. Publ. Bot. 19:129. 1938.
Passiflora pannosa J. E. Sm. in Rees, Cycl. 20: Pond n 28. 181
Passiflora capsularis var. geminiflora DC. Prodr. 3:325. 1828 (as ais.
Meioperis — (J. E. Sm.) Raf. FI. Tellur. 4:105. $
Cieca pannosa (J. E. pozy M. Roem. Fam. Nat. Syn. 2:148. 1846.
Decaloba po i (Juss.) M. Roem. loc. cit. 164. 1846.
Passiflora floribunda Lem. FI. des Serres A e dt 1348.
Passiflora triflora Macf. Fl. Jam. 2:149.
Passiflora miraflorensis Killip, in Journ. Wash, Acad. Sci. 14:109. 1924.
Passiflora isotriloba Cuf. Archivio Bot. 9:1
Plants rather slender, climbing, m n. throughout. Leaves petio-
late, simple, basifixed, subtrapezoid, the top saliently and acutely 2-lobed, occa-
sionally with a smaller central lobe, the base rounded or slightly cordate, 3-9 cm.
long, 4-10 cm. broad; petioles 1-3 cm. long, eglandular; stipules subulate, about
5 mm. long. Inflorescence bearing 2-10 greenish-white, purple-tinged flowers
upon a common peduncle 2-3 cm. long, the associated tendril basal; pedicels 1.0—
1.5 cm. long, subtended by an inconspicuous subulate bract. Flowers 2-3 cm. in
diameter; sepals oblong-lanceolate, 8-15 mm. long, not corniculate; petals linear,
about 10 mm. long; filaments of the corona in 2 series, the outer about as long as
the petals. Fruit subglobose, 5-10 mm. in diameter, greenish yellow.
Southern Florida; Mexico to northern Colombia; Greater Antilles. Clearings
in wet forests, 1300-2000 m. in Panama.

BOCAS DEL 2 Robalo Trail, northern slopes of Cerro Horqueta, am 28.
CHIRIQUÍ: ‘= D Bauer District, Davidson 491; vicinity of Bajo Mon d Gue-
brada Chiqu aa Schery 519; valley of the upper Rio Chiriqui Viejo, > vicinity

of Monte Lirio, Seibert cg Río Ladrillo, above El Boquete, Pittier 3286.

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[Vol. 45
ANNALS OF THE MISSOURI BOTANICAL GARDEN

| M fh W,

SZJ

Fig. 1. Passiflora sexflora

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1958]
FLORA OF PANAMA (Passifloraceae) 7

3. PASSIELORA BIFLORA Lam. Encycl. 3:36. 1789; Killip, in Field Mus. Publ. Bot.
19:185. 1938
zy kę lunata J. E. Sm. Icon. Pl. Rar. 11: bl. 7. 1790, non Juss. nec Vell. necque Poepp.

Passiflora ślebrolu HBK. Nov. Gen. & Sp. 2:135. ;
Cieca glabrata (HBK.) M. Roem. Fam. Nat. Syn. 2:143. 1846.
1846.

i 46.
Passiflora lunata var. costata Mast. in Mart. Fl. e Pe :552. 1872.
i 2;

apere o brighami S. Wats. in Proc. Amer. gu we 1473.1887.

assiflora transversa Mast. in Bot. Gaz. 16:7.

Plants rather slender, climbing, cu; glabrous throughout, the stems con-
spicuously striate. Leaves rather shortly petiolate, simple, basifixed, subtrapezoid,
the top saliently and obtusely 2-lobed, occasionally with a smaller central lobe
or mucro, the base obtuse to broadly rounded, 1-10 cm. long, 5-10 cm. broad,
conspicuously glandular-ocellate; petioles 0.5-3.0 cm. long eglandular; stipules
minutely setose. Inflorescences bearing solitary greenish white or yellow flowers
upon paired peduncles, the associated tendril basal; peduncles about 1.0—1.5 cm.
long, jointed at about the middle or somewhat below; bracts very inconspicuous,
subsetaceous. Flowers 3-4 cm. in diameter; sepals ovate-lanceolate, about 1 cm.
long, not corniculate; petals oblong-lanceolate, slightly shorter than the sepals;
filaments of the corona in 2 series, the outer about as long as the petals. Fruit
subglobose, 1-2 cm. in diameter, greenish yellow.

Mexico to Colombia and Venezuela; Bahamas; from near sea level to about 1,500
m. Widely distributed and common in Panama.

BOCAS DEL TORO: Shepherd Island, Von Wedel 2694, SĘ Water Valley, Von Wedel

998, 1697; Snapper Point, Von Wedel 2645; ; vicinity of Chiriquí Lagoon, Von Wedel 384,
1218, 1305. CANAL ZONE: Ancón Hill, Y oodson, Allen 8 Seibert a ape Fendler

worth & Vestal 307. CHIRIQUÍ: El ide Kill 3608; David, Killip si * cocLé:
Penonorb, ent 226. COLÓN: Porto Bello, Pittier 2461; Catival, Standley 30349.
ANAMÁ: Panamá Vieja to Bella Vista, Allen $. 26; Las Sabanas, Standley 2 25835; Taboga
and, Standley : as, Pittier 3607; San José £ Island, Erlanson 183, 219, Johnston 809,
1189, 549; Tumba Mue de Rood nea isis Standley 29802; Trapiche Island, Miller
1893; Taboguilla Island, "Miller 2011
Popular names of this Wind climber are guate-guate, camacarlata, and cal-
zoncillo.

4. PASSIFLORA LANCEARIA Mast. in Journ. Bot. 23:114. 1885; Killip, in Field
Mus. Publ. Bot. 19:160. 1938.
Passiflora talamancensis Killip, in Journ. Wash. Acad. Sci. 12:260. 1922.
Rather massive climbers, essentially glabrous throughout. Leaves rather long-
petiolate, simple, basifixed, oblong-elliptic and entire to the tip or very broadly

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[Vol. 45
8 ANNALS OF THE MISSOURI BOTANICAL GARDEN

3-lobed with the central lobe stronger than the lateral, obtuse to rounded at the
base, 5—12 cm. long, 2-9 cm. broad, membranaceous to subcoriaceous, rather con-
spicuously glandular-ocellate; petioles 1—4 cm. long, eglandular; stipules subseta-
ceous, inconspicuous. Inflorescences bearing solitary greenish-white, faintly
purple-tinged flowers upon usually paired peduncles, the associated tendril basal;
peduncles about 2-5 cm. long, jointed far above the middle almost directly beneath
the flower; bracts inconspicuously subsetaceous. Flowers 3—4 cm. in diameter;
sepals ovate-lanceolate, 1.0—1.5 cm. long, not corniculate, glabrous to more or less
puberulent within; petals about half to one third as long as the sepals; filaments of
the corona in 2 series, the outer about as long as the petals. Fruit subglobose, 2—5
cm. in diameter.

Costa Rica and Panama, from near sea level to about 1,700 m. altitude.

AS DEL TORO: Old Bank Island, Von Wedel 2120. cmiriquí: Bajo Chorro, Boquete

ha Davidson 336.

This is a representative of a very complex group of species the affinities of which
are far from clear. Perhaps our plants should be referred to P. mollis HBK.

5. PASSIFLORA APETALA Killip, in Journ. Wash. Acad. Sci. 12:255. 1922.

Plants rather slender, climbing, glabrous throughout. Leaves rather long-
petiolate, simple, basifixed, broadly cuneate-subtrapezoid, with 2 strongly ascending
lateral lobes, rarely with a weak central lobe, broadly obtuse or rounded at the base,
3-10 cm. long and broad, inconspicuously glandular-ocellate; petioles 1-5 cm.
long, eglandular; stipules minute and subsetaceous. Inflorescences bearing solitary
yellowish green flowers upon paired peduncles, the associated tendril basal; pe-
duncles about 1.5—4.0 cm. long, jointed above the middle; bracts inconspicuous and
subsetaceous. Flowers about 2 cm. in diameter; sepals narrowly oblong-lanceolate,
about 1 cm. long, not corniculate; petals absent; filaments of the corona in 1 series,
about half as long as the sepals or less. Fruit globose, about 1 cm. in diameter, dark
purple. :

Costa Rica and Panama, in clearings of forests at 1,000—2,000 m.

CHIRIQUÍ: trail from Bambito to Cerro Punta, Allen 316; Chiquero, Davidson 511;

Bajo Mona, mouth of Quebrada Chiquero, Woodson, Allen 65 Scibert 1016; Bajo Mona and
ae ee Chiquero, Woodson & Schery 583; vicinity of "New Switzerl and”, central Vip
Chiriqui Viejo, Allen 1346; valley of upper Rio Chiriqui Viejo, vicinity of Mon
um ‘Seibert 160, G. White 25; Alto m Bro. Maurice 835.

6. PASSIFLORA PANAMENSIS Killip, in Journ. Wash. Acad. Sci. 12:259. 1922.

Plants rather slender, climbing, glabrous throughout. Leaves long-petiolate,
simple, basifixed, broadly 3-lobed, the lobes ascending, acute, and essentially equal,
broadly rounded at the base, 5-12 cm. long and broad, conspicuously glandular-
ocellate; petioles 1.5—3.0 cm. long, eglandular; stipules inconspicuous and sub-
setaceous. Inflorescences bearing solitary greenish lavender-tinged flowers upon
paired peduncles, the associated tendril basal; peduncles 2-4 cm. long, jointed
almost immediately beneath the flower; bracts minute, subsetaceous. Flowers about

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1958]
FLORA OF PANAMA (Passifloraceae) 9

3 cm. in diameter; sepals oblong-lanceolate, 1.0—1.5 cm. long; petals oblanceolate,
about half as long as the sepals; filaments of the corona in 2 series, the outer about
as long as the petals. Fruit globose, about 2 cm. in diameter, purple.

Eastern Panama and adjacent Colombia, in lowland forests.

DARIÉN: along Sambú River, Pittier 5556.

7. PASSIFLORA PUNCTATA L. Sp. Pl. 957. 1753; Killip, in Field Mus. Publ. Bot.
19:183. 1938
Pa ide gis mis era HBK. Nov. Gen. & Sp. 2:136. 1817.
Passiflora miliana Bory, in gus: Gen. Sci. eee Brux. 2:149. pl. 24. 1819.
Passiflora vespertilio Ker, Bot. Reg. 7: pl. 597. 1821, non
Passiflora discolor Link & Otto, iu PL opera 13: 2 5. 1828.
Passiflora retusa Hook. Bot. Misc. 3:3
Decaloba punctata (L.) M. Roem. Baia. "Nat. Po 2:156. 1846.
Cieca discolor (Link & Otto) M. Roem. loc. cit. 140. 1846.
Cieca misera (HE. ) M. Roem. loc. cit. š
a Mast. in Mart. Fl. Bras. 131:556, 593. 1872.

usby,
Passiflora longilobis Mane i in in Link. Tolige. Marta Gs dod; none $; qi 1:71. pl. 63.
1910.

Slender climbers, glabrous or essentially so. Leaves petiolate, simple, basifixed,
much broader than long, with 2 nearly transverse lateral lobes and usually a slender
central mucro or rarely a small central lobe, 0.5—3.0 cm. long, 3-10 cm. broad,
conspicuously glandular-ocellate; petioles 1—6 cm. long, eglandular; stipules minute
and subsetaceous. Inflorescence usually solitary and bearing a single greenish white,
occasionally purple-tinged flower, the associated tendril basal; peduncle 3-10 cm.
long, jointed almost immediately beneath the flower; bracts inconspicuous and
setaceous. Flowers about 2—4 cm. in diameter; sepals ovate-lanceolate, 1-2 cm.
long; petals narrowly oblong, 8—12 mm. long; filaments of the corona in 2 series,
the outer almost equaling the petals. Fruit globose, 1.0—1.5 cm. in diameter,
purple.

Panama to Argentina, at low elevations in thickets and forest clearings.

CANAL ZONE: Barro Colorado Island, Woodworth & Vestal 457, 717, Bailey & Bailey
258, Kenoyer 456, Wetmore 6 Abbe 26, Bangbam 574, Hunnewell 19029; Monte Lirio,
Max pe don between Frijoles and Monte Lirio, Killip 12187; Trinidad River, Pittier 3998.
PANAMA: Mindi, Cowell 169; Rio Mindi, Sutton Hayes 596; Baillemona, Stevens 6

It is questionable whether P. misera can be maintained as a species distinct $e:
P. punctata since their general aspect, variational tendency, and geographical dis-
tribution have so very much in common. ey were kept separate by Killip,
however, who was of the opinion that the inner corona filaments are linear, broadly
dilated at the apex and often lobulate in P. misera and filiform or capillary, rarely
minutely capitellate in P. punctata.

8. PASSIFLORA HAHNII (Fourn.) Mast. in Mart. Fl. Bras. 13':569. 1872.

Disemma Habnii Fourn. in Rev. Hort. 41: pl. 430. 1
Passiflora ui S. Wats. in Proc. Amer. py = 473. 1887.

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[Vol. 45
10 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Rather slender climbers, glabrous throughout. Leaves petiolate, simple, very
eccentrically peltate, broadly ovate, essentially entire or if 3-lobed the central lobe
far stronger and the lateral relatively inconspicuous, the base truncate or broadly
rounded, 5-8 cm. long and broad; petioles 2-3 cm. long, eglandular; stipules large
and foliaceous, inequilaterally reniform, amplexicaul, ciliate-serrate, about 1 cm.
long and 2 cm. broad, deciduous. Inflorescences bearing solitary yellowish-white
flowers upon paired or solitary peduncles, the associated tendril basal; peduncles
1.5-2.0 cm. long, jointed almost immediately beneath the flower; bracts minute
and subulate. Flowers about 4-6 cm. in diameter; sepals oblong, 2-3 cm. long, not
corniculate; petals similar to the sepals; filaments of the corona in 2 series, the outer
about 1.5 cm. long. Fruit globose, 3-4 cm. in diameter, deep purple.

Southern Mexico to Colombia, at low to middle elevations.

COCLE: mountains beyond La Pintada, 400-600 m., Hunter & Allen 629.

9. PASSIFLORA CORIACEA Juss. in Ann. Mus. Hist. Nat. 6:109. pl. 39, fig. 2. 1805;
Killip, in Field Mus. Publ. Bot. 19:83. 1938.

Passiflora diformis HBK. Nov. Gen. & Sp. 2:136. 1817.

Passiflora clypeata J. E. Sm. in Rees, Cycl. 26: Passiflora no. 20. 1819.
Monactineirma coriacea (Juss.) Bory, in Ann. Sci. Gén. Phys. xi 2:138. 1819.
Cieca difformis (HBK.) M. ce Fam. Nat. Syn. 2:140.

Cieca coriacea (Juss.) M. Roem. loc. cit. 148. 1846.

Passiflora obtusifolia Sessé & Moc. pl. Nov. Hisp. 156. 1887.

Rather extensive climbers inconspicuously puberulent to glabrate. Leaves peti-
olate, simple, very eccentrically peltate, deeply 3-lobed, the lateral lobes rather
abruptly acuminate and nearly transverse, as strong as the central lobe to much
stronger, 3-7 cm. long, 6-25 cm. broad; petioles 2-4 cm. long, bearing 2 ocellate
glands near the juncture with the leaf blade; stipules narrowly linear, about 0.5 cm.
long. Inflorescence a terminal 2- to several-flowered raceme, with slightly foli-
aceous, frequently 3-lobed bracts subtended by paired ocellate glands; pedicels
about 5 mm. long. Flowers 2.5-3.5 cm. in diameter, greenish or yellowish white;
sepals oblong-lanceolate, 1.0-1.5 cm. long, not corniculate; petals none; filaments
of the corona in 2 series, the outermost 7-8 mm. long. Fruit globose, 1-2 cm. in
diameter, deep blue.

Mexico to British Guiana and Bolivia, from near sea level to about 2,000 m.
CANAL ZONE: Cocoli Island, Miraflores Lake, P. White 86, G. W bite 133; Las Cascadas,
Standley 29594; Darién Station, Standley 31617; Empire to Mandinga, Piper 5479; Barro

Colorado Island, Shattuck 57, Woodworth & Vestal 501. cocLt: El Valle de Antón,
Allen 1667, 4473. PANAMA: Alhajuela, Pittier 3456.

In some parts of Central America this species is known as murciélago and ala
de murcićlago with reference to the peculiar shape of the leaves.

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1958]
FLORA OF PANAMA (Passifloraceae) 11

10. PASSIFLORA HOLOSERICEA L. Sp. Pl. 958. 1753; Killip, in Field Mus. Publ. Bot.

19:99. ,1998.

Decaloba holosericea (L.) M. Roem. Fam. Nat. Syn. 2:164. 1846.

Rather coarse climbers, densely puberulent throughout. Leaves petiolate,
simple, basifixed, broadly 3-lobed, the lobes rounded and occasionally mucronulate,
the central lobe far stronger, the base rounded to somewhat cordate, 5-10 cm.
long, 4-8 cm. broad; petioles 1-3 cm. long, the glands rather inconspicuous and
broadly poculiform; stipules inconspicuous and filiform. Inflorescences usually
paired, corymbiform, typically bearing 2 or more greenish white or yellow flowers
upon a very short common peduncle, the associated tendril basal; common peduncles
2-4 mm. long; pedicels about 1 cm. long; bracts inconspicuous and filiform
Flowers about 2-4 cm. in diameter, sepals oblong-elliptic, about 1.5 cm. long;
petals broadly elliptic, about 1 cm. long; filaments of the corona in 2 series, the
outer nearly as long as the petals, flushed with yellow and purple. Fruit globose,
about 1.5 cm. in diameter.

Southern Mexico to Colombia and Venezuela; Cuba. Lowland thickets to about
700 m.

CHIRIQUÍ: vicinity of Puerto Armuelles, Woodson & Schery 816.

11. PASSIFLORA AURICULATA HBK. Nov. Gen. & Sp. 2:131. 1817; Killip, in Field
Mus. Publ. Bot. 19:122. 1938.
Passiflora appendiculata G. F. W. Mey. Prim. FI. pei d 223. 1818.
Passiflora cyathophora Desv. in Ham. Prodr. 48.
Passiflora Robrii DC. Prodr. 3:326. 1828.
Passiflora cinerea Poepp. & Endl. Nov. Gen. & Sp. 2:57. n I 77. 1838.
Cieca auriculata (HBK.) M. Roem. Fam. Nat. Syn. 2:14
Cieca appendiculata (G. F. W. Mey.) M. Rowi. loc. cit. ate EE
j 4

Passiflora torta Mast. in Mart. Fl. Bras. 131:548. 187
Passiflora cayaponioides Rusby; in Bull. N. Y. Bot. Gard. 8:107. 1912.
Passiflora AE tt Hoehne, in Comm. Linh. Telegr. Matto Grosso Ann. Bot. 5:76.

Rather coarse climbers, finely puberulent when young, becoming glabrate.
Leaves petiolate, simple, basifixed, broadly ovate to ovate-lanceolate, entire or sub-
entire with the lateral lobes reduced to inconspicuous angles, 4-15 cm. long, 2-10
cm. broad, conspicuously ocellate; petioles 1-3 cm. long, bearing 2 very large and
auriculate glands near the base; stipules setaceous, up to about 1 cm. long. In-
florescences in pairs, bearing a single greenish yellow, purple-tinged flower; peduncle
about 1 cm. long, jointed above the middle; bracts inconspicuous and setose.
Flowers about 2 cm. in diameter, greenish yellow or white and purple-tinged toward
the base; sepals narrowly oblong-lanceolate, about 1 cm. long, not corniculate;
petals linear, about half as long as the sepals; filaments of the corona in 2 series,
the outer about as long as the sepals. Fruit globose, 1.0-1.5 cm. in diameter,
greenish yellow.

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[Vol. 45
12 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Nicaragua to Bolivia and Brazil, from about sea level to 1,200 m.
BOCAS DEL TORO: vicinity of Chiriqui Lagoon, Von Wedel 1202; Water Valley, V
Wedel e. CANAL ZONE: Chagres, Fendler 122; Gatün, Sutton Hayes 463; Agua Clara,
t n, Standley 30949; Agua Clara Reservoir, Stevens 595; Barro Colorado Island,
Standley 4 tenis Zetek 4364. DARIEN: Cana, Williams 964.

12. PASSIFLORA PEDICULATA Mast. in Bot. Gaz. 23: 247. 1897; Killip, in Field
Mus. Publ. Bot. 19:113. 1938.

Rather coarse climbers, glabrous throughout. Leaves petiolate, simple, basifixed,
broadly 3-lobed, the central lobe stronger, more or less deeply cordate at the base,
4-9 cm. long, 5-10 cm. broad, inconspicuously ocellate; petioles 1.5—5.0 cm. long,
rather inconspicuously glandular about midway or above; stipules inconspicuous
and subsetose. Inflorescences solitary or paired, bearing solitary greenish yellow or
white, purple tinged flowers; peduncles about 2-4 cm. long, jointed about midway;
bracts inconspicuous and subsetaceous. Flowers about 3 cm. in diameter; sepals
oblong-lanceolate, about 1.5 cm. long, not corniculate; petals linear, about 1 cm.
long; filaments of the corona in 1 series, somewhat shorter than the petals. Fruit
not seen.

Costa Rica and western Panama, near sea level to about 1,500 m.

BOCAS DEL TORO: Water Valley, Von Wedel 854.

13. PASSIFLORA SUBEROSA L. Sp. Pl. 958. 1753; Killip, in Field Mus. Publ. Bot.
19:88. 1938.

Pa pallida L. loc. cit. 1753.
Pass siflora hirsuta L. loc. cit. 958. 1753.

Passi siflora parviflora Sw. loc. cit. 1788

ranadi s a 80. 1791.
Cieca beterobbylla (Dryand.) Moench, Method. Suppl 101. 1802.
Cieca suberosa (L.) Moench, loc. cit. 102. 180
Cieca minima (L.) Moench, QE 8 de
Passiflora litoralis HBK. Nov. Gen "8 Sp. 2:138. 1817
Baldwinia peltata (Cav.) Raf. Amer. Monthly Aly Mag, 267. 18
Monactineirma angustifolia (Sw.) Bory, in Ann. Gén. Sci. Phys. Brux. 2:138. 1819.
i 819.

onactineirma bederacea (Cav.) Bory, loc. B Eid
Passiflora oliviformis Vell. Fl. Flum. 9: pl. 83.

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1958]
FLORA OF PANAMA (Passifloraceae) i

Passiflora globosa Vell. loc. cit. p. 85. 1827

Passiflora Kobautiana Presl, Fl. Bemerk. 72. 1836.
Passiflora villosa M Fl. dins 2:151. 1837; € Ma
Meioperis peltata (Cav.) Raf. FI. Tellur. 4:10 $

Passiflora Warei Nutt. in Silliman's Journ. ex Torr. T ed Fl. N. Amer. 1:539. 1838.
Passiflora limbata aisi Ind. Se re Hort. Neapo hi

Passiflora flexuosa Gardn. in Lon d dria, Bot. 1:1

Passiflora rato berose Fisch. al Sem. Hort. de aci 9:82, ex Walp. Rep. 2:934.

Cif peltata (Cav.) M. un Fam. Nat. Sme 2:141. 1846.

U
8
n
=
=
&
S
3
=
m
—
“o
N
w

Passiflora suberosa Var. div ricata Griseb. in Bonplandia 6:7.
Passiflora suberosa var. minima (L.) Mast. in Trans i = 630. 1871.
Passiflora suberosa var. peer (L.) MIE. T cit. Tm

Passiflora suberosa var. MI (Sw.) Mast. loc. cit. 1871.
Passiflora suberosa var. pallida (L.) Mast. loc. cit. 1871.
Passiflora suberosa var. bederacea (Cav.) Mast. loc. cit. 1871.
Passiflora suberosa var. lineariloba (Hook. f.) Mast. in Mart. FI. Bras. 131:579. 1872.
Passiflora suberosa var. longiloba Tr. & Planch. in foe Sci. Nat. V. Bot. 17:157. 1873.
Passiflora suberosa var. longipes S. Wats. in Proc. Amer. Acad. 25:149. 1890.
Passiflora calliaquatica Krause, in Bot. Centralbl. Beih. 325; :340. 1914.

Slender climbers, glabrous to densely puberulent, the base of the stems more or
less corky. Leaves petiolate, simple, basifixed, extremely variable, usually definitely
3-lobed, the central lobe longer, rarely almost entire 3-12 cm. long, 2—9 cm. broad,
not ocellate or very inconspicuously so; petioles 1-2 cm. long, inconspicuously
glandular near the juncture with the leaf blade; stipules inconspicuous and linear.
Inflorescences usually paired, each bearing a single small, greenish flower; peduncles
filiform, 1-2 cm. long, jointed almost directly beneath the flower; bracts very
minute. Flowers about 1-3 cm. in diameter; sepals lanceolate, 0.5-1.5 cm. long,
not corniculate; petals absent; filaments of the corona in 2 series, the outer about
half as long as the sepals. Fruit globose, deep purple, 0.5—1.5 cm. in diameter.

Peninsular Florida and southern Texas; Mexico to Brazil and Argentina; Ba-
hamas; Antilles; introduced in the Old World tropics. Lowland thickets. Melon-
cillo; buevo de gallo.

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[Vol. 45
14 ANNALS OF THE MISSOURI BOTANICAL GARDEN

PANAMA: Bella Vista, Standley 25357, Kk 12039; Nuevo San Francisco, Standley
30733. SAN BLAs: Puerto Obaldia, Pittier 4402

The preposterous bibliography testifies to the ubiquity and variability of P.
suberosa. It is odd that it has been so infrequently collected in Panama.

14. PASSIFLORA PULCHELLA HBK. Nov. Gen. & Sp. 2:134. 1817.

Passiflora bicornis Mill. Gard. Dict. ed. 8. Passiflora no. 13. 1768.

Passiflora rotundifolia Jacq. Obs. 2:26. pl. 46, fig. 1. 1767, non L.

Passiflora rotundifolia var. Ng ese DC. Prodr. 5:326. 18
— Jacquini (DC.) M. Roem. Fam. = ae 2:156. 1846.

assiflora divaricata Griseb. in Bonplandia 6
iar ora pulchella var. bifidata Mast. in Engl Bot. "Jahrb. 8:220. 1887.

Rather stout climbers, glabrous throughout. Leaves rather long-petiolate,
simple, basifixed, broadly and cuneately 2-lobed, occasionally with a small central
lobe, 2-6 cm. long, 3-9 cm. broad, rather inconspicuously glandular-ocellate;
petioles 1-3 cm. long, not glandular; stipules linear, 5-10 mm. long. Inflorescences
solitary or rarely paired in the axils of the greatly reduced uppermost leaves, bearing
solitary involucrate blue or lavender flowers, the associated tendrils basal; peduncles
5-8 cm. long, jointed directly beneath the flower; bracts conspicuously involucrate,
broadly obovate, cuneate at the base, entire, 1.0—1.5 cm. long, pale green occasion-
ally tinged with purple. Flowers about 4—6 cm. in diameter; sepals oblong, about
2 cm. long, not corniculate; petals ovate-lanceolate, about 1.5 cm. long; filaments
of the corona in several series, the outer about as long as the petals. Fruit globose,
about 1 cm. in diameter.

Southern Mexico to Colombia and Venezuela; cultivated in Hawaii. Thickets
near sea level.

t Kobe Road, Woodson, Allen & Seibert 1421. PANAMA: Bella
Yin. Killip. ui = Heriberto 218; Las Sabanas, Standley 40781.

This is one of the showiest Panamanian passionflowers because the flowers, al-
though not large, are borne in the very characteristic raceme-like clusters at the
tips of the stems.

15. PASSIFLORA MEMBRANACEA Benth. Pl. Hartw. 83. 1841; Killip, in Field Mus.

Publ. Bot. 19:234. 1938.

Cieca membranacea (Benth.) M. Roem. Fam. Nat. Syn. 2:140. 1841.

Rather stout climbers, glabrous throughout. Leaves rather shortly petiolate,
simple, barely peltate, suborbicular and very indistinctly 3-lobed, the tip and lateral
lobes mucronulate, 5-10 cm. long and about as broad; petioles 2-3 cm. long,
eglandular; stipules — foliaceous, deeply cordate-reniform and odc
amplexicaul, 1.5-3.0 cm. broad. Inflorescences solitary at the nodes, bearin
single very strikingly involucrate greenish white or cream flower; peduncles iis
cm. long, jointed almost immediately beneath the flower; bracts very broadly ovate
or suborbicular, very deeply cordate, 4—5 cm. long and broad, pale green strongly
suffused with rose. Flowers about 7-8 cm. in diameter; sepals oblong-lanceolate,

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1958]
FLORA OF PANAMA (Passifloraceae) 15

about 4 cm. long, not corniculate; petals oblong-elliptic, about as long as the sepals;
filaments of the corona in 2 series, the outer about 1 cm. long. Fruits oblong-
ovoid, tapered at the base, 4-6 cm. long and 2-3 cm. thick, yellowish green.

Southern Mexico to Panama, in | forests.

RIQUÍ: Potrero Muleto to summit, Volcán de Chiriqui, Woodson & Schery 380;

"New. m erland", central valley of Río uM Viejo, Allen 1394.

This is one of the most distinctive and attractive species of the genus by virtue
of the elongate peduncles and large petalaceous bracts which almost equal the
flowers.

16. PASSIFLORA ADENOPODA DC. Prodr. 3:330. 1828; Killip, in Field Mus. Publ.

Bot. 19:222. 1938
Passiflora acerifolia Schlecht. & Cham. in Linnaea 5:89. 183
Dysosmia acerifolia ATENE & Cham.) M. Roem. ję Nat Syn. 2:151. 1846.
Ceratosepalum micrantbum Oerst. Amér. Centr. pl. 17
Passiflora pales te Mast. in Trans. Linn. Soc. 27: si Te i
Passiflora aspera Sessé & Moc. Fl. Mex. 227. 1887

Coarse climbers, minutely hispidulous to glabrate. Leaves long-petiolate, basi-
fixed, deeply (3-)5- to 7-lobed, the central and upper lobes stronger, acuminate,
deeply cordate, 7-15 cm. long, 8-17 cm. broad, membranaceous, hispidulous to
glabrate; petioles 3-8 cm. long, bearing 2 very large prominently stipitate glands
near the juncture with the leaf blade; stipules broadly ovate, amplexicaul, denticu-
late, 1.0-1.5 cm. long and broad, foliaceous, persistent. Inflorescences paired,
bearing a single large purple and white flower; peduncle 3-4 cm. long, jointed
somewhat below the middle and there bearing 3 lacerate subfoliaceous bracts about
1 cm. long. Flowers about 4-7 cm. in diameter; sepals oblong-lanceolate, 3-4 cm.
long, prominently corniculate at the tip; petals narrowly lanceolate, 1.0-1.5
cm. long; filaments of the corona in a single series, about as long as the petals or
slightly longer. Fruit globose, 2.0—2.5 cm. in diameter, greenish yellow, densely
puberulent.

Southern Mexico to Venezuela and Peru at middle to higher elevations.

CHIRIQUÍ: El Boquete, Seemann 1626.
The Seeman collection was found by Killip at both Kew and the British
Museum; it was not examined for the present account.
17. PASSIFLORA VITIFOLIA HBK. Nov. Gen. 8 Sp. 2:138. 1817, Killip, in Field
Mus. Publ. Bot. 19:319. 1938.
Passiflora sanguinea J. E. Sm. in Rees, Cycl. 26: fred no. 45. 1819.
Passiflora puni :
(J. E. Sm.) . cit. vi; chi 8.
Macropbora sanguinea (J. E. Sm.) Raf. Fl. Tellur. 4:103. 1838.
Passiflora servitensis Karst. in Linnaea 30:163. 1859

Tacsonia Bucbanani Lem. Ill. Hort. 14: pl. 519. 1867.
Rather coarse climbers, more or less densely ferruginous-pubescent throughout.

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[Vol. 45
16 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Leaves petiolate, basifixed, rather inconspicuously serrulate, deeply 3-lcbed, the
central lobe stronger, more or less deeply cordate, 7—15 cm. long, 8—17 cm. broad;
petioles 2—6 cm. long, bearing 2 to several rather inconspicuous glands toward the
base; stipules subsetaceous, 3-5 mm. long. Inflorescences solitary, bearing a single
very showy yellow and scarlet flower; peduncle 4-8 cm. long, jointed almost di-
rectly beneath the flower; bracts conspicuous and foliaceous or somewhat petaloid,
lanceolate, glandular-serrate to nearly entire, 1.5—2.5 cm. long. Flowers 10-15 cm.
in diameter; sepals about 6-10 cm. long, corniculate; petals narrowly lanceolate,
nearly as long as the sepals; filaments of the corona in 3 series, the outer 1.5—2.0
cm. long. Fruits broadly ovoid, 3-6 cm. long, greenish yellow, densely puberulent.
Nicaragua to Venezuela and Peru; Cuba and Jamaica; in lowland forests.
Known as guate-guate and pasionaria in Panama.
CANAL ZONE: Las Cruces Trail, Hunter & Allen 456; Barro Colorado Island, Wetmore
& Abbe 76; between Summit and Gamboa, Greenman & Greenman 5220; Chasse, Fendler
118. CHIRIQUÍ: San Bart tolo Woodson & Schery 871. COCLE: mountains beyond La
n° Hunter & Allen 558; El Valle de Antón, Allen I660. COLÓN: Camp Piña, Allen
wu Marraganti, Williams 1150; Chepigana, Terry & Terry 1374; Rio Sabana,
Laud III 126. PANAMA: Tapia River, Juan Díaz region, Maxon & Harvey 6733, 6678,

68 5, Sindy 28101; Twa River, Killip 3313; Orange River, Killip 3335; Pacora,
Bro. Paul 2

The diis flowers of P. vitifolia are frequent and widespread in lowland
thickets and open forests throughout the Republic

18. PassrFLORA WILLIAMSH Killip, in Journ. Wash. Acad. Sci. 12:262. 1922; in
Field Mus. Publ. Bot. 19:355. 1938.

Coarse climbers, softly and densely puberulent throughout. Leaves petiolate,
basifixed, deeply 3-lobed to somewhat below the middle, the central lobe stronger
and somewhat contracted toward the base, inconspicuously serrulate, 8—15 cm.
long, 9-16 cm. broad; petioles 4-6 cm. long, with 2—3 ocellate glands toward the
base; stipules filiform, inconspicuous. Inflorescences solitary, bearing a single
handsome violet flower; peduncle 1.0—1.5 cm. long, jointed directly beneath the
flower; bracts involucrate, united toward the base and completely enveloping the
flower before anthesis, 2.5-3.5 cm. long, densely puberulent, pale green. Flowers
about 6-7 cm. in diameter; sepals oblong, 3-4 cm. long; petals oblong-spatulate,
2.0-2.5 cm. long; filaments of the corona in several series, the outermost less than
half as long as the petals. Fruits unknown.

Known only from Panama, at middle and low elevations.

CANAL ZONE: between Empire and Mandinga, Piper 5481; Barro Colorado Laika Zetek
ay pecu Bismarck, above Penonomé, Williams 585. PANAMA: Las Sabanas, Bro.

This species bears 4 rather conspicuous ocellate glands upon the leaf blade, on
each lobe toward the base of the two sinuses; the central lobe thus with two glands,
the lateral with one apiece.

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1958]
FLORA OF PANAMA (Passifloraceae) 17

19. PASSIFLORA MENISPERMIFOLIA HBK. Nov. Gen. & Sp. 2:137. 1817; Killip,
in Field Mus. Publ. Bot. 19:457. 1938.

Passiflora villosa Dombey, ex Triana & Planch. in Ann. Sci. Nat. ser. 5. 17:154. 1873, in
synon.

Rather coarse climbers, more or less densely pilose to glabrate throughout.
Leaves petiolate, basifixed, broadly 3-lobed to somewhat above the middle, the
central lobe much stronger, usually inconspicuously serrulate toward the cordate
base, 7-15 cm. long, 5-13 cm. broad; petioles 2-5 cm. long, with 2 to several pairs
of slender stipitate glands; stipules conspicuously foliaceous, ovate-subreniform,
deeply amplexicaul, 1.5-3.5 cm. long, glandular-denticulate to subentire. In-
florescences solitary, bearing a single white and purple flower; peduncle 4-6 cm.
long, jointed directly below the flower; bracts rather narrowly lanceolate, 1-2 cm.
long, foliaceous, with rather few glandular denticulations to entire. Flowers about
6 cm. in diameter; sepals narrowly oblong, 2-3 cm. long, corniculate; petals linear-
oblong, about equaling the sepals; filaments of the corona in several series, the
outermost about half as long as the petals. Fruits broadly ovoid, 4-6 cm. long.

Nicaragua to Peru and northern Brazil, from near sea level to about 1500 m.

OCAS DEL TORO: vicinity of Chiriqui Lagoon, Von Wedel 434, 1195, 1406.

ZONE: Madden Dam, Allen 2007; Gatuncillo, Piper 5623; > peg Island, Wood-
worth & Vestal 637. cHIRIQUI: Boquete, Terry 1281. UAS: Sona, Woodson
Seibert & Allen 515.

20. PASSIFLORA SUBPELTATA Ortega, Nov. Rar. Pl. Hort. Matrit. 6:78. 1789;

Killip, in Field Mus. Publ. Bot. 19:436. 1938.
Passiflora alba Link & Otto, Icon. Pl. Rar. 65. pl. 33.
Passiflora adenopbylla n in Mart. Fl. Bras. 13!: e. du
Passiflora atomaria Planch. ex Mast. loc. cit. 570. 187

Rather slender Pu glabrous throughout. Leaves shortly petiolate, basi-
fixed, very broadly 3-lobed to about the middle, the 3 lobes subequal, rounded, with
2-3 pairs of inconspicuous serrulate glands in the sinuses, rounded to subcordate at
the base, 4-9 cm. long, 5-12 cm. broad; petioles 2-6 cm. long, with 1 pair of in-
conspicuous glands at about the middle; stipules conspicuously foliaceous, broadly
ovate, 1-4 cm. long and about half as broad. Inflorescences solitary, bearing a
single purple and white flower; peduncle 2-5 cm. long, jointed directly beneath
the flower; bracts conspicuously involucrate, broadly ovate, about 1.0—1.5 cm.
long and broad, abruptly acuminate, pale green somewhat tinged with rose.
Flowers about 5—7 cm. in diameter; sepals oblong, about 3 cm. long, very con-
spicuously corniculate; petals narrowly lanceolate, 2.0—2.5 cm. long; filaments of
the corona in several series, the outermost about equaling the petals. Fruits sub-
globose, 3-4 cm. in diameter.

Mexico to Venezuela and Colombia, near sea level to about 2,800 m., in
thickets and open forest.

CAN : Miraflores, W bite & W bite 45; Red Tank, Maxon 6574. PANAMA: Juan
Díaz, Standley a: Taboga Island, Standley 27971.

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[Vol. 45
18 ANNALS OF THE MISSOURI BOTANICAL GARDEN

21. PASSIFLORA LIGULARIS Juss. in Ann. Mus. Hist. Nat. 6:113. pl. 40. 1805;
Killip, in Field Mus. Publ. Bot. 19:344. 1938.

Passiflora serratistipula DC. Prodr. 3:328. 1828.

Rather stout climbers, glabrous throughout. Leaves long-petiolate, basifixed,
entire, broadly ovate, deeply cordate, abruptly subcaudate-acuminate, 8—17 cm.
long, 6-15 cm. broad; petioles 5-15 cm. long, with 3-5 pairs of elongate filiform
glands; stipules foliaceous, ovate, 1-3 cm. long. Inflorescences solitary, bearing a
single showy pale green, purple- and lavender-tinged flower; peduncles 2-5 cm.
long, jointed directly beneath the flower; bracts involucrate and foliaceous, ovate,
2-4 cm. long. Flowers about 7-9 cm. in diameter; sepals oblong, 3-5 cm. long;
petals narrowly lanceolate, 3-4 cm. long; filaments of the corona in several series,
the outermost about as long as the petals. Fruits broadly ovoid, 6-8 cm. long,
purplish yellow.

Southern Mexico to Venezuela and Bolivia, in highland forests between 1,000
and 3,000 m

CHIRIQUÍ: vicinity of Finca Lérida, Woodson 6 Schery 238.

22. PASSIFLORA QUADRANGULARIS L. Syst. Nat. 1248. 1759; Killip, in Field Mus.

Publ. Bot. 19:335. 1938.

Granadilla quadrangularis (L.) Medic. Malvenfam. 97. 1787.
Passiflora tetragona M. Roem. Fam. Nat. Syn. 2:165. 1846.
Passiflora macroceps Mast. in Gard. Chron. 1869:1012. 1869.

Coarse climbers with prominently winged stems, glabrous throughout. Leaves
petiolate, basifixed, entire, broadly ovate, abruptly and shortly acuminate, rounded
at the base to broadly cordate, 10-20 cm. long, 8—15 cm. wide; petioles 2-5 cm.
long, with about 3 pairs of ovoid glands; stipules foliaceous, ovate, acuminate, 2-4
cm. long. Inflorescences solitary, bearing a single greenish white flower flushed
with rose or purple; peduncles 1.5—3.0 cm. long, jointed directly beneath the
flower; bracts delicately foliaceous, broadly ovate, cordate, entire or essentially so,
2-4 cm. long and about as broad. Flowers about 8-12 cm. in diameter; sepals
broadly oblong, conspicuously corniculate, 4-5 cm. long; petals ovate-lanceolate,
3-5 cm. long; filaments of the corona in 5 series, about equaling the petals. Fruits
— parca up to 3 dm. long, pale yellow somewhat flushed with rose.

ely cultivated and escaping throughout tropical America; of uncertain
origin.

CANAL ZONE: Chagres, Fendler 110. PANAMA: Juan Díaz, Standley 32057.

This is one of the most popular edible passionfruits, and is popularly known
as granadilla throughout its range in Spanish America and maracujé in Brazil. It
has a rather insipid fruity flavor of indefinite quality, and although capable of

being eaten out of hand it usually is made into a conserve or a thin pudding after
boiling in milk.

(18)

1958]
FLORA OF PANAMA (Passifloraceae) 19

23. PASSIFLORA OERSTEDII Mast. in Mart. Fl. Bras. 13!:562. 1872; Killip, in Field

Mus. Publ. Bot. 19:418. 1938.

Passiflora populifolia Triana & Planch. in Ann. Sci. Nat. ser. 5. 17:150. 1873.
Passiflora Purbusii Killip, in Journ. Wash. Acad. Sci. 12:261. 1922.
Passiflora dispar Killip, loc. cit. 330. 1922.

Rather slender climbers, glabrous throughout. Leaves rather shortly petiolate,
basifixed, entire, ovate or oblong-ovate, acuminate, rounded to deeply cordate at
the base, 6-13 cm. long, 3-9 cm. broad; petioles 1-3 cm. long, bearing several
pairs of small stipitate glands; stipules conspicuously foliaceous, strongly inequi-
lateral, narrowly ovate, caudate-cuspidate, entire, 1-4 cm. long. Inflorescences
solitary, bearing a single showy purple or lavender-rose flower; peduncles 3-6
cm. long, jointed directly beneath the flower; bracts delicately foliaceous, ovate,
rounded or cordate at the base, 1.0—1.5 cm. long. Flowers about 6-8 cm. in di-
ameter; sepals ovate-lanceolate, 3—4 cm. long, conspicuously corniculate; petals
linear-oblanceolate, about 1.5 cm. long; filaments of the corona in many series, the

Y,
O

NY
N
RUS, NS N ZE È
MAR « x A y \ AE Sap
| ES RN NA ża
N KIN iti Bee
LORRY 2222
RY MAN SI 2 27 A
TANI Uk,
TI AGRON AM

7

=,

Fig. 2. Passiflora oerstedii

outermost equaling the petals or somewhat longer. Fruits broadly ovoid, 4-6
cm. long.

Southern Mexico to Colombia and Venezuela, from near sea level to about
1500 m., in open forest.

BOCAS DEL TORO: Water Valley, Von Wedel 1756. cocLé: La Mesa, north of El Valle
de Antón, Allen 2399. cHIRIQUI: Bajo Mona and Quebrada Chiquero, Woodson & Schery
591; Volcán de Chiriquí, Terry 1303.

24. PASSIFLORA AMBIGUA Hemsl. in Curt. Bot. Mag. pl. 7822. 1902; Killip, in
Field Mus. Publ. Bot. 19:363. 1938.

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[Vol. 45
20 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Rather coarse climbers, glabrous throughout or essentially so. Leaves petiolate,
basifixed, entire, ovate to oblong-elliptic, acuminate, obtuse or rounded at the base,
10-20 cm. long, 5-9 cm. broad, rather thickly coriaceous; petioles 2-3 cm. long,
bearing 2 thick ocellate glands toward the base; stipules filiform, relatively incon-
spicuous. Inflorescences solitary, bearing a single showy maroon, rose, or purple
flower; peduncles 5-7 cm. long, jointed directly beneath the flower; bracts free,
ovate, 3-6 cm. long, delicately subfoliaceous or somewhat petalaceous. Flowers
8—12 cm. in diameter; sepals narrowly oblong, 4—6 cm. long, rather inconspicuously
corniculate; petals narrowly lanceolate, 3—4 cm. ong; filaments of the corona in
several series, the outermost about half as long as the petals. Fruits broadly ovoid,
10—12 cm. long, greenish yellow.

Southern Mexico to Panama, from near sea level to about 1000 m., in thickets
and open forest.

BOCAS DEL TORO: Fish Creek Uh Von Wedel 2324; Columbus Island, Vor

Wedel 80. CANAL ZONE: Gatün, Sutton Hayes 430; Barro Colorado Island, Bangbam 466.
COCLE: north rim of El Valle di Antón; Allen 1686.

25. PASSIFLORA NITIDA HBK. Nov. Gen. 8 Sp. 2:130. 1817; Killip, in Field Mus.

Publ. Bot. 19:374. 1938.

Passiflora nympbaeoides Karst. in Linnaea 30:165. 1859.

Rather coarse climbers, glabrous throughout. Leaves rather shortly petiolate,
basifixed, rather distantly serrate or serrulate, at least toward the base, broadly
elliptic to lanceolate, abruptly acuminate, rounded at the base, 9-18 cm. long, 6-
10 cm. broad, subcoriaceous; petioles 1.0—1.5 cm. long, with 2 large ovoid glands
somewhat above the middle; stipules inconspicuous and filiform. Inflorescences
solitary, bearing a single showy white and purple flower; peduncles 3-6 cm. long,
jointed directly beneath the flower; bracts free, ovate, entire, 3-4 cm. long, foli-
aceous or somewhat petalaceous. Flowers 8—9 cm. in diameter; sepals oblong-
elliptic, 4.0—4.5 cm. long, inconspicuously corniculate; petals about as long as the
sepals; filaments of the corona in several series, the outermost about half as long as
the petals. Fruits obovoid, 4—6 cm. long.

Panama to northern Brazil and adjacent Peru, in lowland thickets and open
forests.

CANAL ZONE: Barro Colorado Island, Shattuck 675; Summit, Lindsay 230.

26. PASSIFLORA SEEMANNII Griseb. in Bonplandia 6:7. 1858; Killip, in Field Mus.

Publ. Bot. 19:347. 1938.

Passiflora incana Seemann, ex Mast. in Rasse Linn. Soc. 20:40. 1883, prd ape in synon.
Passiflora orbifolia Planch. & Linden nn. Sci. Nat. ser. 5. 17:150.

Rather stout climbers, ia Pi Leaves Li s entire,
broadly cordate-ovate, abruptly and shortly apiculate-acuminate, 5-10 cm. long,
5-7 cm. broad; petioles 3-7 cm. long, with 2 rather inconspicuous sessile glands
directly below the leaf blade; stipules linear, about 1 cm. long. Inflorescences

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1958]
FLORA OF PANAMA (Passifloraceae) 21

solitary, bearing a single handsome white or cream flower strongly suffused with
purple or violet; peduncles 6-10 cm. long, jointed directly below the flower; bracts
more or less petalaceous, united at the base and completely enveloping the flower
before anthesis, 3-5 cm. long. Flowers about 8—10 cm. in diameter; sepals narrowly
ovate, 4-5 cm. long, rather inconspicuously corniculate; petals oblong-lanceolate,
3-4 cm. long; filaments of the corona in 2 series, the outer about 1 cm. long.
Fruits ovoid, 3-5 cm. long and broad.
Southern Mexico to Colombia, at near sea level to about 1,500 m.

BOCAS DEL TORO: Lincoln Creek, Río Changuinola, Dunlap 422; bic + dera

Lagoon, Von Wedel 1260, 1366; Water Valley, Von Wedel 1679. CANAL Bohio
Maxon 4765; Gamboa, Pittier 2515, Allen 3930; between Summit and res asy
& Greenman 5221; Gatün, Bro. Heriberto 53; between Mt. Hope and Sta. Rita Trail,

Cowell 59; Las Cascadas, near Summit, Standley 29566; East Pirates! Standley 29838;
Quebrada Culebra, Dodge & Allen 17048; Las Cruces Trail, Hunter s. n.; Chagres, Fendler
120; Miraflores, W bite 6 White 42. COLÓN: between [as Field and Catival "Standley
0286. DARIEN: Tucute, Chepigana District, Terry & T 393. o
Campana, Allen 1694; Tapia River, Maxon 6 Harvey pb So pa fo 79, Killip 3297;
Juan Díaz, Standley 30634.

This handsome species has been introduced into cultivation in Mexico, Nicara-
gua, and Hawaii, according to Killip. According to Killip also, the leaves occa-
sionally are lobed, but those of all the specimens cited above are entire.

27. PASSIFLORA FOETIDA L. Sp. Pl. 959. 1753; Killip, in Field Mus. Publ. Bot.
19:481. 1938.
uo vesicaria L. Amoen. Acad. 5:382. 1760.

Passiflora bastata Bertol. Fl, Guat. 427. ;
Dysosmia foetida (L.) M. Roem. Fam. Nat. Syn. 2:149. 1846.
Dysosmia gossybiifolia Wars CY Roem. loc. cit.

ysosmia bastata (Bertol.) M. Roem. loc. cit. 1846.

Dysosmia ciliata (Dryand.) M. R loc. cit. 1846

Dys ensis M. Roem. loc. cit. 150.

Dysosmia nigelliflora (Hook.) M. Roem. loc. cit. TE

P, raquiniana Lem. in Illustr. Hort. 8: pl. 276. 1861

Passiflora Liebmanni Mast. in Mart. Fl. Bras. 131:547. 1872

Passiflora bispida DC. ex Triana & Planch. in Ann. Sci. Nat. ser. 5. 17:172. 1873.
assiflora Moritziana Planch. loc. cit. 175. 1873.

Passiflora muralis Barb. Rodr. in Contr. Jard. Bot. pi 1: wie 1891.

Passiflora Balansae Chod. in Bull. Herb. Boiss. ser. i 2:74
Pokora Lipari Britton, in Bull. Torrey Bot. Club i 19. 1917.
(Also numerous varietal and formal designations).

= slender climbers, commonly more or less yellow-pubescent throughout,
less frequently glabrate or glabrous. Leaves petiolate, basifixed, extremely variable,
in Panama chiefly broadly and acutely 3-lobed to about the middle, the central lobe

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[Vol. 45
22 ANNALS OF THE MISSOURI BOTANICAL GARDEN

far stronger, usually about 4-12 cm. long and wide; petioles usually about 2-6 cm.
long, usually beset with mixed glandular and eglandular hairs; stipules subseta-
ceous, inconspicuous. Inflorescences solitary, bearing a single greenish white or
yellow, more or less purple-tinged flower; peduncles 3-7 cm. long, jointed directly
beneath the flower; bracts foliaceous, complexly 2- or 3-pinnatifid, each filiform
division gland-tipped, usually 2-4 cm. long. Flowers about 4-6 cm. in diameter;
sepals oblong-lanceolate, about 2-3 cm. long, shortly corniculate; petals oblong-
oblanceolate, slightly shorter than the sepals; filaments of the corona in several
series, the outermost about 1 cm. long. Fruits subglobose, yellow or orange, about
2-3 cm. in diameter.

Common tiens tropical America in lowland thickets and open forest,
becoming a wee

BOCAS DEL TORO: Old Bank Island, Von Wedel 2142; Water ey Von Wedel 1718,
1831; vicinity of Chiriqui Lagoon, Von Wedel 1562. CANAL ZONE: Las Cruces Trail,

Hunter 6 Allen 724; near mouth di Rio Chagres, Allen 899; eae -Fondler 117; Ft. San
Lorenzo, jg & Valentine 7013; Matias FERAM, Pittier 6954; G atün d —

Grande: near Culebra, pup 3 2089. CHIRIQUÍ: dei west of ce Allen soak.
ani Bello, Pittier 2474. DARIÉN: trail between Pinogana and bale Allen 292.
PANAMA: Taboga I Island, “Miller 2042, Pittier 3569; Tapia River, Maxon
Pato foetida must be frequent at low elevations in every sivit of
Panama. The species, although unmistakable, is extremely variable in small details,
and Killip recognized no less than 42 named varieties. All of the specimens
enumerated above he referred to var. istbmia Killip.

CARICACEAE

Soft-wooded laticiferous shrubs and small trees, usually dioecious or monoecious
and with disproportionately thick, simple or infrequently branching trunks orna-
mented with large leaf scars. Leaves spiral, usually quite large and long-petiolate,
exstipulate, the blade palmately compound or simple and variously lobed, rarely
entire. Inflorescence axillary at the uppermost nodes, the staminate an extensive
and highly compound thyrse, the pistillate much less compounded and with fewer
and larger flowers, the fruits developed at the base of the young growth and
appearing cauliflorous. Flowers dichlamydeous, 5-merous, regular, hypogynous,
sympetalous but usually imperfectly so in the pistillate; calyx lobes 5, imbricate or
nearly open in aestivation, usually inconspicuous; corolla salverform or infundibuli-
form and with a well-developed tube in the staminate, more or less campanulate
and rather imperfectly sympetalous in the pistillate, the limb 5-lobed with con-
torted aestivation; stamens 10 in 2 whorls inserted near the orifice of the corolla
tube, the antepetalous usually with somewhat shorter filaments than the ante-
sepalous, the filaments occasionally more or less petaloid or with petaloid ap-
pendages, reduced to staminodia or absent in pistillate flowers; pistil superior, 3-
to 5-carpellate, usually 1-celled and with parietal placentation, rarely 3- to 5-celled

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1958]
FLORA OF PANAMA (Caricaceae) 23

and with axile placentation, the ovules numerous, anatropous, the stigmas 3-5,
sessile or essentially so, usually dichotomous, rarely subcapitate, reduced to an
acicular pistillode or absent in staminate flowers. Fruit a very large to rather small
succulent 1- to 5-celled berry, the numerous seeds with pulpy testas and funicles.

The Caricaceae are a small family of four genera, all tropical American except
the bitypic Cylicomorpha of Africa. The plants are of strikingly antediluvian
aspect with their stout, usually simple trunks heavily ornamented with large spiral
leaf scars and their terminal crowns of fern-like leaves. The latex tubes, which
traverse all parts of the plant, are associated with various proteolytic enzymes,
particularly papain.

a. Leaves palmately compound; petals antesepalous; ovary 5-celled............... 1. JACARATIA
aa. Leaves palmately lobed; petals alternisepalous; ovary 1-celled................... 2. CARICA

JACARATIA Marcgr. ex Endl.

JACARATIA Marcgr. ex Endl. Gen. 933. 1839 (as Jaracatia).
Pileus Ramirez, in An. Inst. Med. Nac. Mex. 5:29. 1901.

Dioecious shrubs and trees, branching rather frequently for the family, the
trunk becoming very thick. Leaves palmately compound, with 3-12 petiolulate,
entire or undulate leaflets. Inflorescences in the axils of the uppermost leaves,
thyrsiflorous, the staminate extensive and many flowered, the pistillate reduced and
with few larger flowers. Calyx 5-lobed, inconspicuous. Corolla salverform or
narrowly infundibuliform, the limb 5-lobed, antesepalous. Stamens 10, the fila-
ments and connectives more or less conspicuously petaloid. Ovary 5-celled, con-
taining numerous ovules on axile placentas. Berry usually rather elongate, 5-celled.

Probably about a dozen species y p from Mexico to Paraguay.

a. Stems unarmed; leaflets 3—5, green ben 1. J. DOLICHAULA
. Stems armed with conspicuous spines; dos 5—9, very pale and
adi benea di 2. J. COSTARICENSIS

JACARATIA DOLICHAULA (Donn. Sm.) Woodson, in Ann. Missouri Bot. Gard.

37:404. 1950
Carica dolicbaula Donn. Sm. in Bot. Gaz. 23:247. 1897.

Trees up to 12 m. tall the trunk usually thickened toward the base, the
branches glabrous and unarmed. Leaves palmately compound with 3-5 petiolulate
broadly elliptic or obovate, caudate-acuminate leaflets 6—15 cm. long, 2—6 cm.
broad, green beneath, the petiolules about 1 cm. long; petioles slender, up to about
15 cm. long. Inflorescences much shorter than the subtending petioles, the sta-
minate up to about 30-flowered, the pistillate usually 1- to 3-flowered. Staminate
flowers: calyx lobes broadly ovate, about 1 mm. long; corolla slender and salver-
form, the tube 3—5 cm. long, about 3 mm. in diameter, scarcely dilating toward
the orifice, pale green or greenish white, the lobes oblong-spatulate, obtuse or acute,
1.0-1.25 cm. long, slightly spreading, white occasionally tinged with pink toward
the tip; stamens inserted at the orifice of the corolla tube, exserted, the antepetalous

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[Vol. 45
24 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 3. Jacaratia dolichaula

with very short filaments, the alternating whorl sessile, the anthers narrowly
oblong, about 5 mm. long, the connective petaloid and somewhat longer than the
anther; pistillode narrowly acicular, about 1 cm. long. Pistillate flowers: calyx
lobes broadly ovate, about 1-2 mm. long; corolla shortly salverform, the tube
about 1.5 cm. long, about 5 mm. in diameter at the base, contracting toward the

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1958]
FLORA OF PANAMA (Caricaceae) 25

middle, then dilated to about 7 mm. at the orifice, white, the lobes narrowly oblong,
about 1.5 cm. long, slightly spreading, white; staminodia absent or inserted at the
orifice of the corolla tube, sessile, essentially like the stamens of the staminate
flower but the anthers somewhat smaller and the petaloid connectives somewhat
larger; ovary oblong-ovoid, about 5 mm. long, 5-celled, the stigma with 5 ovate
essentially entire lobes. Berries ovoid-fusiform, about 10 cm. long and 5 cm.
broad, persistently 5-celled, containing numerous compressed ovoid seeds 7-8 mm.
long; fruiting peduncle about 3.5 cm. long.

Costa Rica and Panama, in forests, 200—1000 m. elev. Popular names recorded
by Standley in Costa Rica are papaya, papaya de monte, papayillo, palo de barril,
who explains the last as follows: ‘The palo de barril, which I believe referable to
this species, is common in Guanacaste, where it is a large tree with broad crown
and a very thick trunk. This is so soft or spongy that with a few slashes of a
machete it is possible to fella large tree. If a large trunk is thus cut, and left for
some weeks to dry, it is possible to separate a cylinder of bark. Sections of this
are used in place of barrels, for storing maize and other articles." (Fl. Costa Rica
2:724. 1937).

s DEL TORO: Fish Creek, vicinity of Chiriquí Lagoon, Von Wedel 1084, 2207.

BOCA
VERAGUAS: forested slopes of Cerro Tuté, vicinity of Santa Fé, Allen 4392.

2. JACARATIA COSTARICENSIS I. M. Johnston, in Contr. Gray Herb. 70:79. 1924.

Trees up to 20 m. or more, the branches armed with numerous short stout
spines, glabrous. Leaves palmately compound with 5-9 very shortly petiolulate
oblanceolate shortly acuminate entire leaflets 7-12 cm. long and 2-3 cm. broad,
dark green above, very heavily glaucous beneath, the petiolules about 1.0-1.5 cm.
long; petioles slender, 10-20 cm. long. Inflorescences shorter than the subtending
petioles, the staminate racemose-paniculate, many-flowered, the pistillate erect,
long-pedunculate, 1-flowered. Staminate flowers about 17 mm. long, very glaucous
without, narrowly clavate in bud, the lobes about 7 mm. long, linear, obtuse, the
tube narrowly cylindric, about 1.2 mm. broad, sparsely villous within; stamens
strigose, strongly unequal, the filaments conspicuously connate, the anthers similar,
without a ligulate connective, with a small erect spiniform process. Berry pyri-
form, up to about 7 cm. long and 3 cm. broad, orange, the fruiting peduncle
pendulous, up to about 10 cm. long.

Costa Rica and Panama, in forests near sea level to about 700 m. elev. Tonduz
reports the vernacular name in Costa Rica as papayillo de venado.

COLÓN: vicinity of Camp Piña, alt. 25 m., Allen 3679.

The description of the flowers is taken from Johnston; our specimen is fruiting.
Jacaratia costaricensis is closely related to J. spinosa (Aubl.) DC., as noted by
Johnston. Both species are very glaucous, particularly the lower surface of the
leaves, and the chief distinguishing character obvious from our meagre herbarium
representation appears in the length of the fruiting peduncles, which are only up
to about 2 cm. and suberect in J. spinosa.

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[Vol. 45
26 ANNALS OF THE MISSOURI BOTANICAL GARDEN

2. CARICA L.

CARICA L. Spec. Pl. 1036. 1753.
Papaya [Tourn.] Adans. Fam. PI. 2:357. 1763.
Vasconcellea St. Hil. 2e. Mem. Resed. 13. 1837.

Dioecious or monoecious trees and shrubs, the thick succulent trunk usually
unbranched. Leaves simple and palmately lobed in the Panamanian species, long-
petiolate. Inflorescences in the axils of the uppermost leaves, thyrsiflorous, the
staminate extensive and many-flowered, the pistillate reduced and with fewer
larger flowers. Calyx 5-lobed, inconspicuous. Corolla salverform in the staminate
flowers, campanulate in the pistillate, the limb 5-lobed, alternisepalous. Stam
10, the connective occasionally produced at the tip. Ovary usually 1-celled, sle
septate at the base, containing numerous ovules on parietal placentas. Berry
broadly ovoid-pyriform, 1-celled.

Although the papaya is one of the most common and favorite fruits of the
tropics, appearing to the outlander like some celestial melon, the genus is in a state
of botanical confusion with at least 50 poorly defined "species" ranging from
Mexico to Argentina and in the Antilles. It has been introduced into the Eastern
Hemisphere since long ago and frequently escapes.

a. Leaves with 7 palmate primary veins, usually with 7 deeply and irregu-
larly pinnatifid segments; pistil of pistillate — with 5 dichotomous
lo

or trichotomous stigma ; staminate flowers with broadly deltoid
calyx lobes about 1 mm. long and I or white corollas 3-4 cm. long. 1. C. PAPAYA
aa. Lil with 5 ai e pr imary veins, with 5 shallow segments each with

1-2 broad Leni lobes; pistil d pille poe ith 5 linear un-
divided stigma lobes (presumptive for C. c uensis).
b. Leaves and inflorescences glabrous or paio lly so, the middle seg-
ment of the leaves 3-lobed, the 4 lateral segments unlobed; (staminate
pel with White corollas about 4 cm. long and ovate calyx lobes
out 1 mm. long?) 2. C. CUCURBITIFOLIA
b. Lu and inflorescences rather pacer villous, the middle segment
x the a 3-lobed, the 4 lateral segments with a single broadly
te lo aminate flowers wit i pale gu corollas 1.5—2.5 cm. Du
sad. mew calyx lobes about 3 mm. lon 3. C. CHIRIQUENSIS

o

1. CARICA PAPAYA L. Sp. Pl. ed. 1. 1466. 1753.
Papaya sativa Tussac, Fl. Ant. 3:45. pl. ee 1824.
Carica Mamaja T FI. Flum. 10: pl. 131. 1827.
Papaya vulgaris A. DC. in DC. Prodr. $3 4M. 1864.
Carica bermabbrodita Blanco, FI. Filip. 3:212. 1879.

Dioecious or occasionally polygamous or monoecious trees 2-8 m. tall with
characteristic stout succulent and unbranched trunks, usually glabrous in all parts '
or essentially so. Leaves in a dense terminal crown, long-petiolate; blade simple
but palmately divided into usually 7 more or less irregularly and broadly pinnatifid
segments, variable in size but frequently up to 4—5 dm. in diameter; petiole usually
up to 5-7 dm. long. Staminate inflorescences repeatedly compound, many-flowered,
somewhat shorter than the subtending petioles, spreading or pendulous, the pistillate
far shorter than the subtending petioles, usually only 1- to 3-flowered. Staminate

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1958]

FLORA OF PANAMA (Caricaceae)

27

Fig. 4. Carica papaya

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[Vol. 45
28 ANNALS OF THE MISSOURI BOTANICAL GARDEN

flowers: calyx lobes very broadly deltoid, obtuse, about 1 mm. long; corolla salver-
form white or yellow, the tube about 1.5—2.0 cm. long, about 2 mm. in diameter,
the lobes oblong, 1.0-1.5 cm. long, slightly spreading; stamens inserted at the
orifice of the corolla tube, exserted, the antepetalous anthers about 2.5 mm. long,
nearly sessile and descending basally into the corolla tube, the antesepalous anthers
erect, wholly exserted, about 2 mm. long, with apically 2-lobed filaments of nearly
equal length; pistillode acicular, about 1 cm. long. Pistillate flowers far larger
than the staminate, irregularly campanulate, white or pale cream: calyx lobes
broadly deltoid, about 1 mm. long; corolla essentially polypetalous or very weakly
united, the segments about 5-7 cm. long, 1.5—2.0 cm. broad, broadly acute, irregu-
larly reflexed toward the tips; staminodia lacking, rarely minute and hy nous;
ovary ovoid-ellipsoid, usually about 3 cm. long (including the indefinitely attenuate
style) and 2 cm. broad, the stigma essentially sessile, of 5 thick segments about
1 cm. long fimbriately di- or trichotomous toward the tip. nis extremely
variable, up to 3 dm. long and 2 dm. thick, green or yellow when ri

Papayos are grown throughout the tropics of the world, chiefly at fe eleva-
tions, and tend to become escapes everywhere. In Panama they are one of the
commonest weed trees, and the quality of their fruits varies from the sublime to
the ridiculous. Some of the most inferior produce fruits scarcely larger than a
hen's egg. The vast man-made distribution of Carica papaya is quite at variance
with the very restricted natural ranges of the other published species of the genus,
and even the approximate site of the progenitors of the cultivated papaya is now
impossible to ascertain: it may well be the eastern slopes of the Peruvian Andes
where most of the wild species with similar leaf forms occur today. Under such
circumstances the citation of herbarium specimens is apt to become somewhat
absurd, but is appended to this account for good measure.

Associated with the latex of all parts of the plant are various proteolytic en-
zymes, particularly papain, which are isolated and used for various pharmaceutical
and gastronomic purposes. In Panama leaves of papayo sometimes are cooked with
tough meat, or the raw meat softened by wrapping in green leaves and left over-
night. Hernando Cortés is said to have observed this use of papayo during his
conquest of Mexico in the early eighteenth century, and from this source has
developed the recently booming process of meat tenderizing, in which papain is
a prominent agent.

Carica papaya normally is dioecious. In some districts of Panama the inhabi-
tants adhere to a "superstition" that barren (i.e. staminate) trees of papayo can
be made fruitful by deeply inscribing in the bark the Sign of the Cross with a
machete. I have been shown examples of such incised trees, which do indeed
appear to become polygamous. Local experience also has it that such a tree does
not become fruitful permanently but must be wounded periodically. The response
would appear to have scientific foundation upon the basis of traumatic reversion.

DEL TORO: Careening Cay, Von Wedel 573. CANAL ZONE: Barro Colorado

Island Send 165; low ground, outskirts of Ancón, Greenman t$ Greenman 5073;
Cocoli Island, vicinity of Miraflores Lake, P. W bite 290.

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1958]
FLORA OF PANAMA (Caricaceae) 29

Fig. 5. Carica cucurbitifolia

2. Carica cucurbitifolia Woodson, spec. nov.

Arbores ca. 3-6 m. altae. Folia longe petiolata glabra lamina simplici palmatim

5-divisa basi 5-venosa segmento centrali maximo profunde lateque 3-partito usque
15 cm. longo et lato vel ultra (speciminis nostri juvenalis) segmentis 2 mediis ca

12 cm. longis 6 cm. latis atque 2 basalibus 6 cm. longis 3 cm. latis omnino li

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[Vol. 45
30 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Inflorescentia mascula jam ignota feminea valde contracta usque ca. 7-flora pe-
dunculo ca. 2 cm. longo. Florum femineorum calycis laciniae minute trigonales
ca. 2 mm. longae. Corolla ejusdem generis tubulosa ca. 2.5 cm. longa tubo ca. 3
mm. diam. basi videtur lacteo lobis anguste oblongis ca. aequilongis erectis apicem
versus roseo-tinctis. Staminodia nulla. Ovarium in flore ovoideum ca. 6 mm.
longum glabrum in stylo angusto ca. 3 mm. longo abrupte contractum stigmatibus
5 linearibus integris 10-12 mm. longis. Bacca ignota.

in heavy forest below grass, region north of El Valle, grass ridge known as the
Loma à del Pel alt. 2800 ft., April 8, 1947, P. H. Allen 4478 (Herb. Missouri Bot. Gard.,
HOLOTYPE).

is ovary of C. cucurbitifolia, with its slender style, recalls that of C. s£ylosa
Heilborn of Peru, the leaf segmentation of which, however, corresponds more
closely with that of C. papaya. It is unfortunate that staminate flowers are not
known. In the herbarium of the Missouri Botanical Garden is a staminate specimen
of papayo collected by Skutch (no. 4797) in Costa Rica which appears to be re-
lated to C. cucurbitifolia, although with a differently divided leaf more similar in
outline to that of the succeeding species. The Skutch plant has white corollas
about 4 cm. long and ovate calyx lobes about 1 mm. long. It definitely is not
C. papaya as presently determined.

3. CARICA chiriquensis Woodson, spec. nov.

Arbores usque 7 m. altae vel ultra. Folia longe petiolata lamina simplici pal-
matim 5-divisa basi 5-venosa segmento centrali maximo profunde lateque 3-partito
usque 35 cm. longo nisi ultra segmentis mediis ca. 25 cm. longis 12 cm. latis
margine inferiore late auriculato segmentis 2 basalibus ca. 20 cm. longis 10 cm.
latis margine inferiore acute auriculato membranacea pagina inferiore sparsiuscule
villosula. Inflorescentia feminea jam ignota mascula effuse thyrsiflora pedunculis
gracilibus dense villosulis. Florum masculorum calycis laciniae lanceolatae apice
acuminatae laxe reflexae ca. 3 mm. longae margine minute ciliato. Corolla florum
ejusdem generis pallide viridis magnitudine valde variabilis tubo anguste cylindrico
extus sparse pilosulo ca. 1.5 mm. diam. tum 5 mm. tum 15 mm. longo lobis line-
aribus ca. 1 mm. latis extus apice versus densius pilosulis tum 7 mm. tum 12 mm.
longis laxe patulis. Stamina 10 ad ostium corollae tubi posita exserta antepetalorum
antheris cum appendice apicali ca. 1.5 mm. longis filamento ligulari integro ca. 0.5
mm. longo alternorum vix 1 mm. longis filamento simili sed 2 mm. longo. Pistil-
lodium anguste aciculiforme ca. 8 mm. longum. Bacca ut dicitur ca. 15 cm. longa
13 cm. lata 5-angulata.

CHIRIQUf: Quebrada Velo, vic. Finca Lérida, alt. 5000 ft., July 24, 1947, P. H. Allen
4675 (Herb. Missouri Bot. Gard., HOLOTYPE).

Perhaps it is tempting fate to describe two new dioecious species, apparently
closely related, from such neighboring areas as Chiriquí and Coclé when only the
complimentary sexual forms are known. The forms differ as indicated in the key
and the descriptions, however, and C. cucurbitifolia is glabrous while C. chiriquensis

(30)

1958]
FLORA OF PANAMA (Caricaceae) 31

is villous. The leaf form of chiriquensis is quite similar to that of the Skutch plant
from Costa Rica which was discussed under cucurbitifolia, but the staminate
flowers are strikingly dissimilar.

Fig. 6. Carica cbiriquensis

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[Vol. 45
32 ANNALS OF THE MISSOURI BOTANICAL GARDEN

LOASACEAE

Mostly perennial or annual herbs, sometimes quite massive, erect or clambering,
infrequently twining or climbing, characteristically strongly hispid and with
stinging hairs and glochids. Leaves opposite or alternate, variously incised and
occasionally palmatifid or pinnatifid; stipules lacking or vestigal Inflorescence
terminal or extra-axillary, basically cymose but frequently monopodial and occa-
sionally reduced to a solitary flower. Flowers dichlamydeous, hermaphrodite,
epigynous, 5- or rarely 4-, 6-, or 7-merous; sepals foliaceous, imbricate or open in
aestivation; petals free or united at the base, imbricated or contorted; stamens
rarely 5-10 or fewer, usually very numerous, the filaments more or less united to
the base of the corolla, all fertile or partly staminodial and more or less petaloid;
ovary inferior, usually 3- to 5-carpellate and 1-celled with numerous ovules on
parietal placentas, rarely 1-carpellate with a single pendulous ovule (Gronovia),
the style simple with a capitate or obscurely divided stigma. Fruit a variously
dehiscent capsule.

Except for the peculiar monotypic genus Kissenia of Africa and southern Arabia,
the Loasaceae are an American family most frequent in arid or semi-arid regions
with greatest diversification in Mexico and the southwestern United States and in
Chile. Those who have encountered them in the field remember them henceforth
chiefly because of their painfully stinging hairs which seem to have been specially
contrived to protect the elaborate and frequently very showy flowers.

a. Ovary 3- to 5-carpellate, with numerous ovules or parietal placentas;

stamens 10 to many, fr equently ¿re n part; erect or spraw ing
herbs; leaves variously incised or lobed, sometimes pinnatifid or >
b. Stamens all fer = or Ec outer cun ential petaloi 1»... 1. MENTZELIA
bb. Fertile stamens palous, staminodia ante:
c. Flowers etos | Stamin odia free or carat persa at the base........... 2. KLAPROTHIA
ce. pese > (rarely 7- nre staminodia strongly sated into
r less saccate petaloid scale 3. Loasa
aa. Cus caspa wit a a Du apical pendulous ovule; stamens 5, all
fertile; lead twiners; leaves very deeply and palmately divided............ 4. GRONOVIA

The genus Sclerotbrix extends from southern Mexico to Bolivia and is to be
expected in Panama. It resembles Klaprothia strongly, but may be recognized
readily by its spirally contorted fruits.

1. MENTZELIA L.

MENTZELIA L. Syst. ed. 10. 1076. 1759; Urb. in Nova Acta Abh. K. Leop.-Carol.
Deutsch. Akad. Naturf. 76:22.

Bartonia Pursh, in Sims, Bot. Mag. - t. 1487. 1812, non Muhl.
Acrolasia Presl, Rel. Haenk. 2:39. 5

orreya Eaton, Man. Bot. ed. 7. Bai 836, nec > nec alior.
Creolobiai Lilj. FI. Oefver Sver. Odl. Vest. 67. 1839.
Tracbypbytum Nutt. ex Torr. & Gray, Fl. N. A. 1:533. 1840.
Cbrysostoma Lilj. in Linnaea 15:263.
Hesperaster Cockerell, in Torreya 1:142. 1901.

G2)

1958]
FLORA OF PANAMA (Loasaceae) 33

Y
itis

"
M
ge

e "
| eZ

A e
ua)

Fig. 7. Mentzelia aspera

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[Vol. 45
34 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Herbaceous annuals or perennials, rarely small shrubs or trees, the stems and
foliage scabrous and hispid, but without stinging hairs. Leaves alternate or op-
posite, variously incised or lobed. Inflorescence terminal, cymose, usually few-
flowered. Flowers large to mediocre; sepals foliaceous, 5; petals 5-10, free or barely
united at the base; stamens 10 to very numerous, free or united at the base of the
filaments, wholly fertile or the outermost becoming petaloid and sterile; ovary
inferior, 1-celled, with few to numerous ovules borne upon 3-7 parietal placentas,
the style elongate, filiform, the stigma scarcely dilated, capitate or obscurely divided.
Fruit an apically dehiscent 3- to 7-valved capsule.

A genus of nearly 100 species, principally in the western United States and
Mexico, but extending along the Pacific coast of South America to Patagonia. A
single species is known from Panama.

1. MENTZELIA ASPERA L. Spec. PI. ed. 1. 516. 1753.
Mentzelia stipitata Presl, Rel. Haenk. 2:40. 1831.
Mentzelia pedicellata Pred, Epim. Bot. 246. 1849.
Acrolasia squalida Hook. f. in Trans. Linn. Soc. 20:222. 1851.

. Slender erect or sprawling annuals. Stems slender, dichotomous, up to about
6 dm. long, rather thinly covered with short weak glochidiate hairs. Leaves alter-
nate, rather sparse, shortly but evidently petiolate, the blade rather hastately
3-lobed and irregularly serrate, acuminate, broadly obtuse at the base, about 8 cm.
long and 4 cm. broad or somewhat larger toward the base of the stem, diminishing
gradually toward the tip, lax and thinly membranaceous, weakly and variously
strigillose with simple hairs above and below, the petiole about 5 mm. long. Flowers
in few-flowered leafy-bracted cymes at the tips of the branches, mediocre, pale
orange or salmon, sessile or essentially so; hypanthium narrowly cylindrical, about
1 cm. long and 1 mm. broad at anthesis, densely covered with slender glochidiate
hairs, the calyx lobes broad trigonal at the base, very narrowly subcaudate-
acuminate, nearly as long as the hypanthium, sparsely strigillose but without
glochidiate hairs; petals 5, obovate, up to 1 cm. long and 5 mm. broad, spreading,
pale orange or salmon, slightly united at the base; stamens about 20-30, all fertile,
but the outermost somewhat longer and with enlarged somewhat petaloid filaments,
somewhat shorter than the petals. Capsule narrowly cylindrical but slightly at-
tenuate toward the base, up to about 2.5 cm. long and 5 mm. broad, 3-valved,
covered with the persistent glochidiate hairs.

One of the most widespread species of the genus, occurring from Arizona to
Argentina. Rather frequent in dry thickets upon the Pacific slope of Panama.
CANAL ZONE: Balboa Heights, Greenman & Greenman 50553 Ancón Hill, Greenman &

Greenman 5114, Standley 26360; between Corozal and Ancén, Pittier 2209; Gamboa,
Pittier 4793; Sosa Hill, Balboa, Standley 25256, 32150; Darién "co. Standley 31521.

2. KLAPROTHIA HBK.
KLAPROTHIA HBK. Nov. Gen. & Sp. 6:96. 1823.

Herbaceous annuals, the stems and foliage rather inconspicuously hispidulous to

(34)

1958]
FLORA OF PANAMA (Loasaceae) 35

essentially glabrous, without stinging hairs. Leaves opposite, distinctly petiolate,
serrate. Inflorescence terminal, diffusely cymose, bearing rather numerous small
4-merous flowers. Hypanthium subglobular, the calyx lobes 4, broadly trigonal;
petals 4, free; stamens in antepetalous groups of 4—5, with slender minutely
papillate filaments; staminodia in antesepalous groups of 4—5, with slender pilosu-
lose filaments barely united at the base and incised flabelliform sterile anthers;
ovary 4-carpellary, 1-celled, with 1—2 ovules upon the 4 parietal placentas; style
filiform, the stigma very obscurely divided. Fruit a small septicidally 4-valved
capsule.

A monotypic genus of relatively high elevations extending from Costa Rica to
Venezuela and Bolivia.

Fig. 8. Klaprothia mentzelioides

1. KLAPROTHIA MENTZELIOIDES HBK. Nov. Gen. 8 Sp. 6:96. 1823.

Weak erect or sprawling herbaceous annuals. Stems slender, branching infre-
quently, up to about 6 dm. long, usually less, inconspicuously pilosulose to essen-
tially glabrous. Leaves opposite, rather sparse, lax and long-petiolate, the blade
elliptic to ovate or lanceolate, acuminate, obtuse or somewhat attenuate at the base,
rather evenly serrate, up to about 10 cm. long and 5 cm. broad, delicately mem-
branaceous, minutely strigillose upon both surfaces, the petiole slender, up to about
3 cm. long, commonly horizontally spreading or somewhat reflexed. Inflorescence
terminal, laxly cymose, bearing rather numerous small white shortly pedicellate
flowers. Hypanthium subglobose, about 2 mm. broad, minutely and densely
hirsutulose, the calyx lobes broadly trigonal, about 1 mm. long; petals ea oval
or obovate, somewhat cucullate, 6—8 mm. long. Stamens and stamin e

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[Vol. 45
36 ANNALS OF THE MISSOURI BOTANICAL GARDEN

genus, about as long as the petals. Capsules oblongoid, about 4—6 mm. long and
—4 mm. broad, persistently hirsutulose.
Rather frequent in the moist highland forests at elevations of 1500—2000 m.

S DEL TORO: Robało Trail, northern slopes of Cerro Horqueta, Allen 4917, 5002.
crasionii Bajo Chorro, Woodson t$ Schery 650, Davidson 82.

3. LOASA Adans.
Loasa Adans. Fam. 2:501. 1763.
Ortiga Neck. Elem. 2:400.
Huidobria C. Gay, Fl. Chil. 4 ode 1846.

Annual or perennial herbs or subshrubs, erect or sprawling, rarely twining,
beset in nearly all parts with scabrous, stinging hairs. Leaves alternate or opposite,
variously incised and lobed. Inflorescence cymose, terminal or lateral, occasionally
reduced to a solitary flower. Flowers mediocre to large, usually 5-merous, rarely
6- to 7-merous; sepals foliaceous, open in aestivation; petals valvate or imbricate,
somewhat clawed; stamens numerous, in antepetalous clusters, the filaments elon-
gate, united at the base; staminodia united into conspicuous more or less saccate
antesepalous petaloid scales; ovary inferior, 1-celled, with usually numerous ovules
borne upon 3-5 parietal placentas, the style usually shortly columnar, the stigma
obscurely divided. Fruit a 3- to 5-valved capsule.

75-100 species, extending from Mexico to Patagonia, but particularly numerous
in Peru and Chile. The Loasas sting probably more painfully than any nettle, and
the quality sometimes is not completely lost from herbarium specimens.

a. a a or very Rewa! pcne above; flowers mediocre, about
e petals w 1. L. RUDIS

aa. Lara Era vani: lobed Lt ici
b. Leaves somewhat longer than broad, somewhat pinnately lobed;

flowers about 5 cm. in diameter, > petals pale green 2. L. GRANDIS
bb. Leaves about as long as broad, somewhat palmately lobed; flowers
about 15 cm. in diameter, the ab bright orange 3. L. SPECIOSA

1. LOASA RUDIS Benth. Pl. Hartw. 75. 1839.
Loasa A get je a in una Di n 20
Loasa bipinnata Donn
Loasa triphylla Juss. var. wi rudis (Benth j “Usb. à Gia: in Nova Acta, Abh. K. Leop.-Carol.
Deutsch Akad. Naturfor. 76:239
Weak annual herbs up to "e 1 m. tall. Leaves alternate, rather sparse,
pinnate or very deeply pinnatifid above, the uppermost greatly reduced and simple,
the lower shortly petiolate, up to about 15 cm. long and 9 cm. broad, with 3-5
pairs of oblong-elliptic coarsely serrate leaflets or lobes. Flowers several in laxly
monochasial, leafy-bracted terminal cymes; pedicels 5-10 mm. long; calyx lobes 5,
broadly ovate-trigonal, obtuse, 6 mm. long and broad; petals 5, broadly spatulate,
1 cm. long, widely spreading, white; staminodia profoundly saccate, about 5 mm.
long, striped with red or pink. Capsules erect upon the accrescent pedicels, rather
narrowly campanuate, up to 2 cm. long and 1 cm. broad, densely hispid, the
persistent calyx lobes accrescent and about half as long as the capsule.

(36)

1958]
FLORA OF PANAMA (Loasaceae)

M are
an N ji h DN
ES “4 ROM 1) "
$5 ly = 5

‘ :
A

SIDE

za
= RA
DY E `

ES
Ñ

EN

UI

2

ROLĄ RJ:
A

Sia
E |
=

ki

RS

Fig. 9. Loasa rudis

(37)

es Dl
IN
NS

37

[Vol. 45
38 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Southern Mexico to Panama, where the plant is known locally as pringamoza
or pringamosca. A frequent herb in pastures and forest clearings at elevations of
1500-2000 m.

CHIRIQUÍ: Río Chiriqui Viejo valley, near Cerro Punta, G. White 39; Bajo
Cho orro, Bo diste District Davidson 277, Woodson 6 Schery 645; Casita Alta, Volcán de
Chiriquí E Wolo Allen & Seibert 936.

This plant probably is not conspecific with true L. źripbylla of Ecuador and
Peru which, as the name suggests, has palmately divided leaves. However, our
plant may eventually prove to be conspecific with L. papaverifolia HBK. which
extends from Colombia and Venezuela to Chile; in this case, L. rudis will have to
be superceded by the earlier name.

2. Loasa GRANDIS Standl. in Journ. Wash. Acad. Sci. 17:12. 1927.

Very large coarse herbs up to about 4 m. tall. Leaves alternate, rather sparse,
long-petiolate, broadly oval and divided into 6-8 broad coarsely serrate lobes, up
to 4 dm. long and 3 dm. broad, gradually reduced above, the petiole 3-8 cm. long.
Flowers several or rather few in leafy-bracted, lax terminal cymes; pedicels about
2 cm. long; calyx lobes 5, broadly ovate, about 1 cm. long, sparsely serrate-dentate;
petals 5, pale green, cucullate, broadly obovate, about 2.5 cm. long, 1.5 cm. broad;
scales broadly oval-cucullate, 5-6 mm. long, entire or broadly 3-lobed at the tip
(exappendiculate?) 3-nerved, the opposite staminodia narrowly acicular, long-
acuminate, minutely puberulent immediately above the slightly enlarged base,
about 1 cm. long; stamens in groups of about 12 opposite the petals, the filaments
about 1.5 cm. long, the anthers about 2 mm. long; roof of the ovary about 1 cm
in diameter, densely arachnoid-villous; style about 5 mm. long. Capsules broadly
turbinate, about 2 cm. long, dehiscing with 5 broad valves alternate with the calyx
lobes; seeds innumerable, about 1 mm. long or somewhat less.

Panama and Costa Rica (?), in clearings of moist forest, 70-1000 m. elev.

ZONE: forest along banks of Quebrada La Palma and Cańon of Rio Chagres,
dzi $ “Allen 17336. COCLE: north rim of El Valle de Antón, Allen 1658.

The type of L. grandis comes from Guanacaste, Costa Rica. 1 have not been
able to detect the three small conic appendages below the apex of the scales
described by Standley for the Costa Rica plants.

3. LOASA SPECIOSA Donn. Sm. in Bot. Gaz. 23:8. 1897.

Large coarse weak herbs up to 2 m. tall, abundantly covered with long brown
stinging hairs. Leaves opposite or subopposite, long-petiolate, the blade about as
long as broad, somewhat palmately lobed and serrate in the manner of a maple leaf,
up to 15 cm. long and broad or somewhat larger below, the petiole up to 12 cm.
long. Flowers solitary or in few-flowered terminal cymes, the pedicels up to 6 cm.
long; calyx lobes 5, ovate-lanceolate, acuminate, about 3 cm. long; petals 5, bright
orange, broadly oval, 5-6 cm. long, flat or essentially so, slightly spreading; scales
about 1.8 cm. long, broadly truncate-triangular, the tip unequally 4-lobed, the

(38)

1958]

FLORA OF PANAMA (Loasaceae)

eI rę

E.

}
4 Y
O (PAM
a NI A h M
x M Wi y GY,
Y LA AA
ni 4
A
La
WE
Nt
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Y y
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WI
Y
M
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4

Fig. 10. Gronovia scandens

(39)

39

[Vol. 45
40 ANNALS OF THE MISSOURI BOTANICAL GARDEN

lateral lobes somewhat longer, with 2 broadly oval sacs at the base, the staminodia
2, about 2.5 cm. long, narrowly acicular but greatly bulbose at the base, minutely
and closely aculeolate; stamens in groups of about 14 opposite the petals, the fila-
ments glabrous, about 2 cm. long, the anthers about 2 mm. long; style slender, 1.5
cm. long, minutely hispidulose toward the base, the stigma subcapitate. Capsules
narrowly campanulate, about 2 cm. long and 1 cm. broad, bearing innumerable
reticulate seeds about 1 mm. broad upon usually 3 parietal placentas.

Highland Costa Rica and western Panama, in forest clearings at about 2000—
2500 m. elev.

CHIRIQUi: Bajo Chorro, Boquete District, Davidson 445; vicinity of Cerro Punta, Allen
86.

With its bright orange flowers 15 cm. or more in diameter, this is one of the
most showy herbs of the Chiriqui mountains, but is also one of the most painful
ortigas.

4. GRONOVIA L.
GRONOVIA L. Sp. Pl. ed. 1. 202. 1753.

Slender annual twiners covered with weak uncinate hairs on all parts. Leaves
alternate, long-petiolate, the blade very deeply and palmately divided and coarsely
entate, broadly ovate-reniform, deeply cordate. Inflorescence extra-axillary,
several times dichasial with the ultimate branches monochasial, bearing numerous
rather small sessile flowers. Calyx lobes 5, subfoliaceous, valvate; petals 5, free,
open in aestivation; stamens 5, antesepalous, all fertile; ovary 1-celled, containing
a solitary pendulous ovule, the style rather slender, the stigma subcapitate. Fruit a
small 5-winged cypsela crowned by the persistent calyx.
Two species, extending from Mexico to Venezuela and Peru.

1. GRONOVIA SCANDENS L. Sp. PI. ed. 1. 202. 1753.

Slender annual twiners, weakly uncinate-pilose in all parts. Leaves rather sparse,
long-petiolate, the blade broadly ovate-reniform, deeply cordate, up to 8 cm. long
and 9 cm. broad, deeply and palmately divided into usually five unequal, narrowly
acuminate lobes, the middle somewhat longest, entire, the lateral decreasingly
smaller and with 1 to several broad dentate incisions, the petiole up to 7-8 cm.
ong. Inflorescence about equaling the leaves, compound, bearing numerous rather
small sessile flowers. Flowers greenish yellow; sepals oblong-lanceolate, acute, 5-6
mm. long; petals rather narrowly spatulate, acuminate, about 3 mm. long; stamens
about 4 mm. long, the filaments scarcely longer than the anthers. Fruits pentag-
onal-obovoid, about 3—4 mm. long, decurrently 5-winged from the tip, crowned by
the persistent, somewhat accrescent sepals.

Southern Mexico to Venezuela and Ecuador, in lowland thickets.

PANAMA: vicinity of Old Panamá, Woodson & Schery 948.

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1958]

FLORA OF PANAMA (Begoniaceae)

BEGONIACEAE

By LYMAN B. SMITH AN

D BERNICE G. SCHUBERT

1. BEGONIA L.

BEGONIA L. Sp. Pl. 1056. 1753; A. DC.

Eubetalum Lindl. Nat. Syst. ed. 2, 440. 1836.
Trilomisa Raf. Fl. Tellur. 2:91. 1836.

Platyclinum T. Moore in Moore & Ayers, G

in DC. Prod. 151:278. 1864.

7.
ard. Mag. Bot. 2:153. 1850. n prov.

Spbendantbera Hin. in Versl. Acad. Wetensch pap 4:139. 18
Nepbromischus Kl. in Ind. Sem. Hort. Berol. App. 1
zia i

. 1854.

Pritzelia Kl. in Midi Akad. Berliti 126. 18

Stibadotheca KI. in b fonder. Akad. Berlin M Pru i as i Sralolbece".
854.

Casparya Kl. in Monatsber. Akad. Berlin 127.
ae KI. in Monatsber. Akad. Berlin 127.

i 854, as “Isopteris”.

KI. in Monatsber. agno Berlin 128. 1854

nem Bout. ex A. Dc. in DC. Prod. 15!: :406.

1864.

(41)

41

IVol. 45
42 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Begoniella Oliver, in Trans. Linn. Soc. 28:513. 1873.
Semibegoniella C. DC. in Bull. Herb. Boiss. II. 8:327. 1908.

Annual or perennial herbs, shrubs or rarely small trees. Stem elongate and
succulent or woody or a cylindrical rhizome or sometimes reduced to a tuberous
rhizome, sometimes climbing. Leaves usually alternate or rarely subverticillate,
entire, serrate, lobed or digitately compound, usually asymmetric, sometimes peltate,
stipules 2, free, persistent or deciduous. Peduncles mostly axillary. Inflorescence
usually cymose, sometimes 1-flowered or racemose. Cymes regular or strongly one-
sided, usually dichotomous, unisexual or bisexual. Flowers monoecious. Tepals
free or rarely connate, usually 4 staminate in two pairs and 5 pistillate. Stamens
indefinite, inserted on the receptacle, filaments free or united. Styles usually 3,
free or connate, usually bifid, ovary inferior, placentae axile, simple or divided.
Fruit capsular, usually bearing 3 unequal wings.

a. ice acaulescent or subacaulescent with the leaves clustered near the
of the rhizome, or with slender creeping stolons if the internodes are

saa.
b. Leaves peltat
c. Cymes ied unilateral; capsule-wings slightly unequal; leaves
2—12.5 cm. long . B. CONCHIFOLIA

. Cymes regular; capsule-wings very unequal; leaves 17—40 cm. long. 2. B. NELUMBIIFOLIA
bs. Tarta basifixed.
Cymes po unilateral.

d. Stipules PPE coarsely reticulate-nerved; staminate and
sedium a 2; largest apreni as high as wide................. . B. PLEBEJA
dd. Stipules des ~cariaate with fine nspi icuous ar staminate
tepals est capsule-wing ie thee i igh . B. GARAGARANA

cc. Cymes reg
d. s anches s the cyme 3 or 4 at each node; inflorescence many-
5. B. QUATERNATA

dd. pus of the cyme 2 at each node.
Plants or at least > petioles zada tite.
Indument of the petioles consisting of linear trichomes.
ew di ower „kraje: pals pubescent, the staminate 4, the
ae 3; slender stolons with elongate internodes often

present; capsule reflexed 6. B. CARLETONII
gg. Cyme oe, tepals glabrous, 2 cach; capsule
"yb . B. VILLIPETIOLA

ff. Indument of the petioles consisting of lacerate scales;
pals 2.

be e not at all umbelliform; capsule erec
h.

Larges Boeth narrower than high; leaves entire
or Się 2e serrate, c y ciliate. 8. B. STRIGILLOSA
hh. Largest apela wider than high; anthers much
shorter than the filaments; leaves ciliate- ses iculate..... 9. B. STIGMOSA
gg. Cyme umbelliform with greatly red i
beich: ax i decurved; anthers Saudi than the
m 10. B. vESTITA

ee. Plants completely glabrous.
f. rela: tepals 2 or 3; pistillate bracteoles apparently
ng.

g. Tepals and upper peas | pandena some tepals nearly
epals

equal; stamens on ; pistillate T icti 11. B. BREVICYMA
gg. Tepals and magi p r; ai K pedata very un-
culi apunte nthers longer than the filamen :
E B. MUCRONISTIPULA
s Pistillate. pów 4; aptas bracteoles orbicular, entire;
ses tepals very ual; anthers shorter than the

. B. DAVIDSONIAE

filam
aa. Plants with an erect or ascending stem and distinct internodes.

(42)

d AŚ

1958]
FLORA OF PANAMA (Begoniaceae) 43

Leaves nearly symmetric, ovate 14. B. GLABRA
etri

bb. s: distinctly asymm
. Soft low

annuals; tepals not over 3.5 mm. long
ar Capsule-wings subequal, lunate; pre i. ne; leaves soon
gla 5. B. SEMIOVATA
dd. © renty: very unequal; stamens on a short column; leaves
pressed-pilose 16. B. FILIPES
cc. Cam herbs or kak tepals larg
d. Leaves palminerved, transverse or st “setei oblique

e. Bracts persistent, fimbriate; pistillate tepals 5; staminate tepals

. B. FISCHERI
ee. Bracts „dp ary entire; pistillate tepals 2

f. Cymes few-flowered; peduncles short; se^ cat equal or
iiem: pistillate tep a $
g. Plant pubescent at least on the underside of the leaves;
capsule scarcely d at cu beaked 18. B
gg. Plant glabrous except f ring of trichomes at the apex
the perni bbs slendesy b eaked 19. B
ered; peduncles sai capsule-wings

» OAXACANA

„ UDISILVESTRIS

w d . INVOLUCRATA
gg. Secondary branches moral; rasi distinctly and
regularly cymose; leaves glabr 21. B.
dd. Leaves penninerved, straight o or e diehdy” obli ique.
e. Ovary and capsule e redd ie -horned from the upper part of
nes f

MULTINERVIA

Sy: sle multifi 2-3
ff. Leaves deeply ei at base; anthers subglobose; pistillate
tepals 3; go A bi s B
ee. Ovary bearing 3 wings which are ni aaa the full length of
the angles and are me pe uneq
£; on arca ings subequal; leaves ME. oblanceolate, acu-
2

» URTICAE

» HEYDEI

4. B. TONDUZII

ff. Cine wieża strongly unequal,

g. Leaves subulate-dentate; cymes globose, many-flowered;

stamina ate tepals se subequal . B. OPULIFLORA
gg. Leaves with flat ute not subulate teeth.

h. Pistillate oai 5; styles multifid; staminate tepals 4;
unequal. 6. B. GUADUENSIS

hh. Pill tepek 2 or rarely 3; staminate tepals 2 (or if

styles merely bit
. Interne very short; plants scandent; staminate
2

. B. ESTRELLENSIS
ii. poria elongate; plants usually erect; staminate .

tepals 2 (rarely 3 or
j- Capsule 10-12 m = hab bici of the wings;
cymes moder ately ` Rorifer . B. SEEMANNIANA
jj. Capsule 6-10 mm » cadit of the wings;

ike dd e-denticuleaté, ooo ÓW 29. B. CARPINIFOLIA
kk. Leaves A E: 15 e cm. long, 43. 5 cm. wide, some-
30. B. CONVALLARIODORA

1. BEGONIA CONCHIFOLIA A. Dietr. in Allg. Gartenzeit. 19:258. 1851.

Begonia paa Liebm. in Kjoeb. Vidensk. Meddell. 1852:9. 1853.
Gireoudia concbifolia KI. in oaia cbr. Akad. Berlin 125. 1854.

Gir R pe, lé varii tie Hort. ex A A. DC. in DC. Prod. 151; :338. 1864, nomen in syno
Begonia bumilio Standl. in Ann. Missouri Bot. Gard. 27:322. 1940, non Irmscher, 1929.

Herbaceous. Rhizome oblique, 5 mm. thick, internodes very short. Leaves

(43)

[Vol. 45

di ANNALS OF THE MISSOURI BOTANICAL GARDEN
B conchaefolia B.nelumbiifolia
ni

Fig. 11. Begonia concbifolia and B. nelumbiifolia

peltate, broadly ovate, acuminate, 7—8-nerved, entire or dentate at the ends of the
nerves, 2—12.5 cm. long, nearly or quite glabrous above, rufous-pilose beneath
especially on the nerves, fleshy, petioles 3-10 cm. long, rufous-pilose, stipules per-
sistent, ovate, setose-acuminate, brown, glabrous or sparsely pilose. Peduncles
much exceeding the leaves, to 15 cm. long, rufous-pilose. Cymes bisexual, strongly
unilateral and often much longer than wide, few- to many-flowered. Bracts
deciduous, small, lanceolate to obovate, entire, setose-ciliate. Pedicels 4-12 mm.
long. Staminate tepals 2, suborbicular, 4-5 mm. ong, entire, pink. Stamens few,
free. Pistillate bracteoles elliptic, nearly equaling the ovary. Pistillate tepals 2,
like the staminate. Ovary 3-celled, placentae bilamellate, ovuliferous throughout,
styles connate at base, stigmas lunate-capitate. Capsule erect, subglobose, wings
slightly unequal, sublunate, angled near apex.
Honduras, Costa Rica, Panama.

COCLÉ: epiphytic, vicinity of La Mesa, hills north of El Valle de Antón, alt. 1000 m.,

Allen 2297; vicinity of El Valle de Antón, Allen 2025. PANAMA: on boulders, Cerro
Campana, Allen 2

2. BEGONIA NELUMBIIFOLIA Schlecht. & Cham. in Linnaea 5:604. 1830.

(44)

1958]

FLORA OF PANAMA (Begoniaceae ) 45

Begonia derycxiana Lem. in Hortic. Univ.
5: misc. 355. 18
"moż nelumbiifolia KI. in Monatsber.
in 125. 1854.
iene lbs © DC. in Smithson.
Misc. Coll. 69: no. 12:9. 1919.

Herbaceous. Rhizome repent, short,
15 mm. thick, setose, internodes ex-
tremely short. Leaves peltate, obliquely

very broadly ovate or subelliptic, 7—9- .

nerved, evenly rounded except for the
abruptly acuminate apex or slightly pro-
duced at the ends of the nerves, 17-40
cm. long, 10-28 cm. wide, remotely
denticulate, soon glabrous, the margin
ciliate and sometimes purple, thin, peti-
oles 15-45 cm. long, to 8 mm. thick,
sparsely rufous-hirsute, becoming gla-
brous, stipules persistent, lanceolate, over
m. long, entire, rather firm, pilose.
Peduncle usually exceeding the leaves, to
66 cm. long, 6 mm. thick, soon glabrous.
Cyme regular, much branched, diffuse,
2—5 dm. broad. Bracts deciduous, ovate,
obtuse. Pedicels slender, 9—22 mm. long.
Staminate tepals 2, suborbicular, 6—8
mm. long, white. Stamens few, anthers
narrowly obovate, equaling or longer
than the filaments. Pistillate bracteoles
lacking. Pistillate tepals 2, like the sta-
minate. Ovary 3-celled, placentae bifid,
ovuliferous throughout, styles connate at
base, distinctly divided, the stigmatic
surface linear, spiral, continuous. Cap-
sule erect or somewhat nutant, broadly
ovoid, 6-10 mm. long, wings very un-
equal, the largest ovate, obtuse, to 15
mm. wide, seeds ellipsoid, blunt.
Southern Mexico to Colombia.
NAL ZONE: on rocky bank, forest
dee: the Río Indio de Gatún, near sea level,
Maxon . DARIÉN: Chepigana Th

aa Trail (Camp 1), alt.
Terry 1623.

B.plebeja

A "

ET ASI A
BE RY

Fig. 12. Begonia plebeja

(45)

[Vol. 45
46 ANNALS OF THE MISSOURI BOTANICAL GARDEN

3. BEGONIA PLEBEJA Liebm. in Kjoeb. Vidensk. Meddel. 1852:8. 1853.
Begonia glandulosa sensu J. D. Smith, Enum. Pl. Guat. etc. 4:182. 1895; 5:34. 1899.
Begonia uvana C. DC. in Smithson. Misc. Coll. 69: no. 12:4. 1919

Herbaceous, fuscous-villous when young. Rhizome aly repent, to 8 cm.
long at least, 8-10 mm. thick, internodes very short. Leaves oblique, broadly ovate
or elliptic, acute, palmately 7-8-nerved, denticulate, angulate-dentate or almost
evenly rounded, cordate, 7-20 cm. long, thin, soon glabrous, petioles erect, 6-28
cm. long, usually soon glabrous, stipules persistent, imbricate, lanceolate, acuminate-
setiferous, 1-2 cm. long, entire, heavily and closely reticulate-nerved. Peduncles
exceeding the leaves, up to 22 cm. long. Cymes bisexual, strongly unilateral and
usually much longer than wide, few- to many-flowered. Bracts deciduous, obovate,
ciliate. Staminate tepals 2, broadly ovate, 5-9 mm. long, white or pale pink.
Stamens about 15, filaments short, anthers oblong. Pistillate flowers bracteolate.
Pistillate tepals 2, like the staminate. Ovary 3-celled, placentae bifid, ovuliferous
on all sides, styles short-connate, stigmas lunate-capitate. Capsule suberect, oblong-
ellipsoid, 12 mm. long, wings unequal, the largest subtriangular and as high as wide.

Southern Mexico and Central America

te
& Allen Loo; ; forest along banks of Quebrada Fea, Q. Pura and Cafion of Río Chagres,
alt. 70-100 m., Dodge & Allen I eo forest ing telephone trail between the Río Indio
cacti vicini te rs and the Natu ridge of Rio Puente, Dk & Allen 17482.

es
fer, Sżejezmi & Allen 16683; oh Maestra, alt. 0-25 m., Allen 58; San José
ad. nie on 83, 207, 322, 386, Ex popora Isla de Uva, corn group, Pittier
5109

4. BEGONIA GARAGARANA C. DC. in Smithson. Misc. Coll. 69: no. 12:2. 1919.

Herbaceous, completely glabrous. Rhizome repent, only a small terminal sec-
tion known, internodes very short. Leaves oblique or transverse, palminerved,
strongly asymmetric, broadly ovate or elliptic, abruptly acuminate, shallowly
cordate at base, to 24 cm. long, entire, petioles erect, 25 cm. long, stipules decid-
uous, lanceolate, acute, 2 cm. long, entire, alate-carinate, membranaceous with fine
inconspicuous nerves. Peduncle much exceeding the leaves, to 75 cm. long.
Cymes dichotomous, strongly asymmetric, lax, many-flowered, 15 cm. long.
Bracts deciduous, unknown. Pedicels 24 mm. long. Staminate tepals 4, entire,
glandular-punctate, the outer broadly obovate, 7-9 mm. long, the inner very
narrow. Stamens few on a low torus, filaments very short, anthers narrowly
obovoid, 2.5 mm. long, connective scarcely if at all produced. Pistillate flowers
unknown. Capsule ovoid or ellipsoid, 8—10 mm. long, wings very unequal, the
largest 26 mm. wide, styles deciduous, placentae bifid, ovuliferous throughout, seeds
stalked, ellipsoid.

Endemic.

(46)

19581
FLORA OF PANAMA (Begoniaceae) 47

B.sarasarana B.quaternata

ES E |
A i

Fig. 13. Begonia garagarana and B. quaternata

DARIÉN: Cerro de Garagará, Sambi Basin, southern Darién, alt. 500-974 m., Pittier
pini VERAGUAS: forested slopes of Cerro Tuté, vicinity of Santa Fé, alt. 750 m., Allen
5. BEGONIA QUATERNATA Smith & Schubert, in Journ. Wash. Acad. Sci. 40:244.

fig I, p-s. 1950.

Stemless with the leaves clustered near the apex of the erect densely rooting
rhizome, internodes very short; leaves transverse, very broadly elliptic, obliquely
acuminate and also cuspidate opposite the principal nerves, deeply and narrowly
cordate at base, 17 cm. long, 14 cm. wide, nearly glabrous above, below densely
pilose on the nerves and sparsely elsewhere, petioles slender, 15-18 cm. long, densely
vestite with reflexed lacerate fuscous scales, stipules early deciduous, unknown.
Peduncles 4 dm. long, much exceeding the leaves, slender, glabrous. Inflorescence
regularly cymose, 3—4-radiate, many-flowered, 13 cm. broad, glabrous. Bracts
quickly deciduous, small, elliptic, entire. Flowers white (! Allen) but yellow when

Staminate tepals 2, suborbicular, 5 mm. long, entire. Stamens inserted on
a short column, anthers ellipsoid, slightly shorter than the filaments. Pistillate
bracteoles apparently lacking. Pistillate tepals 2, to 2.5 mm. long. Styles bifid,
ovary 3-winged, wings equal, broadly ovate.

Endemic.

CHIRIQUÍ: vicinity of "New Switzerland", central valley of Río Chiriqui Viejo, alt.
1800-2000 m., January 6-14, 1939, Allen 1336.

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[Vol. 45
48 ANNALS OF THE MISSOURI BOTANICAL GARDEN

B. villipetiola

Fig. 14. Begonia carletonii and B. villipetiola

The general habit of Begonia quaternata is reminiscent of that of B. stigmosa.
However, the flowers of B. quaternata are much smaller, and its cyme has four
primary rays, a character that is very rare if not unique in North American Begonia.

6. BEGONIA CARLETONII Standl. in Journ. Wash. Acad. Sci. 17:314. 1927.

Herbaceous. Rhizome slender with short internodes but also producing very
slender stolons, with internodes 5-8 cm. long, setose-pilose. Leaves asymmetric,
straight or oftener strongly oblique, broadly ovate, abruptly acuminate, cordate at
base, 6—16 cm. long, thin, finely sinuate-dentate, ciliate, sparsely setose-pilose on
both sides, paler beneath, petioles slender, 5-16 cm. long, reflexed-setose, stipules
narrowly lanceolate, 9 mm. long, entire, thin, pilose. Peduncles shorter or longer
than the leaves, slender, pilose. Cymes few-flowered, pilose. Bracts subpersistent,
ovate or elliptic, obtuse, 3-5 mm. long, entire. Pedicels to 18 mm. long. Sta-
minate tepals 4, entire, white, the outer broadly ovate, obtuse, 6-12 mm. long,
sparsely pilose, the inner narrowly oblanceolate, shorter, glabrous. Stamens on a
low torus, numerous, anthers obovate, somewhat shorter than the filaments, con-
nective not produced. Pistillate bracteoles apparently wanting. Pistillate tepals 3,
otherwise like the staminate. Ovary 3-celled, pilose, styles slightly connate at base,
the stigmatic surface lunate. Capsule sharply reflexed, ellipsoid, 7 mm. long, wings
very unequal, the largest reflexed from the lower half of the capsule, oblong, obtuse;
5 mm. wide, the others narrowly lunate.

Endemic.

(48)

SOP. PRISE Do TRA AC a ut al E

ORO

pie ne

a cal

KONA

1958]
FLORA OF PANAMA (Begoniaceae) 49

OCAS DEL TORO: Changuinola Valley, Dunlab 206; locality uncertain, Carleton 206.
clesie Fish oram Mountains, vicinity of Chiriqui Lagoon, von Wedel 2271; Fish
Creek, von Wedel 222

7. BEGONIA VILLIPETIOLA C. DC. in Smithson. Misc. Coll. 69: no. 12:5. 1919.

Herbaceous. Rhizome short with extremely short internodes, rooting densely.
Leaves strongly asymmetric, transversely ovate, abruptly acuminate and often
with a second cusp or lobe more nearly opposite the petiole, cordate at base, 15-21
cm. long, ciliate-denticulate, thin, sparsely pilose above, more densely so beneath
especially on the nerves, petioles slender, 15-25 cm. long, densely villous with linear
trichomes, stipules deciduous, triangular-ovate, acuminate, 15-20 mm. long, pilose.
Peduncle 18-37 cm. long, exceeding the leaves when in fruit, becoming glabrous.
Cymes bisexual, regularly dichotomous, many-flowered, subdense, glabrous except
for the bracts. Bracts quickly deciduous, the lowest large, broadly elliptic, entire,
pilose. Flowers pink or white. Staminate tepals 2, suborbicular, 10 mm. long,
entire. Stamens free, numerous, anthers obovate, shorter than the filaments, con-
nective not produced. Bracteoles lacking. Pistillate tepals 2, like the staminate
but smaller. Ovary 3-celled, placentae bilamellate, ovuliferous throughout, styles
connate at base, stigmatic tissue linear, spiral, continuous. Capsule erect, sub-
globose, 4-5 mm. long, wings distinctly unequal, broadly ovate, the largest to
11 mm. wide.

Endemic.
ANAL ZONE: Mojinga Swamp near mouth of Rio Chagres, alt. 1 m., Allen 864.
kak Bismark above Penonomé, pen: 309; vicinity of El Valle, alt. 600-1000 m. s

Allen 213, 238, 1184, 1664, 2906, 42
8. BEGONIA STRIGILLOSA A. Dietr. in Allg. Gartenzeit. 19:330. 1851.

Gireoudia strigillosa Kl. in Monatsber. Akad. Berlin 125. 1854
Begonia daedalia Lem. in Ill. Hortic. 7: misc. 54. 1860. 8: pl. 260. 1861.
Bonn barbana C. DC M Bull. Soc. Bot. Belg. 35: pt. 1:261.

Herbaceous. Rhizome varying from short and compact sk: very short inter-
nodes and imbricate stipules to slender and branching with distinct internodes.
Leaves oblique, broadly ovate, acuminate, 7—17 cm. long, entire or slightly serrate,
rounded to angulate-dentate, coarsely ciliate, glabrous above, sparsely long-pilose
below especially on the nerves, often spotted or mottled, petioles to 18 cm. long,
often spotted, bearing numerous reflexed fimbriate scales, stipules lanceolate with a
setiferous apex and often fimbriate keel, persistent. Peduncle usually exceeding the
leaves, sparsely pilose, often purple-spotted. Cyme 2-4-branched, subsymmetrical,
mostly glabrous, diffuse. Bracts deciduous, elliptic, entire, mucronulate. Staminate
tepals 2, suborbicular to obovate, 6-9 mm. long. Stamens few, the anthers oblong,
obtuse. Pistillate bracteoles lacking. Pistillate tepals 2, like the staminate. Ovary
3-celled, placentae bifid, ovuliferous throughout, styles persistent, connate at base,
somewhat bifid, stigmas linear. Capsule suberect, ovoid or elliptic-ovoid, locules to
10 mm. long, the two larger wings subtriangular or subobtuse to acuminate.

Guatemala, Costa Rica

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[Vol. 45
50 ANNALS OF THE MISSOURI BOTANICAL GARDEN

B. strigillosa

Fig. 15. Begonia strigillosa and B. stigmosa

Indefinite. Seed no. 7 from Mr. Clayton Kelly of Long Beach, California. Mr.
Rudolf Ziesenhenne who sent us the specimen says that it is called water begonia
for it grows only near streams in Panama. He considers it a distinct species but
so far we are unable to separate it satisfactorily from B. strigillosa. B. strigillosa
has been collected frequently in Costa Rica so that its occurrence in Panama is
to be expected.

9. BEGONIA STIGMOSA Lindl. in Bot. Reg. 31: misc. 32. 1845.

Begonia squarrosa sensu Seemann, Bot. Herald 128. 1853, nomen, non Liebm. 1853.
Gireoudia stigmosa KI. in Monatsber. Akad. Berlin 125. 1854.

Herbaceous. Rhizome repent, to 12 cm. long, 1 cm. in diameter, coarsely
lepidote, internodes very short and covered by the stipules. Leaves oblique and |

(50)

1958]
FLORA OF PANAMA (Begoniaceae) 51

strongly asymmetric, very broadly ovate or suborbicular, abruptly acuminate and
usually with a second cusp more nearly opposite the petiole, cordate at base, 15-30
cm. long, scarcely if at all lobed, ciliate-denticulate, thin, glabrous above or with
a few small white trichomes, the nerves beneath bearing trichomes like those on the
petiole but smaller, petioles erect, to 38 cm. long, 4-10 mm. in diameter, covered
with spreading to reflexed narrowly triangular lacerate pink scales 3-5 mm. long,
stipules tardily deciduous, lanceolate, pilose, membranaceous, 15-20 mm. long.
Peduncle 2-5 dm. long, 7 mm. in diameter, soon glabrous. Cymes bisexual, nearly
regular, few- to many-flowered, 7-22 cm. long, lax, glabrous. Bracts quickly
deciduous, oblong, acute, fimbriate. Pedicels slender, 7-23 mm. long. Flowers
white or pink. Staminate tepals 2, suborbicular, cordate at base, 10—15 mm. long.
Stamens free, very numerous, filaments much longer than the broadly oblong
anthers, connective apiculate-produced. Bracteoles lacking. Pistillate tepals 2 like
the staminate or rarely a smaller one inside the others. Ovary 3-celled, placentae
bifid, ovuliferous throughout, styles bifid, the stigmatic tissue linear, spiral, con-
tinuous. Capsule erect, broadly ovoid, 6-8 mm. long, wings very unequal, the
largest oblong or subdolabriform, 10-15 mm. wide. the others narrowly margini-
orm.

Southern Mexico, Central America and Colombia.

CANAL ZONE: forest along banks of Quebrada La Palma and Sali of Rio Chagres,
70-80 m., Dodge & Allen 17368. cHIRIQUI: forests around El Boquete, alt. 1000-1300
m., Pittier 2896; Maxon 4947; between Hato del Jobo and Cerro Via eastern Chiriqui,
alt. 700—1000 m., Pittier 5411.

10. BEGONIA VESTITA C. DC. in Bull. Herb. Boiss. II. 8:315. 1908.

Herbaceous. Rhizome repent, short, internodes very short. Leaves transverse,
elliptic or ovate, abruptly acuminate, cordate at base, 7-18 cm. long, palminerved,
entire, glabrous above, brown-pilose beneath especially on the nerves, petioles 15-
25 cm. long, subdensely pilose with fimbriate scarious scales, stipules deciduous,
ovate, membranaceous, 10-15 mm. long. Peduncle not much exceeding the leaves
until in fruit, to 35 cm. long, bearing scales like those of the petioles. Cymes bi-
sexual, few-flowered, subumbellate with the ultimate branches greatly reduced.
Bracts deciduous. Pedicels 5-15 mm. long. Flowers white or pink, sometimes
red-punctate. Staminate tepals 2, suborbicular, 5-7 mm. long, rather fleshy,
sparsely pilose. Stamens free, few, anthers oblong, lens than the filaments.
Pistillate bracteoles lacking. Pistillate tepals 2, like the staminate. Ovary 3-celled,
placentae bilamellate, ovuliferous throughout, styles connate at base, bifid, stig-
matic tissue spiral Capsule sharply deflexed, ellipsoid, 10-12 mm. long, pilose,
wings very unequal, the largest subreflexed, broadly ovate, obtuse, to 10 mm. wide,
the others narrowly lunate or marginiform.

Costa Rica, Panama.

CHIRIQUI: rain forest, Bajo Chorro, Boquete District, alt. 1800 m., Davidson 71; epi-

phyte, vicinity of Bajo Chorro, alt. 1900 m., Woodson & Schery War. coek: region
north of El Valle de Asia: alt. 1000 m., Allen 2906.

(51)

52 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 45

B. brevicyma

=
= SS

=
- a
SSS AE ="
Sy
SS EA
Rat w,
=

MAT
*

AL
m ET
SEE
MYR =

cÈ

Fig. 16. Begonias

(52)

1958]
FLORA OF PANAMA (Begoniaceae) 53

11. BEGONIA BREVICYMA C. DC. in Smithson. Misc. Coll. 69: no. 12:3. 1919.

Herbaceous, glabrous. Rhizome 6-7 mm. in diameter, internodes very short.
Leaves oblique, palminerved, strongly asymmetric, broadly subelliptic, abruptly
acuminate with smaller cusps at the end of the principal nerves, cordate at base,
10-14 cm. long, petioles 16-24 cm. long, stipules persistent, broadly ovate, acu-
minate, lacerate near apex, 15 mm. long. Peduncles to 26 cm. long. Cyme regu-
lar, few-flowered. Bracts tardily deciduous, the lower ample and covering the
young flowers, the upper much shorter than the pedicels, glandular. Pedicels to
25 mm. long. Staminate tepals 4, subequal, obovate, 17 mm. long, entire, pinkish
white, glandular, the outer somewhat broader. Stamens on a column, numerous,
anthers obovate, much shorter than the filaments, connective not produced. Bracte-
oles wanting. Pistillate tepals 2, broadly ovate, glandular. Wings of the ovary
very unequal.

Endemic.

CHIRIQUÍ: epi pito, bius of Bajo Chorro, alt. 1900 m., Woodson & Schery 620;
humid forest around L uas Camp, southern slope of Certo: de la Horqueta, Chiriquí,
alt. 1700 m., Maxon 541 zę.

12. BEGONIA MUCRONISTIPULA C. DC. in Smithson. Misc. Coll. 69: no. 12:3.

1919

Herbaceous, completely glabrous. Rhizome 6 mm. in diameter, internodes very
short. Leaves oblique, palminerved, strongly asymmetric, broadly elliptic, apiculate
opposite the principal nerves, otherwise entire, cordate at base, 10 cm. long, petioles
erect, 16 cm. long, stipules persistent, broadly ovate, mucronate, nearly 2 cm.
long (! C. DC.), entire, thin. Peduncle much exceeding the leaves, 37 cm. long.
Cyme regular, few-flowered. Bracts deciduous, unknown. Pedicels to 15 mm.
ong. Staminate tepals 4, entire, the outer broadly obovate, 20 mm. long, one red
and one white, the inner subspatulate, 15 mm. long, white. Stamens free, rather
numerous, anthers oblong-obovate, slightly longer than the filaments. Pistillate
bracteoles lacking. Pistillate tepals 3, the outer suborbicular, 8 mm. long (possibly
immature), the inner one broadly ovate, much smaller. Ovary 3-celled, placentae
bilamellate, ovuliferous throughout, styles 3, connate at base, bifid, stigmatic tissue
spiral. Capsule-wings very unequal, the largest narrowly triangular, much wider
than high, the others narrowly lunate.

Endemic.

IRIQUÍ: between the Rio Ladrillo and Los Siguas Camp, southern slope of Cerro de
la Hause alt. 1200-1700 m., Pittier 3172.
13. BEGONIA DAVIDSONIAE Standl. ex Smith & Schubert, in Journ. Wash. Acad.

Sci. 40:242. fig. I, l-o. 1950

Stemless with the leaves clustered near the apex of the short prostrate densely
rooting rhizome, glabrous, internodes very short; leaves oblique, broadly ovate,
abruptly acuminate and also cuspidate at the ends of the principal nerves, deeply
and narrowly cordate at base, 10 cm. long, 7 cm. wide, petioles slender, 17 cm.

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[Vol. 45
54 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long, stipules deciduous, triangular-ovate, acuminate, 12 mm. long, lacerate toward
apex, thin. Peduncles slender, to 26 cm. long, exceeding the leaves. Cymes regular,
few-flowered, dense. Bracts quickly deciduous. Flowers white or pink (! David-
son). Staminate tepals 4, entire, the outer suborbicular, 12 mm. long, the inner
elliptic, 10 mm. long. Stamens numerous, anthers obovate, much shorter than the
filaments, connective not produced. Pistillate bracteoles orbicular, entire. Pistil-
late tepals 4, like the staminate but smaller. Stigmatic tissue spiral, wings of the
ovary very unequal, the largest oblong, obtuse, much wider than high, the others
narrowly marginiform.

Endemic.

CHIRIQUÍ: rain forest, Bajo Chorro, Boquete District, alt. 1800 m., January 5, 1938,
Davidson 38.

This species appears to be related to Begonia brevicyma and B. mucronistipula
as indicated in the foregoing key, however, it has not been possible to ascertain the
detailed structure of the styles and ovary without danger of seriously damaging the
type, so there may be greater differences than appear.

14. BEGONIA GLABRA Aubl. Pl. Guian. 2:916. pl. 349. 1775.

Begonia scandens Sw. Prodr. 86. ,
Begonia elliptica HBK. Nov. Gen. & Sp. 7:180, pl. 641. 1825.
dem ia lucida Otto & Dietr. in Allg. Gartenz. 16:162. 1848.
Beg moritziana Kunth & Bouché in Ind. Sem. Hort. Berol. 16. 1848.
foe pbysalifolia Liebm. in Kjoeb. Vidensk. Meddell. 1852:19. 1853.
ageneria moritziana Kl. in Monatsber. Akad. Berlin 126. 1854, nomen.
| : à xa

Wageneria in Monatsber. Akad. Berlin 126
Wageneria glabra KI. in Monatsber. Akad. Berlin 126. 1854
Wageneria deflexa KI h. Akad. Berlin 1854:113. 1855

W
Begonia locellata A. DC. in Ann. Sci. Nat. IV. 11:137. 1859.

Succulent herb to 9 m. high. Stem scandent, rooting at the nodes, glabrous.
Leaves nearly symmetrical, broadly ovate, 4-15 cm. long, short-acuminate, rounded
or barely cordate at base, sparsely serrate and ciliate to entire, often undulate,
glabrous, petioles 1-8 cm. long, stipules persistent, ovate-oblong, mucronate, entire,
10-24 mm. long, membranaceous, red-brown. Peduncles axillary, 6-20 cm. long.
Cymes many-flowered, diffuse. Bracts persistent, minute. Pedicels slender, 6-16
mm. long. Staminate tepals 4, white, the outer broadly obovate, 3-8 mm. long,
the inner narrowly elliptic. Stamens free, few, anthers oblong. Pistillate tepals 5,
4-6 mm. long. Styles 3, 2-parted, completely covered by stigmatic papillae, pla-
centae simple. Capsule 6-9 mm. long, largest wing oblong to triangular, 10-14
mm. wide, the other two marginiform, very narrow

Southern Mexico and the West Indies to Guiana, Bolivia and Peru.

CAS DEL TORO: Fish Creek, jak of Chiriqui Lagoon, von Wedel 2204; Fish Creek
Midis von Wedel 2278, 2345: F arm Six, Changuinola id Dunlap 500. CHIRIQUÍ:
Chiri

llanos on slopes of Volcán uí Viejo and der riqui Viejo, alt. 1200 m.,
Allen e COCLE: vicinity of El Valle de Anton, alt. 600-1000 m., Allen 22I, 12944
1655. AMA: forests along Rio Boquerón above Peluca Hydrographic Station, alt. 9

m., Hunter +8 Allen 655. VERAGUAS: forested slopes of Cerro Tutć, vicinity of Santa Vi;
alt. 900 m., Allen 4357.
(54)

1958]
FLORA OF PANAMA (Begoniaceae) 55

B semiovata

B glabra
6)

Fig. 17. Begonia
(Lower right: B. fischeri)

(55)

[Vol. 45
56 ANNALS OF THE MISSOURI BOTANICAL GARDEN

15. BEGONIA SEMIOVATA Liebm. in Kjoeb. Vidensk. Meddel. 1852:22. 1853.
Begonia te DC. in Ann. Sci. Nat. IV. =~ 142. 1859.
142. Di

Begonia guyanensis var. r. glaberrima C. DC. in Bot. Gia ^ 540. 1895.

Herbaceous, slender, 2-6 dm. high. Stem simple or slightly branched, essen-
tially glabrous or glabrescent, prostrate to ascending, sometimes rooting at the
nodes. Leaves strongly asymetric, straight (or rarely transversely ovate) acuminate,
unequally cuneate to cordate at base, subpinnately nerved, occasionally shallowly
lobed with ciliate-serrate margins, 4.5—7 cm. long, 1.5—2 cm. wide, essentially
glabrous on both surfaces, petioles 0.5—2 cm. long, glabrous, stipules persistent,
lanceolate, up to 2 cm. long, entire, glabrous, acuminate. Peduncles axillary, 2-3
cm. long, glabrous. Cymes laxly few-flowered. Bracts persistent, lanceolate to
ovate, subentire to ciliate-serrulate, 1-2 mm. long. Staminate pedicels very slender,
to 6 mm. long. Staminate tepals 2, ovate to suborbicular, 2 mm. long and broad.
Stamens free, few, anthers elliptic or ovate, the connective produced. Pistillate
bracteoles persistent, obovate to orbicular, entire to ciliolate-serrulate to almost
lacerate, about 3 mm. long. Pistillate pedicels to 5 mm. long. Pistillate tepals 5,
elliptic to ovate, 2-3 mm. long. Styles 3, 2-parted with the stigmatic tissue form-
ing a continuous linear spiral band, ovary 3-celled, placentae variable even in the
same ovary. Capsule subelliptic to orbicular, glabrous, 6-14 mm. long, with
subequal rounded wings, seeds stalked, oblong and obtuse, truncate at the base.

Mexico and Guiana to Peru.

s DEL TORO: Carleton 54; von Wedel 370; vicinity of Nievecita, alt. 0—
Woodson, Allen & Seibert 1816; vicinity of Chiriqui Lagoon, von Wedel 1331; iei
297.

nola Valley, Dunlab 167. CANAL ZONE: Chas, Fendler 2
*

16. BEGONIA FILIPEs Benth. Bot. Voy. Sulph. 101. 1845.

Begonia hygrophila C. DC. ex Th. Dur. & Pittier in Bull. Soc. Bot. Belg. 351:265. 1896
Begonia bygropbila var. puberula C. DC. ex Th. Dur. & Pittier i in Bull. Soc. Bot. Belg. 351;
266. 1896.

Begonia mameiana C. DC. in Smithson. Misc. Coll. 69: no. 12:4. 1919.
Begonia leptopoda C. DC. in Smithson. Misc. Coll. 69: no. 12:6. 1919.
Begonia cbeboensis C. DC. in Smithson. Misc. Coll. 69: no. a 8. 1919.
Begonia briza dona kisby, i Descr. New Sp. S. Am. Pl. 66. 192

Herbaceous, 2-6 dm. high. Stem simple to Lo tonde glabrous, finely
ridged. Leaves strongly asymmetric, semiovate to obliquely or transversely ovate,
acuminate, mostly truncate at base, palmately nerved with the chief veins branch-
ing divaricately above and with the margins crenate-dentate and ciliate, 4-10 cm.
long, 2-4 cm. wide, rather abundantly appressed-pilose above with long tapering
hairs, essentially glabrous on lower surface, petiole 0.5—4 cm. long, glabrous, stipules
mostly persistent, oblong-ovate, acute to acuminate to the setulose tip, 6-12 mm.
long, 2-4 mm. wide, very thin, entire. Peduncles axillary, up to 5 cm. long,
glabrous. Cymes few-flowered. Bracts persistent, ovate-acute with setulose tip,

(56)

ME Io RIT VEE m FIA MII TAI MIRI DIETA ANTITESI t ATO E Y RAE TAN a Aa o e A e ee La Yo ego uod eia e e lA e ie te ae Capea ©

1958]
FLORA OF PANAMA (Begoniaceae) 57

up to 7 mm. long and 2 mm. wide. Pedicels glabrous, 4-10 mm. long. Staminate
tepals 2, ovate, to 3.5 mm. long and 2 mm. wide. Stamens few, attached to a
short column, connective slightly produced. Bracteoles obovate, serrate-ciliate, to
3.5 mm. long, more or less persistent. Pistillate tepals 4 or occasionally 5, ovate
to orbicular, ca. 2 mm. long, slightly more than 1 mm. wide. Styles 3, 2-parted,
with the papillate stigmatic surface forming spiral bands. Ovary 3-celled, pla-
centae variable. Capsule glabrous, 4-6 mm. long, the largest wing mostly obtuse,
to 1 cm. wide, the other two subequal, narrow, seeds stalked, oblong, obtuse,
truncate at base.

Costa Rica and Panama to Colombia.

CANAL ZONE: Mamei, alt. 10-30 m., Pittier 2251; forest between Frijoles and Monte
Lirio, alt. 30 m., Killip 12143; along ka | rivulet in forest, Gatün, Ostenfeld 84; Las
Cascadas pla ia. near Summit, S/andley 25706, 25723, 20512; Cerro Gordo, near
Culebra, Standley 26003; hills west of the canal, near Gatún, vetas 27222; un sot
of Frijoles, Standley 27568; Gamboa, Standley 28406; Río Para above East Paraíso,
Standley 29882; vicinity of Fort Sherman, Standley 30952; Obispo, Standley 31743; Barro
Colorado Island, Gatun Lake, Standley 40909, 41020, Woodworth & Vestal 391; | ned
ZE of Quebrada Culebra, alt. 70-75 m., Dodge & Allen s. n.; BRM Mada: am and

Saddle 11, near Alahuela, alt. 90-100 m., Dodge, Steyermark 8 Allen Hi on tree,
geowned ferent along Río Chagres beni junction with Rio Pequeni and Rio Indio, alt.
.» Steyermark a Allen 16795. cHiRIQUI: vicinity of San Felix, alt. 0-120 m., Pittier
pi 5. COLÓN: wooded swam amas France Field and sfatui Es red 30104.
DARIÉN: vicinity b Pinogana, ve 20 m., Allen 937. PANA avaré dee
near Chepo, alt. 50-200 m., Pittier 471 8 "rocky banks of Pacara Lx "Killib 3 156; w
forest, Rio Tapia, Standley 26155, 28234; Taboga Island, Standley 27888; moist pile
Rio Tecumen, Standley 20411; San José blind; Jobnston 10
17. BEGONIA FISCHERI Schrank var. TOVARENSIs (Kl.) Irmscher, Bot. Jahrb. 76:

23: I953
Begonia tovarensis Kl. in Abh. Akad. Berlin 1854:31
Er populifolia sensu Liebm. in Kjoeb. Vidoni Med del 1852:16. 1853, non HBK.

Eos moritziana KI. in Abh. Akad. Berlin 1854:31. 1855, non Kunth & Bouché, 1848.
Begonia nsis B ocanensis A. DC. in DC. Prod. 151:303. 1864.
Bigosie cilibrecieola G: DG. in Sao, Misc. Coll. 69: no. 12:5. 1919.

Herbaceous, 2-12 dm. high, sparsely brown-pilose to glabrous. Stem erect, red,
branches short. Leaves more or less asymmetric, broadly ovate or suborbicular,
acute or rounded, cordate at base, 2-9 cm. long, palminerved, crenate-serrate, cili-
ate, glabrous above, more or less brown-pilose beneath especially on the nerves,
petioles 5-45 mm. long, stipules deciduous, ovate-oblong, to 10 mm. long, ciliate-
serrulate. Peduncles axillary, 15-55 mm. long. Cymes bisexual, few-flowered.
Bracts persistent, ovate, 2-4 mm. long, fimbriate. Pedicels 5-20 mm. long. Sta-
minate tepals 4, the outer orbicular, 8 mm. long, the inner smaller, narrowly
obovate. Stamens free, numerous, filaments short, anthers oblong, the connective
produced, obtuse. Pistillate bracteoles like the bracts. Pistillate tepals 5, obovate,
3-6 mm. long. Styles 3, bifid, the stigmatic tissue linear, spiral, continuous, pla-
centae bilamellate, ovuliferous throughout. Capsule 10-15 mm. long, its wings
decurrent, very unequal, the largest typically ascending and tapering, often hooked,
to 23 mm. wide, seeds fusiform.

(57)

[ Vol. 45
58 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Cuba, Mexico, Central America, Colombia, Venezuela, Peru, Bolivia.

CANAL ZONE: Ahorca Lagarto to Culebra, Cowell 388; on marshy or floating island,
Barro Colorado Island, Kenoyer 460, Bailey 644, Woodworth t$ Vestal 696. cocLE: shaded
margins of marsh near Río Antón, floor of El Valle de Antón, alt. 600 m., Allen 2777.
PANAMÁ: on grassy jams in lake, vicinity of Arenoso, lower Río Trinidad, alt. 26-50 m.,
Seibert 617.

18. BEGONIA OAXACANA A. DC. in DC. Prod. 151:312. 1864.

Begonia oaxacana B pilosula A. DC. in DC. Prod. 151:312. 1864.

Begonia luxii C. DC. in Bot. Gaz. 20:541. 1895.

Begonia serrulatoala C. DC. in Bull. Herb. Boiss. II. 8:321. 1908.

Begonia pubipedicella C. DC. in Smithson. Misc. Coll. 69: no. 12:7. 1919.

Herbaceous to suffruticose, terrestrial or epiphytic, suberect or climbing, very
variable in all its parts, 3-15 . high. Stems 2-12 mm. thick, glabrous to pilose,
green or red. Leaves transverse or oblique or rarely some almost straight, broadly
or narrowly ovate, acuminate, palmately 5—7-nerved, cordate at base, slightly or
usually not at all lobed, 6—18 cm. long, 3-11 cm. wide, serrulate, sparsely hirsute
on both sides or nearly glabrous above, thin, petioles slender, 4-15 cm. long, sparsely
hirsute to glabrous, stipules deciduous, oblong, 12 mm. long, setose-mucronate,
entire. Peduncles axillary, 2-5 cm. long. Cymes few-flowered, bisexual. Bracts
tardily deciduous, broadly ovate or suborbicular, obtuse, 6-12 mm. long, entire.
Pedicels 10-15 mm. long. Staminate tepals 4, the outer suborbicular or broadly
ovate, 10-15 mm. long, pink, glabrous or at times somewhat pubescent, the inner
slightly shorter, obovate, white. Stamens free, numerous, the filaments longer than
the obovoid anthers. Pistillate bracteoles wanting or quickly deciduous. Pistillate
tepals 3, the outer two suborbicular, 8-14 mm. long, the inner one much smaller,
narrowly obovate or oblong. Ovary 2-3-celled, placentae bifid, ovuliferous on all
sides, styles 3, short-connate, bilobed, the stigmatic surface linear, spiral, contin-
uous. Capsule erect, broadly elliptic, 10-15 mm. long, hirsute to glabrous, wings
3 or sometimes up to 6, subequal, narrow, angled at middle or somewhat above
giving the fruit a rhombic or obovate outline, entire to fimbriate-serrulate, seeds
ellipsoid, broadly obtuse, short-stalked.

Southern Mexico, Guatemala, Panama.

CHIRIQUf: humid forest of Cuesta de Las Palmas, southern slope of Cerro de la Hor-
queta, alt. 1700-2100 m., Pittier 3248; rain forest, Bajo Chorro, Boquete District, alt. 1800
m., Davidson 55; vicinity of Casita Alta, Volcán de iriqui, alt. 1500—2000 m., Woodson,

Allen & Seibert 959; Bajo Mono, Robalo Trail, western slopes of Cerro Horqueta, alt. 1500-
2100 m., Allen 4809.

19. BEGONIA UDISILVESTRIS C. DC. in Smithson. Misc. Coll. 69: no. 12:9. 1919.

Herbaceous, 1 m. high, branching, completely glabrous except for a ring of
trichomes at the apex of the petiole. Leaves transverse or oblique, ovate, cordate
at base, long-acuminate, palminerved, to 11.5 cm. long, serrulate, petioles slender,
to 11 cm. long, verruculose, stipules deciduous. Cymes few-flowered, re ,
about equaling the leaves or shorter. Bracts deciduous, entire. Staminate tepals 4,
entire, the outer suborbicular, 5 mm. long, the inner narrowly obovate, shorter.

(58)

1958]
FLORA OF PANAMA (Begoniaceae)

B.oaxacana B.udisilvestris

B.involucrata A sę,

[Vol. 45
60 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Stamens free, anthers oblong, shorter than the filaments. Pistillate tepals 3, the
outer two suborbicular, 5 mm. long, the inner one much smaller and narrower.
Ovary 3-celled, placentae bilamellate, ovuliferous throughout, styles deciduous,
connate at base, bifid. Capsule erect, ovoid, attenuate into a slender beak, wings
equal, very narrow.
Costa Rica and Panama.
CAS DEL TORO: Robalo Trail, northern slopes of Cerro de la Horqueta, alt. 1800-

2100 1 m., Allen 4938. cHIRIQUI: harhid forest of Cuesta de Las Palmas, southern slope of
Cerro de la Horqueta, alt. 1700-2100 m., Pittier 3249; Volcán de Chiriquí, Boquete Dis-
trict, alt. 2700 m., Davidson 1025; trail fein Cerro Punta to headwaters of Rio Caldera,
alt. 2250-2500 m., Allen 1440; Casita Alta to Cerro Copete, alt. 2300-3300 m., Woodson
& Schery aj Portrero Muleto to summit, Volcan de Chiriqui, alt. 3500—4000 m., Wood-
son & Scher

20. BEGONIA INVOLUCRATA Liebm. in Kjoeb. Vidensk. Meddell. 1852:15. 1853.
Begonia PRAC A DC. in Ann. Sci. Mae DI 11:133. 1859.
Begonia laciniosa A. DC. in DC. Prod. 151:340.

Fruticose, 1-2 m. high. Stem RT or becoming glabrous. Leaves
oblique, cordate at base, 2-5-lobed at apex with acuminate points, 15-28 cm. wide,
to 20 cm. high, denticulate, evenly pilosulous above, more densely on the nerves
beneath, petioles 12-18 cm. long, fulvous-tomentose or becoming glabrous, stipules
deciduous, ovate, setose-apiculate, 14-25 mm. long, entire, brown, membranaceous,
glabrous or sparsely pilose. Peduncles axillary, exceeding the leaves, fulvous-
tomentose becoming glabrous. Cymes many-flowered, dense, their primary branches
short and their ultimate branches so nearly aborted that the inflorescence appears

almost umbellate. Bracts deciduous, entire, the outer ovate, ample and completely

enclosing the young inflorescence. Pedicels 5-25 mm. long. Staminate tepals 2,
suborbicular, 6-11 mm. long, entire, sparsely pilose to glabrous, white. Stamens
numerous, anthers oblong, about equaling the filaments, connective produced,
broadly obtuse. Pistillate bracteoles lacking. Pistillate tepals 2, like the staminate.
Ovary 3-celled, placentae bilamellate, styles connate at base, bifid. Capsule
ellipsoid, very unequally 3-winged, the largest wing elliptic or falcate, obtuse,
8-10 mm. wide. :

Guatemala, Nicaragua, Costa Rica, Panama.

CHIRIQUÍ: moist shaded places, valley of the upper Río Chiriquí Viejo, alt. b mens

W bite 34; rain forest, Bajo Chorro, Boquete District, alt. 1800 m., Davidson ;
os; of “New Swi iai, central valley of Rio Chiriqui Viejo, alt 1200-2000. m.,
1349.

21. BEGONIA MULTINERVIA Liebm. in Kjoeb. Vidensk. Meddell. 1852:18. 1853.
Begonia cuspidata C. DC. ex Th. Dur. & Pittier in Bull. Soc. Bot. Belg. 35!:260. 1896.
Suffruticose, 2-3 m. high. Stem flexuous with elongate internodes, brown-
pilose. Leaves transversely elliptic, abruptly acute, cordate at base, palminerved,
12-22 cm. long, entire, glabrous, petioles divergent, 6-16 cm. long, brown-pilose,
stipules deciduous, lance-oblong or narrowly triangular, setose-acute, 2-3 cm. long,

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1958]
FLORA OF PANAMA (Begoniaceae) 61

entire, membranaceous. Peduncles axillary, 15-38 cm. long, exceeding the leaves,
sulcate, soon glabrous. Cymes bisexual, regularly dichotomous, 12—25 cm. broad,
glabrous. Bracts deciduous, suborbicular, entire, the lowest 17 mm. long. Pedicels
7-15 mm. long. Flowers white. Staminate tepals 2, broadly ovate or suborbicular,
8 mm. long, entire. Stamens numerous, anthers obovoid, shorter than the fila-
ments, connective scarcely if at all produced. Pistillate bracteoles lacking. Pistil-
late tepals 2, like the staminate. Ovary 3-celled, placentae bilamellate, ovuliferous
throughout, styles much connate at base, bifid, the stigmatic tissue linear, spiral,
continuous. Capsule subglobose, 7 mm. long, wings very unequal, the largest ovate
or elliptic, to 14 mm. wide, the others narrowly lunate, seeds stalked, ellipsoid.
Costa Rica, Panama.

CHIRIQUf: between Hato del Jobo and Cerro Vaca, eastern Chiriqui, alt. 700-1000 m.,
Pittier 5420.

22. BEGONIA URTICAE L. f. Suppl. 420. 1781.

Begonia urticifolia J. E. Smith, PI. Icon. Ined. 4 È 45. 1790, pro errore.
Beg onia 2 umnaris Benth. PI. Hart ta 131.
rachyptera Benth. PI. Hartweg. 184. "mk
2 trachyptera KI. in Pomo ad Akad. Berlin 127. 1854
Begonia coccinea Ruiz ex Kl. in Monatsber. Akad. owi in 127. 1854, e. p., nomen in synon.
Sassea urticae Kl. in Monatsber. Akad. Berlin 128.
Sassea columnaris Kl. in Monatsber. Akad. Berlin vd
Begonia glabra Ruiz ex Kl. in Monatsber. Akad. Berlin 128. 1854, nomen in synon.
Casparya coccinea Kl. in Abh. Akad. Berlin 1854:128.
Stibadotbeca trachyptera Kl. in Abh. Akad. Berlin 1854: 131. 1855.
Sassea glabra Kl. in Abh. Akad. Berlin 1854:134. 1855.
Begonia cucullata Ruiz ex Kl. in Abh. Akad. on 1854:134. 1855, nomen in synon.
Casparya trachyptera A. DC. in DC. Prod. 151:274. 1864.
Casparya columnaris A. DC. in DC. Prod. 151: we 1864, non Kl. 1854.
Casparya columnaris 8 bia A. DC. i in DC. Prod. 151: :274. 1864.
icae A. DC. in i 4.

Sassea hoffmanniana Kl. ex A. DC. in DC. Prod. 15:275. 1864, nomen in synon.
Begonia monticola C. DC. in Bull. Herb. Boiss. II. 8:325. E
Begonia torresii Standl. in Journ. Wash. Acad Sci. 17:313. 2e

Begonia chiriquensis Standl. in Woodson & Schery in Ann. Mo. Bot. Gard. 27:521. 1940.
pae sowy var. glabra Smith & Schubert in aż. Fl. Peru, in Field Mus. Pub.
Bot. 134:18

is $ to suffruticose. Stem branching, decumbent, rooting at the nodes,
15-40 cm. high or rarely to 2 m., the younger parts usually ferruginous-puberulent.
Leaves straight or nearly so, strongly asymmetric, ovate or elliptic, acute or acu-
minate, base acute on the adaxial side, obtuse and decurrent on the other, 3-8 cm.
long, penninerved, doubly serrate, ciliate, sparsely hirtellous to glabrous above,
densely pubescent on the nerves to wholly glabrous beneath, petiole 2-15 mm. long,
stipules deciduous, ovate, obtuse, setose, 2-6 (rarely to 12) mm. long. Peduncles
axillary, erect, 12-45 mm. long, 1- to few-flowered. Bracts deciduous, elliptic,
setaceous-dentate at apex. Pedicels 6-18 mm. long. Staminate tepals 4, subequal,
3-8 mm. long, the outer elliptic, entire, red, often pilose, the inner obovate, white.
Stamens on a slender column 2 mm. high, filaments short, anthers linear, the con-

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[Vol. 45
62 ANNALS OF THE MISSOURI BOTANICAL GARDEN

nective slightly produced. Pistillate bracteoles deciduous, elliptic. Pistillate tepals
5, subequal, elliptic, 3-5 mm. long, entire, red or white, the outer often pilose.
Styles 3 with many short branches wholly covered by stigmatic papillae, placentae
bilamellate, ovuliferous on all sides. Capsule very variable, broadly turbinate,
obtuse at base, with 3 ascending horns on the angles, the tips of the horns often
deciduous, the apical column well developed, angled or terete, slender, conical or
cylindrical or subclavate, from longer than the horns to rarely shorter.

Costa Rica to Peru.

BOCAS DEL TORO: along streams, Robalo Trail, northern slopes of Cerro de la Horqueta,
ME i m., Allen 4980. cHIRIQUI: rain forest, Bajo Chorro, Boquete District, alt.

Davidson 107; epiphytic, ini from Cerro Punta to headwaters of Rio Caldera,
Va NI m., Allen 1435; vicinity of Cerro Punta, alt. 2000 m., Allen 3484.

23. BEGONIA HEYDEI C. DC. in Bot. Gaz. 20:540. 1895.
Begonia pittieri C. DC. in Bull. Herb. Boiss. II. 8:316. 1908.
Begonia triloba C. DC. in Bull. Herb. Boiss. II. 8:322. 1908.

Erect, to 6 dm. high, branching with elongate internodes, glabrous. Leaves
oblique, cordate at base, narrowly ovate, acuminate, penninerved, 6—10 cm. long,
2—3.5 cm. wide, remotely denticulate, petioles 10-34 mm. long, stipules deciduous,
lance-ovate, subulate, 5 mm. long. Peduncles axillary, 15-28 mm. long. Cymes
few-flowered, exceeded by the leaves. Bracts deciduous. Pedicels 8—10 mm. long.
Staminate tepals 4, entire, obtuse, the outer ovate or elliptic, 7 mm. long, setose
toward base, the inner slightly shorter. Stamens many on a short column, anthers
obovate, nearly equaling the filaments. Pistillate bracteoles elliptic, entire, about
equaling the ovary. Pistillate tepals 3, elliptic or obovate, entire, setose toward
base. Ovary 3-celled, placentae simple, styles deciduous, connate at base, bifid.
Capsule rhomboidal, attenuate at base, bearing three equal slender acuminate horns
9—11 mm. long.

Guatemala, Costa Rica, Panama.

OCLÉ: — vicinity of El Valle de Antón, alt. 600 m., Allen 2078; epiphytic,
La na Allen 2722

24. BEGONIA TONDUZII C. DC. in Bull. Soc. Bot. Belg. 351:264. 1896.
Begonia allenii Standl. in Woodson & Seibert in Ann. Mo. Bot. Gard. 25:830. 1938.
Caulescent, 28-48 cm. high. Stem erect, simple or rarely few-branched, woody
at least toward base, rufous-pilose becoming glabrous with age, internodes mostly
short but distinct. Leaves nearly straight, strongly asymmetric, oblanceolate,
penninerved, acuminate, cuneate, falcate, 6-12 cm. long, 2-5 cm. wide, coarsely
sinuate-dentate near apex, glabrous or pilose on the midnerve beneath, petioles 5-2
mm. long, pilose to glabrous, stipules more or less persistent, broadly ovate, setose-
acuminate, 10-25 mm. long, entire, thin. Peduncles few, axillary, 3-7 cm. long.
Cymes few-flowered, regular, rather dense, pilose at first. Bracts persistent, small,
suborbicular, serrate. Pedicels 5-16 mm. long. Flowers pink or red. Staminate
tepals 2, broadly ovate, 5-8 mm. long, more or less pubescent. Stamens free, ©

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1958]
FLORA OF PANAMA (Begoniaceae) 63

anthers oblong, about as long as the filaments, connective produced, obtuse. Pistil-
late bracteoles like the bracts. Pistillate tepals 5, equal. Ovary 3-celled, placentae
bilamellate, ovuliferous throughout, styles connate at base, stigmatic tissue linear,
spiral, continuous. Capsule erect, subglobose, wings subequal, very variable in
width.

Costa Rica, Panama.

cocLE: El Valle de Antón, along Río Indio trail, alt. 500—700 m., Hunter & Allen
327; north rim, vicinity of El Valle, Allen 234, 1663; banks along trail to La Mesa, hills
north of El Valle de Antón, alt. 800-1000 m., Allen 2286; Cerro Pajita, hills north of El
Valle de Antón, alt. 1000-1200 m., Allen 4174.

25. BEGONIA OPULIFLORA Putz. in Fl. des Serres I. 10: pl. 995. 1854-55.

Suffruticose, branching, over 1 m. high, glabrous. Leaves straight, strongly
asymmetric, ovate or elliptic, acute, 8-9 cm. long, rounded at base, pinnate,
subulate-dentate, petioles much shorter than the blades, stipules oblong-cordate,

Bopuliflora AT

Fig. 19. Begonia opuliflora

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[Vol. 45
64 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long-aristate, membranaceous. Peduncles axillary, about 15 cm. long. Inflores-

cences umbelliform, globose, 50—60-flowered. Pedicels elongate. Flowers white.

Staminate tepals 4, subequal, broadly elliptic, obtuse, entire. Stamens about 30,

slightly connate at base, anthers oblong. Pistillate tepals 5, oblong or lance-oblong,

subacuminate. Styles 3, connate, stigmatic tissue lunate. Fruit 3-winged, the wings

very unequal, the largest much wider than high, twice the width of the others.
Endemic

OCLE: — from cultivation from the former Province of Soto which centered
on Pese

26. BEGONIA GUADUENSIS HBK. Nov. Gen. & Sp. 7:178. 1825.
Begonia ottonis W alpers, Repert. e Syst. 2:212. 1843.

Begonia walbersii Heynh. Nom. 2:6

Donaldia ottonis Kl. in eue Akad Berlin 127. 1854, nomen

Begonia laurina Hort. ex A. D C. Prod. 151:292. 1864, nomen in synon.
Begonia serratifolia C. DC. in Se Misc. Coll. 69: no. 12:7. 1919.

Suffruticose, scandent to 2 m. high, glabrous. Stem erect, branched. Leaves
straight, asymmetric, pinnate-nerved, lanceolate to elliptic-lanceolate, acute or
acuminate, unequal at base with the adaxial side cuneate and the other rounded and
decurrent, 5-9 cm. long, 16-40 mm. wide, doubly crenate-serrate, ciliate, mem-
branaceous, petiole 2-8 mm. long, stipules deciduous or subpersistent, oblong-ovate,
acuminate-setiferous, 8-10 mm. long, scarious. Peduncles terminal and axillary,
4-5 cm. long. Cymes dichotomous, few- to many-flowered, 6-15 cm. in diameter.
Bracts deciduous or subpersistent, ovate, obtuse, acute or mucronate, 4-7 mm. long,
scarious. Pedicels 4-19 mm. long. Flowers pinkish white. Staminate tepals 4,
the outer elliptic-ovate, entire or dentate at apex, 8-15 mm. long, the inner
obovate, distinctly shorter. Stamens free, very numerous, anthers oblong, mostly
shorter than the filaments, connective produced, apiculate or obtuse. Pistillate
bracteoles deciduous, ovate to narrowly obovate, 6 mm. long, entire. Pistillate
tepals 5, subequal with the two outer slightly smaller, elliptic, acute, 6-10 mm.
long. Styles 3, bifid nearly to base, each main branch with 3-4 short spiral
branches, placentae bifid, ovuliferous throughout. Capsule ellipsoid, the largest
wing triangular, horizontal or slightly ascending, 15 mm. wide, the other two
marginiform

Panama, Colombi. Venezuela.

E: side cuts, railroad relocation between Gorgonas and Gatún, alt. 10-50

Pittier sesta drowned forest of Ser Culebra, alt. 70-75 m., Dodge & Allen; a
rm of Quebrada Salamanca, alt. 70 m., Dodge, Steyermark 5 Allen; near Madden D am at
Alahuela, alt. 70 m., Dodge, sali & Allen 16822; forest along telephone tra ail | be-

tween the Río Indio Hydrographic Station and the Natural Bridge of the Río Puente,
Dodge 6 Allen 17481; drowned forest of Quebrada Ancha, alt. 70 m., Dales 8 Steyer-

mat Mews vicinity of San Félix, eastern Chiriquí, alt. 0-120 m., Pittier 51 126.
VERA : hills west of Sona, alt. 500 m., Allen 1034.

27. BEGONIA ESTRELLENSIS C. DC. in Bot. Gaz. 20:540. 1895.

Woody vine. Stem elongate, glabrous. Leaves pinnately veined, ovate-elliptic,

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FLORA OF PANAMA (Begoniaceae) 65

B. Su

aduensis B.seemaniana

B.convallariodora
Es

AGA m)
R ARTO

Fig. 20. Begonia

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[Vol. 45
66 ANNALS OF THE MISSOURI BOTANICAL GARDEN

rather abruptly narrowed to a slender acumination (to 1 cm. long), acute to
cuneate at base with the broader half produced 1 mm. beyond the narrower, re-
motely denticulate, glabrous, foveolate, 5—6.5 (—9.5) cm. long, 1.7-2.3 (-3.3) cm.
wide, petioles to 12 mm. long, glabrous, stipules early deciduous, only scars seen.
Inflorescences pedunculate and cymose, terminal and axillary. Staminate pedicels
to 18 mm. long. Tepals regularly 4, the outer ovate, obtuse, to 6 mm. long and
5.5 mm. broad, the inner elliptic, to 5 mm. long and 2 mm. wide. Stamens numer-
ous, free, the connective produced. Pistillate pedicels slender, glabrous, to 20 mm.
long. Tepals 2, ovate, broadly acute, to 10 (—16) mm. long and 8 mm. wide.
Ovary glabrous, puncticulate. Styles 3, bifid, broadly expanded above the base.
Capsule unequally tri-alate, the largest wing to 10 mm. wide, ascending, acute.
Costa Rica, Panama.

cocLÉ: vicinity of El Valle, north rim, alt. 800-1000 m., Allen 220.

28. BEGONIA SEEMANNIANA A. DC. in Ann. Sci. Nat. IV. 11:133. 1859.
Begonia cbiriquina C. DC. in Smithson. Misc. Coll. 69: no. 12:8. 1919.

Shrub to 3.5 m. high. Stem slender, glabrous, finely ribbed. Leaves asym-
metric, elliptic-obovate, penninerved, serrulate-denticulate with setiferous teeth,
acuminate, unequally obtuse at base with the broader side extended on the petiole
beyond the narrower, glabrous to minutely puberulent throughout, foveolate, 9.5-
14 cm. long, 4.5-6 cm. wide, petioles glabrous, rugose, 5-15 mm. long, stipules
glabrous, ovate to elliptic, obtuse to acuminate and mucronate, truncate to nearly
clasping at base, 15—25 mm. long, 4—6 mm. wide. Peduncles to 7 cm. long. In-
florescences cymose, terminal and axillary. Bracts inconspicuous, the staminate
thin, elliptic, ca. 1 mm. long, the pistillate very early deciduous. Pedicels glabrous,
slender, to 2 cm. long. Staminate tepals 2, ovate-obtuse to semi-orbicular, usually
cordate at base, 5-6 mm. long, 3-8 mm. wide. Stamens numerous, free, with con-
nectives produced and exceeding the shorter outer filaments. Pistillate tepals 2
(rarely a smaller third one present), obovate and obtuse to elliptic, 5-8.5 mm. long,
3—5.5 mm. wide. Ovary glabrous, sometimes with reddish dots on surface. Styles
3, bifid or irregular, essentially free. Capsule tri-alate, the largest wing with
truncate obtuse apex, to 15 mm. wide, one wing marginiform, the third only
slightly broader. Seeds obovoid or ellipsoid, less than 0.5 mm. long.

Costa A Panama.

CHIRIQUÍ: humid forest of Cuesta de Las Palmas, southern slope of Cerro de la Hor-

queta, alt. A m., Pittier; vicinity of El Boquete, alt. 1000-1300 m., Maxon 4948;
Pittier 3152.

29. BEGONIA CARPINIFOLIA Liebm. in Kjoeb. Vidensk. Meddel. 1852:20. 1853.

Herbaceous, suffrutescent or subscandent. Stem glabrous, green to reddish,
finely ridged. Leaves obliquely ovate-acuminate to obliquely elliptic, obliquely
obtuse at base with the broader half produced 1 mm. or more beyond the narrower,

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1958]
FLORA OF PANAMA (Begoniaceae) 67

glabrous, foveolate and prominently veined beneath, prominently serrate-denticulate
with setiferous teeth, 5-7.5 (—11) cm. long, 2.5-3.7 (-4.7) cm. wide, petioles
3.5-10 mm. long, glabrous, stipules persistent, pinkish to greenish when fresh,
linear to lanceolate or narrowly ovate with oblique base and acute apex, 7-13 mm.
long, ca. 5 mm. wide. Inflorescences pedunculate, cymose, axillary and terminal.
Bracts of the peduncles similar to the stipules, secondary bracts and bracteoles
obovate to orbicular, obtuse to truncate. Pedicels glabrous, slender, to 1 cm. long
in fruit. Staminate tepals 2, orbicular, cordate at base, 5-8 mm. long, 6-7 mm.
wide. Stamens numerous, free, filaments of the outer ones shorter, connective pro-
duced. Pistillate tepals 2, chiefly orbicular, 3-6 mm. long, 4.5-5 mm. wide. Styles
3, short-connate. Capsule ca. 6 mm. high, its largest wing ascending, acute to
obtuse, ca. 5 mm. wide, the other two marginiform and subequal. Seeds minute,
slenderly obovoid.
Costa Rica, Panama.

HIRIQUf: vicinity of “New Switzerland”, central valley of Rio Chiriqui Viejo, alt.
1800-2000 m., Allen 1408.

30. BEGONIA CONVALLARIODORA C. DC. in Bot. Gaz. 20:538. 1895.

Herbaceous, 1-15 m., scandent to erect. Stem slender, little branched, ribbed
and glabrous, the internodes of the main axis 5.5-14 cm. long. Leaves pinnately
veined, asymmetric, somewhat denticulate, obliquely elliptic, abruptly acuminate,
the broader half usually produced into a conspicuous lobe at base, 8.5—15 cm. long,
4-7.5 cm. wide, petioles 12-40 mm. long, stipules membranaceous, obliquely
elliptic, entire, acute and apiculate, truncate at base, 25 mm. long, 7 mm. wide.
Peduncles 3-8 cm. long. Cymes regular, multi-flowered. Bracts early deciduous.
Pedicels 6-16 mm. long. Tepals entire, white or pinkish, the staminate 2 (rarely
3 or 4), orbicular, cordate at base, 4-8 mm. long, the inner, when present, elliptic,

1-3.5 mm. long. Stamens numerous, anthers oblong-elliptic, usually equal to or
longer than the filaments, connective produced, obtuse. Pistillate bracteoles soon
deciduous, entire, elliptic, obtuse. Pistillate tepals 2 (rarely 3), orbicular to
rounded-ovate, 6-9 mm. long, the third, when present, smaller. Styles 3, bifid.
Placentae bilamellate, ovuliferous throughout. Capsule ellipsoid, gas. the
largest wing obtuse, 7-12 mm.

Southern Mexico to Panama.

quí: valley of the upper Río Chiriquí Viejo, vicinity of Monte > alt. 1300-
1360 aac m., “Seibert 207; Bajo Mona, mouth of Quebrada Chiquero, along Rio Caldera, alt.
1500-200 0 m., Woodson, Allen & Seibert 1015; vicinity o of Bajo Mona d Quebrada
Chiquero, alt. 1500 m., Woodson & Sche ery 533; vicinity of Bajo Chorro, alt. 1900 m.,
Woodson & Schery 647. COCLE: crize vicinity of El Valle, alt. 600-1000 m., Allen
1188, 1207.

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[Vol. 45
68 ANNALS OF THE MISSOURI BOTANICAL GARDEN

CACTACEAE

More or less massive and succulent, usually leafless terrestrial or epiphytic
plants. Stems usually (except in Pereskia) phylloid and jointed, the joints elongate
to suborbicular, terete to angled or flattened, the nodes with axillary areas (areoles)
usually armed with spines, slender barbs (glochids) or hairs in various numbers and
combinations, occasionally naked. Leaves alternate and laminate in Pereskia, but
in the other genera minute and có or wholly lacking. Flowers pedunculate
and in 1- to several-flowered panicles in Pereskia, in the other genera solitary and
sessile, very large to very small, frequently nocturnal, epigynous (perigynous in
Pereskia); perianth salverform or infundibuliform to rotate, with or without a
conspicuous tube, monochlamydeous, actinomorphic when erect and more or less
zygomorphic by position when horizontal or pendulous, the segments petaloid,
usually white to red or yellow, relatively few to exceedingly numerous, the inner
broader and more petaloid than the outer and lower, the tube naked or with bracts
or areoles or both; stamens usually very numerous, with elongate filaments, inserted
within the perianth tube or upon the ovary in rotate flowers; ovary inferior (sub-
inferior in Pereskia), 1-celled, with few to several parietal placentas usually bearing
numerous ovules, the style more or less elongate and filiform, with an elaborate
radiate terminal stigma. Fruit a berry usually with numerous seeds immersed in
succulent pulp.

The Cactaceae are a typically Western Hemisphere family, with only the genus
Rbipsalis, the Mistletoe Cactus, doubtfully indigenous to the Old World. Species
of Opuntia, however, especially O. ficus-indica, have been introduced since very
early times and become naturalized in Africa and elsewhere. The genera vary in
number according to the interpretation of individual authors, from only 20 (K.
Schumann, in Engl. Nat. Pflanzenfam. III: 6. 1894) to over 100 (cf. Britton
& Rose, in Carnegie Inst. Publ. No. 248. 1919-1923).

Much criticism has been made of the numerous segregate genera proposed by
Britton and Rose and much undoubtedly remains to be learned of their natural
relationships. In Panama, however, their genera usually are quite readily distin-
guishable into apparently natural groups; consequently they are adopted here
without major modification. The controversy of “lumpers vs. splitters” in dealing
with cacti dates at least as early as 1813, when Sims (in Curtis's Botanical Maga-
zine 38: sub bl. 1557) inveighed against Haworth's generic segregates and advo-
cated the return of both Cereus and Opuntia to the Linnaean genus Cactus, now
long abandoned.

In humid Panama the cacti bear little resemblance as a rule to those so familiar
in the landscapes of the arid regions of the United States and Mexico. Here they
are mostly clambering semilianes on rocks, or epiphytes, with only two or three
species of Opuntia and Nopalea, possibly introductions, to represent the Prickly
Pears. Pereskia scarcely resembles a cactus at all, with its shrubby or tree-like habit,
terete stems, and conspicuous leaves. Cacti of the Rhipsalis and Cereus alliances

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1958]
FLORA OF PANAMA (Cactaceae) 69

often are encountered in Panama, but all to frequently in a sterile condition. This,
together with the difficulty of preparing such succulent plants for the herbarium,
accounts largely for the very few specimens found in museums. Actually, the
species are not at all as rare as the few citations would indicate. The total number
in Panama must be considerably greater than that of the present account.
a. Stems not phylloid; leaves ge Lento: a laminate, deciduous; flowers

demum often ‘clus tered, perigyn 1. PERESKIA
aa. Stem ylloid; i docking or inconspicuous and fugacious; flowers

dis peo epigyn

b. Stem joints abbiate to broadly linguiform in our species, entire,

= reoles amphigenous, with glochids; perianth rotate to campan-

È nd ls erect, shorter the stamens; stamen filame not
sensitive, erect at joins po enlarged near the Ls into a i disci.
form stylopodium 2. NOPALEA
cc. Petals to longer than the stamens; stamen filaments sensitive,
spreadin esis and i upon stimulation; style o nly
slightly „kle toward the bas 3. OPUNTIA

bb. Stem joints elongate, terete to ni or flattened and undulate or
lobed, the areoles marginal upon the flowering joints, without glochids.
c. Flowers very large, ra n mediocre, with a definite perianth tube;
stamens adnate o the perianth tube.
d. Flow ing stem pu ribbed or angled, the areoles usually more
or less spiny and bristly.
e. Stem joints sta. 5-angled, with numerous large spines;
plants a. the stems erect at first, but finally spen

and rootin the tips; periant th tube much longer than th
=. nts, a gree bracts inconspicuous and fugacious Ci
rominent spiny areoles 4. ACANTHOCEREUS

Bei joints usually 3-angled, with inconspicuous and infrequent
spines, or unarmed; plants clambering u upon rocks or epiphytic,
rooting la caso from the stem joints; perianth tube shorter
than the segments.
f. Flowers de large; perianth tube with conspicuous and
persistent bracts, not | or hairy 5. HYLOCEREUS
ff. ine ar iocre; nth tube with inconspicuous and
e x ăi noni ei conspicuous persistent bristly
or agrari 6. WEBEROCEREUS
dd. Dag stem joints flat ma vaga lobed or undulate, the

es n to
e Flowers - with an Mal perianth tube, the segm
us; stamens very numerous, tbe filaments pe
lobes n

as teen than the anthers; stigma lol merous 7. EPIPHYLLUM
ee. Flowers mediocre or relatively small, with a mires short peri-
anth tube, the segments about 10-20; stamens a 30,
the Panni not greatly longer a the anthers; stigma lobes
3-5 8. WITTIA
ce: secie” x small, AE a rațe perianth tube or the segments
ite free; stamens adnate (in our species) to the margin of the
h patio e a flowering stem aa terete or essentially so, naked or
with bristly or hairy areoles 8. RHIPSALIS

1. PERESKIA Mill.

PERESKIA Mill. Gard. Dict. ed. 4. 1754; Britton & Rose, in Carnegie Inst. Wash.
Publ. no. 248. 1:8. 1919.
Cortona Neck. Elem. Bot. 2:84. 1790, as Carpophillus.
eireskia Mill. acc. to Steud. Nom. ed. 2. 2:282. 1841, var. typogr.

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70 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Leafy trees or shrubs, sometimes clambering; stems terete, branching, succulent
above, woody below, armed with stout spines in minutely pubescent axillary areoles.
Leaves alternate, sora: broadly laminate, deciduous. Flowers pedunculate, panicu-
late or solitary, terminal and axillary; perianth rotate, with many free or
slightly united id the outer somewhat shorter and less petaloid than the
inner; stamens numerous, inserted in a shallow tube adnate to the base of the
perianth. Ovary subinferior. Fruit a fleshy relatively few-seeded berry bearing
the marcescent remains of the perianth.

A genus of about a dozen species, ranging from Mexico and Florida to Argen-
tina and in the Antilles. Very widely cultivated in the tropics and subtropics of
the world and in greenhouses, and used as an under stock of grafted Zygocacti and
Schlumbergeras. The plants far more resemble the northern Osage Orange in habit
than a true cactus.

a. Mee mies. lianas; spines short and reflexed; berries globose, relatively
1. P. ACULEATA

aa. seem or small trees; spines elongate and straight; berries turbinate,
relatively large. 2. P. BLEO

1. PERESKIA ACULEATA Mill. Gard. Dict. ed. 8. 1768.

Cactus pereskia L. Sp. Pl. 469. 1753.

Cactus lucidus Salisb. Prodr. 349. 1796.

Pereskia longispina Haw. Syn. Pl. Succ. 178. 18

Pereskia aculeata B longisbina (Haw.) DC. Prodr. 3:475. 1828.

Pereskia fragrans Lem. Hort. Univ. 2:40.

Pereskia undulata Lem. Illustr. Hort. 5: Misc. 1t 1858

unione (L.) Karst. Deutsch. Fl. 888. 1882; Britton & Rose, in Carnegie Inst.

Peireskia foetens Spegg. in Weingart, Monatsschr. Kakteenk. 14: ra 1904.
Pereskia godseffiana Hort. Sand. in Gard. Chron. III. 43:257.

Clambering shrubs or lianas; branches terete, ex LE about 3—10 m.
long, with pairs of stout reflexed spines 2-5 mm. long. Leaves shortly petiolate,
the blade variable, usually elliptic to obovate, obtuse to acuminate, the base obtuse
to cuneate, 3—8 cm. long, 1.5—4.0 cm. broad, the petiole 3-5 mm. long. Flowers
usually few or several in terminal or axillary panicles about twice as long as the
subtending leaves; pedicels about 0.5-1.5 cm. long, accrescent in fruit; perianth
white, yellowish, or pink, rotate, about 3-5 cm. in diameter, the segments rather
numerous, the inner obovate-oblong, the outer oblong-linear; stamens numerous,
less than half as long as the perianth segments; style rather stout, slightly longer
than the stamens. Berry globose, 1.5-2.0 cm. in diameter, with inconspicuous
thornless areoles, yellow or orange when ri

Antilles; Panama and Colombia; also reported from southern Florida and Mex-
ico, perhaps as an escape. Frequently cultivated. In the British West Indies the
fruits, known as Barbados Gooseberries, are eaten and made into preserves.

PANAMA: Cerro Campana, alt. 800 m., Allen 3972.

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19581

FLORA OF PANAMA (Cactaceae)

Fig. 21. Pereskia aculeata

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71

[Vol. 45
72 ANNALS OF THE MISSOURI BOTANICAL GARDEN

2. PERESKIA BLEO (HBK.) DC. Prodr. 3:475. 1828; Britton & Rose, in Carnegie

Inst. Wash. Publ. No. 248. 1:17. 1919.
poki bleo HBK. Nov. Gen. & Sp. 6:69. 1823.

Pereskia panamensis Weber, in Bois, Dict. Hort. 739. 1898.

Pues or small trees to 4-7 m. tall; branches terete, rather stout, armed with
few to several slender straight spines usually about 1 cm. long. Leaves rather
long-petiolate, the blade usually elliptic-obovate, acuminate, cuneate, 4-20 cm.
long, 2-7 cm. broad, the petiole 0.5-3.5 cm. long. Flowers usually few or solitary
in terminal or axillary panicles shorter than the subtending leaves; pedicels about
5 mm. long; perianth pink to deep red, rotate, about 5-7 cm. in diameter, the
segments rather numerous, the inner broadly obovate, the outer oblong-linear;
stamens numerous, about half as long as the perianth segments; style rather stout,
slightly longer than the stamens. Berry turbinate, 5-6 cm. long and broad, with
several more or less persistent bracts, yellow when ripe.

Panama and northwestern South America. In Panama the plants are known
as ñajá de culebra and fiajń de espinas. Britton and Rose found that the plants
widely cultivated as P. bleo actually should be referred to P. grandifolia Haw., a
native of Brazil.

RIÉN: trail between Pinogano and Yavisa, Allen 281; Marraganti, Williams 661;
Gene Pittier 1884. PANAMA: Chepo, Pittier 4704. SAN BLAS: Caledonia Harbor, Elmore
L34.

2. NOPALEA Salm-Dyck

NoPALEA Salm-Dyck, Cact. Hort. Dyck. 1849:63. 1850; Britton & Rose, in
Carnegie Inst. Wash. Publ. No. 248. 1:33. 1919.

Massive succulent shrubs; stems phylloid and jointed when young, the joints
fleshy, compressed, oval to oblong-elliptic, bearing amphigenous and rather sparse
areoles armed with hairs and glochids and usually 1 to few stout spines, stoutly
cylindric when older. Leaves inconspicuous, acicular, fugacious. Flowers sessile,
solitary, chiefly marginal upon the young stem joints; perianth shortly campanu-
late, with a short and broad hypanthium, the segments rather few, the outer
somewhat shorter and less petaloid than the inner, erect or very slightly pré
stamens very numerous, the filaments much longer than the perianth, unite
graded series at the base of the hypanthium in a shallow, glandular-dentate cup;
ovary broadly turbinate, deeply concave, more or less tuberculate, the areoles very
prominent; style longer than the stamens, enlarged shortly above the base into a
conspicuous disciform stylopodium. Fruit a fleshy berry with numerous seeds.

About a dozen species of the Antilles and Central America. The two species of
Panama, if they are species indeed, both appear to have been introduced, although
Britton and Rose considered N. dejecta as “perhaps native in Panama.”

a. Areoles aa: spines or nearly so; ovary and berries prominently

tuberculat 1. N. COCHENILLIFERA
aa. Areoles with 2-8 stout spines; ovary and berries scarcely tuberculate....... 2. N. DEJECTA

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73

FLORA OF PANAMA (Cactaceae)

illifera

Nopalea coc.

Fig. 22.

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LVol. 45
74 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. NOPALEA COCHENILLIFERA (L.) Salm Dyck, Cact. Hort. Dyck. 1849:64.
1850, as coccinellifera; Britton & Rose, in Carnegie Inst. Wash. Publ. No. 248.
1:34. 1919.

Cactus cochenillifer L. Sp. PI. 468. 1753

Opuntia cochinelifera (L.) Mill. Gard. Dict. ed. 8. 1768, sphalm.

Stout shrubs to 4 m. tall, the older trunks broadly cylindric, to 2 dm. thick,
the young stem joints elliptic-oblong, to 5 dm. long, green, the areoles without
spines or nearly so. Flowers pink to deep red, the perianth segments erect or nearly
so, the outer 0.5—1.5 cm. long, deltoid, the inner broadly oblong-obovate, 1.5—2.5
cm. long, the hypanthium naked, broadly campanulate or urceolate, 1.0-1.5 cm.
long, about 1 cm. broad at the orifice; staminal filaments 2.5—3.5 cm. long, pink;
ovary 2-3 cm. long, about 1.5 cm. broad, oblong-turbinate, truncate and deeply
concave, tuberculate, the areoles unarmed. Berry oblongoid-turbinate, about 5
cm. long, deep red when ri |

Original habitat unknown, widely cultivated and escaped in tropical America.
Occasionally encountered in cultivation or thickets in Panama, according to both
Standley (Contr. U. S. Nat. Herb. 27:278. 1928) and Britton and Rose, but
only one specimen of definitely Panamanian origin has been found during the
preparation of the present account.

en the Spaniards arrived in Mexico in 1518 they found the natives using a
brilliant purple, red, or orange dye, cochineal (Spanish, cochinilla), which they
brushed from the surface of various cacti, particularly Nopalea cochenillifera.

Their value quickly appreciated, the dye and the plants from which it was gathered

were distributed by the Spaniards to other parts of their empire and to Spain itself

as early as 1523.

It was not until the year 1703 that the microscope revealed the source of the
dye to be a small coccid scale infesting the plants. The dye stuff was obtained by
brushing the insects from the parasitized plant and roasting or boiling their bodies,
which later were dried and pulverized for the vat. Until the invention of the
aniline dyes cochineal was of enormous importance as a dye stuff, more than
6,000,000 pounds valued at about $4,000,000 being produced in 1868 by the Canary
Islands alone. Little use of it is found at present since the dye is not permanent.

CANAL ZONE: Bas Obispo, Verner in 1913.

2. NOPALEA DEJECTA (Salm-Dyck) Salm-Dyck, Cact. Hort. Dyck. 1849:64.

1850; Britton & Rose, in Carnegie Inst. Wash. Publ. No. 248. 1:37. 1919.
Opuntia dejecta Salm-Dyck, Hort. Dyck. 361. 1834.

Shrubs 1-2 m. tall, the old trunks stoutly cylindrical, very spiny, the young
stem joints narrowly elliptic-oblong, to 2 dm. long, bright green, the areoles with
2-8 stout spines. Flowers dark red to pink, the perianth segments erect or nearly
so, the outer deltoid, 0.5-1.0 cm. long, the inner broadly oblong-obovate, 1.5-2.0
cm. long, the hypanthium naked, broadly campanulate, about 1 cm. long and 0.7
cm. broad at the orifice; staminal filaments dark red to pink, about 4 cm. long;

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FLORA OF PANAMA (Cactaceae ) 75

ovary broadly turbinate, truncate and deeply concave, 2.0-2.5 cm. long, 1.0-1.5
cm. broad, scarcely tuberculate, the areoles unarmed or with 1-2 short slender
spines. Berry turbinate, 3—4 cm. long and broad, deep red when ripe.

Commonly cultivated and escaped in tropical America, suggested as “perhaps
native in Panama” by Britton and Rose. Nevertheless, during the preparation of
the present account only a single herbarium specimen from Panama was discovered.

CANAL ZONE: between Empire and Culebra, Cowell 418a.

3. OPUNTIA [Tourn.] Mill.

OPUNTIA [Tourn.] Mill. Gard. Dict. ed. 4. 1754.
Consolea Lem. Rev. Hort. - = 1862.
Tephrocactus ken, Cact. 88

Ficindica St. Lag. in Ann. Sóc. "BAĆ Lyon 7:70. 1880.

RR a small trees, and subshrubs; stems phylloid and jointed when
young, the joints fleshy, compressed and oval to broadly linguiform in our species,
bearing amphigenous areoles armed with hairs, glochids, and spines, very rarely un-
armed. Leaves inconspicuous, acicular and fugacious. Flowers sessile, chiefly
marginal upon the young joints; perianth broadly campanulate, with a short and
broad hypanthium, the segments numerous, the outer progressively shorter and less
petaloid than the inner, widely spreading; stamens very numerous, the filaments
much shorter than the perianth, united at different levels into a shallow glandular
cup, somewhat deflexed at anthesis, sensitive and inflexed upon stimulation; ovary
turbinate to cylindric, concave, the areoles prominent; style slightly longer than
the stamens, terete to fusiform. Fruit a fleshy berry with numerous seeds.

Very many species, all indigenous to the Americas, but some, particularly O.
ficus-indica, introduced and escaping in the dry tropics and subtropics of the East-
ern Hemisphere where their exuberent growth has made them a serious pest in many
areas. In Mexico, where the genus is most diversified and frequent, various species,
particularly O. ficus-indica and O. tuna, are cultivated under the name of nopal
and the fruits, called funas, eaten as a delicacy. Although tunas are marketed in
the United States, chiefly in the southwest, the large usually rather insipid berries
have not attained much popularity.

1. OPUNTIA ELATIOR Mill. Gard. Dict. ed. 8. 1768; Britton & Rose, in Carnegie

Inst. Wash. Publ. No. 248. 1:153. 1919.

Cactus nigricans Haw. Misc. Nat. 1 803.

Cactus elatior (Mill.) Willd. sd "Hort. Berol. Suppl. 34. 1813.

Cactus tuna B elatior (Mill.) S = ne cit. sub bl. 1557. 1813

Cactus tuna ^y nigricans (Haw.) S n Curt. Bot. Mag. 38: pl. 1557. 1813.

Densely branched shrubs to 5 m. tall, the young stem joints narrowly obovate

to „suborbicular, to 4 dm. long, olive-green, the areoles 2—4 cm. apart, minutely

and with 1-8 unequal, terete-acicular brown black-tipped spines 1-7 cm.
long. Flowers about 5 cm. in diameter, deep yellow more or less tinged with
red or salmon, the inner segments obovate, mucronate, about 2 cm. long; staminal

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[Vol. 45
76 ANNALS OF THE MISSOURI BOTANICAL GARDEN

filaments 1.5—2.0 cm. long, deeply tinged with red or salmon, yellow toward the
base; ovary obovoid-turbinate, about 2 cm. long and broad, the areoles unarmed or
nearly so. Berry obovoid, truncate, 4-5 cm. long, deep red when ripe.

Original habitat unknown; said to be frequent in Curacao, coastal Colombia,
and Panama (Britton & Rose), but definitely known stations in Panama at present
are limited to the islands of the Gulf of Panama.

PANAMA: Taboguilla Island, Howe 3792; San Josć Island, on rocks of beach, Erlan-
son 171; locality indefinite, Cbristopberson 20 207.

4. ACANTHOCEREUS (Engelm. ex A. Berger) Britton & Rose

ACANTHOCEREUS (Engelm. ex A. Berger) Britton & Rose, in Contr. U. S. Nat.
Herb. 12:432. 1909; in Carnegie Inst. Wash. Publ. No. 248. 2:121. 1920.

Cereus subsect. Acantbocereus Engelm. ex A. Berger, in Ann. Rep. Missouri Bot. Gard. 16:
Zi 1905.

Succulent erect or clambering shrubs; stems phylloid and jointed, the joints
elongate, rooting adventitiously at the tips, 3- to 5-angled, the areoles marginal
on the angles, shortly pubescent and prominently spiny. Leaves very inconspicuous,
acicular and fugacious or apparently wholly lacking. Flowers nocturnal, sessile,
borne singly at the areoles; perianth large, subsalverform, with an elongate tube,
the segments very numerous, the outer progressively shorter and narrower than
the inner, widely spreading, the tube bearing several minute caducous bracts and
prominent pubescent and spiny areoles; stamens very numerous, the filaments
somewhat shorter than the perianth, united at progressively deeper levels to the
hypanthium; ovary cylindric-ovoid, the areoles numerous and crowded, pubescent
and spiny; style filiform, terete, somewhat shorter than the outer and upper
stamens. Fruit a fleshy berry with numerous seeds.

A genus of about a dozen species, most frequent in Central America. A single
species is recorded from Panama.

1. ACANTHOCEREUS PENTAGONUS (L.) Britton & Rose, in Contr. U. S. Nat.
Herb. 12:432. 1909; in Carnegie Inst. Wash. Publ. No. 248. 2:123. 1920.

Cactus pentagonus L. Sp. Pl. 467. 1753.

Gives sirul Weber ex Gosselin, in Bull. RR His. Nat. Paris 10:384. 1904.
Clambering terrestrial shrubs, erect when young, soon arching and rooting at
the tips of the stems and forming rather extensive clones, the young stem joints

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FLORA OF PANAMA (Cactaceae)

Fig. 23. Acanthocereus pentagonus

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EVol. 45
78 ANNALS OF THE MISSOURI BOTANICAL GARDEN

elongate, rather slender, 3- to 5-angled, the areoles marginal on the angles, shortly
pubescent and with few to several stout straight spines. Flowers very showy, the
perianth tube 15-20 cm. long, about 5 mm. in diameter at the base, abruptly
dilated near the orifice into a short obconic throat about 4 cm. in diameter, armed
with rather sparse spiny areoles, the lobes widely spreading, white or the outer
segments more or less suffused with greenish red, the inner oblanceolate, acute to
acuminate, 4-7 cm. long; ovary cylindric-ovoid, truncate, about 2 ‘cm. long,
densely armed with spiny areoles. Berry oblong-ovoid, 4-5 cm. long, deep red
when ripe, persistently spiny.

Forming thickets at the margins of beaches from the keys of Florida and the
coast of Texas southward to Colombia and Venezuela; sporadically introduced in
the Antilles and perhaps indigenous in Guadeloupe

NE: vicinity of Miraflores Lake, G. White 145. PA

NAM
val bs & Seibert 1490, Maxon & Valentine fon Bella Vista, ma oar ko Poli
Panamá, Hitchcock in 1911; Gorgona Beach, G. W bite 154.

5. HYLOCEREUS (A. Berger) Britton & Rose

HYLOCEREUs (A. Berger) Britton & Rose, in Contr. U. S. Nat. Herb. 12:428.

1909; in Carnegie Inst. Wash. Publ. No. 248. 2:183. 1920.

Cereus subgen. Hylocereus A. Berger, in Ann. Rep. Missouri Bot. Gard. 16:72. 1905.

Succulent shrubby root-climbers, frequently epiphytic; stems phylloid and
jointed, rooting adventitiously at the nodes, the joints elongate, usually 3-angled,
the areoles marginal on the angles, shortly pubescent and with infrequent and
inconspicuous spines. Leaves inconspicuous and fugacious or wholly lacking.
Flowers nocturnal, sessile, borne singly at the areoles; perianth large, broadly in-
fundibuliform, the segments very numerous, the outer progressively shorter and less
petaloid than the inner, the tube rather broad and somewhat shorter than the seg-
ments, bearing rather few conspicuous, persistent foliaceous bracts but without
well-defined areoles; stamens very numerous, the filaments shorter than the peri-
anth, united at progressively deeper levels to the hypanthium; ovary cylindric-
ovoid, with few to numerous persistent accrescent foliaceous bracts but without
well defined areoles; style filiform, somewhat longer than the stamens. Fruit a
fleshy berry with numerous seeds.

About 20 species of Central America, northern South America, and the Antilles.
Several species of Hylocereus are cultivated since they are amongst the most hand-
some of the Nightblooming Cereuses and some have escaped and naturalized in the
tropics of the Old World.

a. —— egens or with few and i inconspicuous spines; — tube and

vary with few and distant bracts; stigma lobes dichotomous................... 1. H. MONACANTHUS
aa. Aide artis several short stout spines; — ray aa ovary with
numerous imbricate bracts; stigma lobes e 2. H. POLYRHIZUS

(Britton Wee a sterile pr collected by =. _in Panama to H. triangularis but remarks
probably native there, however.” The oints of Cowell’s specimen, which » have
ami nai. are capisce lly dol to those of H. soluri aa but bear 8—10 spines at "the areoles rather

hem the 3-5 usual for the latter species.)

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1958]

79

FLORA OF PANAMA (Cactaceae)

f
YON
NS

Fig. 24. Hylocereus monacantbus

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[Vol. 45
80 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. HyLOCEREUS MONACANTHUS (Lem.) Britton & Rose, in Carnegie Inst. Wash.
Publ. No. 248. 2:190. 1920.

Cereus monacanthus Lem. Hort. Univ. 6:60. 1845.

Succulent shrubby root-climbers, usually epiphytic, the young stem joints
elongate, rather stout, usually sharply 3-angled, the areoles marginal on the slightly
undulate angles, very minutely puberulent and occasionally with 1 to few incon-
— spines. Flowers very showy, the perianth tube 12-15 cm. long, about

. in diameter at the base, abruptly dilating to a broadly obconic throat about
7 cm. long and broad, bearing few to several persistent foliaceous bracts 1-2 cm.
long, the segments gradually spreading, greenish white, the inner rather broadly
oblanceolate, subcaudate-acuminate, 12-15 cm. long; ovary cylindric-ovoid, 1.5-
2.0 cm. long, persistent foliaceous bracts 0.2-1.0 cm. long, accrescent; stigma
lobes dichotomous. Berry cylindric-ovoid, about 8 cm. long, unarmed save for the
marcescent bases of the bracts.

Epiphytic in thickets and forested river banks, Panama and Colombia.

on 4689; vicinity of Summit, Bartlett 6 z 17023;

Ai aie. camo e. Pittier i in a I. CHIRIQUf: east of Gualaca, EE Allen
5041; Isla Parida, Maxon 4904. Pinogana, ded = PANA BK io RÓ:
Woodson & Schery 1021; Bella vista, Killip 12030; Tap ver, y Diaz region, Maxon

8 Harvey 6705; Urava Island, Howe 31778; Palo Island, "Cale d 2

2. HYLOCEREUS POLYRHIZUS Cre Britton & Rose, in Carnegie Inst. Wash.
Publ. No. 248. 2:185. 1920

Cereus polyrbizus Weber, in Schum. Gesamtb. Kakteen 151.

Cereus trigonus var. costaricensis Weber, in Bull. Mus. Hist. Ee. “Pais 8:457. 1902.
Hylocereus costaricensis (Weber) Britton & Rose, loc. cit. 186.

Cereus costaricensis (Weber) A. Berger, Kakteen 122.

Succulent shrubby root-climbers, frequently epiphytic, the young stem joints
elongate, usually sharply 3-angled, the areoles marginal on the entire or slightly
undulate angled, armed with several short stout spines. Flowers very showy, the
perianth tube rather stout, 1.0—1.5 dm. long, about 1 cm. in diameter at the base,
gradually dilating to a broadly obconic throat about 6 cm. in diameter at the
orifice, bearing numerous persistent foliaceous bracts particularly imbricating to-
ward the base, the segments gradually spreading, greenish white, the outermost
tinged with purple, the inner rather broadly oblanceolate, acuminate, about 1.5-2.0
dm. long; ovary cylindric-ovoid, about 4 cm. long, the persistent foliaceous bracts
numerous and imbricate, 1-2 cm. long; stigma lobes entire. Berry oblongoid,
about 10 cm. long.

Colombia and Panama. “A common species on barren rocks, islets and cliffs
along the coast [of San José Island], and frequent as an epiphyte, especially on the
branches of Bombax quinatum in the interior of the island [I. M. Johnston, in
ins 8:217. 1949]."

San José Island, Perlas Archipelago, Jobnston 1126, Erlanson 172; Old
Post Rosę & Rose 18506.

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FLORA OF PANAMA (Cactaceae) 81

Both Berger and Britton and Rose distinguish H. polyrbizus and H. costaricensis
solely upon the thickness of the stem joints, whether (according to Britton & Rose)
“only 3 to 4 cm. thick” or “5 to 10 cm. thick,” respectively. ‘The short descrip-
tions of Britton and Rose are remarkably similar, differing almost solely in the
position of identical adjectives. In this regard, Dr. Johnston's comments, partly
quoted above, are very suggestive: "The plant varies in its form and appearance
according to its particular habitat. On flat exposed rock surfaces it may
compact in growth, 1-3 dm. tall, and composed of short, stout, decumbent or
ascending stems 3-6 cm. thick. Stems dangling from high branches or from the
tops of cliffs usually are only 1-3 cm. thick but may become several meters or even
more in length. Stems rooted on branches become flattened, as much as 10 cm.
broad, and tend to wrap themselves about the support. The plant has typically
3-angled stems, but rarely 4-angled ones may be found.” With such testimony, it
appears entirely probable that H. polyrbizus and H. costaricensis are conspecific.

6. WEBEROCEREUS Britton & Rose

WEBEROCEREUS Britton & Rose, in Contr. U. S. Nat. Herb. 12:431. 1909; in
Carnegie Inst. Wash. Publ. No. 248. 2:214. 1920.

Succulent shrubby root-climbers, frequently epiphytic; stems phylloid and
jointed, rooting adventitiously at the joints, the joints elongate, rather indefinitely
winged, occasionally nearly terete or flattened, the areoles marginal on the wings,
very inconspicuously puberulent to essentially naked, seldom with extremely few
and inconspicuous spines. Leaves apparently wholly lacking. Flowers nocturnal,
sessile, of mediocre size in our species, borne singly at the uppermost areoles, very
broadly infundibuliform, the tube very broad and somewhat shorter than the seg-
ments, bearing few to rather numerous broad fugacious bracts and conspicuous
areoles armed with numerous slender subarachnoid bristles and hairs, the segments
very numerous, the outer progressively shorter and less petaloid than the inner,
spreading; stamens very numerous, the filaments shorter than the perianth, inserted
at progressively deeper levels within the hypanthium; ovary cylindric, bearing
crowded fugacious bracts and areoles armed with numerous persistent subarachnoid
bristles and hairs; style filiform, somewhat longer than the stamens. Fruit a fleshy

rry with persistent bristly areoles and numerous seeds.

Three or four questionably distinct species of Panama and Costa Rica.

—

+ WEBEROCEREUS PANAMENSIS Britton & Rose, in Carnegie Inst. Wash. Publ.

No. 248. 2:215. 1920.

Succulent epiphytic root-climbers, the young stem joints elongate, 4-7 mm.
thick, rather indefinitely angled or winged and occasionally flat, the areoles
marginal, inconspicuously indented, very inconspicuously puberulent to naked.
Flowers mediocre, the perianth tube 2-3 cm. long, about 4 mm. in diameter at the
base, immediately dilating to an obconic throat about 1 cm. in diameter at the
orifice, bearing several fugacious deltoid bracts and areoles armed with slender

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[Vol. 45
82 ANNALS OF THE MISSOURI BOTANICAL GARDEN

subarachnoid bristles and hairs, the segments suberect or slightly spreading, pale
yellowish orange tinged with pink, apparently occasionally white, oblong-elliptic,
1.5-3.0 cm. long; ovary oblong-turbinate, truncate, about 1 cm. long, bearing
minute fugacious bracts and persistent subarachnoid-bristly areoles. Berry broadly
obovoid-turbinate, 2-3 cm. long, very prominently tuberculate, bright red when

Fig. 25. Weberocereus panamensis

Lowland streamside forests and thickets, Panama.

COLON: near mA of Rio Chagres, Allen 882; along Río Fato, Pittier 3903. LO-
CALITY UNKNOWN: Hunter 17040, I7041.

The Hunter poa with only the comment "In long strands climbing over
trees" have been determined previously as Selemicereus inermis (Otto) B. & R.
They are quite sterile, however, consisting of older stems which are sharply 4-angled
as a rule. Although we believe that they probably represent W. panamensis it is
possible of course that such actually may not be the case. The flowering stem
joints of W. panamensis which are before us are 2- to 3-angled.

7. EPIPHYLLUM [Hermann] Haw.

EPIPHYLLUM [Hermann] Haw. Syn. Pl. Succ. 197. 1812; Britton & Rose, in

Carnegie Inst. Wash. Publ. No. 248. 4:185. 1923
Pbyllocactus Link, Handb. Erkenn. Gewichse 2:10. 1831.

Pbyllocereus Miq. in Bull. Sci. Phys. Nat. Bruxelles 112. 1839.

Succulent shrubby plants, usually epiphytic root-climbers; stems phylloid and
jointed, usually of two intergrading types: the primary terete or subterete, usu-
ally rooting, the secondary (flowering) flattened and leaf-like, lobed or undulate,
the areoles marginal, naked or rarely minutely puberulent. Leaves lacking. Flow-
ers nocturnal or ephemeral, sessile, borne singly at the areoles; perianth usually
large, infundibuliform, the segments very numerous, the outer progressively shorter
and less petaloid than the inner, reflexed or spreading, the tube relatively slender,
usually longer than the segments, bearing usually few and sparse, inconspicuous

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1958]
FLORA OF PANAMA (Cactaceae) 83

bracts, usually without definite areoles; stamens very numerous, the filaments
somewhat shorter than the perianth and united at progressively deeper levels to the
hypanthium; ovary frequently cylindric-ovoid, with few to numerous naked or
rarely minutely puberulent areoles; style filiform, somewhat longer than the
stamens. Fruit a fleshy berry with numerous seeds.

About 20 species of tropical America, with the greatest concentration in Cen-
tral America. These are the species of Nightblooming Cereus usually cultivated
as house plants in the United States. A characteristic growth pattern of most
Epiphyllums is the differentiation of the stems into more or less terete, rooting
primary joints and flattened, leaf-like secondary joints upon whic

h the flowers
eventually are borne.

Areoles appear with geometric regularity upon the secondary
Joints, but are erratic upon the primary. e two basic types of joint may be
combined in any way: a secondary proceeding from a primary, vice versa, a pri-
mary from a primary, or a secondary from a secondary.

a. Flowers tes elongate and slender, the perianth tube 6 or more tim

longer than the segments; secondary (flo wering) stem joints rather narrow-
nd callose-marginate in dessication, the areoles naked

. E. PHYLLANTHUS
aa. Flowers usually ‘broad and massive, the perianth tube 2—3 (or less) times
- nger than the segments.
Hess relatively small and inconspicuous, about 8—15 cm. long, the
oen ianth tube times longer than the segments; secondary stem
= rather rote setce, thick and callose-marginate in die
ion, the areoles naked or indefin itely papillate 2. E
` Flowers ire and landa the periant th tube less than twice as long
s the segmen

» PITTIERI

y
o

c SP. jaa about 15-25 cm. long: primary stem joints rather slender
and teret

d. Sicilie stem joints rather enh yap z agri) m m
the ids minutely puberulent; flowers about m. long. 3. E. LEPIDOCARPUM
dd. rin. c stem joints deeply erenate, the ae "ibid
iiu: flowers about 20—25 cm. lon 4. E. MACROPTERUM
cc. Flowers about 40 cm. long; primary stem “joi ints very stout and
e or less winged or angled (secondary stem joints unknown)..... 5. E. GIGAS

1. EPIPHYLLUM PHYLLANTHUS (L.) Haw. Syn. Pl. Succ. 197. 1812; Britton &

Rose, in Carnegie Inst. Wash. Publ. No. 248. 4:187. 1923

Pbyllocactus pr nr Der Link, H Handb. Erkenn. Gewichse 2:11. ;
Rbibsalis pbyllantbus oh ae Sch. in Mart. FI. Bras. 4?:298. 1890, in jig as to basinym.
Epipbyllum gaillardae Britton & Rose, in Contr. U. S. Nat. Herb. 16:2 19
P bri gaillardae a & Rose) Vaupel, in ra ct Kakteznk. 23: 87. 1913.
Succulent epiphytic shrubs, the primary stem joints terete, elongate, about 5
mm. thick when young, probably thicker when old, the secondary stem joints
narrowly oblong-elliptic to linear-elliptic, usually obtuse to rounded at the tip,
cuneately decurrent into a terete subpetiolar base, 3-5 dm. long, 2-8 cm. wide,
rather narrowly serrate, thick and callose-marginate in dessication, the areoles 3-5
cm. distant, naked or indefinitely papillate, borne superficially upon the upper
broad margin of the serrations. Flowers nocturnal, very elongate and slender, white,

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ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 26. Epipbyllum pbyllantbus

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[Vol. 45

1958]
FLORA OF PANAMA (Cactaceae) 85

the perianth tube about 18-25 cm. long, narrowly tubular, about 3 mm. in di-
ameter at the base, abruptly dilated into an extremely short conic throat about
8 mm. broad at the orifice, bearing very few distant and inconspicuous external
bracts, the perianth segments very narrowly oblong-elliptic, acuminate, spreading,
the inner about 3-4 cm. long, the ovary oblongoid, inconspicuously tuberculate.
Berry oblong-turbinate, truncate, somewhat tuberculate-angulate, about 7-8 cm.
long and 2 cm. broad, bright red when ripe.
Costa Rica to Brazil, Bolivia and Paraguay.

NAL ZONE: Quebrada Culebra, Madden Lake, Dodge 6 Allen V dicc COCLE: hills
m of El Valle, alt. 1000 m., Allen 2334; Penonomé, Williams 423. ÓN: Porto
sesta Pittier in 1912. DARIEN: headwaters of Rio Chico, alt. 500-750 E we 4609.

PANAMÁ: vicinity of Pacora, alt. 40 m., Allen 2241; Juan Díaz, Maxon 6 Har rvey 6706.

2. EPIPHYLLUM PITTIERI (Weber) Britton & Rose, in Contr. U. S. Nat. Herb.

16:258. 191
Pbyllocactus pittieri Weber, in Bois, Dict. Hort. 2:957. 1898.

Succulent epiphytic shrubs, the primary stem joints terete or weakly angulate,
5-10 mm. thick when young, probably thicker when old, the secondary stem
joints linear-elliptic, acute to obtuse at the tip, decurrent to a short terete sub-
petiolar base, about 2-10 dm. long and 2-6 cm. broad, rather narrowly serrate,
thick and callose-marginate in dessication, the areoles 3-6 cm. distant, naked or
indefinitely papillate, borne superficially upon the upper broad margin of the serra-
tions. Flowers nocturnal, relatively small and inconspicuous, white, the perianth
tube about 6-10 cm. long, narrowly tubular, about 4 mm. in diameter at the base,
abruptly dilated into an extremely short conic throat about 7 mm. broad at the
orifice, bearing few and distant inconspicuous petaloid bracts, the perianth segments
linear-elliptic, acuminate, spreading, the inner about 4-5 cm. long, the ovary
narrowly oblongoid, inconspicuously bracteate and tuberculate. Berry oblongoid,
about 2 cm. long, deep red when ripe.

Costa Rica and Panama.
TORO: vicinity of Chiriqui Lagoon, Von Wedel 1294; Water Aa Von

Wedel 1. I 1528, 1580, 17684; brada Nigua, Almirante Bay, Von Wedel 7; Garay Creek,
Von Wedel ago 0. rii: i. Drayton Ts Barro Colorado Midland Shattuck 603.

CHIRIQUf: forest, Puerto Armuelles, Darid II

3. EPIPHYLLUM LEPIDOCARPUM (Weber) Britton & Rose, in Contr. U. S. Nat.

Herb. 16:257. 1913.

Phyllocactus lepidocarpus Weber, in Bull. Mus. Hist. Nat. Paris 8:462. 1902.

Succulent epiphytic shrubs, the primary stem joints weakly 3-angled, up to 1
cm. thick when young, becoming terete and thicker when old, the secondary stem
joints oblong- to narrowly ovate-elliptic, obtuse at the tip, decurrent to a very
short weakly angled subpetiolar base, about 2-7 dm. long and 3-7 cm. broad,
rather narrowly serrate, remarkably thin and more or less venose in dessication, the
areoles 4-5 cm. distant, minutely puperulent, borne superficially upon the upper

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[Vol. 45
86 ANNALS OF THE MISSOURI BOTANICAL GARDEN

broad margin of the serrations. Flowers nocturnal, rather broad and handsome,
white with the outer segments flushed with rose, the perianth tube about 7—8 cm.
long, tubular, about 4 mm. in diameter at the base, rather gradually dilating to a
narrowly conical throat about 1.5 cm. broad at the orifice, bearing a few incon-
spicuous deciduous linear bracts subtending puberulent areoles toward the base, the
segments 7-8 cm. long, the outer linear, flushed with rose, the inner narrowly
obovate-elliptic, white, the ovary narrowly turbinate, bearing crowded persistent
ovate bracts and conspicuous puberulent areoles. Berry turbinate-fusiform, occa-
sionally conspicuously beaked, 6-9 cm. long, 2-4 cm. thick, covered with the
accrescent hyaline bracts and puberulent areoles, purplish pink when ripe.

Highland Costa Rica and western Panama, epiphytic or upon rocks.

CHIRIQUÍ: valley of the upper Rio Chiriquí Viejo, vicinity of Monte Lirio, alt. 1300-
1900 m., Seibert 206; Bajo Chorro, Boquete District, alt. 6000 ft., Davidson 253

A species particularly noteworthy because of its peculiar scaly fruits.

4. EPIPHYLLUM MACROPTERUM (Lem.) Britton & Rose, in Carnegie Inst. Wash.
Publ. No. 248. 4:193. 1923

Pbyllocactus macropterus Lem. Illustr. Hort. 11: Misc. 73. 1864.

Pbyllocactus tbomasianus K. Sch. in Monatsschr. Kakteenk. 5:6. 1895,

Pbyllocactus costaricensis Weber, in Bull. Mus. Hist. Nat. Paris 8:463. 1902.

Pbyllocactus macrocarbus Weber, loc. cit. 464. 1902.

Epiphyllum costaricense (Weber) Britton & Rose, in Contr. U. S. Nat. Herb. 16:256.

1 i
Epipbyllum tbomasianum (K. Sch.) Britton & Rose, loc. cit. 259. 1913.

Massive succulent epiphytic shrubs, the primary stem joints terete or weakly
4-angled and 5-8 mm. thick when young, becoming stouter and terete in age, the
secondary stem joints narrowly obovate- to oblong-elliptic, obtuse to rounded at
the tip, broadly decurrent to a short weakly angled subpetiolar base, about 6-8 dm.
long and 5-10 cm. broad, rather broadly and regularly crenate, rather thin and
venose in dessication, conspicuously callose-marginate, the areoles 2-5 cm. distant,
minutely puberulent, borne within the deep indentations of the crenations. Flowers
nocturnal, very large and handsome, the perianth tube 12-15 cm. long, tubular,
arcuate, about 6 mm. broad at the base, gradually dilating to a conical throat about
3 cm. broad at the orifice, pale greenish rose, bearing rather conspicuous persistent
subfoliaceous bracts, the segments about 8-12 cm. long, slightly spreading, the
outer linear-lanceolate, pale greenish rose, the inner obovate-elliptic, white; ovary
oblongoid, about 3-4 cm. long, bearing many subfoliaceous bracts. Berry oblong-
fusiform, usually beaked, about 10-12 cm. long and 4-5 cm. thick, slightly
tuberculate at the minutely puberulent areoles, red when ripe

Costa Rica and Panama, epiphytic and upon rocks, apparently usually at low
altitudes.

BOCAS DEL TORO: vicinity of Chiriquí Lagoon, Von Wedel 1164; Snapper Point, Von
Wedel 2030; Water Valley, Von Wedel 736; Shepherd Island, Von Wedel 2722. CHIRI-
Qui: Puerto Armuelles, Davidson 1190.

An aS: showy plant, but surpassed by the one which follows:

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1958]
FLORA OF PANAMA (Cactaceae) 87

5. EPIPHYLLUM gigas Woodson & Cutak, spec. nov.

Frutices succulenti scandentes radicantes, ramis ut dicitur ultra pedes centum
in longitudinem metientibus; articulis rami radicantibus validissimis ancipitibus
leviter undulatis nostris 2—3 cm. crassis; articulis florigeris infeliciter ignotis.
Flores maximi speciosissimi ut creditur albi noctiflori; perianthii tubo ca. 22-25
cm. longo basi ca. 7 mm. lato apicem versus gradatim dilatato ostio ca. 2.5-3.0 cm.
diam. extus bracteas linearis paucas gerente; perianthii segmentis exterioribus line-
aribus longe acuminatis 15 cm. longis vel ultra patulis, interioribusne latioribus in
exemplo nostro vix bene visis; ovario late cylindrico ca. 1.5 cm. longo inconspicue
bracteato areolas parvas paucas minute puberulentes gerente. Bacca ignota.
Species inter species generis jam cognitas omnibus partibus longe maxima.

AMÁ: Cerro Trinidad, alt. 800—1000 m., climbing rock faces of main peak, Oct.
20, 1946, P. H. Allen 3772 (Herb. Missouri Bot. Gard., HOLOTYPE).

This is a tremendous species, far surpassing in size of all parts any other known
Epiphyllum; as such it is impossible to ignore, even though one quite naturally
would desire the flowering stem joints and fruits for a complete description. The
rather regular undulations of the primary stem joints and the puberulent areoles of
the ovary suggest an affinity with E. macropterum and E. oxypetalum. Mr. Allen's
statement that the holotype specimen was a “giant liana, to several hundred feet
in length” might seem totally incredible were it not for the gigantic flower which
he preserved.

8. WITTIA K. Sch.

Wrrria K. Sch. in Monatsschr. Kakteenk. 13:117. 1903; Britton & Rose, in

Carnegie Inst. Wash. Publ. No. 248. 3:206. 1923.

Pseudorbipsalis Britton & Rose, loc. cit. 4:213. 1923.

Succulent shrubby root-climbers, usually epiphytic; stems phylloid and jointed,
usually of two intergrading types as in Epipbyllum: the primary terete or weakly
winged, usually rooting, the secondary (flowering) flattened and leaf-like, lobed
or undulate, with marginal areoles covered with few to several minute scales and
indefinitely papillate or naked. Leaves lacking. Flowers ephemeral (or noc-
turnal?), sessile, borne singly at the areoles; perianth mediocre or relatively small,
tubular to tubular-salverform or campanulate, the tube about as long as the seg-
ments or shorter, bearing a few inconspicuous deltoid bracts, the segments about
10—15, erect or spreading; stamens about 30—45, the filaments few to several times
longer than the anthers, inserted at progressively deeper levels within the perianth
tube, the outermost conspicuously shorter than the innermost; ovary ovoid-
fusiform, minutely tuberculate-bracteate. Fruit a fleshy berry with rather few
seeds,

It is extremely difficult to justify the separation of Pseudorbipsalis from Wittia
upon the basis of our scanty herbarium material, and surely it is nearly impossible
to do so even from the meagre descriptions and rather irrelevant comments of

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[Vol. 45
88 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Britton and Rose. Granted that there is an obvious transition between the sub-
tribes Epiphyllanae and Rhipsalidanae, the purpose of illustration as well as that of
convenience would be served better by fewer gradient genera than those recognized
by Britton and Rose.

Britton and Rose enumerate two species each for Wittia and Pseudorbipsalis.
In addition to the two species in Panama, W. amazonica is known from extreme
northeastern Peru and adjacent Colombia, and Ps. ala/a from Jamaica.
a. Flowers tubular-salverform, white flushed with rose, the — seg-

m

ents widely spreading; flowering stem joints narrow vende 1. W. HIMANTOCLADA
a. Flowers tubular, deep pink tipped with lavender, the perianth segments
te

m

2. W. PANAMENSIS

1. Wrrria himantoclada (Roland-Gosselin) Woodson, comb. nov.

Rbipsalis bimantoclada Roland-Gosselin, in Bull. Soc. Bot. France 55:694. 1908.

Wittia costaricensis Britton & Rose, in r. U. S. Nat. Herb. 16:261. 1913.
Pseudorbibsalis bimantoclada (Roland-Gosselin) Britton & Rose, in Carnegie Inst. Wash.
l. No. 248. 4:213. 1923.

Succulent epiphytic root-climbers, the primary stem joints terete or weakly
winged, up to about 1.5 m. long, the secondary stem joints narrowly lanceolate or
oblanceolate, 2—5 dm. long,1-5 cm. broad, obtuse to rounded at the tip, cuneately
decurrent into a subpetiolar base, narrowly repand, thick and coriaceous in dessi-
cation, the areoles 1-2 cm. distant. Flowers tubular-salverform, white flushed
with rose or orange without, the perianth tube about 5-6 mm. long, about 3 mm.
in diameter at the base, the perianth segments 10-15, narrowly elliptic to linear,
about 1 cm. long and up to 3 mm. broad, widely spreading; stamens about 45,
slightly exserted to barely included; ovary broadly cylindrical, about 7 mm. long,
deep rose. Berry unknown.

Costa Rica and Panama.

cocLÉ: north rim of El Valle, Allen 1897.

2. WITTIA PANAMENSIS Britton & Rose, in Contr. U. S. Nat. Herb. 16: 241.
1913; in Carnegie Inst. Wash. Publ. No. 248. 4:207. 1923.

Succulent scandent shrubs, epiphytic or upon rocks, the primary stem joints
terete or weakly ancipitous, the secondary stem joints very narrowly oblong-
anceolate, rounded at the tip, cuneately decurrent into a very short subpetiolar
base, 3-5 dm. long, 3-5 cm. broad, very narrowly and closely cuneate-serrate,
thick and callose-marginate in dessication, the areoles 1.0—2.5 cm. distant, naked
or indefinitely papillate beneath the minute imbricate scales, borne superficially
upon the upper margin of the narrow serrations. Flowers ephemeral, rather small,
tubular, pink tipped with lavender, the perianth tube about 1.5 cm. long, about
2 mm. in diameter at the base, scarcely dilated toward the orifice, the perianth
segments about 10, about 1 cm. long, erect or essentially so; ovary about 3 mm.
long, minutely tuberculate-bracteate. Berry not seen.

Eastern Panama; Colombia and Venezuela.

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1958]
FLORA OF PANAMA (Cactaceae) 89

Fig. 27. Wittia panamensis

DARIEN: Marraganti, Williams 698; trail between Pinogana and Yaviza, Allen 278.
PANAMA: Chepo, Pittier 4571. SAN BLAS: Caledonia, Elmore L40.

Wittia panamensis is distinguished from W. amazonica by Britton and Rose
solely upon the fruit, whether "smooth" or "roughened by small tubercles" re-
spectively. I have seen no fruit of W. panamensis as yet, but the ovaries of the
flowers certainly are minutely tuberculate and I rather suspect that the fruit of
the two species may have much in common.

9. RHIPSALIS Gaertn.

RHIPSALIS Gaertn. Fruct. Sem. 1:137. 1788; Britton & Rose, in Carnegie Inst.

Wash. Publ. No. 248. 4:219. 1923, nom. conserv.

Hariota Adans. Fam. 2:243. 1763, nom. rejic.

Rather slender succulent root-climbers, usually epiphytic; stems phylloid and
jointed, terete or essentially so in our species, the areoles spiral, naked to abundantly
hirsute or bristly. Leaves lacking. Flowers very small, sessile or somewhat sunken
within the stem joints, solitary; perianth rotate, the segments free or essentially so,
relatively few, sometimes only 5, usually spreading; stamens about twice as many
as the perianth lobes, the filaments inserted on the margin of the hypanthium.
Berry small, usually white, with rather few small seeds.

A perplexing genus of 50 or more species in tropical and subtropical America;
also with several species described from the Old World, at least some of which may
be introduced from America. Known as Mistletoe Cactus in the United States.

Besides the ubiquitous Rb. cassytha there are two very distinct species in
Panama which are known only from sterile specimens. Both probably are un-

ribed, but of course cannot be provided with names until flowering plants
are found.

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ANNALS OF THE MISSOURI BOTANICAL GARDEN

da, FA
A RO
E AP

nta 8 Na = >
sea ARMIN
a Ai:

3 GZK

Fig. 28. Rhipsalis cassytba

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[Vol. 45

19581

FLORA OF PANAMA (Cactaceae) 91
a. Sterile > minutely puberulent to essentially naked.
b. Flowering areoles minutely puberulent to a 7 naked 1. RH. CASSYTHA
bb. Pity areoles very conspicuously floc 2. RH. SP.
aa. Sterile areoles with conspicuous tufts of anda bristles 3. RH. sp.

1. RHIPSALIS CASSYTHA Gaertn. Fruct. Sem. 1:137. 1788; Britton & Rose, in

Carnegie Inst. Wash. Publ. No. 248. 4:225. 1923, as cassutba.

Cactus pendulus Sw. Prodr. 77.
Rbipsalis parasitica Haw. Syn. PI. Suse. 187. 1812
Hariota cassytha (Gaertn.) Lem. Cact. Gen. Nov. Sp. 75. 1839.

Pendulous, usually epiphytic succulents up to 9 m. long; stem joints terete,
dichotomous or in false whorls, yellowish green, about 2-3 mm. thick when young,
the areoles very inconspicuous, glabrous to minutely puberulent. Flowers lateral,
solitary, greenish white, the perianth segments about 2 mm. long. Berry globose,
white (rarely pink), about 5 mm. in diameter.

hroughout tropical America, apparently also in Ceylon and tropical Africa.
Very frequent in Panama.

OCAS DEL TORO: Water Valley, Von Wedel 649, 1670; Little d Vire Wedel 2529;
Pons River, Von Wedel 2584; = > Von Wedel 2924. E: vicinity of
re dden Dam and Alahuela, alt. 80-100 m., Hunter & Allen xa fano "Colorado Island,

Shattuck 447; Bas Obispo, PO s.m.; Gatun Lake, Rowlee & Rowlee 405; Balboa,
Standley 28546.

2. RHIPSALIS SP.

This species has stem joints which are about half as thick as is usual for those
of Rb. cassytba. The sterile areoles are glabrous or essentially so, but the young
flowering areoles are covered with exceedingly dense tufts of yellow hairs 2-3 mm.
long. If this is a species of series FLOCCOSAE, as seems probable, it is the most
northern representative of it.

BOCAS DEL TORO: vicinity of Chiriqui Lagoon, Von Wedel 1393.

3. RHIPSALIS SP.

This little plant resembles a Lycopodium at first glance. It is juvenile, appar-
ently, and the stems, which are somewhat thicker than those of RP. cassytha, have
the sterile areoles beset with conspicuous tufts of spreading bristles 1-2 mm. long.
Its general aspect agrees in general with the illustration of Rb. aculeata given by
Britton & Rose (in Carnegie Inst. Wash. Publ. No. 248. 4: pl. xxiv, fig. 8). On
the other hand, a good many Rhipsalis seedlings appear bristly or spiny and later
more mature growth becomes almost glabrous.

BOCAS DEL TORO: Pumpkin River, Von Wedel 2572.

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MISSOURI BOTANICAL GARDEN

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Economic
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T. and Editor GEORGE B. VAN SCHAACK,
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Research Associate Senior Taxonomist

BOARD OF TRUSTEES

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< JoHN S. LEHMANN

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p bone of n vp dad of the Academy oi
= Missouri È Science of St. Louis
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RAYMOND R. TUCKER, ©
umie roza: St. Lovis. >

Oscar E Grama, r

v XLV Number 2

Annals
of the

Missouri Botanical
Garden

MAY, 1958

= of Panama. Part VII, Fascicle : 2 > (fiale to Myrtaceae) roe
eee! oy oe 93-201

i paw UARIERLY AT + aa IEEE A Si:
BY rum BOARD OF TRUSTEES OF P THE MISSOURI BOTANICAL GARDEN A

Annals
of the

Missouri Botanical Garden

A Quarterly Journal containing Scientific Contributions from the
Missouri Botanical Garden and the Henry Shaw School of Botany of
Washington University in affiliation with the Missouri Botanical
Garden.

„Information
The ANNALS OF THE MISSOURI BOTANICAL GARDEN appears four times
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GARDEN are listed in the Agricultural Index, published by the H. W. Wilson
Company.

FLORA OF PANAMA

BY

ROBERT E. WOODSON, JR.
AND
ROBERT W. SCHERY

AND COLLABORATORS

PART VII

Fascicle 2

THYMELAEACEAE

LYTHRACEAE (L. I. Nevling)
LECYTHIDACEAE
RHIZOPHORACEAE (D. P. Gregory)
COMBRETACEAE (A. W. Exell)
MYRTACEAE (G. J. H. Amshoff)

ANNALS
OF THE

Missouni BOTANICAL GARDEN

Vol. XLV MAY, 1958

No.

FLORA OF PANAMA

Part VII. Fascicle 2*

THYMELAEACEAE

Shrubs and small trees, usually with soft wood and leathery flexible branches.
Leaves alternate, spiral, exstipulate, pinnately veined. Inflorescence terminal or
lateral, determinate, simple, or compound with the ultimate divisions umbelliform
or capituliform. Flowers perigynous, perfect or (in our genera) unisexual and
dioecious, regular or rarely somewhat zygomorphic; perianth dichlaymdeous or (in
our genera) monochlamydeous through loss of the corolla, usually tetramerous,
tubular to salverform or subrotate, more or less petalaceous, usually lepidote or
puberulent without, the limb (calyx) 4- to 5-parted; stamens in 1 or 2 cycles, the
outer antesepalous and inserted upon the perianth lobes, the inner antepetalous and
inserted within the perianth tube; pistil usually 1-carpellate, superior, usually borne
upon a short gynophore and subtended by an inconspicuous disc, containing a single
pendulous ovule upon the ventral placenta, the stigma capitate, usually sessile or
subsessile. Fruit a dry nutlet or small drupe (our genera); seed exalbuminous or
essentially so, the embryo with thick convex cotyledons.

Thymelaeaceae are a rather small family chiefly abundant in South Africa and
Australia and very poorly represented in the northern hemisphere, particularly in
America. Although the small flowers include only vestigial petals at most, occa-
sionally they are attractive because of petalaceous pigmentation of their hypanthium
(perianth tube), aggregation into dense clusters, and sweet scent.

The wood of the branches is soft and the young twigs are remarkably leathery
and flexible as a rule, hence the popular name Leatherwood for Dirca palustris, the
unique representative of the family in the eastern United States. This quality of
the stems is due in part to the anomalous development of the phloem which fre-
quently forms an abundant interwoven fabric of soft fibers. In fact, Standley
(FI. Costa Rica 2:759. 1937) reports the vernacular name mastate for Daphnopsis
seibertii in Costa Rica, which suggests the making of bark cloth.

a. Stamens 8, sessile or subsessile; pistillate flowers with staminodes...........-- 1. DAPHNOPSIS
aa. Stamens 4, widely exserted upon slender filaments winged at the base;
pistillate flowers without staminodes 2. SCHOENOBIBLUS

* Assisted by a grant from The National Science Foundation.

Issued May 27, 1958. (93)

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[Vol. 45
94 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. DAPHNOPSIS Mart. 8 Zucc.
DAPHNOPSIS Mart. & Zucc. Nov. Gen. 8 Sp. 1:65. 1824.

Hargasseria Schiede & Deppe, ex C. A. Meyer, in Bull. Acad. St. Petersbourg 1:356. 1843.
Nordmannia Fisch. & C. A. Meyer, loc. cit. 355. 1843.
Coleophora Miers, in Ann. & Mag. Nat. Hist. ser. 2. 8:197. 1851.

Shrubs or small trees. Leaves alternate, simple, exstipulate, membranaceous to
coriaceous. Inflorescence terminal or axillary, cymose or paniculate, the ultimate
branches usually umbelliform or capituliform. Flowers apetalous, dioecious,
tetramerous; perianth salverform to campanulate, more or less petalaceous, usually
densely puberulent, the limb deeply 4-lobed. Staminate flowers: stamens 8, sessile
or subsessile, the outer cycle antesepalous, inserted at the base of the perianth lobes,
barely exserted, the inner cycle antepetalous, inserted within the perianth tube;
pistillode present, surrounded at the base by a delicate entire or lobed disc. Pistil-
late flowers usually smaller than the staminate: staminodes 8, 4, or absent; pistil
l-carpellate, containing a single pendulous anatropous ovule; stigma capitate,
sessile or subsessile, usually slightly exserted; disc entire or lobed, hypogynous,
usually delicately membranaceous. Fruit a small slightly fleshy drupe.

Ż
Za

a
CRY
Y di

Fig. 29. Dapbnopsis seibertii

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1958]
FLORA OF PANAMA (Thymelaeaceae) 95

About 25 species in southern Mexico, Central and South America, and the
Antilles. A single species in Panama.

1. DAPHNOPSIS SEIBERTII Standl. in Ann. Missouri Bot. Gard. 24:192. 1937

Small dioecious trees 3-10 m. tall. Young stems minutely appressed-puberulent,
soon glabrate, the bark pale grayish yellow, with small and crowded verrucose
lenticels. Leaves alternate, petiolate, the blade elliptic to somewhat oblong- or
oblanceolate-elliptic, acuminate to subcaudate-acuminate, rather narrowly attenu-
ate at the base 6—15 cm. long, 2-5 cm. broad, subcoriaceous, glabrous above,
essentially glabrous or minutely and indefinitely puberulent beneath; petiole 1.0—
1.5 cm. long. Inflorescence terminal, the peduncle repeatedly dichotomous, densely
and minutely appressed-puberulent, ebracteate, about as long as to somewhat
shorter than the leaves, the flowers in capituliform clusters of several at the ends
of the ultimate branches. Staminate flowers pale cream: perianth campanulate,
densely appressed-puberulent without, the tube about 1.5 mm. long, about 2 mm.
broad at the orifice, glabrous within, the lobes ovate-suborbicular, very slightly
unequal, about 2 mm. long, widely spreading at anthesis; anthers about 0.5 mm.
long, the outer inserted somewhat below the middle of the perianth lobes, the inner
inserted just beneath the orifice of the perianth tube; pistillode about 0.75 mm.
long, the stigma almost attaining the insertion of the inner stamens, the disc about
half as long as the ovary, cupuliform, crenate; pedicel about 2.5 mm. long. Pistil-
late flowers pale cream; perianth narrowly urceolate, minutely appressed-puberulent
without, the tube about 1.5 mm. long, about 1 mm. broad at the orifice, the lobes
ovate-trigonal, strongly unequal, the outer about 1 mm. long, slightly spreading;
pistil about 2 mm. long, the ovary ovoid, glabrous, the style short and stout, the
stigma capitate, abundantly papillate; staminodia 8, about 0.5 mm. long; pedicel
about 2 mm. long. Drupe ovoid, about 6 mm. long, with a very thin pulp, white,
essentially sessile.

dar Costa Rica and Panama, in highland woodland 60—1000 m. elev.

between Las Margaritas and EI Valle, ME ru Allen & Seibert 1281, 1764;
El Valle de Antón and vicinity, Seibert 416, 444. HERRERA: Pesé, Allen 795.

Daphnopsis seibertii is quite distinct from * other species of northern Central
America and Mexico, and is most closely allied to D. caribaea Griseb. of Puerto Rico
and the Lesser Antilles.

2. SCHOENOBIBLUS Mart. & Zucc.

SCHOENOBIBLUS Mart. & Zucc. Nov. Gen. & Sp. 1:65. 1824.

Shrubs or small trees. Leaves alternate, simple, exstipulate, membranaceous to
coriaceous. Inflorescence terminal, paniculiform, the ultimate branches umbelli-
orm. Flowers apetalous, dioecious, tetramerous; perianth rotate, more or less
petalaceous, usually densely puberulent, the limb deeply 4-lobed. Staminate flow-
ers: stamens 4, widely exserted upon elongate filaments, antesepalous, inserted at

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LVol. 45
96 ANNALS OF THE MISSOURI BOTANICAL GARDEN

\
LEE
PIE
|

Y

Y

Fig. 30. Schoenobiblus panamensis

the base of the perianth lobes; pistillode minute, stipitate, subtended by a minute,
usually eccentric disc. Pistillate flowers (unknown from Panama): somewhat
smaller than the staminate; staminodes not reported; pistil 1-carpellate, containing
a single pendulous anatropous ovule; stigma capitate and obscurely 2-lobed, elevated
upon a short stout style, exserted, subtended by an inconspicuous disc. Fruit a
slightly fleshy drupe. :

Perhaps half a dozen poorly defined species of northern South America, with
one species each in Trinidad and Panama. The few specimens collected almost
invariably have been staminate, and structural details of the pistillate flowers are

own almost solely from the description of S. ellipticus Pilger in which, however,
reference to staminodia is not made, intentionally or unintentionally. It is un-
fortunate that Pilger was not more explicit in this respect.

1. SCHOENOBIBLUS PANAMENSIS Standl. & L. O. Wms., in Ceiba 3:33. 1952.

Small dioecious trees 2-3 m. tall, the branches slender, essentially dichotomous,
glabrous or glabrate, grayish brown. Leaves alternate, the blade subsessile,

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1958]
FLORA OF PANAMA (Lythraceae) 97

lanceolate, narrowly subcaudate-acuminate, narrowly cuneate at the base, 12-20
cm. long, 3.5—5.0 cm. broad, rather thinly membranaceous, prominently venose,
glabrous. Staminate inflorescences terminal and in the upper leaf axils, paniculi-
form with few umbellate branches, the secondary peduncles 2.0—2.5 cm. long,
minutely appressed-puberulent, bearing about 8—12 small pink and white flowers,
the pedicels very slender, about 1.7 cm. long, minutely appressed-puberulent.
Flowers densely appressed-puberulent without, rotate, the tube about 1 mm. long
and broad at the orifice, pink, the lobes oblong-elliptic, spreading, white, about
4 mm. long and 1 mm. broad; stamens exserted, the filaments about 2 mm. long,
the lower half concrescent to the base of the perianth lobes, the anthers about 1
mm. long; pistillode minutely fusiform, somewhat less than 1 mm. long, sparsely
pilosulose, the disc flabelliform, eccentric, about half as long as the pistillode.
Pistillate inflorescences and flowers, and fruit unknown.

Panama, in lowland forests.

BOCAS DEL TORO: vicinity of Chiriqui Lagoon, Von Wedel 410.

The natural affinities of this plant, the specimen of which is rather fragmentary,
are quite obscure. It seems to be most closely related to S. peruviana of the upper
Amazon of Peru which, however, has more copious inflorescences and flowers nearly
twice the size of those of S. panamensis.

LYTHRACEAE

By LORIN I. NEVLING, JR.

Herbs, shrubs, or trees. Leaves opposite, whorled or very rarely alternate,
simple, entire, pinnately veined, sessile or petiolate; stipules small and caducous or
absent. Flowers dichlamydeous, perfect, sometimes dimorphic or trimorphic, in
l- to several-flowered cymes gathered into lateral or terminal clusters, perigynous,
actinomorphic or zygomorphic. Hypanthium free from the ovary, tubular,
urceolate or campanulate, usually appendiculate between the calyx lobes. Petals
inserted near the orifice of the hypanthium, usually of the same number as the
calyx lobes or reduced, generally spatulate, crumpled in aestivation. Stamens
usually twice the number of calyx lobes or less, rarely more, sometimes hetero-
morphic, occasionally alternately unequal, posterior stamen sometimes reduced to a
hypogynous gland; anthers included or exserted, generally versatile, dehiscing
longitudinally. Ovary superior, sessile or stipitate, sometimes heteromorphic,
generally bicarpellate, the placentation axile to free-central; style simple, sometimes
heteromorphic, elongate or short; stigma capitate or bilobed. Fruit a dehiscent or
indehiscent capsule; seeds sometimes winged, without endosperm, the embryo
straight,

3 A family of 22 genera and about 500 species. Cosmopolitan but most abundant
in tropical America.

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[Vol. 45
98 ANNALS OF THE MISSOURI BOTANICAL GARDEN

a. Leaves and flowers without prominent black punctate glands; herbs,

shrubs or trees.

b. Flowers relatively small, delicate, 3- to 6-merous; fruit a small mem-

branaceous capsule; seeds wingless; herbs or shrubs.

FI s3 5-merous, actinomorphic or rarely obscurely zygo-

morphic, axillary; hypanthium turbinate or campanulate, about as

long as broad, not gibbous or spurred; stamens inserted at one level;
h

rbs.
d. Flowers usually solitary; leaves obtuse to attenuate at the base;

capsule septicidally dehiscent
dd. Flowers usually in clusters of 3, rarely 1 or 5; leaves auriculate
at the base; capsule dehiscing irregularly. 2. AMMANNIA

1. ROTALA

s, medianl
thium tubular, longer than broad, often gibbous or spurred at the
base; stamens inserted at two levels; herbs or small shrubs... 3. CUPHEA
bb. Flowers massive, fleshy, 12- to 16-merous; fruit a large woody capsule; i
seeds winged; trees 4. LAFOENSIA
aa. Leaves and flowers with prominent black punctate glands; shrubs or
small trees............. =

5. ADENARIA

Members of two Asian genera have been introduced into Panama and are
sometimes found as escapes. These genera are Lawsonia and Lagerstroemia. Law-
sonia inermis L. is cultivated for ornament and because of the henna dye which is
extracted from the leaves. Lagerstroemia indica L. and Lagerstroemia speciosa
Pers., known as Crepe Myrtle, are widely cultivated as ornamentals because of their
showy pink, purple or white flowers.

1. ROTALA L.
ROTALA L. Mant. 175. 1771.
Suffrenia Bellardi, in Act. Taurin. 7:445. 1794.
inia Raf. Aut. Bot. 9. 1817, non Nutt.
Winterlia Spreng. Syst. 1:519, 788. 1825.
Ameletia DC. in Mém. Soc. Phys. Genève 32:82. 1826.
Nimmoia Wight, in Madr. Journ. Sci. 5:311. 1837.

„2: t. 1007. 1843,
Hypobrichia Benth. in Griseb. Cat. Pl. Cub. 108. 1886.

Aquatic or palustrine herbs, glabrous throughout, simple or sparsely branched.
Leaves opposite or verticillate, sessile. Inflorescence of 1-flowered axillary cymes.
Flowers usually 3- or 4-merous, bisexual, actinomorphic. Hypanthium appendic-
ulate or exappendiculate, generally urceolate, membranaceous, the calyx lobes
generally equal, valvate. Petals as many as the calyx lobes or absent. Stamens as
many as the calyx lobes or fewer, equal, included, the anthers ovoid or oblong-ovoid,
the filaments inserted about one-third from the base of the hypanthium. Ovary
3- or 4-carpellary, sessile or with a short gynophore, 1-locular, the placentation
free-central; ovules many; stigma capitate. Fruit a horizontally striate septici
capsule; style and hypanthium persistent; seeds minute.

About 38 species. America, Europe, Africa and Australia.

a. Plants palustrine; leaves and flowers decussate; hypanthium appendic-

ulate, 2.5—5.0 mm. long; petals Dent. i | 1. R. RAMOSIOR
aa. Plants aquatic or palustrine; at least the upper leaves and flowers ternate;

hypanthium exappendiculate, 0.5-1.0 mm. long; petals absent................. 2. R. MEXICANA

(98)

1958]

FLORA OF PANAMA (Lythraceae)

Fig. 31. Rotala mexicana

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99

[Vol. 45
100 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. ROTALA RAMOSIOR (L.) Kochne, in Mart FI. Bras. 132:194, £. 39, fig. I. 1877.

mmannia ramosior L. Spec. Pl. ed. 1. 1:120. 1753.
Ammannia bumilis Michx. Fl. Bor.-Amer. 1:99. 1803.
18

825.

mmannia catbolica Cham. & Schlechtd. in Linnaea 2:378; 1827.

Ammannia occidentalis DC. Prod. 3:78. 1828.

Ammannia monoflora Blanco, FI. Filip. ed. 1. 1:64. 1837.

Ammannia dentifera A. Gray, PI. Wright. 2:55. 1853.

Rotala ramosior var. interior Fernald & Griscom, in Rhodora 37:169, fab. 345, figs. I 6 2.
3

Relate ramonor var. dentifera (A. Gray) Lundell, in Bull. Torr. Bot. Club 69:395. 1942.

Erect palustrine herbs to 3.5 dm. tall; branches somewhat angular at least
toward the apex. Leaves elliptic or elliptic-oblanceolate, obtuse at the apex, cuneate
to attenuate-cuneate at the base, 0.7-3.7 cm. long, 0.1-0.8 cm. wide. Flowers
sessile, subtended by 2 linear bracts to 5 mm. long, 0.75 mm. wide. Hypanthium
urceolate, 2.75—3.0 mm. long, 1.5—2.0 mm. broad at the orifice; calyx lobes broadly
deltoid. Petals 0.7—1.25 mm. long, 0.6-0.75 mm. wide, white. Stamens as many
as calyx lobes, minute, the anthers ovoid, about 0.3 mm. long, 0.3 mm. wide, the
filaments to 1 mm. long. Ovary globose, 1.5-2.0 mm. long, sessile; style 0.2-0.5
mm. long. Seeds orbicular, about 0.25 mm. in diameter.

The United States, Central America, Antilles, Ecuador, Brazil, Philippine
Islands. ;

CANAL ZONE: along the old Las Cruces Trail, between Fort Clayton and Corozal,
Standley 29053. PANAMÁ: near Matias Hernández, Standley 28936, 28987; between
Matias Hernandez and Juan Diaz, Standley 31996, 32013; Juan Diaz, Standley 30512,
Killip 3273.

2. ROTALA MEXICANA Cham. & Schlechtd. in Linnaea 5:567. 1830.

Rotala mexicana subsp. typica var. a forma media Koehne, loc. cit. 1877.
Rotala mexicana subsp. ty pica var. a forma major Koehne, loc. cit. 1877.
otala mexicana subsp. tybica var. spruceana (Griseb.) Koehne, loc. cit. 1877.
Rotala mexicana subsp. tybica Koehne, in Engl. Bot. Jahrb. 1:151. 1880.
Rotala mexicana su sp. bierniana Koehne, loc. cit. 1880.
Rotala mexicana subsp. by pica var. chamissoana Koehne, in Engl. Pflanzenr. 4:30. 1903.
Rotala mexicana subsp. pusilla (Tul.) Koehne, loc. cit. 1903.

Creeping aquatic or palustrine herbs to 3 dm. tall, the branches obscurely
angular. Leaves ternate, the submerged leaves linear, 3-18 mm. long, 0.5-3.0 mm.
wide, the emersed leaves 3-8 mm. long, 1-2 mm. wide, both obtuse or retusé
cuneate-attenuate at the base. Flowers sessile, subtended by 2 setose bracteoles.

ypanthium urceolate, 0.6-1.0 mm. long, 0.5-0.6 mm. broad at the orifice; calyx

(100)

1958]
FLORA OF PANAMA (Lythraceae) 101

lobes narrowly deltoid, sometimes unequal. Petals absent. Stamens commonly 2,
the anthers oblong-ovoid, about 0.1 mm. long, 0.25 mm. wide, the filaments about
0.4 mm. long. Ovary globose, abruptly narrowed into a short gynophore, 0.4 mm.
long; style about 0.1 mm. long. Seeds suborbicular, about 0.3 mm. long, 0.25
mm. wide.

Central and South America, Asia, Africa and Australia. Found at low eleva-
tions in periodic streams and potholes.

CANAL ZONE: along the old Las Cruces Trail, between Fort Clayton and Corozal,
Standley 29222; Darićn Station, Standley 31539. cHIRIQUI: lower portion of valley and
marshes along Rio Antén, El Valle de Antén, about 500 m., Hunter & Allen 366. PANAMA:
near Matias Hernández, Standley 28080; near the big swamp east of the Río Tecumen,
Standley 26715; vicinity o Juan Franco Race Track, near Panama, Standley 27817; along
road between Panamá and Chepo, Dodge, Hunter, Steyermark, t$ Allen 16721; Las Sabanas,
Bro. Heriberto 151; San José Island, Perlas Archipelago, East Harbor, Johnston 830, Erlanson
450.

Since this species is often found in periodic waters, two habits are found: an
emersed and a submerged. Rotala mexicana flowers when the plants become
emersed. Considerable difficulty is encountered in attempting to identify the
submerged specimens. Identification, by means of vegetative characters only, is
best accomplished by use of the obtuse or retuse leaf apex character.

2. AMMANNIA L.
AMMANNIA L. Sp. Pl. ed. 1. 1:119. 1753.

Cornelia Ard. Animadv. Specim. 2:9. 1764
Crybłotbeca Blume, in Bijdr. 1128, 1129. 1826
Ditbeca Miq. nd. Bat. 1:61

Diplostemon Miq. loc. cit. 1855.

Hapalocarpum Mi 1855

iq. loc. cit. ;
Ammannella Miq. loc. cit. 618. 1855.

Annual herbs, simple or sparsely branched, the young branches obscurely
tetragonal. Leaves opposite, sessile. Inflorescence of (1- to) several-flowered
Cymes gathered into axillary clusters; flowers (5-, 6- or) 4-merous, bisexual,
actinomorphic. Hypanthium urceolate, appendiculate or exappendiculate, the
calyx lobes valvate. Petals present or absent. Stamens of the same or twice the
number of calyx lobes or less, inserted about a third from the base of the hypan-
thium. Ovary generally 4-carpellary, superior, sessile, the placentation generally
axile; ovules many; style sometimes obsolete; stigma capitate. Fruit an irregularly
dehiscent ovoid capsule, the hypanthium and style persistent; seeds many, minute.

About 20 species. America, Europe, Asia, Africa, Hawaiian and Mariana
Islands. Only one indigenous and one introduced species in Panama.

1. A. COCCINEA

a. Petals present; hypanthium appendiculate; stamens exserted
2. A. LATIFOLIA

aa. Petals absent; hypanthium exappendiculate; stamens included.........-...-.--

l. AMMANNIA COCCINEA Rottb. in Pl. Hort. Univ. Havn. Programm. 7. 1773.

Ammannia ramosior acc. to L. Mant. 2:332. 1771, not L. Spec. PI. ed. 1.
mmannia purpurea Lam. Encl. 1:131. 1783.

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[Vol. 45
102 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ammannia sanguinolenta Swartz, Fl. Ind. Occ. 1:272. 1797.
Ammannia octandra acc. to Cham. & Schlechtd. in Linnaea 2:376. 1827, not L.
Ammannia stylosa Fisch. & Mey. in Ind. Sem. Hort. Petrop. 7:41. 1841.
nia A i
Ammannia texana Scheele, in Linnaea 21:588. 1846.
Ammannia sanguinolenta subsp. purpurea Koehne, in Mart. FI. Bras. 13?:207, t. 40, f. 4.

1877.
Ammannia sanguinolenta subsp. robusta Koehne, loc. cit. 208, £. 40, f. 4b. 1877.
Ammannia sanguinolenta subsp. longifolia Koehne, loc. cit. 208. 1877.

Ammannia coccinea subsp. robusta (Heer & Reg.) Koehne in Engl. Bot. Jahrb. 1:250.

1880.

Ammannia coccinea subsp. longifolia Koehne, loc. cit. 1880.
Ammannia pubiflora Sosn. in Bull. Mus. Cauc., Tiflis, 8:165. 1915.

Erect slender herbs to 4 dm. tall, glabrous throughout, the stems angular. Leaves
sessile, narrowly oblong, acute at the apex, auriculate-amplexicaul at the base, 1.5-
6.5 cm. long, 0.2-1.3 cm. wide, the veins obscure. Inflorescence spiciform, leafy-
bracted, the flowers in subsessile 1- to 3-flowered minutely bracteate cymes. Hy-
panthium urceolate, exappendiculate, becoming globose in fruit, 2.5—3.5 mm. long,
about 2 mm. broad at the orifice, the calyx lobes generally unequal. Petals as many
as calyx lobes, about 1.5 mm. long, 1.25 mm. wide, violet. Stamens exserted, the
filaments glabrous, the anthers about 0.5 mm. long. Ovary ovoid, about 1.5-2.5
mm. long, 1.5 mm. in diameter, glabrous; style 1.5—2.5 mm. long.

The United States, Central and South America, Antilles; Hawaiian, Mariana,
and Philippine Islands. Believed introduced into Panama in hay from Texas.

PANAMA: San José Island, Perlas Archipelago, Johnston 12 39.

2. AMMANNIA LATIFOLIA L. Spec. PI. ed. 1. 1:119. 1753.

Ammannia lytbrifolia Salisb. Prodr. 65. 1796.
Isnardia subbastata Ruiz & Pav. Fl. Peruv. 1:66, t. 86. 1798.

mmannia sagittata DC. Prodr. 3:80. 1828.
Ammannia bastata DC. loc. cit. 78. 1828.
Ammannia bumilis var. 8 Torr. & Gray, in FI. N. Amer. 1:480. 1838-1840.
Ammannia catholica Hook. & Arn. Bot. Beechey's Voy. 289. 1841, acc. Seemann.
Ammannia lingulata Griseb. Cat. PI. Cub. 106. 1866.
Ludwigia scabriuscula Kellogg, in Proc. Calif. Acad. 7 :78. 1876.

Erect slender herbs to 4.5 dm. tall, glabrous throughout. Leaves sessile, nar-
rowly oblong-lanceolate, acute at the apex, auriculate-amplexicaul at the base, 3.07
8.5 cm. long, 0.4-1.3 cm. wide, the primary lateral veins and submarginal vein
obscure. Inflorescence spiciform, leafy-bracted, intercalary, the flowers in rela-
tively distant, subsessile, 1- to 3-flowered minutely bracteolate cymes. Hypanthium
urceolate, appendiculate, becoming globose in fruit, 2.4-4.0 mm. long, 1-2 mm.
broad at the orifice, the calyx lobes broadly deltoid. Petals absent. Stamens
included, the filaments glabrous, the anthers about 0.25 mm. long. Ovary ovoid,
1.5-3.5 mm. long, 1.0—2.5 mm. in diameter, glabrous; style about 0.5 mm. long.

Southern United States, Central and South America, Antilles.

PANAMÁ: Bella Vista, at sea level, Killip 12031.

(102)

103

1958]
FLORA OF PANAMA (Lytbraceae)

A

os
\

Ammannia latifolia

(103)

Fig. 32.

[Vol. 45
104 ANNALS OF THE MISSOURI BOTANICAL GARDEN

3. CUPHEA P. Br.
CUPHEA P. Br. Nat. Hist. Jam. 216. 1756.
Melanium P. Br. loc. cit. 215. 1756.
Parsonsia P. Br. loc. cit. 199. 1756.
Balsamona Vell. in Vandelli, Fasc. DI 151271.
u 62

129%:
Melvilla Anders. in Journ. Arts & Sci. 25:207. 1807.
Bergenia Raf. Sylv. Tellur. 102. 1838.
j it. 1838.

Melfona Raf. lue. de 1838.
Maja Klotzsch, in Schomb. Fl. Fauna Guian. 1191. 1848.

Annual or perennial herbs or shrubs, generally branched, sometimes rhizomatous.
Leaves opposite or approximate, sessile or petiolate; the stipules setose. Inflores-
cence of 1- to several-flowered interpetiolar cymes gathered into terminal bracteate
or leafy paniculiform or racemiform clusters. Flowers 6-merous, bisexual,
medianly zygomorphic. Hypanthium tubular, appendiculate, more or less gibbous,
sometimes spurred, ribbed, occasionally colored, the calyx lobes equal or rarely
unequal, valvate. Petals spatulate, 6 or rarely reduced to the posterior 2, inserted
at the orifice of the hypanthium, red to violet. Stamens 1 1, the 2 posterior inserted
lower in the hypanthium than the 9 anterior, included, the 9 anterior equal or when
unequal the antesepalous stamens longer than the antepetalous, included or exserted,
the posterior antesepalous stamen modified into a basal hypogynous, unilateral,
Ovate to ovate-reniform or more rarely strongly involute-cupuliform disc, the
anthers oblong, the filaments glabrous or pubescent. Ovary bicarpellary, superior,
sessile, bilocular toward the base, unilocular above, the placentation incompletely
axile; ovules few to many; style slender; stigma capitate. Fruit a unilaterally
loculicidal capsule enclosed within the persistent hypanthium which also is ruptured
by the reflexed protuberant placenta. Seeds lenticular, sometimes marginate.

A very perplexing genus. The United States, Central and South America,
Galapagos and Hawaiian Islands.

a. Hypanthium at anthesis 4-10 mm. lone.
b. Flowers ebracteolate; disc cupuliform; seeds more than 20................. 1. C. UTRICULOSA
bb. Flowers bracteolate; disc unilateral; seeds less i
C= Gy there - terminal bracteate racemiform or paniculi-
el

; disc A
d. Style and sema included... a 2. C. SETOSA
dd. Style and at least the antesepalous stamens greatly exserted......... C. EPILOBIFOLIA
cc. Cymes gathered into terminal foliate racemiform or paniculiform
clusters; disc erect to horizontal.
e. Hypanthium not spurred, merely gibbous; seeds emarginate;
plants often rhizomatous... aa a . C. CALOPHYLLA

O MUT UN e 5. C. CARTHAGENENSIS
INFUNDIBULUM

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1958]
FLORA OF PANAMA (Lytbraceae) 105

Cuphea rotundifolia Koehne, resembling C. calophylla but with more rotundate
leaves, has been reported from Panama but I have not seen specimens of this species
from that area.

1. CUPHEA UTRICULOSA Koehne, in Mart. Fl. Bras. 132:452. 1877.

Cupbea gracilis acc. to Seemann, Bot. Voy. Herald, 121. 1852—57, not Benth.
Cuphea panamensis Hemsl. Diagn. Pl. Nov. Mex. 3:52. 1880.

Cupbea utriculosa var. panamensis (Hemsl.) Koehne, in Engl. Bot. Jahrb. 1:452. 1881.
Cupbea utriculosa var. donnell-smitbii Koehne, in Bot. Gaz. 18:203. 1893.

Cuphea utriculosa var. a forma communis Koehne, in Engl. Pflanzenr. 4:108. 1903.

Cuphea utriculosa var. a forma ciliifolia Koehne, loc. cit. 1903.

Parsonsia utriculosa Standl. in Contrib. U. S. Natl. Herb. 23:1017. 1924.

Erect slender herbs to 4 dm. tall, sometimes subligneous near the base; branches
with minutely tomentose lines decurrent from the paired nodes above. Leaves
oblong-elliptic, acute at the apex, cuneate-attenuate at the base, 0.4—6.0 cm. long,
0.2-1.1 cm. wide, sessile or subsessile, both surfaces glabrous, the primary lateral
veins inconspicuous, the margin minutely ciliate. Flowers decussate, solitary;
pedicels 4-13 mm. long, bracteoles absent. Hypanthium subampullaceous, 5.5—
8.0 mm. long, median diameter 1.5—2.5 mm., transverse at the orifice, gibbous at
the base, minutely puberulent, the posterior calyx lobe enlarged. Petals 6, the 2
posterior 2.5—4.5 mm. long, 1.5-2.0 mm. wide, the 2 anterior 3.0—5.5 mm. long,
1.5-2.5 mm. wide; violet. Stamens 11, the anterior 9 equal, included, the lower
half of the filaments pubescent; disc cupuliform, split anteriorly. Ovary broadly
ovoid, 2.5-3.25 mm. long, villous; style 1-2 mm. long, villous, included. Seeds
24-36, suborbicular, around 1 mm. ong, 0.75 mm. wide, emarginate.

Southern Mexico to Panama in moist habitats.

mark & Allen 17233; drowned forest of Quebrada Ancha, 70 m., Steyermark & Allen s. n.;
Rio Paraíso above East Paraiso, Standley 29877. cocLÉ: vicinity of Ola, alt. 100-350 m.,
Pittier 5032; Bismarck, 2000-3000 ft. elev., above Penonomé, Williams 264. PANAMÁ:
Río La Maestra, 0-25 m., Allen 30.
2. CUPHEA sETOsA Koehne, in Mart. Fl. Bras. 132:223. 1877.
Cupbea rigidula acc. to Seemann, Bot. Voy. Herald, 121. 1852-57, not Benth.
Cuphea setosa var. a forma seemannii Koehne, in Engl. Bot. Jahrb. 1:456. 1881.
Cuphea setosa var. glabrescens Koehne, loc. cit. 1881
Erect herbs to 3.8 dm. tall, subligneous at least near the base, the branches with

brown hispid recurved hairs in lines decurrent from the paired nodes above, area

tween lines grayish-strigose. Leaves elliptic to ovate-elliptic, acute at the apex,
attenuate at the base, 3-9 cm. long, 1-3 cm. wide, both surfaces strigose with
interspersed setose hairs, the primary lateral veins arcuate, almost forming a sub-
marginal vein, the margin ciliate; stipules setose. Cymes gathered into terminal,
bracteate, racemiform, rarely paniculiform, clusters. Flowers decussate, solitary,
infra-interbracteal; bracts elliptic, to 2.5 mm. long, 1 mm. wide, coarsely ciliate,

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[Vol. 45
106 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 33. Cupbea utriculosa

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1958]
FLORA OF PANAMA (Lytbraceae) 107

caducous; peduncle 0.5-1.0 mm. long; pedicel 1.25-2.0 mm. long; bracteoles 2,
minute, sometimes caducous. Hypanthium ampullaceous, 5.0—6.5 mm. long, median
diameter about 1.5 mm., oblique at orifice, spur straight, hispid without, tomentose
within, sometimes insect-infested and becoming swollen and bladdery. Petals 6,
1.5-2.0 mm. long, 0.75—1.0 mm. wide, violet. Stamens 11, the anterior 9 alter-
nately unequal, included, the anthers oblong, to 0.5 mm. long, the filaments
tomentose with the exception of the 3 anterior antesepalous; disc unilateral,
strongly reflexed. Ovary narrowly ovoid, 1.0—1.25 mm. long, 0.5—0.75 mm. in
diameter, tomentose; style 1.5—2.5 mm. long, tomentose, included. Seeds 4, almost
cordiform, 1.5 mm. long, 1.5 mm. wide, emarginate,

Central America, Colombia, Peru, Bolivia.

CANAL ZONE: forest between Peluca Hydrographic Station and Quebrada Peluca, along
Rio Boquerón, 70— m., Steyermark & Allen 17258.

3. CUPHEA EPILOBIFOLIA Koehne, in Mart. Fl. Bras. 132:223. 1877.

Cuphea epilobifolia var. costaricensis Koehne, in Engl. Bot. Jahrb. 1:457. 1881.

Cupbea ebilobifolia var. costaricensis forma canescens Koehne, in Bot. Gaz. 20:4. 1895.

Cupbea ebilobifolia var. costaricensis forma tonduzii Koehne, in Engl. Bot. Jahrb. 39:156.
1900.

Cupbea epilobifolia var. venezuelana Koehne, in Engl. Pflanzenr. 4:112. 1903.

4 forma lindenii Koehne, loc. cit. 1903.

Pte sporto var. costaricensis forma boffmannii Koehne, in Engl. Pflanzenr. 4:112.
Cuphea ebilobifolia var. costaricensis forma endresii Kochne, loc. cit. 113. 1903.

Cuphea epilobifolia var. caquetae Sprague, in Ann. Bot. 17:161, £. II, f. 1-6. 1903.
Parsonsia ebilobifolia Standl. in Contrib. U. S. Natl. Herb. 23:1018. 1924.

Woody shrubs to 2 m. tall, apical portions of branches densely wooly or at least
with tomentose lines decurrent from the paired nodes above. Leaves elliptic, sessile
or subsessile, acute at the apex, attenuate at the base, 3-14 cm. long, 0.8-3.0 cm.
wide, both surfaces puberulent to strigose, the costa glabrous to brown-strigose
beneath, the submarginal vein sometimes conspicuous, sometimes bearing brown
strigose hairs, the margin entire; stipules somewhat persistent. Cymes gathered
into terminal bracteate racemiform or paniculiform clusters; flowers decussate,
infra-interbracteal; bracts lanceolate, to 4 mm. long, 1.5 mm. wide, puberulent,
coarsely ciliate, caducous; peduncle 0.75—1.0 mm. long; pedicel 1-5 mm. long;
bracteoles 2, minute. Hypanthium ampullaceous, 4.0-7.5 mm. long, median
diameter 0.7 5-1.5 mm., oblique at the orifice, the spur straight to recurved, densely
strigose or hirsute, sometimes insect-infested and becoming swollen and bladdery.
Petals 6, 1.25-2.0 mm. long, 0.25-1.25 mm. wide, pink to violet. Stamens 11,
the anterior 9 alternately unequal, antesepalous exserted, antepetalous included to
slightly exserted, the filaments villous except for the 3 anterior antesepalous, or

glabrous; disc unilateral, strongly reflexed. Ovary broadly ovoid, 1.5—2.5 mm.
long, villous; style 2.75-5.5 mm. long, villous, exserted. Seeds 4—6, suborbicular
to orbicular, 1.01.5 mm. long, 1.0-1.5 mm. wide, emarginate to slightly marginate.

Costa Rica, Panama, Venezuela.

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[Vol. 45
108 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Rowlee 379. cocLé: vicinity of El Valle, 800— , north rim, Allen 215; north

of El Valle de Antén, near Cerro Turega, alt. 650-700 m., Wo. & Schery 161; E
Valle de Antón, floor, vi ca Tomás Arias, alt. 600 m., Allen 3624; El Valle de
Antón, along Río Indio trail, 500— s Hunter & Allen ZONE: forest

9. AL
etween Peluca Hydrographic Station and Quebrada Peluca, along Rio Boquerón, 70- m.,
Steyermark & Allen 17236.

4. CUPHEA CALOPHYLLA Cham. & Schlechtd. in Linnaea 2:361. 1827.

Cupbea plumbaginea Mart. in Flora Beibl. 212:62. 1838.

Cupbea antisypbilitica acc. to Seemann, Bot. Voy. Herald, 121. 1852-57, not HBK.

Cuphea orthodisca Koehne, in Mart. Fl. Bras. 132:224. 1877.

Cuphea microstyla Koehne, loc. cit. 1877.

Cuphea calophylla var. calophylla Koehne, in Engl. Bot. Jahrb. 2:138. 1881.

Cuphea calophylla var. calophylla forma deformis Koehne, loc. cit. 1881.

Cuphea calophylla var. orthodisca Koehne, loc. cit. 18

Cuphea calophylla var. microstyla Koehne, loc. cit. 139. 1881.

Cuphea calophylla var. calophylla forma breuningii Koehne, in Engl. Bot. Jahrb. 29:156.
00

1900.
Cuphea calophylla var. calophylla forma blumbaginea (Mart.) Koehne, in Engl. Pflanzenr.
4:116. 1903

Parsonsia calophylla Standl. in Contrib. U. S. Natl. Herb. 23:1018. 1924.

Erect or sprawling herbs to 5.5 dm. tall, rhizomatous, the branches wooly
interspersed with occasional large red hairs, scabrous. Leaves lanceolate to elliptic-
lanceolate, acute at the apex, obtuse at the base, 0.8—4.0 cm. long, 0.1-2.0 cm.
wide, reduced toward the apex, above glabrous to puberulous and strigose, beneath
strigose and scabrous, margin minutely ciliate, primary lateral veins slightly arcuate;
petiole 1-5 mm. long. Cymes gathered into terminal foliate racemiform or panicu-
liform clusters, the flowers alternately interpetiolar, solitary; peduncle 3.5-7.0 mm.
long; pedicel about 0.25 mm. long; bracteoles 2, minute. Hypanthium sub-
ampullaceous, 4-7 mm. long, median diameter about 1.25 mm., transverse at the
orifice, gibbous at the base, white puberulent with interspersed red hairs. Petals 6,
2.5—3.0 mm. long, 1.5—2.0 mm. wide, violet. Stamens 11, the anterior 9 alter-
nately unequal, included, the filaments densely villous; disc unilateral, not reflexed.
Ovary broadly ovoid, 1.5—3.0 mm. long, greatly elongating in fruit, glabrous;
style 1.0-1.5 mm. long, glabrous or villous, included, becoming exserted in fruit.
Seeds 5-9, suborbicular, 1.0—1.8 mm. long, 1.0—1.5 mm. wide, emarginate.

Central America, Brazil.

CANAL ZONE: Chagres, Fendler III, 223; between Chagres Batteries and Fort San
Lorenzo, Fort Sherman Military Reservation, Maxon t$ Valentine 7025; rocky banks of
Chagres at El Vigía, Pittier 2376, 2377; drowned forest of Quebrada below Río Indio
Hydrographic Station, 70 m., Steyermark 17396; forest between Peluca Hydrographic
Station and rada Peluca, along Río uerón, 70— m., Steyermark & Allen 172315
around Gamboa, alt. 20—1 da, m., Pittier 4794; along the old Las Cruces Trail, between Fo

ey

26; Cerro Ancón, Bro. Heriberto 127; Mount Hope Cemetery, Standley 28764; Juan VE
È: i elev., William

750 ft., Allen 4611. PANAMÁ: forests of the upper Miibòni River, alt. 150-400 m»
Pittier 4481; Orange River Valley, vicinity of Juan Diaz, Killip 3330.

|
|

1958]
FLORA OF PANAMA (Lytbraceae) 109

5. CUPHEA CARTHAGENENSIS (Jacq.) Macbr. in Field Mus. Publ. Bot. 8:124.
1930

Lytbrum cartbagenensis Jacq. Stirp. Amer. Hist. 148. 1763.
Balsamona pinto Vand. Fasc. pl. 15. 1771.

Cuphea balsamona Cham. & Schlechtd. in Linnaea 2:363. 1827.
Cuphea elliptica var. a Koehne, in Engl. Bot. Jahrb. 2:145. 1881.
Parsonsia binto Heller, in Minn. Bot. Stud. 9:862. 1897

Erect herbs to 4.5 dm. tall, sometimes subligneous near the base, simple to
alternately branched, the branches generally glandular-hirsute with interspersed
white tomentose hairs becoming hispid toward the apex. Leaves lanceolate to ovate
or elliptic, acute at the apex, cuneate or attenuate at the base, 1.0—5.5 cm. long,
0.4-2.1 cm. wide, both surfaces strigose and scabrous, upper surface sometimes also
puberulent, the margin ciliate, the primary lateral veins arcuate; petiole 0.1—0.3
mm. long; stipules minute. Cymes gathered into terminal foliate racemiform or
paniculiform clusters, the leaves subtending the flowers gradually reduced above,
setose-ciliate, both surfaces strigose and scabrous. Flowers interpetiolar, generally
solitary; peduncle 0.5—1.2 mm. long; pedicel 0.25-0.75 mm. long; bracteoles 2,
minute. Hypanthium subampullaceous or ampullaceous, 2.5-7.0 mm. long, the
median diameter around 1.25 mm., transverse at the orifice, spur straight, with
glandular hirsute pubescence generally restricted to the ribs, sometimes shortly
villous toward the base. Petals 6, 1.25—2.0 mm. long, about 0.5 mm. wide, violet
to purple. Stamens 11, the anterior 9 alternately unequal, included, the filaments
glabrous or villous; disc not reflexed. Ovary broadly ovoid, 2-3 mm. long,
glabrous; style 0.5-1.0 mm. long, included, glabrous. Seeds 3-7, orbicular to
elliptic, 1.0-2.5 mm. long, 0.75—2.0 mm. wide, marginate.

The United States, Central and South America; Antilles; Hawaii. The presence
of this species in Hawaii is probably the result of introduction.

BOCAS DEL TORO: Isla Colón, alt. 0—120 m., southwest of Bocas at Macaw Hills, Von
Wedel 525; Macaw Hill, swampland, Von Wedel 4; vicinity of Chiriqui Lagoon, Von Wedel
2404. AL ZONE: Gatún, Piper 5996; near Gatún, Standley 27316; Barro Colorado
Island, Bailey & Bailey 500, Aviles 60, Starry 272, Kenoyer 464, Duchassaing s. n.; be-
tween Las Cascadas and Bas Obispo, along the railroad, altitude 50 m., Pittier 3747; Old
Panamá, Riley 150, 151. cHIRIQUI: Boquete, el. 3800 ft., Davidson 577, 673; vicinity of

Río Tecumen, Standley BA Sabanas, Bro. Paul 51; Isla Taboga, ca. 0-186 m., Woodson,
Allen & Seibert 1433; San José Island, Perlas Archipelago, Johnston 988, Erlanson 499.

Examination of the type material of Cupbea elliptica reveals much disparity
between the specimens and the original description, but it does fall well within the
range of measurements of C. carthagenensis. I have been unable to find any char-
acter or combination of characters to distinguish C. elliptica var. a from C. cartha-
Senensis. The characters formerly employed can be shown to be unreliable. I feel
certain that only one species is represented.

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[Vol. 45
110 ANNALS OF THE MISSOURI BOTANICAL GARDEN

6. CUPHEA INFUNDIBULUM Koehne, in Mart. Fl. Bras. 132:236. 1877.
o appendiculata acc. to Seemann, in Bot. Voy. Herald, 121. 1852— ka not Benth.
Cupbea infundibulum var. foliosa Kochne, in Engl. Bot. Jahrb. 7:43. 18

Erect slender herbs to 2.5 m. tall, subligneous at least near "- base, branches
white-puberulent interspersed with yellow hispid hairs at least near the apex. Leaves
broadly elliptic, acuminate, obtuse and abruptly decurrent at the base, 3—13 cm.
long, 1.5—4.5 cm. wide, above puberulent and scabrous, beneath strigose, with a
well-developed submarginal vein, margin minutely ciliate; petiole to 5 (-12) mm.
long. Flowers in young plants solitary, interpetiolar, transitional in fully mature
plants to cymes gathered into terminal bracteate paniculiform clusters; bracts
elliptic, 2.5—5.0 mm. long, 1.0-2.5 mm. wide, both surfaces sericeous, caducous.
Peduncle 1—4 mm. long; pedicel 0.75—1.5 mm. long; bracteoles 2, minute. Hypan-
thium infundibuliform, 23-35 mm. long, median diameter 2.6—3.0 mm., transverse
at the orifice, at most slightly gibbous at the base, the lower third yellowish-green,
puberulent and hirsute, the upper two-thirds salmon-red, puberulent. Petals 2,
posterior, 2.5—3.25 mm. long, 1.0-1.75 mm. wide, red. Stamens 11, the anterior
9 alternately unequal, exserted, the filaments glabrous; disc unilateral, semi-
cupuliform at the base, not reflexed. Ovary very narrowly ovoid, 3-8 mm. long,
glabrous; style 2.3-3.0 mm. long, glabrous, exserted. Seeds 5-7, suborbicular,
about 2.5 mm. long, 2 mm. wide, emarginate.

Costa Rica, Panama.

m., Allen abd; trail from Paso Aa to Monte bios upper duy of Río aude Vigo,
alt. 1500-2000 m., Allen 1510; rocky plains about 5 miles south of Boquete, alt. 3000 ft.,
Allen 4710; Bajo Mon o, Boquete ae eis el. 4500 ft., Davidson 489; El Done E 1450
m., Killip 3552; vicinity of El Boquete, alt. 900-1200 m., Bro. Maurice 689; forests along
Río Ladrillo and vicinity, above E Badian, alt. 1200-1300 m., Maxon peius pastures
around E] Boquete, alt. 1000-1300 m., Piźtier 2883; be e Cer rro Vaca and Hato del
Loro, eastern Chiriqui, alt. 850-1100 M Pittier 5383; valley of the upper Rio Chiriquí
Viejo, vicinity of cvm Lirio, 1300—1900 m., Seibert 304; valley of the upper Río Gariché,
ip © AR m., Seibert 333; vicinity of Casita Alta, Volcán de Chiriquí, ca. 1500-20 000

Allen t$ Seibert 862; Finca Lérida to Boquete, ca. 1300-1700 m., Woodson,
Allen 8 3 Seibert 1133.

4. LAFOENSIA Vand.
LAFOENSIA Vand. Fl. Lusit. & Bras. Specim. 33. 1788.
Calyplectus Ruiz & Pav. Peruv. & Chil. Prodr. 73. 1794.

Small trees, the young branches more or less tetragonal, becoming terete with
age. Leaves opposite, petiolate, glandular, stipulate. Inflorescence of 1- flowered
cymes gathered into terminal leafy racemiform or rarely paniculiform clusters
Flowers usually 12- to 16-merous, bisexual, actinomorphic. Hypanthium generally
campanulate with alternate coriaceous teeth between the equal, conduplicate-
valvate calyx lobes, the appendages absent. Petals inserted slightly below the orifice
of the hypanthium, as many as the calyx lobes, spatulate, white to yellow. Stamens
generally twice the number of calyx lobes, exserted, the anthers oblong-linear, the

(110)

XU At ad e Lea, Cep ues HEA WRP OZ quodi tica

1958]

111

FLORA OF PANAMA (Lythraceae)

"AE R = d
R R SY

=

=

TT -

Sy PEA}
$ Yip LE. 2
EG GI

Fig. 34. Lafoensia punicifolia

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[Vol. 45
114 ANNALS OF THE MISSOURI BOTANICAL GARDEN

filaments united at the base into a fleshy perigynous cup continuous with the
hypanthium. Ovary with a gynophore, bicarpellate, unilocular, the placentation
free-central; ovules many; stigma capitate. Fruit an indehiscent capsule; see
winged.

About 10 species. Mexico to Brazil. Only 1 species indigenous to Panama.

1. LAFOENSIA PUNICIFOLIA DC. in Mem. Soc. Phys. Genève 32:86, £. I. 1828

(as bunicaefolia).

Small trees to 15 m. tall, glabrous throughout. Leaves elliptic to obovate,
obtusely subcaudate-acuminate at the apex, obtuse to rounded at the base, 5-9 cm.
long, 1.0—3.5 cm. wide, the gland at distal end of the costa opening on the lower
surface by means of a small pore, the submarginal vein well developed; petiole 5-6
mm. long. Peduncle 19-27 mm. long; pedicel 2.5—5.0 mm. long; bracteoles 2, 5-7
mm. long, 8—12 mm. wide, coriaceous, caducous. Hypanthium campanulate, 2.5-
4.0 cm. long, 2.5—3.5 cm. broad at the orifice, coriaceous except the more or less
deltoid calyx lobes. Petals inserted about 5 mm. below the orifice of the hypan-
thium, 2.7—3.0 cm. long, 0.8—1.5 cm. wide, sometimes erose. Stamens equal,
greatly exserted, the anthers 7-8 mm. long, the filaments 7-9 cm. long, inserted
at one level about one-third above the base of the hypanthium, contorted in bud,
red. Ovary turbinate, 5-10 mm. long, about 5 mm. in diameter, somewhat
abruptly narrowed into a cuneate gynophore 5-11 mm. long, glabrous; style to
10 cm. long, exserted. Fruit a hard thick-walled capsule to 6.5 cm. long, 4 cm.
in diameter. Seeds oblong-lanceolate, to 40 mm. long, 5-15 mm. wide; cotyledons
ovate.

Mexico, Guatemala, Panama, Venezuela, Colombia, Bolivia. A yellow dye is
extractable from the leaves of this species. The wood is used in cabinet work.
Known as amarillo in Panama.

ONE: Barro Colorado Island, Aviles 28b, Shattuck LR Ag 3616; Barro

Colorado ‘tae shore of cove S. of Fr. Lock site, Woodworth & tal 475. DARIEN:
al de Sta. Marta, South Darién, Pittier 6985; forests around ka ee southern Darićn,

Pittier 69854. PANAMA: along road between Panamá and Chepo, Dodge, Hunter, Steyer-
mark 6 Allen 16705; between ix. Sabanas and Marias Hernia. Standley 31907; per
near Chepo, 30 m., eg id Allen 24; near rs Kluge 18; Chepo, alt. about 60 m»

Pittier 4763; near the big swamp east of the Río Tecumen, Standley 26556, Eus Río

Tecumen, Standley 29437; jacy of Pacora, sit ca. 35 m., 1006.

5. ADENARIA HBK.

ADENARIA HBK. Nov. Gen. & Spec. 6:185, f. 549. 1823.

Shrubs or small trees, black punctate-glandular generally, the young branches
tetragonal, becoming terete with age. Leaves opposite, petiolate; stipules setose-
Cymes gathered into axillary umbelliform clusters. Flowers 4- or 5-merous, bi-
sexual, heterostylic, actinomorphic. Hypanthium campanulate, not ribbed, the
calyx lobes more or less equal, valvate; exappendiculate. Petals inserted at the
orifice of the hypanthium, as many as calyx lobes, rose or white. Stamens usually

(112)

"psc cen e Es
A UELUT lere BPO A RON

1958]

FLORA OF PANAMA (Lythraceae) "PD
A GF
EN ( ZEŃ
|

MIA

Fig. 35. Adenaria floribunda

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[Vol. 45
114 ANNALS OF THE MISSOURI BOTANICAL GARDEN

twice the number of calyx lobes, included or exserted, the filaments united at the
base to form a somewhat fleshy perigynous ring at the base of the hypanthium.
Ovary bicarpellary with a gynophore, bilocular, the placentation axile; ovules
many; stigma bilobed. Fruit an indehiscent ovoid capsule, the style and hypan-
thium persistent; seeds small.

Mexico to Argentina. Monotypic. The tree Adenaria floribunda HBK. has small
rose or white flowers which are highly aromatic. This fragrance makes it a highly
desirable addition to gardens; hence, it is sometimes cultivated.

1. ADENARIA FLORIBUNDA HBK. Nov. Gen. & Spec. 6:188. 1823.
Adenaria parjure a HBK. loc. cit. 185. 1823.

Adenaria lanceolata Beurling, i in Akad. Handi Stockh. 124. 18
Adenaria floribunda var. a forma purpurata (HBK.) Lei in Mart. Fl. Bras. 132:344.

Adenaria floribunda var a forma grisleoides (HBK.) Koehne, loc. cit

pes ria purpurata var. australis Grise b. in Lorentz, Veget. Noe ŚR 116, 135.
87%.

sia floribunda var. parviflora Koehne, in Engl. Bot. Jahrb. 4:406. ME

Adenaria floribunda var. micropbylla Koehne, in Engl. Pflanzenr. 4:247. 1903.

Erect shrubs or trees to 2.5 m. tall, young branches shortly ns and glan-
dular, the bark becoming shredded whitish-silver. Leaves lanceolate, acute to
acuminate at the apex, obtuse to obtusely attenuate at the base, 2.0—13.5 cm. long,
0.6-5.0 cm. wide, above dark green to purplish red, glabrous to minutely puber-
ulent, paler beneath, villous on veins, the primary lateral veins forming an indistinct
and sometimes interrupted submarginal vein; petiole 0.5—5.0 cm. long; stipules to
1.5 mm. long. Cymes gathered into axillary umbelliform clusters; peduncle 1-7
mm. long; bracts in two superposed pairs; pedicel 5-7 mm. long, occasionally
subtended by a minute bracteole. Flowers trimorphic: short-styled, intermediate-
styled and long-styled. Intermediate-styled flowers: hypanthium campanulate 2-4
mm. long, 3.25-4.0 mm. broad at the orifice, tomentose and glandular without,
tomentose within, the calyx lobes more or less deltoid; petals 3.5—4.0 mm. long,
1.0-1.5 mm. wide, glandular without, rose or white; stamens equal or occasionally
alternately unequal with antesepalous stamens one-third longer than the ante-
petalous, exserted, the anthers 0.5 mm. long, the filaments (when equal) 4.5-5.5
mm. long, glabrous, united at the base; ovary turbinately globose gradually nar-
rowed into a cuneate gynophore, 1.75—2.0 mm. long, glabrous or villous, upper |
glandular, the placentation axile; style 1.0-1.5 mm. long, glandular toward the
base. Short-styled like the intermediate-styled except: hypanthium 2.0—3.25 mm.
broad at the orifice; petals 2.0—2.75 mm. long, 0.5—1.25 mm. wide; stamens equal,
exserted, the filaments 3.0—4.25 mm. long, glabrous; ovary narrowly ovoid, 1.0-
1.25 mm. long, diameter 0.25—0.5 mm., upper half villous and glandular; sty
0.5—1.0 mm. long. Long-styled flowers like the intermediate-styled except:

(114)

1958]
FLORA OF PANAMA (Lecytbidaceae) 115

2.5 mm. long, 1.0-1.5 mm. wide; stamens equal, included, the filaments 1.5-2.5
mm. long; ovary turbinately globose, 1.5-2.5 mm. long, diameter 1-2 mm., upper
half villous and glandular, the gynophore reduced; style 1.5-2.0 mm. long. Fruit
an indehiscent ovoid capsule; seeds about 1 mm. long, 0.5 mm. wide.

Known as fruta de pavo in Panama.

. & Fort
Sherman Military Reservation, Maxon & Valentine 6995; Old Experiment Station, 3 miles

Ancón Hill, Standley 25155; Barro Colorado Island, Kenoyer 402,
Barro Colorado Island, Miller Trail, Shattuck 481. PANAMA: between Matias Hernández
and Juan Diaz, Standley 31970; roadside near Chepo, Dodge 10720.

This species has been reported as being rarely 3- or 6-merous. This is not
surprising in view of the variation between 4- and 5-merous flowers within the
same cluster. The panamanian material examined does not demonstrate the 3- or
6-merous condition.

Unusual stamen number is often found, that is, one more stamen than usual.
This is due to chorisis, generally in the antesepalous whorl.

LECYTHIDACEAE

Trees, usually massive and tall. Leaves alternate, exstipulate, simple, pinnately
veined, entire or somewhat incised, usually large and clustered toward the tips of
the branches. Inflorescence variously racemose or paniculate, usually few-flowered,
terminal or lateral, occasionally cauliflorous. Flowers perigynous to epigynous,
perfect, regular or zygomorphic, usually large and showy; perianth dichlamydeous
but the calyx frequently reduced to inconspicuous teeth or a low rim, the petals
4-12, rather fleshy; stamens numerous and in several centripetally decreasing series,
the filaments frequently more or less fleshy and petalaceous, united at the base into
a common androphore which may be radially symmetrical and wholly fertile or
strongly zygomorphic with a very prominent posterior, usually staminodial hood
involute over the low disc of the anterior fertile stamens, the anthers basifixed,
dehiscing longitudinally or rarely by terminal pores; pistil 2- to 6-carpellate, the
ovary inferior or subinferior, 2- to 6-celled, with 2 to numerous anatropous ovules
borne apically, laterally, or basally upon the axile placenta, the style usually very
short, the stigma radiate to capitate. Fruit very rarely a large 1-seeded berry with
thin subcoriaceous flesh, but usually a large woody or fibrous pyx, indehiscent or
dehiscent by a circumscissile operculum; seeds characteristically large and massive,
rarely compressed and samaroid, sometimes accompanied by a more or less arillate

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[Vol. 45
116 ANNALS OF THE MISSOURI BOTANICAL GARDEN

funicle and fetid pulp derived from the placenta and endocarp, exalbuminous, the
embryo differentiated or undifferentiated.

The Lecythidaceae (sensu stricto) are wholly American in distribution and are
particularly abundant in the Amazonian rain forest or hylaea, extending northward
in sharply decreasing numbers as far as British Honduras. Fifteen genera are
recognized by Knuth in his treatment of the family for Engler's ‘Das Pflanzenreich
(479?*:1-146. 1939). The trees include some of the tallest and most massive
of the forest and consequently are poorly collected. In Panama they occur mostly
as incidental specimens rather than in aggregated stands or groves.

Although the flowers are most attractive and curious, the thick-walled fruits
are even more striking, particularly those of Lecytbis, Eschweilera, and Couratari,
which resemble nothing so much as crude earthenware pots. The characteristically
large hard-shelled seeds represent the family the world over in the Brazil-nut of
commerce (Bertholletia excelsa HBK.).

As with several other intricate families of plants, the name of Lecythidaceae is
inseparably associated with that of John Miers, who as an English mining engineer
spent eighteen of his early years in Chile, Argentina and Brazil. During his resi-
dence in South America, quite without the benefit of special training in the subject,
without libraries or herbaria, botany became for Miers an avocation quite as
absorbing and exacting as his profession of mineralogy and metallurgy, and he pro-
ceeded to accumulate an incredible mass of original notes and sketches of various
critical plant families to the elaboration and publication of which he devoted the
remainder of his life in England until his death in 1879 at the age of ninety-one
years.

Thirty-six years after his return to Britain and at the age of eighty-five, Miers
published his memoir ‘On the Lecythidaceae’ (in Trans. Linn. Soc. 30?:157-318.
1874) which is the embodiment of his voluminous personal information discovered
in the field in South America and supplemented by subsequent studies in the
libraries and museums of Europe. Whatever criticism may be directed against the
occasional unreality of his taxonomic concepts, the fact remains that little has been
added to our biological knowledge of the family in intervening years.

The recent treatment of Knuth, to which reference already has been made, is an
improvement over that of Miers in that it includes analytical keys and the citation
of considerably more numerous specimens, as might be expected. It is a compila-
tion without much original insight, nevertheless, and depends in the final analysis
upon the first-hand knowledge of Miers. The keys, too, present a constant problem
since they are divided into geographic couplets for the most part and because of the
unfortunate fact that most species are represented by only one or a very few
herbarium sheets either in flower or in fruit but seldom both. The possibility of
replicate species therefore is great. Of chief aid in the study of the species of
Panama is the short study by Pittier (in Contr. U. S. Nat. Herb. 26:1-14. 1927)
on the Lecythidaceae of Central America which has the advantages as well, perhaps,
as some of the disadvantages of extensive field study.

(116)

1958]
FLORA OF PANAMA (Lecythidaceae) 117

a. Androphore radially symmetrical, wholly fertile.
b. Flowers large and showy; petals pw anthers dehiscing by apical
gad : bins

pores; Fruit a mediocre pyx containing few to several seeds, dehiscing
through deliquescence or irregular shredding of the operculum........... 1. GUSTAVIA
bb. Flowers small or mediocre; petals 4; anthers dehiscing 2
aji a oe coriaceous or fleshy indehiscent berry usually c
itary seed 2. GRIAS

nin soli
aa. Androphore strongly bilat erally —— produced into a prom-
inently sip”; te, usually stamin e posteri 5008,
b. Stamen pla wholly mi Wire celled; fruit a large
coriaceous indehiscent kocia the seeds vein, lateral, vedi
etid 3. COUROUPITA

bb. Stamen s of ri ite st kacie at least in par
c. Hood à imply involute, wholly stam inodial, the lower stamens
“iaia I gia 4-celled; fruit "Ade woody subglobose
dehiscent eeds several, di a es thick fe angular,
with gry e um. d f eshy funicles
cc. Hood spirally ipee - _—, staminodial; ovary 2-celled; fruit a
ni adr us depressed-campanulate dehiscent
pyx, the seeds s bud: Wien sess thick and angular
ccc. Hood spirally involute then nn revolute, wholly —
ovary 3-celled; fruit esta iaceous narrowly campanulate
dehiscent pyx, the MM sere ań very thin ind dl ‘winged 6. COURATARI

. LECYTHIS

>

5, ESCHWEILERA

1. GUSTAVIA L.

GUSTAVIA L. Pl. Surinam. 18. 1775, nom. conserv.
Japarandiba Aa Fam. 2:448. 1763, nom. rejic

Pirigara Aubl. Hist. PI. Guian. 1:487. 1775, pro parte.

Teichmeyeria sel: Introd. 267.

Spallanzania Neck. Elem. 2:79. 1790.

Trees, frequently of gigantic size, rarely shrubby, the leafy twigs ordinarily
stout. Leaves usually crowded toward the tips of the twigs, petiolate to sessile,
usually conspicuously serrate and large, occasionally rather small for the family
and entire or subentire. Flowers usually lateral and cauliflorous, occasionally
terminal or subterminal. Hypanthium broadly urceolate, smooth or indefinitely,
rarely conspicuously, winged or ribbed. Calyx manifest or reduced to a more or
less obscurely 4- to 6-lobed ring. Petals 6-12, rather unequal, large and showy.

drophore radially symmetrical, wholly fertile, the elongate inflexed fleshy fila-
ments of the innumerable stamens coherent at the base into a shallow perigynous
ring adnate to the base of the corolla; anthers narrowly cylindrical, dehiscent by
2 apical pores. Ovary 4- to 8-celled, containing numerous lateral anatropous
ovules on a stout axile placenta; style conic, the stigma 4- to 8- radiate. Fruit a
mediocre pyx containing few to several seeds, dehiscing through deliquescence or
irregular shredding of the operculum. Embryo differentiated, with fleshy coty-
ledons and minute hypocotyl and plumule.

About 40 to 50 species from Costa Rica to Peru and Brazil; one or two species
in the Lesser Antilles.

a. Calyx lobes clearly differentiated, E. to Te war to the hypanthium,

I eous, accrescent is ; fruit relatively small,

pe ruit
ongitudinally ribbed or vi ib dry and dehiscent through
Ceres of the operculum

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[Vol. 45
118 ANNALS OF THE MISSOURI BOTANICAL GARDEN

b. Flowers supported by a slender 2-bracteolate pedicel, conspicuously

ferruginous-tomentellous, the petals 1.5-2.0 cm. long; pyx turbinate,

y i broad as long, densely ferruginous-tomentellous;
leaves essentially entire 1. G. PLEUROCARPA

. Flowers without a proper pedicel, the hypanthium immediately sub-

tended by 2 small bracteoles, indefinitely ferruginous-papillate, the

l about as long as

c
c

petals 4.0—4.5 cm. long; pyx cylindric-campanulate,
broad, glabrous or essentially so; leaves rather obscurely undulate-
serrate.
aa. Calyx reduced to a more or less obscurely 4- to 6-lobed ring, obscure an
not accrescent in fruit; fruit relatively large, not clearly ri
winged, probably more or less pulpy and dehiscent through deliquescence
f the operculum.
b. Leaves mediocre, up to about 4 cm. long, with proportionally long
petioles, oblong-lanceolate; flowers about 6 cm. in diameter; pyx not
ut 5 mm. thick, the operculum large, as
broad as the body of the fruit 3. G. NANA
arge, up to about 10 dm. long, with proportionally short
petioles, elliptic-oblanceolate; flowers 10-15 cm. in diamete
t i 1

2. G. BRACHYCARPA

F
pt
3
$

r; pyx very
eshy, the pericarp about 5-10 mm. thick, the operculum small, about
one third as broad as the body of the fruit 4. G. SUPERBA

In the strict sense, Linnaeus” Gustavia would embrace only the last two species
of the key and Aublet's Pirigara the first two. It is indeed a great temptation to
separate the two groups generically, but it is resisted here for the very cogent fact
that we must qualify the description of the fruits by the adverb "probably."
Aublet himself, however, describes the fruit of Pirigara bexapetala (= Gustavia
hexapetala (Aubl.) Sm.), which would coincide with the former group, as “sèche”,
while Miers (loc. cit. p. 159) reports that in Gustavia, supposedly of the latter
group, "on the ripening of the fruit, the dissepiments and placentae become re-
solved into a pulp, which envelopes the seeds, all escaping together with tbe decay
of tbe operculum [italics ours]."

Whether the fruits differ or no, it is clear that such recent authors as Eyma (in
Fl. Surinam. 31:121. 1934) and Pittier (loc. cit. 2. 1927) err in characterising
the fruits as indehiscent without any qualification. It is impossible to understand
why Pittier, particularly, could actually key Gustavia upon such a statement im-
mediately before describing G. pleurocarpa Pitt. and G. brachycarpa Pitt., the type
collections of which, gathered by Pittier himself, include abundant dry, empty
pyxes with perfectly round opercular openings. I have also seen similar fruits with
very definite opercular openings upon specimens of G. bexapetala, as has my friend
Noel Y. Sandwith, of Kew, for G. laciniosa.

Insofar as floral characters distinguishing Pirigara and Gustavia are concerned,
other than the natures of the calyx, I have been able to observe only that
anthers of Gustavia are proportionally more elongate and deeply sulcate than those
of Pirigara. The flowers themselves of Gustavia, however, are most usually densely
cauliflorous while those of Pirigara are more in smaller terminal or subt rminal
clusters. In short, the problem appears insoluble upon our present evidence, and is
commended to the attention of future collectors in the field.

1. GUSTAVIA PLEUROCARPA Pittier, in Contrib. U. S. Nat. Herb. 26:4. 1927.

Trees up to 20 m. tall, with stout straight trunks; leafy twigs slender. Leaves

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1958]
FLORA OF PANAMA (Lecythidaceae) 119

rather distant, distinctly petiolate, the blade elliptic to elliptic-obovate, rather
abruptly acuminate, obtusely cuneate and somewhat decurrent to the petiole, 8—20
cm. long, 4-8 cm. broad, glabrous, firmly membranaceous, essentially entire; petiole
1.5-2.0 cm. long. Flowers terminal or subterminal, solitary or paired; peduncle
about 2 cm. long, rather stout, 2-bracteate at about the middle, densely ferruginous-
tomentellous; hypanthium broadly turbinate, about 7 mm. high and 10 mm. wide,
densely ferruginous-tomentellous, longitudinally winged alternate with the calyx
lobes; calyx lobes 6, broadly triangular, acute, about 8 mm. long, thickly foliaceous,
densely ferruginous-tomentellous; petals 6, yellowish white, broadly ovate, nearly
equal, 2.5-3.0 cm. long, densely ferruginous-tomentellous without and within
minutely so. Somewhat immature fruits turbinate, about 8 mm. high and 18 mm.
in diameter, crowned by the somewhat accrescent persistent calyx lobes, densely
ferruginous-tomentellous without, not very prominently winged.

Lowland forests of the Atlantic watershed, Panama, Colombia.

COLÓN: Loma de la Gloria, near Fató (Nombre de Dios), Pittier 3857, 4104, 4426.

I doubt very much that this species actually is distinct from G. bexapetala, but
I do not feel competent to make the reduction.

Fig. 36. Gustavia brachycarpa

2. GUSTAVIA BRACHYCARPA Pittier, in Contrib. U. S. Nat. Herb. 26:3. 1927.

Trees 5-15 m. tall, with stout straight trunks; leafy twigs slender. Leaves
rather distant, distinctly petiolate, the blade obovate-elliptic, rather abruptly and
narrowly acuminate, obtusely cuneate and somewhat decurrent to the petiole, 9-22
cm. long, 3-8 cm. broad, glabrous, firmly membranaceous, rather obscurely
undulate-serrulate; petiole 1-2 cm. long. Flowers terminal or subterminal, solitary
or in pairs; peduncle rather slender, 1-2 cm. long in flower, accrescent and decur-
rently winged in fruit; hypanthium turbinate, broadly winged, about 5 mm. high

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[Vol. 45
120 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and 8 mm. broad, minutely ferruginous-papillate; calyx lobes 6, broadly triangular,
about 12 mm. long, thickly foliaceous, ferruginous-papillate; petals 6, white,
broadly oval, nearly equal, 4.0—4.5 cm. long, minutely ferruginous-papillate with-
out and within. Mature fruits cylindric-campanulate, about 3 cm. long and 2 cm.
broad, gradually decurrent to the accrescent winged peduncle, crowned by the
accrescent persistent calyx lobes, essentially glabrous without.

Lowland forests and swamps of the Pacific watershed, Panama.

CHIRIQUÍ: vicinity of San Félix, Pittier 5269; west of Remedios, Allen 787.

The beautiful and informative fruits of Pittier 5269 have been discussed above.
Allen 787 is in flower. This must correspond to Seemann’s (Bot. Voy. H.M.S.
Herald 126. 1853) record of Lecytbis coriacea DC. from Remedios, which he calls
ollito (little pot-tree), doubtless with reference to the dry dehiscent fruits. With
great significance from the standpoint of our argument Seemann reports the ver-
nacular name of the true Gustavias as membrillo (quince tree), which is in wide
use in Panama to the present day.

3. GUSTAVIA NANA Pittier, in Contrib. U. S. Nat. Herb. 26:5. pl. 3-4. 1927.
Gustavia rbodantba Standl. in Field Mus. Publ. Bot. 4:239. 1929.

Shrubs or small trees to about 9 m. tall; leafy twigs rather slender. Leaves
congested seasonally toward the tips of the branches, of mediocre size for the genus
and with proportionately long slender petioles, the blade rather narrowly oblong-
lanceolate, up to about 4 dm. long and 8 cm. broad, rather gradually and acutely
acuminate, more abruptly cuneate at the base, rather obscurely and distantly
undulate-serrulate, membranaceous, glabrous; petiole variable in length, about 3-15
cm. long. Flowers mostly densely cauliflorous beneath the terminal cluster of
leaves; peduncle corymbosely several-flowered; hypanthium depressed-turbinate,
about 7 mm. broad and 5 mm. long at anthesis; calyx reduced to an obscurely
4-lobed ring 1.5-2.0 cm. in diameter, minutely puberulent-papillate; petals 6-8,
more or less unequal, oblong-obovate, deep pink, 2.5—3.5 cm. long, densely papillate.
Mature fruits subglobose, up to about 7 cm. in diameter, the opercular region of
about equal diameter, not very fleshy, the pericarp about 5 mm. thick.

Apparently endemic to Panama, in lowland forests.

ON: Loma de la Gloria, near Fató (Nombre de Dios), Pittier 4093. DARIÉN: near

COL
mouth of Rio Yapć, Allen 349; Marraganti and vicinity, Williams 655. SAN BLAS: Permé,
Cooper 239.

4. GUSTAVIA suPERBA (HBK.) Berg, in Linnaea 27:444. 1856.

Pirigara superba HBK. Nov. Gen. & Spec. 7:261. 1825.
ustavia insignis Linden, ex Hook. Bot. Mag. bl. 5069. 1858.
Gustavia superba var. salviniae Hemsl. Biol. Centr.-Am. Bot. 1:413. 1880.
Japarandiba superba (HBK.) O. Ktze. Rev. Gen. 240. 1891.
Trees to about 20 m. tall, with stout straight trunks, branching infrequently;
leafy twigs rather stout. Leaves congested seasonally toward the tips of
branches, usually very large and with proportionally short and stout petioles, che

(120)

19581
FLORA OF PANAMA (Lecytbidaceae) 121

KC

===

|

I)
/

i

=

j

Fig. 37. Gustavia superba

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[Vol. 45
I4 ANNALS OF THE MISSOURI BOTANICAL GARDEN

blade elliptic-oblanceolate, up to about 10 dm. long and 2 dm. broad, rather
abruptly and acutely acuminate, narrowly cuneate and more or less decurrent to
the petiole, coarsely to rather obscurely undulate-serrate, membranaceous, glabrous;
petiole 2-9 cm. long. Flowers mostly densely cauliflorous beneath the terminal
cluster of leaves; peduncle stout, usually about 6 cm. long, corymbosely several-
flowered; hypanthium turbinate, about 1.5 cm. long and broad at anthesis; calyx
reduced to a more or less obscurely 4-lobed ring 1.5-2.0 cm. in diameter, densely
puberulent-papillate; petals 6—12, more or less unequal, oblong-obovate, white,
usually yellowish at the base and more or less tinged with pink or lavender toward
the tip, 4—7 cm. long, minutely papillate. Mature fruits depressed-globose, up to
about 8 cm. broad and 6 cm. long, the opercular region about one third as broad,
very fleshy, the pericarp about 5-10 mm. thick.
Costa Rica to Ecuador, in lowland forests. Membrillo.

, $56; Monte
Lirio, Christopherson 128; Empire, Christopherson 197; Barro Colorado Island, Standley
II

R
93. NAMA: Pacora, Bro. Pau 341; Juan Díaz region, Maxon t$ Harvey 67464; Taboga
Island, Standley 27934; Taboguilia Island, Miller 1998; Trapiche, Perlas Islands, Allen
2610, Miller 1874; San José Island, Erlanson 495, Jobnston 282.

The enormous leaves crowded toward the ends of the branches render the
membrillo very conspicuous. Although the trees do not pass through consecutive
leafy and leafless stages, as do some in the tropics, and always are in leaf, their
growth is in definite cycles and the leaves are produced in well marked “flushes.”
Preceeding such a flush, large terminal buds resembling those of the temperate
hickories are produced. As the first leaves develop from these buds the nodes and
subsequent leaf scars are conspicuously congested.

Freshly cut wood and wilting leaves of the membrillo produce an offensive odor
of putrifaction which has earned for the trees the suggestive name of palo de
muerte in some localities. This is apparently a quality shared by other Gustavias,
one species of Venezuela having been named G. fustis-mortui by Pittier. The fruit
is produced toward the end of the dry season from February to April on the Pacific
slope. It consists of a brown or greenish rind enclosing numerous large brown
seeds surrounded by an orange pulp which is reputedly edible, as the name mem-
brillo (quince) suggests, but according to Dr. I. M. Johnston is scarcely palatable.

4. GRIAS L
GRIAS L. Syst. ed. 10. 1075. 1759.
Pirigara Aubl. Hist. Pl. Guian. 1:487. 1775, pro parte.

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1958]

FLORA OF PANAMA (Lecythidaceae)

Fig. 38. Grias pittieri

(123)

123

[Vol. 45
124 | ANNALS OF THE MISSOURI BOTANICAL GARDEN

Large trees, the leafy twigs stout and fistulose. Leaves usually crowded toward
the end of the twigs, ordinarily sessile or subsessile. Flowers few to several in dense
umbelliform cauliflorous clusters. Hypanthium urceolate. Calyx nearly entire to
broadly and irregularly 4-lobed, not prominent in fruit. Petals 4, rarely 5, sub-
equal, rather small for the family. Androphore radially symmetrical, wholly fertile,
the elongate inflexed fleshy filaments of the innumerable stamens coherent at the
base into a shallow perigynous ring; anthers small and ovoid, dehiscing longitudi-
nally. Ovary 4-celled at anthesis, each cell containing 2-4 pendulous ovules, but
usually only 1 cell and 1 ovule developing in fruit; stigma sessile, 4-radiate, inserted
on a prominent square disc. Fruit a rather mediocre coriaceous or fleshy pyriform
indehiscent berry usually containing a single seed, the opercular region and calyx
obscure.

About a dozen species, according to Knuth, extending from British Honduras
to the Guianas and Peru, and with one species in Jamaica. The trees are rather
similar to those of Gustavia in their stature and large leaves.

a. Leaves very large, about 6-10 dm. long and 2-4 dm. broad; flowers
relatively large, the petals about 1.5-2.0 cm. long. Atlantic slope........... 1. G. FENDLERI
a. Leaves medi

m. long and 5-16 cm. broad; flowers

10C —4.5 >
relatively small, the petals about 8-10 mm. long. Pacific slope................. 2. G. PITTIERI

»

1. GRIAS FENDLERI Seem. Bot. Voy. H.M.S. Herald 126. 1854.

Trees to abont 13 im. tall Lester eile or subsessile, the blade obovate-
spatulate, shortly and abruptly acuminate to broadly rounded at the tip, gradually
attenuate from above the middle to a decurrent subpetiolar base, about 6-10 dm.
long and 2-4 dm. broad, subcoriaceous, glabrous. Flowers in small umbelliform
cauliflorous clusters; hypanthium about 3 mm. long and 2 mm. broad; calyx
frequently nearly entire in bud, but at anthesis usually splitting into 4 rather broad
lobes about 2 mm. long; petals 4, white or cream, about 1.5—2.0 cm. long. Fruit
unknown.

BOCAS DEL TORO: Fish Creek hills, vicinity of Chiriqui Lagoon, Von Wedel 2423.
CANAL ZONE: Chagres, Fendler 187.

2. GRIAS PITTIERI R. Knuth, in Engl. Pflanzenr. 42192:29, 1939.

Trees up to about 10 m. tall. Leaves sessile or subsessile, the blade oblanceolate-
spatulate, shortly and acutely acuminate, gradually attenuate from near the tip t0
a decurrent subpetiolar base, about 4.0—4.5 dm. long and 5—16 cm. broad, sub-
coriaceous, glabrous. Flowers in small umbelliform cauliflorous clusters; hypan-
thium about 2 mm. long and 1.5 mm. broad; calyx nearly entire in the bud but
at anthesis usually splitting into 4 irregular broad lobes about 1.5 mm. long; petals
4 or rarely 5, white, about 8-10 mm. long. Fruit pyriform, 2-3 cm. long, 2.0-2.5
cm. broad, gradually attenuate to a persistent pedicel of nearly equal length,
greenish brown.

Pacific coast of Panama, in lowland forests. Cooper and Slater report the
vernacular names jaguey and membrillo.

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1958]
FLORA OF PANAMA (Lecythidaceae) 125

HIRIQUÍ: vicinity of Puerto Armuelles, Woodson 6 Scbery 901; Progreso, Cooper 6
Slater sej DARIÉN: around Pinogana, Pittier 6552; trail between Pinogana and Yavisa,
273:

It is unfortunate that intermediate stations between Chiriqui and Darićn have
not been found. While the leaves of the two populations are quite similar, both
specimens from Chiriqui are in fruit and both from Darićn are in flower only, and
their correlation thus is in doubt to that degree.

3. COUROUPITA Aubl.
COUROUPITA Aubl. Hist. Pl. Guian. 708. 1775.
Pontopidiana Scop. Introd. 195. 1777.
Elsholtzia Neck. Elem. 2:256. 1790.
Pekea Juss. i 249. 1798, pro parte.

Trees of moderate to gigantic size, the leafy twigs ordinarily stout. Leaves
crowded toward the tips of the twigs, shortly petiolate, broadly obovate-cuneate
and rather small for the family, entire to serrulate. Flowers in extensive sub-
terminal to cauliflorous racemes and panicles. Hypanthium broadly obovoid.
Calyx deeply 6-lobed, inconspicuous in fruit. Petals 6, more or less unequal, white
more or less suffused with red. Androphore strongly bilaterally symmetrical,
wholly fertile, produced into a prominent simply involute posterior hood, the
stamens of the hood somewhat larger and with longer filaments than those of the
central disc; anthers very small and ovoid, dehiscing longitudinally. Ovary 6-
celled, containing numerous lateral anatropous ovules; stigma sessile, 6-radiate,
inserted on a prominent round disc. Fruit a large globose indehiscent coriaceous
berry, the seeds several, compressed, of moderate size, immersed in fetid pulp, the
embryo differentiated and with fleshy cotyledons.

About 19 species according to Knuth, extending from Nicaragua to Amazonian
Peru and Brazil and in Jamaica and the Lesser Antilles. Couroupita guianensis
Aubl., known in English as Cannon-ball Tree, is occasionally cultivated in tropical
parks and gardens. The handsome, usually reddish flowers vary from 3 to 9 cm.
in diameter, and the large globular fruits may be 2 dm. in diameter.

In prefacing his account of the Couroupitas of Central America Pittier wrote,
Owing to the scarcity of the cannon-ball trees, it has been found difficult to obtain
satisfactory specimens. For this reason the three following species are incompletely
described." Knuth, although presenting detailed descriptions, generates even less
confidence by his key to the species of Central America, which is reproduced below

or the possible use of the reader:

B. Species oz nae. ; A
cè > a; Costarica: flos ad 7 cm diam 15. C. micaraguensis
a. Sk SEM dif s ^ 7 NA aa Date ao io iaca lia aa e ac 16. C. Cutteri
B. Flos 7-8 cm diam . 17. C. darienensis
T Fo4-—*ocomdan —— — — —. e". CIE C. odoratissima
8. Flos 3 cm diam iii 19. C. parviflora

In his key to the three Central American species recognized by him, Pittier

(125)

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

AE i
Na et
A

A
E Po,
VC
drop e
EA

nm Na
-— i
Kł T

mr ARE

(N
y A B i,
7

2

- N
LUTTE: 7 j $
Ą \ 7 TS

\S

IE
NE

NG

27 Vm 1/77; ) ZA

SN
a 7 \
„eg;

J Y
Zs y) M == 74
UL
RA

W/M
(MD

ey 7, Uf A
M Ą \

Fig. 39. Couroupita guianensis

(126)

1958]
FLORA OF PANAMA (Lecythidaceae) 127

distinguishes C. odoratissima Seem. as a "low spreading savanna tree, branched
almost from the base" and C. nicaraguensis DC. and C. darienensis Pittier as "high
forest trees"; yet Seemann assigns a height of 60—80 feet to the type trees of C.
odoratissima. Couroupita cutteri Morton & Skutch and C. parviflora Standl. both
are reported only through single collections in the immediate vicinity of the
Chiriquí Lagoon.

Since a total of only three herbarium specimens is available for our study at
present, it appears quite impossible to treat the species in the usual way since the
leaves are nearly, if not quite, identical and the dimensions of the flowers analysed
by Knuth present a continuum (the flowers of the type specimen of C. parviflora
obviously are not fully expanded). The three specimens examined for the present
account are as follows:

S DEL TORO: i. from Almirante, Skutch 19 (type of C. cutteri in U.S. Nat.
Herb.); | Chatuninala wine Cooper & Slater 11 (type of C. parviflora in U. S. Nat.
osa ‘nt forests around Pinogana, Pittier 6563 (type of C. darienensis in U. S.

The type specimen of C. odoratissima was collected by Seemann in the forests
of Rio de Jestis, between Santiago and Puerto Mutis, Veraguas. According to
Pittier, the species since has been seen by Dr. Otto Lutz in forests of Hato de San
Juan, between San Lorenzo and San Félix, eastern Chiriqui, but the photograph to
which he refers is not included in the U. S. National Herbarium.

An adequate herbarium representation of Couroupita is one of the outstanding
desiderata for a complete Flora of Panama. Seemann was informed by local resi-
dents that C. odoratissima was to be found only in a small area at the type locality.
Failure of recent collectors to find more ample material suggests that the species,
one or several, must be extremely local and infrequent.

5. LECYTHIS Loefl.
LECYTHIS Loefl. It. Hisp. 189. 1758.
Bergena Adans. Fam. 2:345. 1763, nec Neck. nec Raf.

Large trees. Leaves large and coriaceous, or unusually small for the family,
entire to crenulate, usually not congested at the tips of the twigs. Flowers in
subterminal or axillary panicles. Hypanthium obovoid. Calyx deeply 6- lobed,
becoming a more or less prominently thickened and approximately median rim in
fruit. Petals 6, unequal. Androphore strongly bilaterally symmetrical, produced
into a prominent posterior simply involute wholly or partially staminodial hood
concealing the low staminiferous central disc; anthers ovoid, dehiscing longi-
tudinally. Ovary 4-celled, with several pendulous anatropous ovules; stigma sessile,
4-radiate. Fruit a large woody subglobose dehiscent pyx, the seeds several, large
and angulate, pendulous from the greatly enlarged fleshy funicles, the embryo
undifferentiated.

About 50 species, according to Knuth, extending from Costa Rica to southern
Brazil, and in the Lesser Antilles. 'The trees are amongst the largest of the tropical

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[Vol. 45
128 ANNALS OF THE MISSOURI BOTANICAL GARDEN

American forests, and the hard wood is used structurally. The large fruits resemble
aboriginal pottery, hence the Spanish vernacular olla de mono and the English
Monkey-pot. The trees and their timber are known as cocobolo in Panama and
Costa Rica.

Pittier (in Contrib. U. S. Nat. Herb. 26:8. 1927) recognized four species of
Lecythis in Panama, basing them chiefly upon rather obscure differences seen in the
fruits. In an earlier paper on the Lecythidaceae of Costa Rica (loc. cit. 12:95-
101. 1908), however, he included a photograph of several fruits of L. costaricensis
which displays striking variance and casts considerable doubt upon the reality of
his subsequent species. Only two species are recognized here.

a. Leaves large and coriaceous, about 20-50 cm. long, sa or essentially

so; pyx with the margin of the orifice Lu. rough and prestati

muricate with the bases of hę sagra veins, s calycine rim in

definitely erose or poet ire. Pacific slop 1. L. TUYRANA

rabini ia about ioe m. long, finely ges:
with the m a of the or rifice è concave, Paso. vely smooth and with

little evidence of abscissed veins, the calycine rim conspicuously Ma
Atlantic slope . L. AMPLA

m
E]
m
©
2

.8
a
B
p
=
p

1. LECYTHIS TUYRANA Pittier, in Contrib. U. S. Nat. Herb. 26:9. pl. 7. 1927.
Lecytbis melliana Pittier, loc. cit. 8. pl. 6. 1927.

Large trees to about 30 m. tall, the leafy twigs stout and thinly rimose. Leaves
large and coriaceous, broadly oblong to oval, abruptly and shortly acuminate,
rounded and inconspicuously decurrent at the base, about 20-60 cm. long and 6-20
cm. broad, entire, glabrous; petiole stout and canaliculate, 1-2 cm. long. Inflor-
escence paniculate, many-flowered, the stout puberulent peduncle about as long as
the subtending leaves. Flowers sessile; calyx lobes broadly ovate to reniform,
obtuse or rounded, 6-8 mm. long; petals oval, 2.5—3.0 cm. long, yellow. Fruit
subglobose, deep brown and roughly verrucose, 10—15 cm. long and broad, the
calycine rim indefinitely erose to subentire, median or submedian, the margin of
the orifice convex, rough and irregularly muricate with the bases of the abscissed
veins.

Pacific slope of eastern Panama, in lowland forests; sometimes planted for
ornament in the Canal Zone.

AL ZONE: near firehouse, Balboa, Standley 26898; Balboa, Standley 30867, Joban-
sen de DARIÉN: Pinogana, Pittier 6 576, 6538.
2. LECYTHIS AMPLA Miers, in Trans. Linn. Soc. 30:204. pl. 43, figs 1-2. 1874
Lecythis armilensis Pittier, in Contrib, U. S. Nat. Herb. 26:9. pl. 8. 1927.

Large trees to about 40 m. tall, the leafy twigs remarkably slender. Leaves
relatively small and membranaceous, broadly elliptic, abruptly and rather shortly
subcaudate-acuminate, obtuse at the base, about 8—10 cm. long, finely crenulate,
glabrous; petiole slender, about 5 mm. long. Flowers unknown to us. ruit
globose-subampullaceous, yellowish brown and finely granular, not verrucose, 127
20 cm. long, 12-15 cm. broad, the calycine rim at the upper third and distinctly
6-lobed, the margin of the orifice concave, relatively smooth and with little
evidence of the abscissed veins.

(128)

1958]
FLORA OF PANAMA (Lecythidaceae)

|
iy

UM

tl

N WN AN N

Fig. 40. Lecythis tuyrana

(129)

129

[Vol. 45
130 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Atlantic lowlands of Panama; Colombia.
SAN BLAS: hills of Sperdi, near Puerto Obaldia, Pittier 4343.

I have not seen the old fruits, also from Puerto Obaldia, upon which Pittier

based L. armilensis, but the photographs appear to be essentially similar to those of
L. ampla and well within the range of expected variance.

6. ESCHWEILERA Mart. r
ESCHWEILERA Mart. ex DC. Prodr. 3:293. 1828.

Large trees. Leaves rather large and coriaceous, entire in our species, usually
not congested at the tips of the twigs. Flowers in subterminal or axillary panicles
or racemes. Hypanthium obovoid. Calyx deeply 6-lobed, usually less than half
adnate to the fruit in our species, becoming more or less prominently thickened in
fruit or rarely (E. calyculata) retaining their coriaceous texture. Petals 6, more
or less unequal (of abnormally thin texture in E. panamensis). Androphore
strongly bilaterally symmetrical, produced into a prominent spirally involute
staminodial posterior hood concealing the relatively few minute fertile stamens of
the central disc. Ovary 2-celled, each with several nearly sessile erect basal ovules;
stigma capitate and essentially entire, sessile or borne upon a prominent stylo-
podium. Fruit a mediocre, woody or subcoriaceous, depressed-campanulate,
dehiscent pyx, the seeds several, basal, sessile, thick and angulate, the embryo
undifferentiated.

This is the largest genus of Lecythidaceae, consisting of over 90 species accord-
ing to Knuth, and extending from Costa Rica to central Brazil. ‘The trees are
generally similar to Lecytbis, but do not attain such great size in Panama.
a. n Wy small, =

1.5-2.0 cm. in diameter, the petals n
in tex the calyx 1 Shei long; peduncles slender and pani
late, rari da ached toward the iei leaves seas sat
t 6-12 cm. lo tlantic seri Blas 1. E. PANAMENSIS
aa. Flowers — large, Spa 5—3.5 cm. n due ter, the petals thick
xture, the calyx lobes long; pelle: thick

mm. and racemose,
characteristically + a PF leaves rather large, usually about 15—35
cm

. Pya very thickly woody, the — about 1 cm.

calyx lobes „ei to about midw

prominent and woody.

hick or more, the
of the Pyx an becoming very
Pacific slo ară Chiriq 2, E. PITTIERI
bb. Pyx thin and subcoriaceous, the pericarp sare more than 2 mm.
thick the calyx lobes adnate to the base o
lobes strongly adnate, not t accrescen li oming vestigial in
grees leaves dicta te x rrowly and acutely subcaudate-
i ific slo Darié

bă arićn
cc. Calyx lobes weakly adnate, accrescent and prominent in fru

lea obtuse or roun ida to abru ae and da
Atlantic slope: Bocas del Toro and Coló

. E. GARAGARAE
acuminate.

. E. CALYCULATA

1. ESCHWEILERA PANAMENSIS Pittier, in Contrib. U. S. Nat. Herb. 26:12. 1927.
Eschweilera reversa Pittier, loc. cit. 11. 1927, pro parte?

Trees up to 25 m. tall, the leafy twigs relatively slender. Leaves decir
petiolate, oval, very abruptly and shortly acuminate or obtuse at the tip, broa T

(130)

1958]
FLORA OF PANAMA (Lecythidaceae) 131

obtuse or rounded at the base, about 8—12 cm. long, 3—8 cm. broad, coriaceous,
glabrous; petiole rather stout, about 5 mm. long. Inflorescence subterminal and
axillary, paniculate, many-flowered, somewhat longer than the subtending leaves,
the peduncle slender and minutely puberulent. Flowers relatively small, about
1.5-2.0 cm. in diameter; calyx lobes deltoid, 1-2 mm. long; petals unequal, about
1 cm. long, yellow, thin in texture for the family. Fruit (according to Pittier)
“globose-depressed, about 4 cm. long and 5 cm. in diameter, 2-celled, slightly
attenuate at base, with rather thin walls; calycinal zone bearing the persistent
enlarged lobes of the calyx, the interzonal band 1.5—2.0 cm. broad.”

Lowland forests of Atlantic slope, Panama.

SAN BLAS: hills back of Puerto Obaldia, Pittier 4338.

The fruit accompanying the type specimen apparently has been lost or mis-
placed in the U. S. National Herbarium.

The type sheet of E. reversa (Pittier 4394 in U. S. Nat. Herb.) from the Plain
of Sperdi, near Puerto Obaldia, seems to be a mixture consisting of a leafy branch,
possibly a Rinorea, and a detached pyx and operculum of an Eschweilera. Although
it would be odd for such a veteran as Pittier to mis-associate two such unrelated
plants and although I cannot be positive of the species of the leaves, I feel certain
that the leaves are not from an Eschweilera, particularly because of their prom-
inent linear stipules. The fruit resembles those of E. pittieri, of the Pacific slope
of Chiriqui, very closely.

In his discussion of E. reversa Pittier states that the pyx and the operculum of
his type sheet of the species were found detached under trees, presumably from
which his leafy branch was obtained, "but since there were trees of Eschweilera
banamensis scattered in the forest all around, the source of that lid is not abso-
lutely certain.” Under such circumstances, E. reversa cannot be regarded as more
than a nomen dubium at the present time.

2. EscHWEILERA PITTIERI R. Knuth, in Engl. Pflanzenr. 4?!?^:95. 1939.
sent verruculosa Pittier, in Contrib. U. S. Nat. Herb. 26:13. pl. 12. 1927, non
1ers,

Trees up to 20 m. tall, the leafy twigs relatively slender. Leaves shortly peti-
olate, elliptic-oblong, abruptly and rather shortly and acutely acuminate, broadly
obtuse or rounded at the base, 12-30 cm. long, 6-12 cm. broad, subcoriaceous,
glabrous; petiole rather stout, 1.0-1.5 cm. long. Inflorescence racemose, several-
to many-flowered, much shorter than the subtending leaves, the peduncle stout and
glabrous. Flowers relatively large, about 3.0—3.5 cm. in diameter; calyx lobes
ovate, obtuse, approximately lI CI long; petals yellow or white, obovate-oblong,
more or less unequal, about 2 cm. long. Fruits very thickly woody, the pericarp
about 1 cm. thick or more, conspicuously brown-verruculose, particularly in the
calycine zone, depressed-globose, about 2.5—5.0 cm. long and 3.5—6.0 cm. broad, the
calyx lobes adnate to about midway of the pyx and becoming very prominently
woody, the operculum conspicuously domed and nearly hemispheric, almost half as
deep as the body of the pyx.

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[Vol. 45
132 ANNALS OF THE MISSOURI BOTANICAL GARDEN

/

d

JA

4 y Do m
KD. mia «T
/ W

| Í
Ny ull
Mp
| U

AA

Y)

7

A

Fig. 41. Eschweilera pittieri

(132)

1958]
FLORA OF PANAMA (Lecytbidaceae) 133

Lowland forests, Pacific slope of Panama.

cHIRIQUÍ: vicinity of San Félix, Pittier 5290, 5738; Rio Tinta, along main highway,
Woodson, Seibert 6 Allen 402; San Félix to Cerro Flor, Allen 1920; vicinity of Remedios,
Allen 3476.
3. ESCHWEILERA GARAGARAE Pittier, in Contrib. U. S. Nat. Herb. 12:12. 1927.

Trees up to about 35 m. tall, the leafy twigs relatively slender. Leaves elliptic-
oblong, narrowly and acutely subcaudate-acuminate, broadly rounded or obtuse at
the base, about 12-35 cm. long and 3-12 cm. broad, membranaceous to subcoria-
ceous, glabrous; petiole about 1.0—1.5 cm. long. Inflorescence racemose, several-
flowered, the peduncle rather stout and inconspicuously puberulent, much shorter
than the subtending leaves. Flowers relatively large, about 3.0—3.5 cm. in diameter;
calyx lobes broadly ovate, acute, about 6-9 mm. long; petals yellow, somewhat
unequal, broadly oval, about 2 cm. long. Fruits depressed-globose, about 5 cm.
long and 5.5—6.0 cm. broad, the pericarp rather thin and subcoriaceous, scarcely
more than 2 mm. thick, the calyx lobes strongly adnate to the very base of the
PX, not accrescent, becoming obscure and vestigial, the operculum rather flat and
not conspicuously constricted at the orifice, about one third as deep as the body
of the pyx.

Lowland forests of the southeastern Pacific slope of Panama.

DARIÉN: foothills of Garagará, Sambü basin, Piźtier 5616; vicinity of Pinogana, Allen
920; Marraganti on the Tuyra R., about El Real, Pittier 6590.
4. EscHWEILERA CALYCULATA Pittier, in Contrib. U. S. Nat. Herb. 12:97. pis.

I-2. 1908

Trees up to 10 m. tall, or apparently sometimes more than twice as tall (ac-
cording to Von Wedel), the leafy twigs relatively stout. Leaves rather narrowly
obovate-oblong, obtuse or rounded to abruptly and obtusely acuminate, broadly
and obtusely cuneate, 13-35 cm. long, 5-15 cm. broad, coriaceous, glabrous;
petiole about 1-2 cm. long. Inflorescence racemose, several-flowered, the peduncle
stout and glabrous, less than half as long as the subtending leaves. Flowers rela-
tively large, about 3.5 cm. in diameter; calyx lobes broadly ovate, acute, about 6
mm. long, greatly accrescent in fruit but retaining their coriaceous texture and
only weakly adnate to the base of the pyx (according to Pittier); petals oval-
obovate, about 2 cm. long, rather unequal, yellow. Fruit (according to Pittier and
the cited illustrations) “7 cm. in diameter and about 6 cm. high, depressed-globose,
thin-walled, rather smooth, with persistent sepals becoming twice larger than in
flower”, the subhemispheric operculum nearly twice as deep as the body of the pyx,
not conspicuously contracted at the orifice.

Lowland forests of the northwestern Atlantic slope of Panama, and adjacent
Costa Rica. Reported as mata cansada by Cooper and Slater.

BOCAS DEL TORO: Isla Colón, Chiriquí Lagoon, Vom Wedel 2804, 408; Almirante
ari Cooper & Slater 57; Changuinola valley, Dunlap 493. CANAL ZONE: near Fort

olph, Maxon & Harvey 6537, Standley 28675.

I have not been able, as yet, to examine well-authenticated fruits of E. caly-

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[Vol. 45
134 ANNALS OF THE MISSOURI BOTANICAL GARDEN

culata. The fruits labeled such in the U. S. National Herbarium to accompany
Cooper & Slater 57 certainly are Couroupitas. Since Cooper and Slater also col-
lected Couroubita in the Almirante region, the fruits probably should accompany
Cooper & Slater II, the type sheet of C. parviflora Standl.

7. COURATARI Aubl.
COURATARI Aubl. PI. Guian. 2:723. 1775.
Lecythopsis Schrank, in Denkschr. Akad. Muench. 7:241. 1821.

Gigantic trees. Leaves usually of moderate or rather small size for the family,
usually membranaceous and more or less pubescent, entire or inconspicuously cren-
ulate, usually not congested at the tips of the twigs. Flowers rather small, in
terminal panicles. Hypanthium obconic. Calyx deeply 6-lobed to subentire, not
accrescent, vestigial in fruit. Petals 6, subequal, usually pink or rose. Androphore
strongly bilaterally symmetrical, produced into a prominent, posterior hood spirally
involute then reversely revolute, wholly staminodial and concealing the rather few
minute fertile stamens of the central disc. Ovary 3-celled, containing numerous
basal ovules; stigma sessile or subsessile, minutely 3-radiate. Fruit a mediocre
subcoriaceous narrowly campanulate dehiscent pyx; seeds several, basal, very thin
and broadly winged, the embryo subcentral.

A genus of approximately 20 described species, extending from Costa Rica to
central Brazil.

1. COURATARI PANAMENSIS Standl. in Field Mus. Publ. Bot. 4:239. 1929.

Gigantic trees up to about 30 m. tall, the leafy branches moderately stout,
prominently angled and minutely ferruginous-hirtellous when young. iis
broadly oval, obtuse to rounded or very abruptly and obtusely cuspidate, broadly
rounded at the base, 7-18 cm. long, 5-10 cm. broad, membranaceous, more or less
conspicuously ferruginous-hirtellous on the midrib and veins; petiole about 1 cm.
ong. Inflorescences terminal, paniculate, several- to many-flowered, the stout
minutely hirtellous peduncles somewhat longer than the subtending leaves. Flow-
ers unknown to us. Fruits cylindrical-campanulate, occasionally somewhat gibbous,
5-10 cm. long, about 3.0—3.5 cm. in diameter at the orifice, dark brown and con-
spicuously lenticellate; operculum strongly trigonal, permanently attached to the
stout elongate placenta; seeds oblong-oblanceolate, thinly membranaceous, about
7 cm. long and 1.5 cm. broad, rich brown.

Lowland forests of the Atlantic slope of Panama; perhaps also in lowland
forests of eastern Costa Rica. Barbour reports the vernacular names of coco, coc0

e mono and coquito in Panama.

BOCAS DEL TORO: Cricamola valley, region of Almirante, Cooper 542. CANAL ZONE:
near Battery VII, area west of Limén ay, Gatun Locks and Gatün Lake, Johnston s$."
PANAMA: Rio Indio drainage, about 9 miles east of trans-isthmian highway, Barbour
1059.

These three specimens, in addition to one other from Costa Rica, are the only

(134)

1958]
FLORA OF PANAMA (Lecythidaceae) 135

EA
kopi
a "A

mi pi

lim pa A
Vri AI
Rm N NE |

Fig. 42. Couratari panamensis

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[Vol. 45
136 ANNALS OF THE MISSOURI BOTANICAL GARDEN

records of Couratari from Central America. The three Panamanian specimens
show fruit only; the Costa Rican (Allen 5686, from the vicinity of Rio Esquinas,
Puntarenas province) is in flower. Whether a single species is represented is con-
jectural of course. The flowers of Allen 5686 are pink and about 1.5-2.0 cm. in
diameter; they were used in the composition of the accompanying illustration.

RHIZOPHORACEAE

By DAVID P. GREGORY

Trees or shrubs, often with swollen nodes. Leaves opposite, simple, entire or
serrate, with lanceolate, foliose, interpetiolar, caducous stipules. Inflorescence
axillary; flowers perfect and regular, sometimes solitary but usually cymose, perigy-
nous to epigynous. Calyx 3- to 14-merous, the calyx tube adnate to the ovary or
free, valvate, persistent. Petals equal in number to the sepals, free, often unguicu-
late and sometimes elaborately fimbriate, convolute or inflexed in the bud. Stamens
from twice the number of sepals to more numerous, in one whorl, inserted on a
lobed disc; anthers usually 2-celled, sometimes many-celled (Rhizophora). Ovary
superior, half-inferior or inferior, the locules and carpels 2-5, the septa sometimes
failing to develop and resulting in a single locule; placentation axile; ovules usually
2 per locule; style single, the stigma lobes usually equal in number to the carpels.
Fruit a leathery or fleshy berry crowned by the persistent calyx, usually indehiscent
but sometimes septicidally dehiscent, with one locule and one seed or 2-5 locules
each 1-seeded. Seeds sometimes germinating while the fruit is still on the tree
(Rhizophora), with or without endosperm.

This family has two groups of genera, one maritime and the other of upland
forest genera. The representatives in Panama include one genus of each group,
Rbizophora of the tidal zone and Cassipourea of inland areas. Rhizophora is the
more important economically. ‘The mangroves are a factor in land building, their
prop roots forming a lattice that collects silt and organic debris. The bark of these
plants contains from 20 to 45 per cent of tannins, although their tannin is appar-
ently inferior to that of some other plants used in the process of tanning. Ri
dyes also are obtained from the young shoots. Other uses are as firewood and
charcoal, for piling, for cabinet work and for manila and craft pulp. ;

Economic uses of Cassipourea are fewer, most important being the use of its
strong, flexible wood for such things as canoe paddles.

a. Leaves coriaceous; ultimate branchlets thick; inflorescences axillary and

cymose; stamens 8, cuneate, sessile or subsessile, the anthers many-

locular; ovary half-inferior, 2-locular; fruit a dry indehiscent berry, the

eed germinating in attached fruits; plants of mangrove swamps with
characteristic adventitious aerial and prop roots
Leaves membranaceous; ultimate branchlets slim; flowers fascicled in the
leaf axils or solitary; stamens 15-25 on slender filaments, the anthers
4-locular it a fl
capsul

1. RHIZOPHORA

>»
m
H

; OVary nearly superior, 3-locular; fruit a fleshy septifragal
e; not plants of the mangrove association, roots subterranean......... 2. CASSIPOUREA

(136)

1958]
FLORA OF PANAMA (Rbizophoraceae) 137

1. RHIZOPHORA L.
RHIZOPHORA L. Sp. PI. 1:443. 1753.

Mangle Pluk. ex Adans. Fam. PI. 2:445. 1763.
ngium Rumph. ex Scop. Intr. Hist. Nat. 218. 1777.
Asophora Neck. Elem. Bot. 2:361. 1790.

Maritime trees and shrubs with adventitious aerial and prop roots, branchlets
stout and furrowed, with prominent leaf and stipule scars. Leaves opposite, simple,
entire, evergreen, coriaceous, often spotted dorsally; stipules foliose, sessile, cadu-
cous. Inflorescence axillary, cymose, persistently bracteate. Flowers perfect,
regular. Sepals 4, valvate, distinct, coriaceous, reflexed and persistent in fruit.
Petals 4, alternate with the sepals, involute, often with hairy margins, caducous.
Stamens 8—12, introrse, sessile, in one whorl, cuneate, areolate, a membrane covering
the areolae opening at the inner edge to free the pollen. Ovary half-inferior,
bilocular, the superior portion conical, surrounded by a fleshy disc; style terete, the
stigma bifid or rarely trifid; ovules 2 per locule. Fruit a dry berry crowned by
the persistent calyx, indehiscent. Seed 1 by abortion, suspended, germinating while
the fruit is still on the tree.

Several species of tidal flats and river estuaries widely distributed between the
Tropic of Cancer and the Tropic of Capricorn, girdling the world.

a. Shrubs reaching 5 meters (rarely trees reaching 25 meters); inflorescence

2- to 6-flowered; floral buds >. broadest at the base, acu-

minate at the tip; sepals 8-15 mm. g at anthesis; petals 5-11 mm.
long; style 3-7 mm. long; bracteoles truncate-trapezoid. Either coast.
b. Flowers relatively large at anthesis, the sepals 11-15 mm. long, the
1 mm. long, the style 5-7 mm. long. Atlantic coast........- 1. R. MANGLE
m

l
petals 5-8 mm. long, the style 3.5—4.0 mm. long. Pacific coast......... 2. R. SAMOENSIS

h ol
anthesis 8.0—8.5 mm. long; petals 6-7 mm. long; style 4.0—4.5 mm. long;
bracteoles deltoid. Pacific BNAGE. ||| “oi s pu un OU die Vice aa miei si R BREVISTYLA

1. RHIZOPHORA MANGLE L. Sp. PI. 1:443. 1753.
Rbizophora americana Nutt. N. Amer. Sylv. 1:95. 1842.

Trees to 25 m. tall. Leaves obovate or elliptic, the apex obtuse, the base cune-
ate, slightly decurrent on the petiole, 7-15 cm. long, 2.0-8.5 cm. broad, glabrous,
coriaceous; petiole 1.0-2.5 cm. long. Inflorescence once-dichotomous and 2-
flowered, rarely trichotomous and 3-flowered; peduncles 1.5—3.5 cm. long; pedicels
0.5—2.0 cm. long; bracteoles truncate-trapezoid. Floral buds ampullaceous, broad-
est at the base, acuminate at the tip; sepals 11-15 mm. long, 4 mm. wide; petals
involute, 9-11 mm. long, 2.0-2.5 mm. wide; stamens 8, 5.5-7.0 mm. long; ovary
bilocular, half-inferior, the conical upper half terminating in a bifid style, the
length of ovary from base of disc to tip of stigma 5-7 mm. Fruit 1.9-2.6 cm.
long, 1.0-1.3 cm. wide, the protruding seedling up to 40 cm. long. ;

This species occurs along the Atlantic coast from Florida to Brazil and in the
West Indies, and also occurs, though not so commonly, along the west coast of

(137)

[Vol. 45
138 ANNALS OF THE MISSOURI BOTANICAL GARDEN

|]

Mim

€ È KL]

(| |

U

Fig. 43. Rhizophora samoensis

(138)

1958]
FLORA OF PANAMA (Rbizophoraceae) 139

Africa. It grows only along the coast, either within the high tide zone or in lakes
behind beaches or river estuaries in which it can become established. Under suitable
conditions it forms dense stands in shallow water and apparently contributes to
land building once it is established.

BOCAS DEL TORO: Nances Cay Island, Von Wedel 577; Water Valley, Von Wedel 994.
CANAL ZONE: Cristóbal, Salvoza 1010; Manzanillo Island, Hayes 46. cHIRIQUI: vicinity
of San Bartolomć, Peninsula de Burica, Woodson 6 Schery 934.

2. RHIZOPHORA SAMOENSIS (Hochr.) Salvoza, in Nat. and Appl. Sci. Bull.
Philipp. 5:220. 1936.

Rhizophora mangle var. samoensis Hochr. in Candollea 2:447. 1925.

Shrubs to 5 m. tall. Leaves lanceolate, elliptic or ovate, the apex obtuse, the
base cuneate, slightly decurrent on the petiole, 7.5—15.5 cm. long, 2.5-6.5 cm.
broad, glabrous, coriaceous; petiole 0.5—2.5 cm. long. Inflorescence once or twice di-
chotomous and 2- to 4-flowered, rarely trichotomous and 3- to 6-flowered; primary
peduncles 1.0—4.5 cm. long; pedicels 3-8 mm. long; bracteoles truncate-trapezoidal;
sepals 8-10 mm. long, 3-4 mm. wide; petals involute, hairy at the margins, 5-8
mm. long, 1-2 mm. wide; stamens 8, 4.5—6.0 mm. long; ovary bilocular, half-
inferior, the conical upper half terminating in a bifid style, length of ovary from
base of disc to tip of stigma 3.5-4.0 mm.; disc fleshy, 1 mm. high. Fruit 1.5-2.5
cm. long, 1.0-1.5 cm. wide, the protruding seedling 25-35 cm. long.

This species occurs on the Pacific coast of Mexico, Central America, and South
America from lower Baja California to Ecuador. It is also found on some of the
Pacific islands. This plant is similar in size to the smaller form of R. mangle but,

ike R. mangle and R. brevistyla, it never grows to a large tree. Otherwise its
ecology is probably similar to that of R. mangle, to which it seems closely related.

CANAL ZONE: Bella Vista, Salvoza 1006.

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[Vol. 45
140 ANNALS OF THE MISSOURI BOTANICAL GARDEN

3. RHIZOPHORA BREVISTYLA Salvoza, in Nat. and Appl. Sci. Bull. Philipp. 5:211.
1936.

Rbizophora racemosa Hieron. in Engl. Bot. Jahrb. 20:61. 1895, non Mey.

Trees to 20 m. tall. Leaves elliptic to broadly elliptic, the apex obtuse, the base
cuneate, slightly decurrent on the petiole, 9.5-15.0 cm. long, 4.5-7.0 cm. broad,
glabrous, coriaceous; petiole 1.5—2.0 cm. long. Inflorescence several times dichot-
omous with from 10—50 flowers; primary peduncles 1.5—3.5 cm. long; pedicels
0.3—1.0 cm. long; bracteoles deltoid. Floral buds ovoid, broadest near the middle,
acute to obtuse at the tip; sepals 8.0—8.5 mm. long, 3.5—4.0 mm. wide; petals
involute, 6—7 mm. long, 1.5 mm. wide; stamens 8, 5.5-6.0 mm. long. Ovary
bilocular, half-inferior, the superior portion from the base of the disc to the tip
of the bifid style 4.0—4.5 mm. long. Fruit 2.7-3.5 cm. long, 1.1—1.6 cm. broad,
the protruding seedling 30—45 cm. long.

This species occurs on the Pacific coast from Panama to Ecuador. As described
by I. M. Johnston (in Sargentia 8:222. 1949.) on San José Island, Panama, there
are two types of growth form. The most common is a shrub reaching 6 meters in
height, resembling R. mangle in general appearance and also resembling R. samo-
ensis which grows with it in Panama and south along the Pacific coast. The other
growth form is a tree reaching 20 meters in height. Johnston also points out the
correlation between growth form and number of flowers per inflorescence in this
species, the smaller plants having greatly reduced numbers of flowers, as few as
4—6 per inflorescence in the smallest ones.

CANAL ZONE: Bella Vista, Salvoza 1007; locality lacking, Allen 1726. PANAMÁ

M an
José Island, Pearl Archipelago, Harlow 33, Jobnston 253, 1357; Saboga Island, Pearl
Islands, Miller 1963.

2. CASSIPOUREA Aubl.
CASSIPOUREA Aubl. Hist. Pl. Guian. 1:528. 1775

Tita Scop. Introd. 219. 1777.
Legnotis Sw. Prod. 84. 1788.
Ricbaeia Thou. Gen. Nov. Madag. 25. 1806.
Richeia Poir. Encycl. Supp. 4:680. 1816.
8

Trees or shrubs, with slim branchlets. Leaves opposite, coriaceous Or POR
branaceous, simple, with caducous interpetiolar stipules. Flowers perfect E
regular, solitary to fascicled in the axils of the upper leaves, bracteate, with articu
lated pedicels. Calyx coriaceous, 4- to 7-lobed, valvate, hairy or glabrous, 3
panulate to spreading. Petals unguiculate and often elaborately fimbriate a
lacerate, inflexed, pubescent or glabrous. Stamens 8-40, inserted on or outside uit
disc, filaments equal to or longer than the calyx. Ovary 2- to 4-loc 4
hypogynous or slightly adnate to the base of the calyx, surrounded by a fleshy 0

(140)

141

ha
7

^
f
4

í

y

===, XS

1958]

FLORA OF PANAMA (Rbizopboraceae)

Fig. 44. Cassipourea elliptica

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[Vol. 45
142 ANNALS OF THE MISSOURI BOTANICAL GARDEN

membranaceous disc, each locule with 2 pendulous ovules; style filiform or stout.
Fruit a fleshy capsule, septifragal; seeds 2—4, arillate.

About 60 species of mountains, river banks and sandy shores; tropical America,
Africa (where the species are most numerous), Ceylon and southern India.

1. CASSIPOUREA ELLIPTICA (Sw.) Poir. Dict. Suppl. 2:131. 1811.
Legnotis elliptica Sw. Prod. 84. 1788.
Cassipourea alba Griseb. in Gótting. Abh. 7:223. 1857.
Cassipourea elliptica B alba Griseb. Fl. of B. W. Indies 274. 1860.
Cassipourea elliptica y pauciserrata Griseb. loc. cit. 1860.
Cassipourea cubensis Urb. Symb. Antill. 7:293. 1912.
Cassipourea macrodonta Standl. loc. cit. 242. 1929.

rubs or trees to 17 m. tall. Leaves elliptic or ovate, the apex acuminate or
acute, the base cuneate, 5-16 cm. long, 2.0—6.5 cm. wide, the upper half sometimes
serrate, glabrous, coriaceous or membranaceous; petiole 3-10 mm. long; stipules
4—5 mm. long, appressed pilose dorsally, soon caducous. Flowers numerous,
occasionally solitary; pedicels 2-5 mm. long, articulated just beneath the flower,
minutely bracteate at the base, strigose to glabrous. Calyx fleshy, campanulate, to
5 mm. long, with 4—5 lobes 2 mm. long, often stipitate at the base, sericeous within.
Petals 4—5, spatulate, laciniate, pilose to glabrous. Stamens 15—25, the slender
filaments sometimes united at the base, inserted on the outside of the fleshy disc.
Ovary 3-locular, sericeous above at the base of the style; style slender, 4—5 ps
long, sericeous at the base or nearly to the stigma; stigma capitate. Fruit ovoid,
crowned by the persistent calyx. ;

This species is widespread in the West Indies and in Central America, growing
along rivers, near beaches, and in forested areas.

OCAS DEL TORO: Old Bank Island, vicinity of Chiriquí Lagoon, Von V edel 283
Little Bocas, vicinity of Chiriqui Lagoon, Von Wedel 2490; Fish Creek Mts., vicinity del
Chiriquí Lagoon, Von Wedel 2244; Fish Creek, vicinity of Chiriquí Lagoon, Von Wede

ooper 440; Wa "

qui Lagoon, Von Wedel 1554, 1658, 1659; Changuinola Valley, Dunlap 447; Bastimentos,
Mariano Creek, vicinity of Chiriqui Lagoon, Von Wedel 2897; vicinity of Chiriqui page

on Wedel 1268, 1268A; locality lacking, Von Wedel 386. CANAL ZONE: Manz s
Island, Hayes 51; near Fort Randolph, Standley 28640; south of Fort Sherman, Jobns íi
1767; Barro Colorado Island, Bangham 566, 416, Salvoza 840, 970, Wetmore & Abbe pe
Woodworth & Vestal 527. cHIRIQUI: vicinity of San Felix, Pittier 5741. Ey. usi
Chagres, Fendler 19%. DARIÉN: vicinity of La Palma, Pittier 5483, 6617; Cana-

characteristics. This plant is not clearly enough defined from C. elliptica to war”

: : : it
rant specific rank. Standley's C. podantha is not even an extreme variant, and

cannot be distinguished from C. elliptica.
(142)

1958]
FLORA OF PANAMA (Combretaceae) 143

COMBRETACEAE

Bv A. W. EXELL

Trees, shrubs or lianas, rarely subherbaceous (never in Panama). Leaves
opposite, verticillate, alternate or spiral, exstipulate, simple, almost always entire.
Flowers perfect or perfect and staminate in the same inflorescence, usually actino-
morphic, sometimes slightly zygomorphic, in axillary or extra-axillary, elongated
or subcapitate spikes or racemes or in terminal and sometimes axillary panicles.
Receptacle (hypanthium) usually in two distinct parts, the lower receptacle sur-
rounding the ovary and adnate to it and the upper receptacle (calyx-tube) pro-
duced beyond it to form a short or long tube terminating in the (sometimes poorly
developed) calyx-lobes, caducous or persistent. Calyx-lobes 4-5 (rarely 6-8 or
almost obsolete) sometimes accrescent (not in Panama genera). Petals 4-5 or
absent, conspicuous or very small. Stamens usually twice as many as the sepals or
petals, borne inside the upper receptacle, usually in two series, exserted or included,
anthers versatile or more rarely adnate to the filaments and immobile. Disk usually
present, often hairy, intrastaminal. Style free in American genera (partly attached
to the upper receptacle in Quisqualis). Ovary completely inferior in all American
genera (semi-inferior in the West African genus Strephonema), unilocular, with
usually 2 (sometimes up to about 6) pendulous ovules of which only one usually
develops. Fruit (pseudocarp) very variable in size and shape, fleshy or dry, usually
indehiscent, often variously winged or ridged, 1-seeded.

The Combretaceae, an entirely tropical and subtropical family, found in rain-
forest, deciduous forest, savanna and mangrove formations, comprise 18 genera of
which two, Combretum and Terminalia, contain the great majority of species.
Some species of Terminalia yield useful timber and the ‘myrobalans’ used for dye-
ing in Asia. A few species of Combretum are cultivated in the tropics and as
stove-plants in temperate countries. Terminalia catappa L., a native of tropical
Asia and Polynesia, is often planted in towns as an avenue tree and Quisqualis indica
L., the Rangoon Creeper, is a climber often grown in tropical gardens.

In the course of their evolution the Combretaceae seem to have developed under
the influence of two principal tendencies: (1) the attraction of pollinating insects

y the massing of relatively small flowers in dense or rather dense inflorescences,
though with a number in individual exceptions (such as Combretum cacoucia) ;
(2) dispersal of the fruits by means of laterally extended wings (again with numer-
ous exceptions). The progressive development of inflorescences and fruits along
these lines leads to obvious space difficulties as the more the flowers are crowded
into pseudo-capitate spikes the less room there is for the development of wings to
the fruits. Several of the genera recognized in the family represent different solu-
tions to this problem. In Panama we have one of the most ‘advanced’ solutions in
the genus Conocarpus where the flattened pseudocarps, perhaps after a secondary
reduction in the wings, are tightly packed into a cone-like aggregate fruit. The
act that isolated species of Combretum, belonging to quite different sections, have

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[Vol. 45
144 ANNALS OF THE MISSOURI BOTANICAL GARDEN

wingless or nearly wingless fruits, often distributed by water, may also indicate
that wingless fruits have developed from winged ones.

In the various forms which the winging of the fruit may take we have an
interesting combination of two factors. From the inferior ovary, probably origi-
nating from a single carpel, embedded in receptacular tissue, there is apparently
what we might call an ‘internal’ tendency to produce a 2-winged fruit resulting
from the bilateral symmetry of the carpel. From the 4-merous or 5-merous calyx,
on the other hand, there seems to be an ‘external’ tendency to produce a 4-winged
(Combretum) or 5-winged (Combretum and Terminalia) fruit. In Terminalia,
in particular, it is interesting to see the effect of these two tendencies. We find,
for example, in addition to numerous species with either 2-winged or 5-winged
fruits respectively, certain species, such as T. amazonia in Panama, with primarily
2-winged fruits in which three secondary wings or ridges are also developed. If
this interpretation is correct it would seem that the great plasticity in the winging
of the fruit is due to the surrounding, in an inferior fruit, of a bilateral ‘core’ by
a 4-merous or 5-merous “skin”.

a. Detecta (hypanthium) without adnate bracteoles; petals present or
bre

Petals rei (in Panama species) 1. COMBRETUM
bb. Petals abse
ruits sdi ry, not closely aggregated.
d. ree versatile.
Calyx michel tensili upper meet. nad calyx caducous

"a nama es) before the fruit rip 2. TERMINALIA
ee. Calyx, only slightly toothed; upper r receptacle and calyx per-
sistent on the fruit or only tardily dee ————— Ü—
E a ers adnate to the filaments, im ile
. Fruits aggregated into a cone-like structure
aa. Receptacle Agli thium) with two small adnate bracteoles near the `
present

BUCIDA
. BUCHENAVIA
NOCARPU

LA >

6. LAGUNCULARIA

TÉ o d genus Quisqualis (receptacle without adnate bracteoles; petals
present) is separated from Combretum by the adnation of the style for part of its
length to the wall of the upper receptacle (calyx-tube). The species cultivated,
Q. indica, can at once be distinguished from all the indigenous Combretaceae by
its much elongated, narrowly infundibuliform upper receptacle (calyx-tube), UP
to 8 cm. long when the flowers are fully developed.

1. COMBRETUM Loefl.
CoMBRETUM Loefl. Iter. Hispan. 308. 1758, nom. conserv.

Grislea L. Sp. Pl. 348. E > phe 164. 1754.
Aetia Adans. Fam. 2:84,

a rgera z RZ
Cristaria Sonner. Voy. Ind. Orient. 2:247. f. = 1782.
Hambergia Neck. Elem. Bot. 141. 1790.
Schousboea Willd. in L. Sp. Pl. 2:578. 1799. Obs.
Poivrea Comm. [ex ż Nae a ida 320. 1789, in syn. (sphalm. “Pevraea”)] ex Thou.
A Il. Austr. Afr. 28.
Gonocarpus Ham. Prodr. pl. "Ind. Coc. 39. 122
Forgaria Vell. FI. Flum. 4:152. 1825; ^ 13: un

(144)

1958]
FLORA OF PANAMA (Combretaceae) 145

Chrysostachys Pohl, Fl. Bras. 2:65. £. 143.
Sheadendron Bertol. f. in Mem. Acad. dola. $ :574. E. 4. 1850.
Ii Jena Bl. Mus. Lugd. “i Hun

Świcie Reichb. ex Oliv. Fl. Tro day k > 424. 1871 in syn., - Adans. (1763).
Campylochiton Welw. ex Hiern, im Afr. Pl. Welw. 2:353. 189

Trees, shrubs or woody climbers, very rarely subherbaceous. Leaves opposite,
verticillate or more rarely alternate, usually petiolate, almost always entire, often
conspicuously lepidote. Petioles sometimes persisting after leaf-fall, forming
thorns. Flowers usually perfect, actinomorphic or rarely slightly zygomorphic, in
elongated or subcapitate, axillary or extra-axillary spikes or racemes or in terminal
or terminal and axillary, often leafy panicles. Receptacle usually clearly divided
into a lower part (lower receptacle) surrounding and adnate to the ovary and an
upper part (upper receptacle) varying from patelliform to elongate-infundibuli-
form terminating in the calyx-lobes. Upper receptacle often further differentiated
into a lower part containing the disk and a usually more expanded upper part.
Calyx-lobes usually 4 or 5 (rarely more) deltoid to filiform or sometimes scarcely
developed. Petals usually 4 or 5 (very rarely absent but never in Panama species
except perhaps by abortion) small and inconspicuous or showy and exceeding the
calyx-lobes, white, yellow, orange, red or purple, glabrous or hairy, occasionally
scaly. Stamens usually twice as many as the petals, inserted inside the upper
receptacle, often biseriate, usually exserted; anthers dorsifixed, versatile. Disk
intrastaminal, glabrous or hairy, sometimes very small or absent. Ovary com-
pletely inferior, probably of 1 carpel, unilocular with usually 2 (occasionally up
to 6) pendulous ovules of which only 1 develops. Style simple, free, usually ex-
serted, rarely very short. Fruit (pseudocarp) 4- to 5-winged, -ridged or -angled,
sessile or stipitate, usually indehiscent, 1-seeded; pericarp usually thin and papery,
sometimes leathery, more rarely fleshy.

ut 250 species throughout the tropics (except Australia) ; extending in the

New World from Mexico to Argentina.

The Panama species are classified as follows:

Sect. EucoMBRETUM Don.: C. fruticosum (Loefl.) Stuntz; C. sambuense Pittier
Sect. CACOUCIA (Aubl.) Engl. & Diels.: C. cacoucia Exell.

Sect. COMBRETASTRUM Eichl.: C. laxum Jacq.

Sect. SPINOSAE Exell.: C. decandrum Jacq.; C. spinosum Bonpl.

orze

»

. oo meni
Flow rs measuring at least 7 mm. from the ~~ to the tips of the
calyx-lobes, conspicuous "wę scaly; disk well developed, margin pilose
- Scales of inflorescence eign golde-brown (in dried Lins
Petals rarely more than 1.5 mm. long 1. C. FRUTICOSUM
c= a; a of inflorescence red (in dad specimens). Petals 4-5 mm.

2. C. SAMBUENSE

me gla broas. | — .— 0057007 4. C. LAXUM
pen

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[Vol. 45 |
146 ANNALS OF THE MISSOURI BOTANICAL GARDEN i

„45. Combretum: Diagrams of Flower Structure. (a) C. wywi M: c cacoschi
e T — (d) C. spinosum; (e) & (f) C. decandrum. By courtesy of Society
Lon

(146)

1958]

FLORA OF PANAMA (Combretaceae) 147
c. Lower receptacle, nearly glabrous or patent-pubescent, slightly
pedicellate and narrowed at the apex 5. C. DECANDRUM
cc. Lower receptacle appressed-pubescent, sessile, scarcely narrowed

at the apex 6. C. SPINOSUM
bb. Flowers slightly zygomorphic, relatively large, up to 3 cm. from the
rhachis to the tips of the calyx-lobes. Fruit narrowly winged or
ridged 3. C. CACOUCIA

1. COMBRETUM FRUTICOSUM (Loefl.) Stuntz, in U. S. Dept. Agric. Bur. Pl. Ind.
Seeds & Pl. Import. no. 31:86. 1914.

Gaura fruticosa Loefl. Iter. Hispan. 248. 1758.
Combretum occidentale L. Syst. Nat. 2:999. 1759, nom. illegit.
Combretum secundum Jacq. Enum. Syst. Pl. Ins. Carib. 19. 1760.
Combretum laxum sensu L. Sp. Pl. 1:496. 1762, pro parte, non Jacq.
Combretum formosum Don, in Trans. Linn. Soc. Lond. 15:420. 1827.
Combretum micropetalum DC. Prodr. 3:19. 1828

ombretum aurantiacum Benth. in Hook. Lond. Journ. Bot. 2:222. 1840
Combretum loeflingii Eichl. in Mart. Fl. Bras. 14?:110. 1867, nom. illegit.
Combre j i it. 1867.
Combretum benthamianum Heurck & Mill. Arg. Obs. Bot. 220. 1870.
Combretum gloriosum Rusby, in Mem. Torr. Bot. Club 6:35. 1896.

u

Combretum lepidopetalum Pittier, loc. cit. 245. 1917.

Combretum multidiscum Rusby, Descr. New Sp. S. Amer. Pl. 69. 1920. i

Combretum loeflingii subsp. ornithophilum Suesseng. in Mitteil. Bot. Staatssamml. Münch.
14. 1950.

For an explanation of some complicated points in the synonymy see Exell, in
Journ. Linn. Soc. Lond. 55:117. 1953.

Shrub of 3-8 m. or liana, branchlets somewhat quadrangular, scaly at first but
otherwise glabrous (in Panama specimens). Leaves papyraceous or chartaceous,
opposite, petiolate; petioles scaly, 7-10 mm. long; lamina elliptic to broadly elliptic,
5-12 X 3-7 cm., minutely verruculose above, conspicuously scaly beneath, scales
usually golden or golden-brown, occasionally reddish, dense but scarcely marginally
contiguous, usually glabrous or nearly so in Panama but variously hairy in other
regions, rather bluntly acuminate at the apex, cuneate or rounded at the base.
Flowers yellow or greenish-yellow, 4-merous, sessile or subsessile in terminal panicles
of spikes and lateral spikes, rhachis scaly. Lower receptacle (ovary) quadrangular,
2-3 mm. long, densely scaly; upper receptacle clearly differentiated into two parts,
a lower narrowly infundibuliform part containing the disk and an upper broadly
infundibuliform or campanulate part terminating in the calyx-lobes, measuring
about 7-8 mm. from the apex of the ovary to the tips of the calyx-lobes, with
Conspicuous golden or golden-brown (rarely reddish) scales otherwise nearly

rous (in Panama). Calyx-lobes broadly ovate, about 2 mm. long. Petals
yellow, usually glabrous, very variable in size and shape but usually not more than
1-1.5 mm. long, shorter than the calyx-lobes. Stamens 8, filaments 15-20 mm.
long, anthers about 1 mm. long. Disk infundibuliform with a pilose margin. Style
15-20 mm. long, glabrous. Fruit subspherical in outline, 14-18 mm. in diameter,
reddish-brown or purple, with 4 flexible wings about 6 mm. wide, scaly, shortly
stipitate,

(147)

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

148

So
„e. TEEN
< IAS /

AU
SN
\ SY Y
SS
SS

>

— s
==

AOS
S

„SUN

A

ZA ER
La M

WA

^

Combretum fruticosum

Fig. 46.

(148)

1958]
FLORA OF PANAMA (Combretaceae) 149

Widespread in the New World from Mexico to Argentina.

For an account of the variability of this species see Exell, in Journ. Linn. Soc.
Lond. 55:121. fig. 3. 1953.

Specimens of this species, as here delimited, have often been named C. farinosum
Kunth, a species which, in my opinion, extends only from Mexico to Costa Rica.
It is not easy to give cenventional key characters to separate these two species,
especially in view of the wide range of variation I have included in C. fruticosum.
Unfortunately I have never had the opportunity of examining living specimens,
but really well-preserved herbarium specimens of the two species are quite different
in appearance. In C. farinosum the flowers (receptacles) look silvery-grey, the
petals dry red or dark red, the golden or reddish filaments make a considerable
contrast in color with the receptacle and the dried anthers are about 1.5 mm.
long. In C. fruticosum the inflorescences of modern, carefully prepared specimens
are of a greenish golden-brown color (reddish in older specimens), the petals dry
yellow and the dried anthers do not exceed 1 mm. in length.

It is possible to have a merry time making ‘species’ out of this complex for
there is an extraordinary range of variation in the petals and in the indumentum.
Add to this a number of ancient nomenclatural tangles and it is easily understand-
able how this species became so richly endowed with synonyms.

COCLÉ: between Aguadulce and Río Chico, Pittier 5006; near Penonomé, Williams 72.
CANAL ZONE: near Cristóbal, Colón, Broadway 162, 589; near Salamanca Hydrographic

The specimen collected by P. & G. White, at Miraflores, is noted as being a
‘large tree. This may be an error.

2. COMBRETUM SAMBUENSE Pittier, in Contrib. U. S. Nat. Herb. 18:242. 1917.

Woody liana, branchlets glabrous, slightly quadrangular. Leaves chartaceous
to subcoriaceous, opposite, petiolate; petiole 5-10 mm. long, scaly; lamina ovate to
broadly elliptic, 11-18 X 5.5-9.5 cm., glabrous, minutely verruculose above with
conspicuous golden-yellow scales beneath, scales rarely marginally contiguous,
usually acuminate at the apex and rounded at the base. Flowers reddish or purpli h,
4-merous, sessile, in terminal panicles of spikes and lateral axillary spikes, rhachis

nsely rufous-scaly. Lower receptacle (ovary ) quadrangular, 5 mm. long, densely
rufous-scaly; upper receptacle fleshy, broadly infundibuliform, slightly narrowed
towards the base, measuring about 10 mm. to the tips of the calyx-lobes, densely
rufous-scaly outside, villous within. Calyx-lobes ovate, 4-5 mm. long. Petals 4,
yellow, broadly elliptic, acute, 4-5 mm. long, glabrous, slightly shorter than the
calyx-lobes. Stamens 8, inserted at the margin of the disk, filaments 25-35 mm.

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[Vol. 45
150 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long, anthers 2 mm. long. Disk infundibuliform, pilose at the margin. Style 35-40
mm. long, glabrous. Fruit crimson, suborbicular, 1.8—2.5 cm. in diameter, scaly
and Meus. pubescent.

N: near Pinogana, alt. 20 m., Allen 4275; Río Sambú, Pittier 5548; Tuculi, Che-
= = 16 m., Terry & Terry 1375.
3. COMBRETUM CACOUCIA Exell, apud Sandw. in Bull. Misc. Inf. Kew, 1931:469.

193%;

Cacoucia coccinea Aubl. Pl. Guian. 1:450. £. 179. 177
Schousboea coccinea (Aubl.) Willd. in L. Sp. PI. Lo Ee
Warsa cacoucia Baill. Hist. Pl. 6:275. 1877, nom. illeg

retum coccineum (Aubl.) Engl. & Diels, in Engl. Mow, Afr. Pflanz. -Fam. & -Gatt.
eo 110 1899, nom. illegit., non Lam. (1785
Shrub or liana, branchlets rufous-pilose. Leaves chartaceous, opposite, petiolate,
petiole 5—8 mm. long, pilose, lamina broadly elliptic to oblong-elliptic, up to 20 X
10 cm., often pilosulose or pubescent on the midrib and principal nerves beneath,
otherwise glabrous or nearly so, not conspicuously lepidote, acuminate at the apex
and subcordate at the base, lateral nerves 7—8 pairs rather widely spaced. Flowers
red, 5-merous, slightly zygomorphic, shortly pedicellate, pedicel 3-5 mm. long
tomentellous, in stout terminal racemes up to 40 cm. long, rhachis tomentellous or
velutinous, bracts ovate-acuminate or lanceolate, up to 15 mm. long, caducous.
Lower receptacle (ovary) tomentellous, about 2.5 mm.; upper receptacle
(calyx-tube) campanulate or broadly infundibuliform, slightly curved, up to 15
X 11 mm. sericeous outside. Calyx-lobes triangular, up to 5 mm. long. Petals 5,
ovate or elliptic, mucronate, 10 X 7 mm., minutely pubescent on the outside or
nearly glabrous. Stamens 10, exserted, filaments about 20 mm. long, anthers E
1.5 mm. long. Disk well-developed with margin produced into an annular ridge
which nearly closes the upper receptacle. Style 25 mm. long, glabrous. Fruit
narrowly oblong-ellipsoid, longitudinally 5-ridged, up to 6 X 2 cm., velutinous at
first, eventually nearly glabrous.
British Honduras to Brazil.
BOCAS DEL TORO: Water Valley, Von Wedel 1628, 1902; Old Bank Island, Von Wedel
1979; Fish Creck lowlands, Von Wedel 2373. CANAL ZONE: Majunga Swamp, neighbor-
ood of Río Chagres, Allen 854; Gatün, Hayes 8, s. n.; Lion Hill Station, Hayes 545
atún, Heriberto 110; Rio Chagres, below Ga atún, Maxon 4798; Frijoles, pee 2692.
COLÓN: Camp Pińa, 0-50 m., Allen 3432; Río Culebra, above Sta. Isabel, Pitt er 4169.
For a discussion of the significance of the geographical distribution of Com-
bretum Sect. CACOUCIA, to which this species belongs, see Exell, in Journ. Linn:
Soc. Lond. 55:104. 1953.

4. COMBRETUM LAXUM Jacq. Enum. Pl. Carib. 19. 1760.
Combretum puberum Rich. in Act. Soc. Hist. Nat. Par. 1:108. 1792.
Combretum obtusifolium Rich. loc. cit. 1792.
Combretum mexicanum Bonpl. Pl. SEPE 2:159, 1, 132. 1809:
ta, ss laevis Vell. Fl. Fiam. 152. 1825.

m odoratum Pav. ex boa i Tees Ba Soc. Lond. 15:430. 1827.
dA cordatum Don, loc. cit. 440. 1827.
Combretum ferrugineum Don, lx cit. 1827.

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1958]
FLORA OF PANAMA (Combretaceae) 151

Combretum bugi Cambess. in St.-Hil. Juss. & Cambess., Fl. Bras. Merid. 2:247. £. 130.

Chrysostachys ovatifolia Pohl, Pl. Bras. 2:66. £. 143. 1831.
ombretum variabile Mart. in Flora 22, Beibl. 1:62. 1839.

Combretum pulchellum Mart. loc. cit. 64. 1839

Combretum ferrugineum Hoffmansegg, ex Mart. loc. cit. 1839.

. 1843.
Combretum jacquinii Griseb. Fl. Brit. W. Ind. 275. 1860, nom. illegit.
Combretum viscidum W right, ex Griseb. Cat. PI. Cub. 109. 1866.

Combretum accedens Heurck & Muell. Arg. in Heurck. Obs. Bot. 234. 1870.

Combretum odoratissimum Sessć & Moc. Fl. Mex. 90. 1894.

Combretum epiphyticum Pittier, in Contrib. U. S. Nat. Herb. 18:247. 1917.

Combretum oblongifolium Rusby, Descr. New Sp. S. Amer. Pl. 70. 1920.

Combretum marchii Fawc. & Rendle, in Journ. Bot. 63:115. 19

Combretum brunnescens Gleason, in Bull. Torrey Bot. Club 53:291. 1926.

The infra-specific taxa have been omitted. They are cited by Exell, in Journ.
Linn. Soc. Lond. Bot. 55:127. 1953.

Liana (sometimes a shrub ?), branchlets rufous- or tawny-tomentose to almost
glabrous. Leaves chartaceous to subcoriaceous, opposite, petiolate, petioles 2-7 mm.
long, tomentose to almost glabrous, lamina oblong or oblong-elliptic to broadly
ovate or suborbicular, 6-18 X 3-7 cm., not conspicuously scaly, varying from
tomentose to nearly glabrous, usually acuminate sometimes subcaudate at the apex,
rounded or subcordate at the base. Flowers white or yellow, small, fragrant, sessile,
4-merous, in terminal or terminal and axillary panicles, rhachis rufous- or fulvous-
tomentose, tomentellous or pubescent. Lower receptacle 1-1.5 mm. long, tomen-
tose, tomentellous, pubescent or nearly glabrous; upper receptacle cupuliform, 1-2
X 1.5-2.5 mm., densely pubescent to nearly glabrous. Calyx-lobes deltoid, about
0.7 mm. long. Petals 4, broadly obovate to transversely elliptic or subreniform,
about 1 mm. in diam., glabrous, exceeding the calyx-lobes. Stamens 8, filaments
about 4 mm. long, anthers 0.5 mm. long. Disk small, glabrous. Style 2.5-4 mm.
long. Fruit oblong, narrowly ovoid or suborbicular, 4-ridged or -winged, 2.4 X
1.5-2 cm., pubescent or glabrous.

Widespread in the tropics and subtropics of the New World from Mexico to
northern Argentina.

S 7, s. n.; Frijoles, Hayes 54, s. n.; Cano Quebrado, Pittier 6668, 6819; Rio Chagres,

CANAL ZONE: Quebrada Tranquilla, alt. 70-80 m., Dodge & Allen 17501; near Gatún,
ayes 7, s. H A
Steyermark & Allen 16774. PROVINCE UNKNOWN: Hayes 771; Seemann I 658.

As in the preceding species a very broad view has been taken here. Those who
wish for a narrower delimitation of the species will find names available in the
synonymy for at least many of the taxa they may wish to recognize. In Panama,
Dodge & Allen 17501, Seemann 1658 and Steyermark 6 Allen 16774 on the one
hand and Hayes 54 on the other represent extremes of variation and could be
separated specifically with the greatest of ease. The three former specimens, in
the event of such segregation, would seem to agree well with the Mexican material
of C. mexicanum Bonpl. It is only when the species is studied over the whole
hemisphere, especially with regard to the extraordinary range of variation in Brazil,

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[Vol. 45
152 ANNALS OF THE MISSOURI BOTANICAL GARDEN

that the problem arises. The present conception of the species is, however, more a
temporary convenience than a final solution. It is necessary to have far more
fruiting material, at present very poorly represented in the herbaria. Included in
the ‘species’, in its present conception are both narrowly angled and quite broadly
winged fruits. If these differences could be correlated in any way with differences
of leaf-shape and indumentum we should be on the way towards a more satisfactory
system. Judging from what occurs in other species of Combretum and Terminalia
in other parts of the world, one would expect to find ridged and angled fruits in
flooded regions, along river-banks, etc. where distribution is by water, while species
with broad-winged fruits are found in hill-slope and mountain forests, savannas
etc. Field observations are badly needed.

5. COMBRETUM DECANDRUM Jacq. Enum. Pl. Carib. 19. 1760.

Combretum alternifolium Pers. Syn. 1:412. 1805, nom. illegi
Gonocarbus jacquinii Hamilt. Prodr. Fl. Ind. Occ. 39. 1825, nom. illegit.

Combretum nicoyanum Pittier, loc. cit. 18:247.
Combretum idrico A Rusby, Descr. New Sp. ^ Amer. Pl. 69. 1920. È
Liana, branchlets pubescent or glabrous, spiny. Leaves alternate or sometimes
subopposite, papyraceous, petiolate, petiole 2-5 mm. long, pubescent to tomentose,
persistent and somewhat accrescent forming a spine, lamina elliptic to broadly
elliptic, 5-14 X 3-7.5 cm., pubescent, especially on the nerves beneath, or nearly
glabrous, often with hairy domatia in the axils of the nerves beneath, bluntly
acuminate at the apex, blunt, rounded or subcordate at the base. Flowers white,
subsessile or shortly pedicellate, 5-merous, in large terminal panicles, often with
reduced leaves at the points of branching, rhachis tomentose, pubescent or glabrous.
Lower receptacle about 1 mm. long, glabrous, nearly glabrous or loosely patent-
pubescent, usually somewhat constricted at the apex and often with a short pedicel
up to about 0.5 mm. long; upper receptacle campanulate to cupuliform, 2.5-3 X
1.5—2 mm., glabrous. Calyx-lobes 5, broadly deltoid, about 0.5 mm. long. Petals
5, obovate or obovate-elliptic, 2-3 X 1.5—1 mm., hairy. Stamens 10, filaments
3-5 mm. long, anthers 0.4 mm. long. Disk small, fleshy, glabrous, s surrounding
the base of the style. Style about 4 mm. long. Fruit suborbicular in outline, 1.5
cm. in diam., usually glabrous, 5-winged, wings thin, flexible, 5-6 mm. broa d
From Mexico to Colombia.
ANAL ZONE: Victoria Fill, near Miraflores Locks, Allen 1746; Empire Station, Hayes
is near Gorgas Memorial Laberstocy, P. White 82. DARIEN: Patino, Pittier 6607.

6. COMBRETUM sPINOSUM Bonpl. PI. Aequinox. 2:161. 1809.
Combretum guayca Humb. Rel. Hist. 3:3. gi nom. nud.
Poivrea eriopetalum DC. Prodr. 3:18
Combretum eriopetalum (DC.) Don Gai S» 2:665.
Combretum punctulatum Pittier, in Contrib. U. S. Nat. "Herb. 18:248.
This differs from the preceding species only in having baee dier pa:
flowers with the lower receptacle (ovary) appressed-pubescent and scarcely

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1958]
FLORA OF PANAMA (Combretaceae) 153

stricted at the apex. One specimen, from Venezuela, with 4-merous flowers has
en seen.

Panama, Venezuela and in the West Indies from Cuba to Trinidad.

CANAL ZONE: Rio Trinidad, Pittier 4002.

It may prove necessary to unite this with C. decandrum when more material
becomes available. Allen 1746, placed under C. decandrum, which I have now
seen for the first time, has some appressed pubescence on the lower receptacle
though it agrees in other respects with C. decandrum. The distribution areas of
the two species, if such they be, overlap in Panama and there may well have been
some hybridization.

2. TERMINALIA L.

TERMINALIA L. Syst. Nat. 2:674 (err. 638). 1767; Mant. Pl. 21. 1767, nom.
conserv.

Adamaram Adans. Fam. 2:445. 1763, proparte excl. Hort. Malab. 4: 7. 5. 1682.

Panel Adans. loc. cit. 447. 1763, pro parte quoad Hort. Malab. 4: £. TO. 1682.

Myrobalanifera Houtt. Handleid. Pl. Kruidk. 2:485. £. 10, fig. 2. 1774.

Tanibouca Aubl. Pl. Guian. 1:448. t. 178. 1775.

Pamea Aubl. loc. cit. 2:946. t. 359. 1775.

Aristotelia Comm. ex Lam. Dict. Encycl. Bot. 1:349. 1783.

Fatraea Juss. in Ann. Mus. Par. 5:223. $

Pentabtera Roxb. [Hort. Beng. 34. 1814 nom. nud.] Fl. Ind. 2:437. 1832.

Vicentia Allem. PI. Nov. Bras. cum tab. 1844.

Chichharronia A. Rich. Ess. FI. Cub. 529. £. 43. 1845.

i Trees, sometimes of great stature, frequently buttressed, rarely shrubs, branch-
ing often sympodial. Leaves usually spirally arranged, often crowded in pseudo-
whorls at the ends of the branchlets, usually petiolate, entire, often minutely
verruculose and pellucid-punctate, often with domatia, frequently with two or
more glands at or near the base of the lamina or on the petiole. Flowers actino-
morphic, 5-merous (rarely 4-merous) usually in axillary spikes with staminate
flowers towards the apex and perfect flowers towards the base of the spike, or all
perfect, more rarely in terminal or terminal and axillary panicles; staminate flowers
stalked, stalks resembling pedicels but equivalent to the lower receptacle with
abortion of the ovary; perfect sessile. Receptacle divided into a lower part (lower
receptacle) enclosing and adnate to the ovary and often narrowed above it, and an
Upper part, often scarcely developed, expanded into a shallow cup terminating in
the calyx-lobes. Calyx-lobes deltoid, ovate or triangular. Petals absent. Stamens
usually 10, exserted; anthers dorsifixed, versatile. Disk intra-staminal, usually
pilose or barbate, sometimes glabrous or poorly developed. Style simple, free,
exserted. Ovary completely inferior, unilocular with 2 (rarely 3-4) pendulous

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154

ANNALS OF THE MISSOURI BOTANICAL GARDEN

A.

udi
U
Li T 7 4
X p t ad

Fig. 47. Terminalia cbiriquensis

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[Vol. 45

1958]
FLORA OF PANAMA (Combretaceae) 155

ovules of which only one develops. Fruit (pseudocarp) very variable in size and
shape, often fleshy and drupe-like, sometimes dry and leathery or corky, often 2-
to 5-winged, usually with the endocarp at least partially sclerenchymatous (thus
distinguishing it from Combretum).

About 200 species throughout the tropics.

In addition to the two species listed below, T. catappa L., an Asiatic and Poly-
nesian species with large obovate leaves and drupaceous fruits, is commonly planted.
a. Leaves obovate and abruptly acuminate. Fruit 5-winged, two wings

transversely elongated and much longer than the others 1. T. AMAZONIA
eaves elliptic. Fruit with 2 broad lateral and 1 inconspicuous carinate
adaxial wings 2. T. CHIRIQUENSIS

l. TERMINALIA AMAZONIA (J. F. Gmel.) Exell, in Pulle, FI. Surin. 3:173. 1935.

>»
LE

Chuncoa amazonia J. F. Gmel. in L. Syst. Nat. 2:702. 1791.

Gimbernatea obovata Ruiz & Pav. Fl. Peruv. Chil. Prodr. 138. 1794.

Chuncboa obovata (Ruiz & Pav.) Pers. Syn. 1:486. 1805.

Chuncoa obovata (Ruiz & Pav.) Poir. in Encycl. Méth. Bot., Suppl. 2:258. 1811.

Terminalia obovata (Ruiz & Pav.) Steud. Nomencl. Bot. 2:668. 1841, non T. obovata
Cambess. (1829).

Terminalia odontoptera Heurck & Müll. Arg. Obs. Bot. 217. 1870.

Myrobalanus obovatus (Ruiz & Pav.) O. Kuntze, Rev. Gen. 237. 1891.

Terminalia bayesii Pittier, in Contrib. U. S. Nat. Herb. 18:239. 1917.

Large tree up to 30 m. high with buttresses and sympodial branching. Branch-
lets rufous-pilose or rufous-sericeous, soon glabrescent. Leaves chartaceous to
subcoriaceous, spirally arranged in pseudo-verticels at the ends of the branchlets,
petiolate, petiole 1-10 mm. long, rufous-pubescent, lamina obovate or cuneate,
abruptly acuminate at the apex, cuneate at the base usually with two glands, 5-10
X 3.5 cm., usually rather sparsely appressed-pubescent or appressed-pilose with
hairy domatia in the axils of the principal veins beneath. Flowers white, cream or
yellowish-green, apparently all perfect, 5-merous, sessile, in axillary spikes 6—7 cm.
long, rhachis tomentellous. Lower receptacle 2-2.5 mm. long, sericeous; upper
receptacle shallow-cupuliform, 0.8 X 2 mm., appressed-pubescent. Calyx-lobes
deltoid, 0.8 mm. long, usually not reflexed. Stamens 10, filaments 1.8-2 mm. long,
anthers subglobose, 0.4 mm. in diam. Disk densely pilose. Style 2 mm. long,
glabrous. Fruit about 5 X 10 mm., sparsely pubescent, 5-winged, two wings
transversely elongated to about 5 mm., the other three usually rudimentary (or
vestigial) not more than 1 mm. in width.

From Mexico to Guiana, Trinidad, Brazil and Peru. The wood is said to be
very hard and of good quality.

CANAL ZONE: Victoria Fill, near Miraflores Lock, Allen 1754; Paraiso Station, Hayes
718, s. n. PANAMA: near Cerro Jefe, alt. 600 m., Allen 3430.

In most American botanical works this species is called Terminalia obovata
(Ruiz & Pav.) Steud (1841) but this name is preoccupied by T. obovata Cambess.
(1829). The epithet is also antedated by Chuncoa amazonia J. F. Gmel.

2. TERMINALIA CHIRIQUENSIS Pittier, in Contrib. U. S. Nat. Herb. 18:238. 1917.
Tree up to 30 m. tall. Branchlets rufous-sericeous when young, soon glabres-

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[Vol. 45
156 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 48. Bucida buceras

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1958]
FLORA OF PANAMA (Combretaceae) 157

cent. Leaves spirally arranged, often pseudo-whorled at the ends of the branchlets,
membranaceous to chartaceous, petiolate, petiole 2-18 mm. long, rufous-sericeous
when young, lamina elliptic, 10-18 X 4-6 cm., rufous-sericeous when young and
retaining some pubescence on the nerves when adult, otherwise nearly glabrous,
minutely verruculose above, pellucid-punctate at least in some stages, acuminate
at the apex, cuneate at the base. Flowers yellowish-green, sessile, apparently all
perfect, in axillary spikes 8-11 cm. Fruit transversely oval, about 1.5-2.0 X
2.5-3.5 cm., pale greenish yellow, glabrous, subsessile, samaroid, 3-winged, the 2
lateral very broad and pronounced, about 1.5 cm. wide, the 1 adaxial merely
carinate and less than 1 mm. wide.

Panama, Costa Rica.

BOCAS DEL TORO: Almirante, Cooper 6 Slater 55, Seibert 1534.

The description of the fruit is taken from Skutch 4252 (MO), collected in the
vicinity of El General, Prov. San Josć, Costa Rica, the flowering companion sheet

of which (Skutch 3990, MO) agrees well with Seibert 1534 (MO).

3. BUCIDA L.

Bucma L. Syst. Nat. 2:1025. 1759, nom conserv.
Buceras P. Br. Hist. Jam. f. 23. 1756, nom rejic.

Trees. Leaves spirally arranged, usually pseudo-whorled. Upper receptacle
remaining attached to the fruit and calyx-lobes scarcely developed. Fruit not
winged, slightly oblique. Other characters as in Terminalia.

wo or three species from Florida, Central America, Guiana (fide Pulle) and
in the West Indies.

Grounds for separating this genus from Terminalia are slender; Bentham and
Hooker and a few other authors have not maintained it. The apparently ex-
cellent character of the persistent upper receptacle suffices adequately to distinguish
the two genera in the New World but one or two Malayan-Polynesian species of
Terminalia, quite unrelated to Bucida, show the same feature. There seems, how-
ever, no grave disadvantage in maintaining the status quo.

1. Bucipa BUCERAS L. Syst. Nat. 2:1025. 1759.

Tire bucida Crantz, Inst. Rei Herb. 1:133. 1766.
Fils ires buceras (L.) Wright, in Sauv. Fl. Cub. 38. 1868.
yrobalanus buceras (L.) O. Kuntze, Rev. Gen. 237. 1891.

Medium-sized tree, 5—20 m. high. Branchlets at first sericeous, sometimes
spiny (but apparently rarely in Panama); growth sympodial Leaves coriaceous,
Pseudo-whorled at the tips of the branchlets, shortly petiolate, petiole 2-10 mm.
long, sericeous, eventually glabrescent, lamina elliptic, narrowly elliptic or obovate-
elliptic, 2-9 X 1-4 cm., pilose-sericeous when young, later appressed-pilose,
appressed-pubescent or nearly glabrous, blunt or rounded at the apex, cuneate at
the base and often with two not very conspicuous glands. Flowers pale green (or

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[Vol. 45
158 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 49. Buchenavia cabitata

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1958]
FLORA OF PANAMA (Combretaceae) 159

violet?), 5-merous, sessile, mostly perfect, in axillary spikes 2-8 cm. long, rhachis
appressed-pubescent. Lower receptacle 1—1.5 mm. long, sericeous-tomentose;
upper receptacle shallow-cupuliform, 1 X 2.5 mm., appressed-pubescent, persistent.
Calyx-lobes little developed. Stamens 10, filaments 4 mm. long, anthers 1 mm.
long. Disk 5-lobed, round the base of the style. Style 3.5 mm. long, glabrous.
Fruit ovoid or pyriform, up to 4-6 X 2.5—4.5 mm., slightly and somewhat un-
equally 5-lobed, often slightly contorted, not winged, densely appressed-pubescent,
crowned by the persistent upper receptacle (calyx), often galled.

Florida to Guiana (fide Pulle) and in the West Indies.

o: southwest of Bocas at Macaw Hill, bois Wedel 5573 ag of
Chiriquí pasti Shia Colón, Vos Wedel 2929. CANAL ZONE: Aspinwall, H.

This species is very variable in size and shape of leaf dA presence or y. of
spines. As far as can be judged from the small amount of material seen, it is
mainly the larger-leaved spineless form which occurs in Panama. In the West
Indies forms with very much smaller leaves and well developed spines occur and
Bucida spinosa Jennings (Terminalia spinosa Northrop, non Engl.) from the Ba-
hamas, with leaves only about 1 cm. long and numerous spines, has been separated
as a distinct species. The sterile specimen Von Wedel 557 from Macaw Hill has
narrow leaves and one or two spines. It is described, possibly erroneously, as having
violet flowers.

he timber is said to be useful.

4. BUCHENAVIA Eichl.

BUCHENAVIA Eichl. in Flora 49:164. 1866.
Hudsonia A. Robinson, ex Lunan, Hist. Jam. 2:310. 1814, not L. (1767).

Trees or shrubs. Leaves alternate or pseudo-whorled. Flowers 5-merous,
Perfect and staminate together in elongated or subcapitate axillary spikes. Petals
absent. Stamens 10, anthers adnate to the filaments, immobile. Fruit fleshy, 5-
ridged or 5-angled, endocarp sclerenchymatous.

12-15 species, throughout tropical America.

+ BUCHENAVIA CAPITATA (Vahl) Eichl. in Flora 49:164. 1866.

kał cabitata ur Eclog. Amer. 1:50. £. 8, fig. I
Hudsonia arborea A. Robinso inson, ex Lunan, Hist. Jam. 2:310. 1814
Terminalia obova ta Cambess. in St.-Hil. Juss. vignes Fl. Bras. Merid, 2:241. 1829.
Terminalia — Steud. Nomencl. Bot. 2:668. 184

ree about 8 m., branchlets iii when very young, soon
ia growth sympodial. Leaves pseudo-whorled at the tips of the branch-
lets, duttécspns to subcoriaceous, petiolate or subsessile, petiole up to about 6 mm.
long, rufous-pilose at first, later glabrescent, lamina obovate-cuneate, usually
rounded at the apex and decurrent into the petiole, 2-9 X 1—3.5 cm., tomentose
when young becoming glabrous. Flowers mostly perfect, 5-merous, sessile, in sub-
Capitate spikes, peduncles 1.5—7 cm. long, rufous-tomentose when young, glabres-
Cent. Lower receptacle 1-3 mm. long, narrowed and slightly twisted at the apex,

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[Vol. 45
160 ANNALS OF THE MISSOURI BOTANICAL GARDEN |
|

RE ELA
SEN
oes NA ARE

Fig. 50. Conocarpus erectus

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1958]
FLORA OF PANAMA (Combretaceae) 161

densely sericeous; upper receptacle shallow-cupuliform, 1 X 2.5 mm., sparsely
pubescent or nearly glabrous. Calyx-lobes obsolete. Petals absent. Stamens 10,
biseriate, filaments about 2 mm. long, anthers about 0.6 mm. long, adnate to the
filaments. Disk patelliform with a free, densely pilose margin. Style subulate, 3.5
mm. long, glabrous. Fruit greenish-yellow (black when dried), ellipsoid, somewhat
flattened, more or less 5-ridged, 2-3 X 1-1.5 cm., mesocarp fleshy, endocarp hard
and stony.
Panama to Bolivia and northern Brazil, and in the West Indies.

PANAMA: Rio La Maestra, 0—25 m. alt., Allen 35.

5. CONOCARPUS L.
CoNocanPus L. Sp. Pl. 1:176. 1753; Gen. Pl. 81. 1754.
Rudbeckia Houst. ex L. loc. cit. 1754, in syn. non L. (1753).
Rudbeckia Adans. Fam. 2:80, 599. 1763, nom. illegit. non L.

Small trees or shrubs, erect or prostrate. Leaves alternate. Flowers sessile, 5-
merous, perfect and staminate in the same inflorescences, densely aggregate into
pseudo-capitula. Petals absent. Stamens usually 10, sometimes fewer by abortion,
anthers dorsifixed, versatile. Disk intrastaminal Ovules 2, pendulous. Fruits
(pseudocarps) laterally compressed, 2-winged, densely aggregated in a cone-like
mass.

A genus consisting of two littoral species, often constituents of mangrove
swamps, one in tropical and subtropical America and Africa, the other in Somali-
land and Arabia.

1. CONOCARPUS ERECTUS L., Sp. PI. 1:176. 1753, Var. ERECTUS.

Conocarpus procumbens L. loc. cit. 177. 1753.

Conocarpus supinus Crantz, Inst. Rei Herb. 1:355. 1766.

Conocarpus acutifolius Humb. & Bonpl. ex Roem. & Schult. in L. Syst. Veg. 5:574. 1819.

Conocarpus erectus var. arboreus et procumbens DC. Prodr. 3:16. 1828.

Fanqrerpai pubescens Schumach. in Kongel. Dansk. Vid. Selsk. Naturvid. & Math. Afh.
315

Terminalia ericta (L.) Baill. Hist. PI. 6:266, 275. fig. 240. 1877.

This has been placed under var. erectus for nomenclatural reasons, not with any
intention of considering that the erect form differs from the prostrate form but
because var. sericeus (not occurring in Panama) may be a distinct taxon.

Small tree up to about 8 m. high, sometimes sprawling or procumbent, with
stilt-roots and flattened, slightly winged branchlets, usually appressed-pubescent at
first, soon glabrescent. Leaves alternate, shortly petiolate or subsessile, petiole up
to 4 mm. long, pubescent or glabrous with two prominent glands near the apex or
at the base of the lamina, papyraceous, chartaceous or subcoriaceous, somewhat
fleshy, lamina narrowly elliptic, up to 11 X 3.5 cm., often much smaller, usually
glabrous, rarely very sparsely pubescent, acute and slightly acuminate at the apex,
Cuneate at the base and often decurrent into the petiole; midrib fairly prominent
on the under surface, lateral nerves and reticulation rather inconspicuous. Flowers

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[Vol. 45
162 ANNALS OF THE MISSOURI BOTANICAL GARDEN

greenish-white, in the axils of small ovate or lanceolate ciliate bracts, aggregated in
subglobose capitula, 3-5 mm. in diam., with sericeous peduncles 5-15 mm, long,
forming a terminal panicle and capitula also sometimes solitary in the axils of the
upper leaves. Lower receptacle (ovary) 1 mm. long, sericeous; upper receptacle
(calyx) cupuliform, 1 X 1.5 mm., nearly glabrous, with lobes 0.5 mm. long.
Stamens normally 10, biseriate, filaments 1.5-2 mm. long, anthers 0.5 mm. long.
Disk pilose. Style 2 mm. long. Fruits squamiform, 3 X 3-3.5 mm., somewhat
sericeous, recurved at the apex and often bearing the remains of the calyx, imbri-
cated in a subglobose or cone-shaped structure, 1-1.5 X 0.7-1.3 cm.

Tropical and subtropical America and Africa.

BOCAS DEL TORO: Isla Colón, Von Wedel 500; Old Bank Island, Von Wedel 2037;
Careening Cay, Chiriqui Lagoon, Von Wedel 2813. CANAL ZONE: without precise local-
ity, Allen 1725. cHIRIQUI: San Bartolomé, Peninsula de Burica, 0.5 m., Woodson
Schery 938. PANAMA: salt-marshes, Barclay 2512; Perlas Archipelago, San José Island,
oi 339; vicinity of Bejuco, edge of mangrove-swamp, Woodson, Allen © Seibert
1683.

6. LAGUNCULARIA Gaertn. f.
LAGUNCULARIA Gaertn. f. in Gaertn. Fruct. 3:209. t. 217. 1807.

Horau Adans. Fam. 2:80, 585. 1763.
Spbenocarbus Rich. Demonstr. Bot. 92. 1808.
Rbizaeris Raf. Sylv. Tellur. 90. 1838

The name Laguncularia Gaertn. f. is antedated by Horau Adans. and should
be conserved.

Small trees and shrubs. Flowers sessile, 5-merous, mostly perfect with occa-
sional staminate in terminal panicles of elongated spikes with additional spikes in
the axils of the upper leaves. Receptacle infundibuliform, scarcely produc
beyond the ovary, with 2 small adnate bracteoles near the apex and crowned with
the persistent calyx. Petals 5, small. Stamens 10, biseriate, not exserted. Disk
10-rayed. Style subulate, capitate at the apex. Ovary unilocular with 2 pendulous
ovules. Fruit almost sessile, longitudinally ribbed or narrowly winged, pericarp
leathery. :

A genus comprising one or perhaps two species inhabiting mangrove-swamps e
tropical America and western tropical Africa and often dominant in such habitats.

1. LAGUNCULARIA RACEMOSA (L.) Gaertn. f. in Gaertn. Fruct. 3:209. t. 217.
1807.
Conocarpus racemosus L. Syst. Nat. 2:950. 1759.
Schousboa commutata Spreng. Syst. Veg. 2:332. 1825, nom. illegit.
Rbizaeris alba Raf. Sylv. Tellur. 90. 1838, nom. illegit.
Laguncularia obovata Miq. in Linnaea 18:752. 1844
Small tree up to about 10 m. high or bush, with glabrous branchlets usually
chestnut-brown when dried, slightly swollen at the nodes. Leaves opposite; :
sate, petiolate, petiole 1—1.5 cm. long, glabrous, with two conspicuous sia di
p ,

mm. below the apex, lamina chartaceous to subcoriaceous, elliptic or oblong

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1958]

FLORA OF PANAMA (Combretaceae)

iy
SS ^
wate
MA
ITAL
LEONA
\ NN
AD

Fig. 51. Laguncularia racemosa

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163

[Vol. 45
164 ANNALS OF THE MISSOURI BOTANICAL GARDEN

up to 10 X 5.5 cm., glabrous, usually blunt or rounded at the apex and slightly
rounded at the base, lateral nerves rather numerous but inconspicuous, reticulation
just visible. Flowers white or greenish-white, sessile, 5-merous, in terminal panicles
up to about 20 cm. long of elongated spikes up to 7-8 cm. long, rhachis densely
appressed-pubescent, bracts suborbicular to ovate, pubescent, 1.5 X 1.3 mm., soon
caducous. Receptacle infundibuliform, elongating as the fruit develops, tomentel-
lous, 2.5 X 1.5 mm., with two small obversely elliptic bracteoles, 1.5 X 2 mm.,
near the apex, expanded at the apex into a persistent cupuliform calyx with 5 broad
lobes, imbricate in bud. Petals 5, suborbicular, appressed-pubescent. Stamens 10,
filaments very short, inflexed in bud, inserted near the margin of the disk. Disk
fleshy, 10-rayed. Style 1 mm. long, glabrous, stigma capitate. Fruit flattened-
obovoid-ellipsoid, at first densely later sparsely appressed-pubescent, 12-20 X
6-10 mm. when mature, more or less longitudinally winged or ribbed, wing up to
2 mm. broad but usually less than 1 mm., crowned by the persistent remains of
the calyx.

Florida to Peru and Brazil, also in West Indies, Galapagos Islands and Fernando
do Noronha; in western tropical Africa from a to ae

boe Allen 25 ‘Isla Coló ón, s ew a of Bocas, Macaw Hills Va wd 522; Chiri-
quí Lagoon, Von We 061; vicinity of Chiriquí Lagoon, Nances Cay, Von Wedel 2801,
037; CANAL ZONE: betw anamá Corozal, alt. 20-30 m., Pittier 4441; banks 0

R;
= sula de Burica, Woodson & Schery 936. PANAMA: Isla Taboga, 0-186 m., on rocky
a, "Woodion, Allen 6 Seibert 1534.

I have seen specimens with glabrous flowers and fruits from southwest Florida
(Rugel 239) and Cuba (Jack 5255) but not, as yet, from Panama. These may
well be Laguncularia glabriflora Presl (Reliq. Haenk 2:22. 1831) though the
latter was described as having red flowers (perhaps erroneously). The type is
from Guayaquil. It would be interesting to study populations of L. racemosa to
see whether glabrous-flowered plants occur. As far as the present meagre evidence
goes it would seem that L. glabriflora could be treated as a glabrous variety of L.
racemosa but without any well-marked geographical distribution.

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1958]
FLORA OF PANAMA (Myríaceae) 165

MYRTACEAE

By G. J. H. AMSHOFF

Shrubs or trees with bicollateral vascular tissue and intraxylary phloem. Leaves
opposite, rarely subalternate, simple, entire or at most crenulate, pellucid-punctate,
these aromatic glands usually present also in other parts of the plant; stipules
usually wanting. Inflorescence racemose or cymose, often paniculate, axillary or
subterminal in the axils of the upper leaves before the development of the terminal
bud or rarely truly terminal. Flowers 4- or 5-merous, hermaphrodite, regular, sub-
tended by two opposite bracteoles. Sepals either free and usually imbricate in the
bud or concrescent, calyptriform and circumscissile from the base at anthesis.
Petals 4 or 5, rarely minute or wanting. Stamens usually numerous, inserted on
the margin of the receptacle in 1 to several series; filaments incurved in the bud,
rarely straight, often conspicuous by their length and color or at least by their
number; anthers dorsifixed, rarely basifixed, 2-celled as a rule, dehiscing longi-
tudinally, the connective usually terminating in a gland, the pollen trigonous,
3-porous. Ovary inferior, rarely subinferior, 2- to several-celled, the receptacle
often produced above; placenta mostly axile; ovules 2 to several in each cell; style
simple, filiform, the stigma small. Fruit a berry or a drupe or, in some Asiatic
and Australian genera often cultivated in other countries, a loculicidal capsule;
seeds without endosperm, the embryo straight or incurved with free or connate
cotyledons (in the first case the radicle also free and the cotyledons either folia-
ceous and contort-plicate or thick and plano-convex).

Distribution: Species about 3000, chiefly in the tropics and in Australia. In
Europe only Myrtus communis. The Central American genera fall into three
subtribes.

A. kory ie foliaceous, more or less sont ge d radicle elongate,
about as lon the cotyledons. Pen mostly 5-merous, are culate;
ovary 2. to pim ai ed, rind gale 2 in each cell. Bey mostly r 2-
seeded. (Subtribe MYRCII

te Calyx concrescent, at ides. propo and falling away like a

lid, remaining attached at one point, at last quite deciduous. Indu-

Sepals fr often formed by deki gór ROSATO 1. CALYPTRANTHES

j s free, imbricate in the bu: Ak Haire simple... nitro zine eee ceceno eee

2a. Receptacle cup-like, ei a ve the ovary; berry globose..........--- . AULOMYRCIA

Fr ceptacle not produced prenda the ovary; berry dud m ipod or

5

3, MYRCIA

X Embryo straight, either homogeneous (the cotyledons and radic le con-
nate) or with free plano-convex cotyledons and short radicle. Flowers
mostly 4-merous, in more or less abbreviated racemes, often glomerate,
rarely solitary or cymose; ovary 2(-3)-celled, the ovules 2 to several
in each cell. Berry mostly 1-seeded, dieere several-seeded. (Sub-
E A
y re pr c fl the expanded cluster of stamens
perdi ae OE emen ch i ICE 4. SYZYGIUM
b. Infor rescences glomerate or racemose, rarely solitary or cymose;
flowers small, the expanded cluster of stamens less than 1 cm. in à
i. 77:177 a — 0 100. RE ULM e a tea
2a. Receptacle produced above the ovary, circumscissile base. Ovules “
r 4, ascending, in each cell of the ovary. Flowers iens DESEE 5. MYRCIARIA

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[Vol. 45
166 ANNALS OF THE MISSOURI BOTANICAL GARDEN

b. pea, not po above the ovary. Ovules usually more than
2 ine ell of the o RE = owers in racemes, fascicles or glomer-
ules, sgh wakes or 6. EUGENIA
C. Embryo incurved, the r sacle dosi much longer than x minute

cotyledons. Inflorescence usu 1- to 7-flowered DES = on

long peduncle (or pedicel if sę mh Uva y 2- to died;

ovules several to numerous in each cell. Seeds omis: numerous, small.

eese MYRTINAE)

closed, sepals not reflexed or spreading until anthesis..................... 9. PsIDIUM
ru Bu d open, -— reflexed or incanta before anthesis 2
2a. ig incurved in the , filiform; eher dorsifixed. Leaves

iocre 7. CALYCOLPUS

b. e zaa akon | or nearly so in the bud, flattened; anthers basifixed.

Leaves very s 8. UGNI

In a recent publication by E. Kausel (in Ark. Bot. n. s. 3:491-516. 1956) the
following information of systematic importance concerning the germination of
the seed is given:

Myrcioideae (in Panama—Myrcia, Aulomyrcia and Calyptranthes): Germina-
tion epigaeous, the large foliaceous, contort-plicate, ensheathing cotyledons become
the first assimilating leaves of the young seedling.

Myrtoideae (in Panama—U gni, Calycolpus and Psidium): Germination epi-
gaeous, the cotyledons very small.

Eugenioideae (in Panama—Eugenia and Myrciaria): Germination hypogaeous,
the cotyledons remaining in the seed under the ground.

Beside the genera enumerated in the key, representatives of the Australian
genera Melaleuca and Eucalyptus are cultivated in Panama. Melaleuca leucoden-

ron L. is an ornamental shrub or small tree with narrowly lanceolate, alternate
leaves with conspicuous longitudinal nerves and flowers in terminal spikes. It is
cultivated in the Plant Introduction Garden at Summit, Canal Zone, and probably
elsewhere.

The most frequently cultivated species of Eucalyptus is E. globulus Labill.
planted as a shade tree. Pimenta officinalis, the Allspice, is planted in various parts
of temperate and hot regions of tropical America, but has not yet been reported
from Panama.

1. CALYPTRANTHES Sw.

CALYPTRANTHES Sw. Prodr. Veg. Ind. Occ. 79. 1788, nomen conservandum.

Cbytraculia P. Br. Hist. Jon: 239. 3796,
Cbytralia Adans. Fam. 2:80. 1763.
Suzygium P. Br. ex Adi. le cit. 244. 1763.
Cal bibranthes Raeuschel, Nom. ed. 3. 144. 1797.
Caly ptranthus Juss. Dict. Sci. Nat. 6:274. 1817

Shrubs or trees, the indument if present partly or entirely composed of di-

brachiate hairs. Inflorescence cymose-paniculate, several-flowered, rarely only 3
flowered, usually subterminal. Receptacle distinctly produced above the ovary.

Calyx concrescent, circumscissile at anthesis but the calyptra remaining ip
at one point. Petals minute or wanting. Stamens numerous, incurved in the buc
anthers ovate, 2-celled, dorsifixed, opening by longitudinal slits. Ovary >>

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1958]
FLORA OF PANAMA (Myrtaceae) 167

3-celled; cells 2-ovulate, the ovules ascending; stigma simple. Berry globose,
crowned by the persistent free margin of the receptacle, 1- to 2-seeded, cotyledons
free, contort-plicate, the radicle elongate.

Distribution: Species about 70 in tropical and subtropical America.
la

Inflorescence 1- to 3-flowered, on a long slender peduncle 3. C. HYLOBATES
Inflorescence cymose-paniculate, several- or many-flowered
la. Leaves large 12-28 cm. long; flower buds globose, about 2.5 mm. high. 1. C. TUMIDONODIA
. Leaves much smaller, less than 15 cm. long
3a. Inflorescence and flower buds pubescent
. Inflorescence and flower buds quite glabrous, the ultimate flowers not
more than 3 together, mostly pedicellate ... 2. C. SCHIEDEANA
. Ultimate flowers glomerate, sessile, up to 5 together; inflorescence
shortly villous with stalked hairs .. 4. C. CHYTRACULIA
Ultimate flowers partly pedicellate, usually not more than 3 together;
inflorescence finely sericeous with sessile hairs 5. C. COSTARICENSIS

Calyptrantbes pittieri Standl. also recorded from Panama, but which I have not
seen, seems to be characterized by its thick and coriaceous leaves less than 10 cm.
long, and pubescent cymose-paniculate inflorescence with sericeous flower buds
about 3.5 mm. long.

.
.

a

c

E
p

Sg

1. CALYPTRANTHES TUMIDONODIA Schery, in Ann. Missouri Bot. Gard. 30:215.

1943.

Calyptrantbes urophylla Standl. & Williams, in Ceiba 3:215. 1953.

Shrubs about 2.5 m. high, or small trees, nearly glabrous. Twigs terete, with
thickened nodes. Leaves elliptical or elliptical-oblong, long acuminate at the apex,
acute at the base, glabrous above, sparsely pubescent or glabrate beneath, 12-28
cm. long and 5-9 cm. wide, membranaceous, the midrib impressed above, prom-
inent beneath, the lateral nerves in about 25 pairs, slender, inconspicuous on bot
sides, the marginal nerve at about 2-3 mm. distant from the margin; petiole about
8 mm. long, thick, canaliculate. Inflorescence cymose-paniculate, about 9 cm.
long, terminal and in the upper axils, lax and many-flowered, brown-pubescent with
sessile dibrachiate hairs. Flower buds brown-pubescent, about 3 mm. igh.
Flowers whitish. Endemic. In the original description of C. tumidonodia both
specimens of Von Wedel are cited. Von Wedel 2195 was afterwards, apparently
in haste, redescribed as C. uropbylla.

BOCAS DEL TORO: Fish Creek hills, Von Wedel 2195, 2223.

2. CALYPTRANTHES SCHIEDEANA Berg, in Linnaea 27:28. 1855.

Myrcia aromatica Schlechtend. loc. cit. 13:415. 1839, non Calyptranthes aromatica St.

Hil. (1828).

Calyptrantbes schlechtendaliana Berg. loc. cit. 27:29. 1855.

Trees or shrubs, glabrous throughout, the twigs slender, subterete. Leaves
Petiolate, broadly elliptical to lanceolate- oblong, 4.0-7.5 cm. long and 2.0—4.5 cm.
wide, acute or obtuse at the base, acuminate at the apex, chartaceous, the midrib
impressed above, prominent beneath, the lateral nerves numerous, prominent on
both sides, joining the marginal nerve near the margin; petiole about 4 mm. long.
Inflorescence paniculate, many-flowered, about as long as the leaf, the ultimate

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[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

168

La da i "
x VAT

ZY UP i

Fig. 52. Calyptranthes chytraculia

NS == NI:
N (SUE

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1958]
FLORA OF PANAMA (Myríaceae) 169

flowers sometimes 3 together, sessile, but for the greater part pedicellate. Flower
buds globose, glabrous, 1.5-2.0 mm. high. Fruit globose, 4-6 mm. in diameter.
Mexico and Panama. Originally described from Mexico and quite common in
that country.
PANAMA: Taboga Island, Barclay s. n. VERAGUAs: Coiba Island, Seemann 624.
3. CALYPTRANTHES hylobates Standl. spec. nov. ined. in herb.

Arbores parvae ca. 8 m. altae trunco ca. 10 cm. diam. ramulis gracilibus multo
divisis apice quadrangularibus vel anguste alatis glabris. Folia oblonga mem-
branacea glabra basi acuta apice longe acuminata 5-7 cm. longa 2.0-2.5 cm. lata
nervo medio supra impresso subtus prominente nervis lateralibus numerosis tenuibus
uterque superficieibus prominulis ca. 1 mm. ab margine in nervum marginalem
collictis; petiolo brevi ca. 2 mm. longo. Inflorescentia 1- vel 3-flora pedunculo
gracili usque 3.5 cm. longo glabro flore centrale sessile sive lateralibus usque 1 cm.
longe pedicellatis. Flores adulti ignoti. Bacca globosa ut dicitur fulva; cotyledon-
ibus foliaceis contorto-plicatis.

Panama and Costa Rica.

BOCAS DEL TORO: region of El Almirante, Cooper 366 (HOLOTYPE, Herb. Chicago Nat.
Hist. Mus.).

4. CALYPTRANTHES CHYTRACULIA Sw. Prodr. Veg. Ind. Occ. 79. 1788.

Shrubs or low trees, the young twigs subterete, sparsely pubescent, glabrescent.
Leaves elliptical, acute at base, acuminate at apex, glabrous, chartaceous, up to 12
cm. long and 4.5 cm. wide, the midrib impressed above, prominent beneath, the
lateral nerves numerous, prominulous on both sides, joining the marginal nerve
Very near the margin at about 1.5 mm. distance from the margin; petiole canalicu-
late, up to 8 mm. long, pubescent, glabrescent. Panicles nearly as long as the leaf,
rarely shorter, brown-pubescent with dibrachiate stalked hairs, the branches 2-4
together, the ultimate flowers mostly sessile, sometimes partly pedicellate, 1-5 to-
gether in small glomerules. Flower buds obovoid, about 2 mm. high, brown-
puberulous, opening with a shallow cup-like lid. Petals none. Filaments up to 4
mm. long; anthers small, at length twisted. Berry globose, glabrous; cotyledons
contort-plicate.

Yucatan Peninsula, Guatemala and Panama; Jamaica.

BOCAS DEL TORO: Changuinola valley, Cooper & Slater 143.

The Panama specimen quite agrees with specimens from British Honduras
distributed as C. millspaugbii Urb. The inflorescence is usually larger than in
Urban's description.

5. CALYPTRANTHES COSTARICENSIS Berg, in Linnaea 27:20. 1855.

Trees of 427 m., the crown dense and rounded, the branchlets puberulous,
Compressed, Leaves elliptical, acute at the base, shortly acuminate at the apex,
chartaceous, glabrous except the midrib, 6-10 cm. long and 3-5 cm. wide, the

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[Vol. 45
170 ANNALS OF THE MISSOURI BOTANICAL GARDEN

midrib impressed above, prominent beneath, striate-nerved with numerous lateral
nerves, these prominulous on both sides and joining the marginal nerve very near
the margin; petiole slender, canaliculate, about 5 mm. long. Panicles often longer
than the leaf, puberulous with dibrachiate sessile hairs, the branches opposite or
ternate, the ultimate flowers at most 3 together, sessile or pedicellate. Anthers 1
mm. long. Flower buds subglobose, about 3 mm. long, puberulous, opening with
a petaloid concave lid. Berry globose, about 1 cm. in diameter.

The type specimen collected by Warscewicz "in Costa Rica et Veraguas" has
not been seen by us, and apparently no other collections have since been made in
Panama. Standley (Fl. Costa Rica 2:770. 1937) feels that the species may be
endemic to Costa Rica and reports it as "Common in forests and thickets of the
Meseta Central, at 1,100—2,000 meters.”

2. AULOMYRCIA Berg
AULOMYRCIA Berg, in Linnaea 27:35. 1855.

Myrcia subgen. Aulomyrcia (Berg) Niedenzu, in Engl. & Prantl, Nat. Pflanzenfam. 37:75.
1893.

Shrubs or small trees. Inflorescence paniculate, axillary. Bracts and bractlets
small, deciduous. Flowers mostly small, 5-merous or sometimes (not in the Central
American species) 4-merous. Flower buds mostly obconic. Receptacle cup-like
produced above the ovary, glabrous or pubescent outside, generally glabrous inside.
Sepals imbricate, often very unequal, with the inner sepals much larger than the 3
outer sepals. Petals white, orbiculate or obovate. Stamens numerous, inserted on
the free margin of the receptacle; filaments incurved in the bud; anthers ovate,
dorsifixed, opening by longitudinal slits. Ovary 2- to 3-celled; ovules 2 in eac
cell, ascending. Berry globose, crowned with the cup-like, persistent, free margin
of the receptacle and with persistent sepals, 1- to 3-seeded; cotyledons free, more or
less contort-plicate; radicle elongated, inserted near the apex of the embryo.

Distribution: Species about 200 in tropical America. The genus is nearly
allied to and often united with Myrcia DC. The cup-like produced receptacle of
Aulomyrcia, a character so useful in distintinguishing other Myrtaceous genera,
seems to be the only constant character to distinguish it from Myrcia.

la. Leaves large, up to 30 cm. long........................ 1. A. ZETEKIANA
b. Leaves small, 4—6 cm. log. 0 2. A. TOMENTOSA

1. AULOMYRCIA zetekiana (Standl.) Amsh. comb. nov.
Eugenia Zetekiana Standl. in Journ. Wash. Acad. Sci. 15:286. 1925.

Shrubs 2.5—4.5 m. high, with few branches, the branchlets densely ferruginous
tomentose. Leaves lanceolate or narrowly oblong-lanceolate, about 30 cm. long;
6.5-9.5 cm. broad, attenuate at apex, rounded at base, coriaceous, glabrous above,
tomentose beneath along costa, or glabrate, the venation reticulate} prominent 02
both sides, the costa stout, the lateral nerves about 20 on each side (young leaves
dark purple, glabrous or nearly so); petiole swollen, nearly 1 cm. long. Flowers m

(170)

(IZI)

T

Fig. 53. Aulomyrcia tomentosa

(av22v14CWI) VWVNVd AO VUOTA

IZI

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[Vol. 45
172 ANNALS OF THE MISSOURI BOTANICAL GARDEN

terminal, simple or branched racemes 7—8 cm. long, densely velvety-tomentose.
Flowers not known. Receptacle produced shortly beyond the ovary, tomentose
outside. Sepals 5, subequal, semiorbicular, broadly rounded at apex (the shriveled
remains in our specimens about 0.3 mm. long). Petals not seen. Berry globose,
densely brown-tomentose, when dry about 1.5 cm. in diameter, green and brown,
crowned by the persistent cup-like receptacle and sepals, 1- or 2-seeded. Cotyledons
rather thick, plicate; radicle free, less than half the circumference of the see
(about one third).

Endemic, in wet forests.

CANAL ZONE: hills north of Frijoles, Standley 27503; hills west of the canal, near
Gatún, Standley 27192.

2. AULOMYRCIA TOMENTOSA (Aubl) Amsh. in Rec. Trav. Bot. Néerl. 39:153.

1942.

Eugenia tomentosa Aubl. Pl. Guian. 1:504. £. 200. 1775
Myrcia tomentosa ( Aubl.) DC. Prodr. 3:245. 1828.
Eugenia vallis Standl. in Ann. Missouri Bot. Gard. 27:323. 1940.

Shrubs or small trees to 12 m. tall, the bark smooth, reddish brown, the young
twigs densely villous. Leaves obovate, rounded, retuse, obtuse or shortly and
obtusely acuminate at the apex, narrowed toward the base, membranaceous to
chartaceous, at first pilose on both sides, later glabrescent especially above, 4-6
(-10) cm. long and 2-4 (—5) cm. broad; petiole pubescent, 2-5 mm. long.
Panicles densely flowered, villous. Flowers white, pubescent, the buds 2.0—2.5 mm.
high. Receptacle shortly produced above the ovary, white-villous outside, glabrous
inside. Sepals unequal, the smallest about 1 mm. long and broad, the largest about
1.5 mm. long and 2 mm. broad. Petals suborbiculate, crenulate, about 2 mm.
long. Ovary 2-celled. Berry globose, orange, 5-8 mm. in diameter, 1-seeded.
Cotyledons contort-plicate, with elongate radicle.

Distribution: Panama to Brazil; Trinidad.

COCLÉ: vicinity of El Valle de Antón, alt. 600—1000 m., Allen 1773, 2512, 2578.
PANAMA: San José Island, Perlas Archipelago, Johnston 801, 747, Erlanson 284.

3. MYRCIA DC.

Myrcia DC. ex Guillemin, Dict. Class. Hist. Nat. 11:378. 1826.
Aguava Raf. Sylva Tellur. 107. 1838.
Calycampe Berg, in Linnaea 27:129. 1856.

Shrubs or trees. Inflorescence paniculate, axillary or subterminal. Bracts
bractlets mostly small Flowers 5-merous, in the bud more or less constri¢
under the calyx. Receptacle not or scarcely produced above the ovary, pubescent
outside. Sepals imbricate, equal or slightly unequal. Petals white, orbiculate a
obovate, pubescent outside. Stamens numerous; filaments incurved in the DUS
anthers ovate, dorsifixed, opening with longitudinal slits. Ovary 2- to 4-celled,
terminated by a sericeous disc; ovules 2 in each cell, ascending. Berry 1- di
ellipsoid, or 2- to 3-seeded and obovoid or subglobose when ripe, crowned by

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1958]
FLORA OF PANAMA (Myrtaceae) 173

persistent sepals. Seeds with cartilaginous shining testa; cotyledons free, longi-
tudinal (i.e. so placed in the fruit that they are parallel with the greatest length
of the fruit), contort-plicate; radicle elongate, half-encircling the embryo.
Distribution: About 100 species in tropical America.
In Panama three rather distinct species can be distinguished. However the
genus is badly in need of revision, since the differences are small and in some cases
not constant. Four or five other species are known from Central America.

la. Leaves large, 7—17 cm. long, RE: aeroplano, the aei nerves distant,

at last impressed above and pro t beneath; twigs glabrous; flowers

relatively large in lax and few- rad pr ee de reli 2. M. COUMETA
b. Leaves smaller, Brody ot more than . long, densely reticulate,

the lateral nerves prominulogź to obsolete above; twigs densely pubescen

at least when young; flowers smaller, in dense more or less pubescent

pena :
2a. Leaves coarsely reticulate-veined, the young twigs and inflorescences

more or jm densely pubescent with spreading hairs 1. M. COSTARICENSIS
b. Leaves fin ely ain A GRAŁ i jo young twigs and inflorescences

sparsely and minutely pubescen 5. M. GATUNENSIS

l. Myrcia COSTARICENSIS Berg, in Linnaea 27:104. 1854.

Shrubs or small trees 2-8 m. high, the crown dense and rounded, the young
branchlets densely pubescent with spreading, somewhat brownish hairs. Leaves
petiolate, lustrous, lanceolate-oblong or elliptical-oblong, commonly 4—7 cm. long,
obtuse or rounded at the base, acuminate at the apex, at first sparsely pubescent on
both sides, later glabrate, the lateral nerves prominulous on both sides; petiole about
3 mm. long. Panicles densely pubescent, densely flowered. Flowers white, fragrant.
Receptacle sericeous, about 1.5 mm. high. Sepals about 1 mm. long, rounded.
Petals about 3 mm. long, pubescent outside. Berry oblong, red to black.

Distribution: Costa Rica and Panama.

quí Viejo, 1200 m., Allen 100 LÉ: between Las Margaritas pag p Ha Woodson,
Allen 8 a 1304; hills south a El Valle de Antón, alt. about Allen 2519,

PANAMA: Isla Taboga, ca. 0-186 m., Woodson, Allen & Seibert I 1446; La Chor-
rera, AS 200 ft., Bro. Paul 818. ri Eger Cafiazas, Allen 163.

This species, known locally as arrayán (myrtle) is the most common Myrcia
species in Panama, several times collected in nearly all provinces.

2. MYRCIA COUMETA (Aubl) DC. Prodr. 3:245. 1828.
Mi Coumeta Aubl. PI. Guian. 1:497. t. 196. 1
cia Coumeta (Aubl.) Berg, in Linnaea 27: Ae 1855.

Shrubs or small trees, the twigs almost glabrous. Leaves ovate or ovate-oblong,
acute or obtuse at the base, acuminate at the apex, 7-17 cm. long and 4-6 cm.
wide, the Principal lateral nerves distant, at last impressed above and prominent
beneath, the veins laxly reticulate; petiole robust, about 5 mm. long. Panicles

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ANNALS OF THE MISSOURI BOTANICAL GARDEN

174

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1958]
FLORA OF PANAMA (Myrtaceae) 175

nearly glabrous, distantly flowered, with 3-5 pairs of 1- to 9-flowered branches,
the pedicels slender, up to 8 mm. long. Flowers larger than in the two other
species, the sepals typically up to 2 mm. long, the petals about 4 mm. long. Berry
globose in the Panama specimens.

The typical form in French Guiana, Surinam, and also in Costa Rica; the
following Panama specimens seem to belong to this species although their fruit is
globose instead of oblong and the flowers are not yet fully developed. In Costa
Rica the species has been confused with Myrcia carnea (Mey.) DC., described
from British Guiana but still imperfectly known.

BOCAS DEL TORO: Fish Creek hills, Von Wedel 2441, 2871, Cooper 570.
3. MYRCIA GATUNENSIS Standl. in Publ. Field Mus. Bot. 4:154. 1929.

Twigs and inflorescences minutely and finely pubescent. Leaves lanceolate-
oblong or elliptical, rounded or obtuse at the base, rather long-acuminate, 7-10 cm.
long, chartaceous, the lateral nerves numerous, delicate, prominulous on both sides
or obsolete above, the veins densely and finely reticulate, not conspicuous; petioles
mostly about 2 mm. long, sometimes longer. Inflorescence slender, shorter than the

af. Flowers minute, the petals 1.5—2.0 mm. long.

Apparently endemic.

CANAL ZoNE: Barro Colorado Island, Kenoyer 468; Barro Colorado Island, Barbour
Point to next point south, Bangbam 498; Barro Colorado Island, shores of Gatün Lake,
south of Laboratory, Bangham 451.

Two other species sometimes are recorded for Panama. The first, M. oerstede-
ana Berg, described from Costa Rica, has leaves similar to those of M. gatunensis
but is more pubescent; the Panamanian specimens previously referred to that species
agree better with M. costaricensis. 'The second, M. plicato-costata Berg, is known
from Costa Rica, but the record for Panama is dubious. It is questionable whether
M. costaricensis, M. gatunensis, and M. oerstedeana are distinct species.

4. SYZYGIUM Gaertn.

SYZYGIUM Gaertn. Fruct. 1:166. 1788.
Opa Lour. Fl. Cochinch. 308. 1790.

Jambosa DC. Prodr. 3:286. 1828.

Microjambosa Blume, Mus. Bot. Lugd.-Bat. 1:117. 1849.

Shrubs or trees. Inflorescence usually centrifugal, cymose, more or less panicu-
late. Flowers 4-merous. Receptacle produced above the ovary. Sepals persistent,
mostly semi-orbiculate, sometimes distinct. Petals free or coherent. Anthers
dorsifixed, opening with longitudinal slits. Ovary 2(-4)-celled, the cells many-
ovulate. Berry globose or pear-shaped, often carnose and edible, mostly 1-seeded.

esta adhering to the pericarp. Cotyledons free, carnose; radicle short, hidden
between the cotyledons. :

Distribution: Species about 200 in tropical and subtropical Asia, the following
Cultivated or sometimes naturalized in Panama.

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[Vol. 45

176 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1a. Flowers red; inflorescence mostly lateral, often cauliflorous 1. S. MALACCENSE
white; i i rminal

b. Flowers inflorescence chiefly te
2a. Leaves lanceolate, acu t the base 2. S. JAMBOS
b. Leaves elliptical-oblong, rounded at the base 3. S. SAMARAGNENSE

1. SYZYGIUM MALACCENSE (L.) Merrill & Perry, in Journ. Arn. Arb. 19:215.

1938
Eugenia malaccensis L. Sp. Pl. 470. 1753.

Jambosa malaccensis (L.) DC. Prodr. 3:286. 1828.
Jambosa domestica Bl. Mus. Bot. Lugd.-Bat. 1:91. 1849.

Low trees with very dense rounded crown. Leaves large, oblong-elliptical,
obtuse or acute at the base, acute or acuminate at the apex, coriaceous, glabrous,
20-30 cm. long and 8-18 cm. wide, the lateral nerves in about 10-14 pairs. In-
florescences lateral, few-flowered. Flowers red. Receptacle glabrous. Sepals 4-8
mm. long. Petals orbiculate, red, 14—18 mm. long. Stamen filaments red, greatly
exserted; anthers yellow. Berry obovoid, about 7 cm. long, whitish or yellowish
or tinged with red.

Native of Asia, sometimes planted as a shade tree. The bright purple red
flowers are exceptionally beautiful. They are borne amongst the leaves however
and are not conspicuous when the tree is viewed from a distance; but the petals
fall upon the bare soil beneath and form a lovely carpet. The fruit is called
marañón de Curasao, a not inappropriate name, since the fruit strongly suggests, in
shape and color, that of Anacardium occidentale L., called marañón throughout
Central America. Another name is Malay apple or manzana de Faiti.

2. SYZYGIUM JAMBOS (L.) Alston, in Trimen, Handb. Fl. Ceylon 6: Suppl. 115.
I.

Eugenia jambos L. Sp. Pl. 470. 1753.
Jambosa vulgaris DC. Prodr. 3:286. 1828.

Small trees though often 15 m. high, with a dense crown of green foliage.
Leaves lanceolate, attenuate toward the acute apex, acute at the base, glabrous,
coriaceous, 15-20 cm. long and 4-6 cm. wide, the lateral nerves in about 8 pairs,
the marginal nerve at about 4 mm. from the margin. Inflorescence few-flowered,
terminal, glabrous. Flowers, including the many filaments, about 5 cm. in di-
ameter Receptacle glabrous, 1.5 cm. high. Sepals 4, semi-orbiculate. glabrous,
about 7 mm. long. Petals about 1 cm. long, white. Berry pear-shaped or SWZ
globose, about 3 cm. in diameter and 3.5—5.0 cm. long, pinkish or yellowish, 2- or
several-seeded, crowned with the persistent incurved sepals.

Native of the East Indies and widely cultivated as a shade tree throug
tropical America because of the dense foliage, showy flowers and edible fruits.
fruit is not much esteemed in Central America, although elsewhere it is much
for the making of jellies. Known as manzana and manzana rosa, and in Panama
the tree is usually called pomarosa; in English Rose apple, Malabar plum and Plum
rose. It frequently escapes and naturalizes in Panama.

hout

The

3. SYZYGIUM SAMARAGNENSE (Blume) Merrill & Perry, in Journ. Arn. Arb. 19:
115. 7238

(176)

|

1958]
FLORA OF PANAMA (Myrtaceae) 177

Eugenia javanica Lam. Encycl. 3:200. 1789, non Sysyzium javanicum Miq.
Myrtus samaragnensis Blume, Bijdr. 1084. 1826.
Jambosa samaragnensis (Blume) DC. Prodr. 3:286. 1828.

Trees. Leaves elliptical-oblong, rounded or subcordate at the base, mostly obtusely
acuminate at the apex, 10—24 cm. long and 4-11 cm. wide, chartaceous, glabrous,
aromatic. Inflorescence terminal or lateral, in the first case often many-flowered,
the flowers usually in clusters of 3. Receptacle cup-like, produced above the ovary,
about 12 mm. high. Sepals 4, semi-orbiculate, glabrous, about 4 mm. long. Petals
and stamen filaments yellowish white, the petals about 1 cm. long, concealed by
the elongate stamens. Berry pear-shaped or subglobose, about 3-5 cm. long,
pinkish or white, carnose, insipid.

Native to the East Indies but cultivated throughout the tropics. In Panama
it is less common than the two other species.

5. MYRCIARIA Berg
MYRCIARIA Berg, in Linnaea 27:320. 1856.

Shrubs or trees. Flowers subsessile, glomerate, axillary and lateral. Bractlets 2,
enclosing the ovary, often connate at the base. Receptacle distinctly produced
above the ovary, at last circumscissile. Sepals 4, imbricate in the bud and enclosing
the petals. Petals small, fimbriate. Stamens numerous; filaments incurved in the
bud; anthers ovate, dorsifixed, opening with longitudinal slits. Ovary 2- to 3-
celled; cells 2- or 4-ovulate, the ovules ascending. Berry globose, crowned with
the disc only. Embryo undivided.

Distribution: Species about 40 in tropical America.

la. Leaves 2-5 (-7) cm. long, acute at the base 1. M. FLORIBUNDA
b. Leaves 5-12 cm. long, rounded to cordate at the base 2. M. VISMEAEFOLIA

1. MYRCIARIA FLORIBUNDA (West, ex Willd.) Berg, in Linnaea 27:330. 1856.

Eugenia floribunda West, ex Willd. Sp. Pl. 2:960. 1800.
Eugenia O'Neillii Lundell, in Bull. Torrey Bot. Club 64:555. 1937.
Myrciaria O'Neillii (Lundell) I. M. Johnston, in Sargentia 8:228. 1949.

Shrubs or small trees 2-10 m. high. Trunk 3-25 cm. in diameter, covered
with a tight smooth light-brown exfoliating bark that is practically indistinguish-
able from that of its common associate Calycolpus (cf. I. M. Johnston, loc. cit.
1949). Twigs densely foliate, the internodes 1-2 cm. long, at first puberulous,
soon glabrate. Leaves elliptic- to lanceolate-oblong or lanceolate, long-acuminate
to caudate at the apex, acute at the base, 2-5 (—7) cm. long and 1.0-2.5 cm. wide,
chartaceous, at first pubescent on the margin and midrib, soon glabrate, the midrib

at above, prominent beneath, the lateral nerves numerous, delicately prominulous
to nearly obsolete on both sides, the marginal nerve 0.5—1.0 mm. within the margin;
Petiole slender, 2-5 mm. long. Flowers glomerate, 2-5 together. Receptacle
glabrous, about 1.5 mm. long, at last circumscissile above the ovary. Sepals 4,
imbricate, about 1 mm. long, deciduous with the receptacle. Petals obovate,
fimbriate, slightly longer than the sepals. Ovary glabrous outside. Berry globose,

(177)

[Vol. 45

ANNALS OF THĘ MISSOURI BOTANICAL GARDEN

178

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Fig. 55 Myrciaria floribunda

(178)

1958]
FLORA OF PANAMA (Myrtaceae) 179

8-15 mm. in diameter, 1-seeded, edible. Embryo homogeneous; radicle broadly
adnate.
Southern Mexico to Panama; Guiana and Brazil; West Indies. A common
element in the understory of more or less shady forest at low elevations.
CANAL ZONE: hills north of Frijoles, Standley 27521; between France Field and Cati-
i ri

val, Prov. Colón, Standley 30192; without precise locality, Mrs. Epplesbeimer anno 1910
027. LE: vicinity of Nata, Allen ; Bismarck,

2. MYRCIARIA VISMEAEFOLIA (Benth.) Berg, in Linnaea 27:536. 1856.
Eugenia vismeaefolia Benth. in Hook. Journ. Bot. 2:320. 1840.

Shrubs or small trees 4-6 m. high, the trunk up to 1 dm. in diameter, the twigs
glabrous and with distant foliage. Leaves oblong to lanceolate-oblong, 6-12 cm.
long and 2-4 cm. wide, long-acuminate and acute at the apex, rounded or obtuse
at the base, coriaceous, glabrous, the midrib slightly elevated above, prominent
beneath, the lateral nerves numerous, delicate; petiole about 5 mm. long. Flowers
glomerate, axillary and lateral, whitish, the bractlets small, connate, ciliate. Re-
ceptacle glabrous, at last circumscissile above the ovary. Sepals glabrous outside,
puberulous inside, imbricate in the bud with 2 smaller and 2 larger. Petals 4, about
2 mm. long, pubescent, fimbriate. Ovary 2-celled, 4-ovulate, the ovules ascending.
Berry globose, up to 1 cm. in diameter, 1- to 3-seeded, blackish blue when dry,
edible. Embryo homogeneous.

Distribution: Panama, British Guiana and Surinam. The description as given
here refers to the Panamanian variety, which differs from the typical form of the
Guianas by its firmer leaves which are never (?) cordate at the base, its longer
petioles and its slightly larger flowers.

CANAL ZONE: Summit, Zeżek 3520.

6. EUGENIA L.

EUGENIA L. Sp. Pl. 470. 1753.
Catinga Aubl. Hist. Pl. Guian. 1:511. #. 203. 1775.
Gre G 8.

ggia Gaertn. Fruct. 168. 7. 33. 17
Guapurium Juss. Gen. 324. 1789.
Olynthia Lindl. Collect. t. 19, in obs. 1821.
Ossinia Comm. ex DC. Prodr. 3:237. 1828

erocarbus Hassk. in Flora 25:2 Beibl. 36. 1842.
Syllysium Meyen & Schau. in Nova Acta Acad. Nat. Cur. 19: Suppl. 1. 334. 1843.

Myrcianthes Berg, loc. cit. 315. 1856.
. 1856.

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[Vol. 45
180 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Anamomis Griseb. Fl. B. W. I. 240.
Psidiastrum Bello, in Anal. Soc. punt Hist. se 10:272. 1881.
Myrtopsis O. Hoffm. in Linnaea 43:133. 18

Oxydiastrum Niedenzu, in Engl. & Prantl, Nat. Pflanzenfam. = 2105. 1893.
Ch eas Pierre, in Bull. Soc. Linn. Paris, n. ser. 71.

Shrubs or trees. Flowers tetramerous, in soe i or Mr racemes, fascicles
or glomerules, rarely solitary or cymose. Receptacle scarcely or not produced above
the ovary. Sepals free, imbricate, the two inner mostly distinctly larger than the
two outer, persistent. Petals free, often caducous. Stamens numerous; filaments
incurved in the bud; anthers dorsifixed, opening with longitudinal slits. Ovary
2 (-3) -celled, the cells several- to many-ovulate, sometimes only 2-ovulate. Berry
globose or ellipsoid, mostly 1-seeded, rarely few-seeded. Embryo homogeneous, the
radicle broadly adnate to the wholly connate cotyledons, or sometimes cotyledons
with a short split between them (or in one species in Panama, also aberrant
by its cymose inflorescence, with cotyledons free and radicle short but distinct).

Distribution: Species numerous, probably more than 1000, in the tropics and
subtropics, chiefly in America. The genus sometimes is united with Syzygium
Gaertn., and then a few hundred additional species must be added.
la. Inflorescence cymose, 1- to 7- flowered; cotyledons fre I E
. Inflorescence various, but never cymose; cotyledons connate
2a. wits ey nb; 4
b. B ooth or at most faintly ribbed.................. o qe
3a. Luna. ovate; ua solitary in the wew ot bracts at the base of med

shoots, or in facile of > de peter’ 1.5 cm. long or more; berr

JĄ

. FRAGRANS

o:

m

. UNIFLORA

berry RAA. ng, up to
4a. Leaves linear, less than 5 mm. cyk 4
pedicels.

ANDERE 17. E. OCTOPLEURA
owers solitary, on long slender

— DS iN AM DU s SER . E. ALFAROANA
b. Leaves o about 1 cm. wide or more; flowers glomerate, fasciculate, A
OF FACCNIOB. porri OL ID Hu NN
5a. Flowers precocious, deo, the Len cels cem the ovary
densely puberulous; leaves ena mes t 5 cm. long, the marginal nerve
at some distance from the margin... E. NESIOTICA 6
b. Flowers present at the same NR a as fully developed leaves... ooo pi 7
6a. de pare: mostly less than 5 cm. long, subsessile if up t m. long... nà
Leaves larger, a rule more than 5 cm. long, or definitely en 9
if shorter: Liri e A A a
7a. Flowers glomerate; leaves almost sessile, oblong to lanceolate................... 6. E. PITTIERI
b. Flowers in fascicles or in abbreviated r acemes, long-pedicellàte..............................r?
8a. Inflorescence an abbreviated 1- -flowered raceme; the pedicels 4—10
long; leaves oblong or tauceolate- SR 7. E. CHEPENSIS
b. Inflorescence an axillary 6- to 15-flowered fascicle, the pedicels 4-6 mm.
long; lave dlipial. <<“ 0. ua a 8. E. COSTARICENSIS
9a. Leaves ta NEN te sy agree sparsely béo. cd to
glabrescent on both s , Up to - long; inflorescence and o
pubescent; flow arene i . E. BIFLORA 10
b. Lam elliptical oe lr e MM T
10a Tn glabrous or desires leaves and ovary always glabrous (see 11
o Ei nos i) ii I
b Yong twigs densely pd: ent with patent hairs; is is pipes long- 17
pubescent or sericeous at >> beneath; ovary pu t ot glabrous... etm
lla. Flowers distinctly racemose (som metimes deri in. the xim Nr 12
longer than the pedicels; riban acu

1958]

FLORA OF PANAMA (Myrtaceae) 181
b. Flowers fasciculate or in more or less abbreviated racemes: leaves obovate,
rounded or obtuse and scarcely acuminate at the apex 13
12a. Flowers small, the petals about 3 mm. long; rachis and pedicels very
slender 11. E. OERSTEDEANA
b. Flowers larger, the petals about 6 mm. long; plant more robust in all
arts. 10. E. COLORADENSIS
13a. Leaves very large, 20—25 cm. long 12. :
b. Leaves much smaller .14

mm.
ong; inflorescence an abbreviated raceme with slender rachis and pedicels,
artly iculat

13. E. ACAPULCENSIS
4

b. Flowers larger, the petals 5—7 mm. long; inflorescence more robust
15a. Leaves obovate, rounded, obtuse or subtruncate at the apex; inflorescence
poorly known but possibly cymose, 3-flowered (but embryo homo-

14. E. HIRAEIFOLIA

geneous)
b. Leaves elliptical, ovate or oblong
16a. Sepals very short, about 1 mm. long, even in the bud much shorter
than tals : . 16. E. TAPACUMENSIS

peta A
b. Sepals larger, unequal, 3-5 mm. long, covering the petals in the bud. 18. E. AUSTIN-SMITHII

17a. Leaves sericeous beneath with a shining golden, eventually fading
ind tum; flowers pedicellate.........................---// e 19. E. CHRYSOPHYLLUM

b. Leaves pubescent with spreading hairs, glabrescent or glabrous; twigs
s

riaceou:
glabrous; inflorescence and flowers poorly known a E MAMA ADI
b

b. Leaves membranaceous, the nerves distinct, glabrous... Mmmm

19a. Flowers small, sessile or nearly so, glomerate, the sepals not more than

1 i PEEP T Lagé dir E OR MUR oe te ee

b. Flowers larger, long-pedicellate (up to 1 cm.), fasciculate, the sepals up
to 4 mm. long

9. E. ORIGANOIDES

20. E. OREINOMA

The most common species are E. acapulcensis and E. origanoides which are
probably to be found in all provinces. The first species is glabrous and easily
recognizable by its minute flowers in slender abbreviated racemes; the second species
is pubescent and has equally small but glomerate flowers. E. biflora, particularly
frequent on Taboga Island but probably scattered throughout the country, has
narrowly lanceolate leaves and pubescent few-flowered racemes. E. fragrans is
particularly frequent in the Chiriquí region, and is the only species with a dichot-
omously branched inflorescence. E. oerstedeana and E. coloradensis, both with
flowers in long glabrous racemes, E. chepensis, with small leaves and flowers in
abbreviated, nearly glabrous racemes, and E. nesiotica, with pubescent, fasciculate,
precocious flowers and small leaves, are most common in the Canal Zone. All other
species have been collected but once or twice, some in recent years and others in
the last century.

l. EUGENIA ALFAROANA Standl. in Journ. Wash. Acad. Sci. 14:240. 1924.

Shrubs 1-5 m. high, densely branched, the branchlets very slender, thinly pilose-
sericeous. Leaves linear or nearly so, often broadened near the base, 5-6 cm. long,
1.5-4.0 mm. wide, acute at the base, acute and aristate-mucronate (bristle about
1.5 mm. long) at the tip, thin, when young thinly sericeous-pilose with whitish
hairs, soon glabrate; petioles 2-4 mm. long, tomentose and pilose. Flowers axillary,
solitary on long, slender, appressed-pilose pedicels 1.5-2.0 cm. long. Bractlets 2
at the base of the calyx, filiform, 2-3 mm. long. Sepals ovate, subulate-acuminate,

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[Vol. 45
182 ANNALS OF THE MISSOURI BOTANICAL GARDEN

2-3 mm. long, tomentose or glabrate. Petals broad, white, 3 mm. long, ciliolate,
copiously gland-dotted. Ovary 2 mm. high, densely white-tomentose, 2-celled,
each cell with 3-4 relatively large ovules. Berry purple-black, 6-7 mm. in
diameter, glabrate, crowned with the persistent sepals, 1-seeded; seed about 6 mm.
in diameter, the embryo homogeneous.

El Salvador, Panama. Escoba (in Panama).

CHIRIQUf: Sabana de El Boquete, alt. 700-1100 m., Piźtier 3302.

The specimen cited is sterile; flowering material is needed for more sure
determination.

2. EUGENIA UNIFLORA L., Sp. Pl. 470. 1753.
Eugenia michelii Lam. Encycl. 3:203. 1789.
Stenocalyx michelii (Lam.) Berg, in Linnaea 27:310. 1856.

Shrubs or small trees 3-10 m. high. Leaves ovate to ovate-lanceolate, shortly
acuminate, obtuse to rounded or subcordate at the base, 2.5—6.0 cm. long and 1.5-
3.5 cm. broad, chartaceous, glabrous, lateral nerves prominulous on both sides,
arcuate-anastomosing at about 3 mm. within the margin, the veins laxly reticulate;
petiole about 2 mm. long. Flowers solitary in the axils of bracts at the base of
young shoots or sometimes in fascicles of 4-8; bracts scarious, up to 5 mm. long.
Pedicels slender, glabrous, 1.5—2.0 cm. long. Sepals oblong, ciliate, up to 4 mm.
long, glabrous. Petals obovate, unguiculate, 8-12 mm. long. Ovary glabrous,
8-ribbed, 2-celled; cells several (10—14-) ovulate; placenta peltate, thickened in
the middle. Berry depressed-globose, 8-ribbed, 1-seeded, 2.5—3.0 cm. in diameter,
red, edible. Cotyledons partly free.

Cultivated in the tropics and subtropics, native of tropical America. The
fruits are edible, sweet, aromatic and of excellent flavor. The shrubs are especially
well-adapted to cultivation as a hedge plant (Standley). Vernacular name:
Surinam Cherry.

CANAL ZONE: Barro Colorado Island, Shattuck 71.

3. EUGENIA FRAGRANS (Swartz) Willd. Sp. Pl. 2:964. 1800.

Myrcia seleriana Donn. Sm. in Bot. Gaz. 27:332. 1894

Small trees 3-8 m. high, almost glabrous. Leaves petiolate, elliptic to obovate,
acute at the base, obtuse or very shortly acuminate at the apex, 2-6 cm. long an
1.5-3.0 cm. wide, thick and coriaceous, glabrous, the midrib impressed above
prominent beneath, the lateral nerves usually prominulous on both sides. Flowers
in axillary 3- to 7-flowered cymes, the central flower(s) sessile, the lateral pedicel-
late. Peduncle up to 3.5 cm. long. Flowers 4-merous, white, fragrant. Sepals
about 2 mm. long, glabrous except at the margin. Ovary with sparse hairs, 2-
celled, many-ovulate. Petals glabrous, about 3 mm. long. Anthers orbiculate.

(182)

1958]
FLORA OF PANAMA (Myrtaceae) 183

Fig. 56. Eugenia fragrans

(183)

[Vol. 45
184 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Berry globose or ellipsoid, glabrous, orange, 1- or 2-seeded (or rarely more), 8
mm. long or larger. Cotyledons free with a short radicle between them.

West Indies and Central America. The Central American form is always with
4-merous flowers, the West Indian sometimes with 5-merous flowers. In Panama
at altitudes of 1000 m. or more.

CHIRIQUf: outskirts of wood, Volcán de Chiriqui, Seemann 1150; Río Chiriqui Viejo

valley, G. Wbite 108, P. Wbite 217, 211; Boquete, Davidson 460, 774, 821, 1062; vicinity
of Cerro Punta, alt. 2000 m., Allen 1563.

Seemann 1150 was named Eugenia arayan by Seemann (Bot. Voy. Herald 125.
1852-59) in mis-association with Myrtus arayan HBK. Nov. Gen. & Sp. 6:133.
1823, a Peruvian plant. Allen r563 is a rather distinct looking specimen with
suborbiculate, very rigid leaves with impressed nerves agreeing with Eugenia
rigidissima Cuf. from a volcano in Costa Rica.

4. EUGENIA BIFLORA (L.) DC. Prodr. 3:276. 1828.
Myrtus biflora L. Syst. 1056. 1759.
Eugenia sericiflora Benth. Bot. Voy. Sulphur 89. 1844.

Shrubs 3—4 m. high; twigs terete, at first brown-tomentose intermixed with
white hairs, later glabrate. Leaves lanceolate, acute at the base, long-acuminate
and mucronate at the apex, up to 7 cm. long and 1.0—2.5 cm. wide, subcoriaceous
or membranaceous, sparsely white-pilose on both sides, glabrescent, the midrib im-
pressed above, prominent beneath, the lateral nerves numerous, inconspicuous >
both sides. Inflorescence racemose, 3- to 7-flowered, tomentose. Flowers white.
Bracts setaceous, 2 mm. long; bractlets also setaceous, longer than the ovary.
Sepals ovate, apiculate, about 2 mm. long. Petals about 4 mm. long, sericeous
outside. Ovary 2-celled, sericeous; ovules about 6-10 in each cell, arranged in a
semicircle open above. Berry subglobose, 7 mm. long, smooth or faintly ribbed,
edible. ;

The species is distributed throughout tropical America in several varieties.
Vernacular name: coralillo.

COCLE: Penonomć and vicinity, Williams 63, 188, 355, 2776. cmiriquí: Cerro Vaca,
Pittier 5316. PANAMA: Taboga Island, Maxon 6920, Macbride 2776, Barclay 977; Stand-
ley 27024, 27886, 29732, Miller 1843, 2034, Pittier 3525. vERAGUAs: Santiago de Vera-
guas, Seemann 1149.

5. EUGENIA NESIOTICA Standl. in Field Mus. Publ. Bot. 17:203. 1937.

Densely branched shrub flowering precociously while defoliated; young tW!8$
grayish-puberulous, terete, densely foliate. Leaves elliptical-oblong, acute at the
base, obtusely acuminate at the apex, 4—5 cm. long, 1—2 cm. wide, glabrous except
the midrib, midrib prominulous above and prominent beneath, the veins finely BASS
ulate on both sides; petiole about 2.5 mm. long. Flowers fasciculate, the pedicels
white-pubescent, 5—6 mm. long. Sepals puberulous, about 2 mm. long, reflexed at
anthesis. Petals orbicular, 3.5 mm. long. Ovary densely puberulous, 2-celled, the
cells about 6-ovulate. "Berry resembling a huge cherry in color and texture, 25-
3.0 cm. in diameter, the flesh about 0.5 cm. thick, the taste acid, edible.
single, sphaerical, covered with fibre (Zetek).”

(184)

1958]
FLORA OF PANAMA (Myríaceae) 185

Endemic. Vernacular name: guayabillo.

CANAL ZONE: Barro Colorado Island, Zetek 3646, 3835, 3482, Starry 169, Standley
40987; Obispo, Standley 31712.

This is one of the very few Myrtaceae in which the flowers develop before the
leaves. The small reticulate leaves with the midrib elevated above characterize this
species even when sterile. A species with rather similar leaves is E. chepensis
Standl., in which the reticulation is more obscure and the marginal nerve single
and very close to the margin.

6. EUGENIA PITTIERI Standl. in Field Mus. Publ. Bot. 8:145. 1930.

Slender shrubs 1.0—4.5 m. high; young twigs densely and minutely pubescent,
densely foliate, the internodes 1.0—2.5 cm. long. Leaves more or less distichous,
almost sessile. oblong to oblanceolate, acute at the base, obtusely acuminate at the
apex, 4-6 cm. long, 1-2 cm. wide, membranous, glabrous except on midrib and
margin, densely glandular-punctate, the midrib flat or slightly elevated above,
prominulous beneath, the principal lateral nerves several but distant, diverging at
an angle of about 60” with the midrib, delicate, the marginal nerve very close to
the margin, the reticulation obscure throughout. Flowers minute, glomerate, few,
axillary and lateral. Sepals nearly 1 mm. long. Petals obovate, 2 mm. long.
Ovary glabrous, 2-celled, the cells 4- to 6-ovulate. Berry subglobose, sessile, 5—7
mm. long, red, crowned by the persistent sepals.

Panama and Costa Rica, at low altitudes.

PANAMA: banks of Mamoni R., below La Capitana, Pittier 4484.

The second Panamanian specimen cited by Standley, Mrs. A. P. Epplesheimer
s.n. is a specimen of Myrciaria floribunda.

7. EUGENIA CHEPENSIS Standl. in Field Mus. Publ. Bot. 8:144. 1930.

Slender shrubs 2-3 m. high, the young twigs densely and minutely pubescent,
densely foliate. Leaves more or less distichous, oblong or lanceolate-oblong, attenu-
ate at both ends, up to 4.5 cm. long and 1-2 cm. wide, membranous, glabrous,
the midrib flat or slightly elevated above, prominulous beneath, the lateral nerves
several but distant, diverging at an angle of about 60° from the midrib, the
marginal nerve very near the margin, the reticulation obscure throughout; petiole
about 2 mm. long. Flowers axillary, in a 1- to 4-flowered abbreviated raceme, the
Peduncle and pedicels minutely pubescent, the pedicels slender, 4-10 mm. long.
Petals not seen. Fruit a small globose berry, green to purple, about 6 mm. in
diameter, crowned by the 4 persistent semi-orbiculate sepals.

Nicaragua and Panama, at low elevations.

ation and Quebrada a

NE: forest between Peluca Hydrographic e je

CANAL ZO
Steyermark & Allen 1 ; P d cańon of Rio
7250; forest between Quebrada La Palma an
Dodge & Allen 1 7362; upper Río Chilibre, Seiber? 1501; Gatún, Hayes 77, 236, 610
Chagres, Fendler 189; without locality, Hayes 255. PANAMA: along Rio Charara, near
Chepo, Pittier 4716.

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[Vol. 45
186 ANNALS OF THE MISSOURI BOTANICAL GARDEN

8. EUGENIA COSTARICENSIS Berg, in Linnaea 27:213. 1856.

Slender shrubs, the young twigs minutely pubescent, soon glabrate, densely
foliate. Leaves more or less distichous, elliptical, acute at the base, obtusely
acuminate at the apex, 3-4 cm. long and 1-2 cm. wide, chartaceous, glabrous,
finely reticulate, the midrib impressed above and prominent beneath, the lateral
nerves several but distant, delicate, the marginal nerve very near the margin;
petiole slender, 4-6 mm. long, glabrous. Flowers in axillary 6- to 15-flowered
fascicles, the pedicels slender, sparsely pubescent, 4-6 mm. long, the peduncle
practically absent. Sepals 4, the larger about 1.5 mm. long. Petals suborbiculate,
about 2.5 mm. long. Style glabrous. Ovary glabrous, many-ovulate. Berry not
seen.

Costa Rica and Panama, at higher elevations, to 1900 meters.

VERAGUAS: locality unknown, Warscewicz s. n. (not seen).

In Berg's day, "Veraguas" included all of western Panama. From the descrip-
tion, this appears to be a very distinct species, but more recent collections have
failed to include it.

9. EUGENIA ORIGANOIDES Berg, in Linnaea 29:229. 1858.
Eugenia banghamii Standl. in Journ. Arnold Arb. 11:125. 1930.

Shrubs or trees 2.5—6.0 m. high; twigs densely brown-pubescent with spreading
hairs, glabrescent. Leaves elliptical to oblong, acute to rounded at the base, shortly
acuminate at the apex, 5-10 cm. long and 2-5 cm. wide, at first pubescent on both
sides, at length glabrate except on the midrib, the midrib scarcely impressed above,
prominent beneath, the lateral nerves about 10 on each side, prominulous on bo
surfaces, passing into a marginal nerve at about 2-3 mm. distance from the margin;
petiole robust, 3 mm. long, pubescent. Flowers glomerate, axillary, small, sessile
or nearly so, white or pale yellow. Sepals ciliate, up to 1 mm. long. Petals obovate,
3 mm. long, glabrous. Ovary sparsely pubescent, 2-celled; cells about 5-ovulate.
Berry globose, not more than 6 mm. in diameter, red to purplish-black, with thin
juicy pericarp, 1-seeded.

Distribution: Mexico to Colombia.

¡bert

406, Wood.
Shattuck 1043, Aviles s.n. cocLé: Penonomé and vicinity, Williams 214; DOC SB
Margaritas and El Valle, Woodson, Allen & Seibert 1727; Llano Bonito, north of Las

d e:

Archipelago, Johnston 137, 387, 741, 779, 489, Erlanson 66; near Punta Paitilla, Standley
26275; vicinity of Juan Franco Race Track, near Panamá, Standley 27790, 27719-
10. EUGENIA COLORADENSIS Standl. in Trop. Woods 52:27. 1937.
Eugenia melanosticta Standl. in Journ. Arnold Arb. 11:126. 1930, non Koorders & Valeton

Glabrous shrubs or trees. Leaves oblong, or elliptical- to obovate-oblong, pet
at the base, shortly and acutely acuminate at the apex, 8-17 cm. long, about A
cm. wide, quite variable in size, membranaceous to coriaceous, glabrous, the mi

(186)

1958]
FLORA OF PANAMA (Myrtaceae) 187

impressed above, prominent beneath, the lateral nerves in about 10 pairs, incon-
spicuous above, prominulous beneath, arcuate, anastomosing near the margin.
Inflorescences axillary, few-flowered, racemose, the racemes often gathered into a
panicle, the rachis and the pedicels rather robust, the pedicels 4-7 mm. long. Sepals
semi-orbiculate, unequal, the larger up to 2.5 mm. long. Petals oblong, nearly
7 mm. long. Ovary glabrous, 2-celled; ovules about 4-5 in each cell, pendulous.
Berry ovoid-globose, 10-12 mm. long, crowned by the conspicuous sepals.
Costa Rica and Panama. Known as guayabito de monte.

CANAL ZONE: Lion Hill station, Hayes s. n.; Barro Colorado Island, Aviles 21, Shattuck
1118, Bangham 445, Salvoza 909, Zetek 4508.

Perhaps not distinct from the following:

11. EUGENIA OERSTEDEANA Berg, in Linnaea 27:285. 1856.

(?) Eugenia balancensis Lundell, in Phytologia 1:481. 1941.

Glabrous trees; twigs slender, much branched. Leaves ovate or oblong, acute
at the base, long-acuminate at the apex, 4-8 cm. long and 2-4 cm. wide, membrana-
ceous, glabrous, the midrib impressed above, prominent beneath, the lateral nerves
up to about 12 pairs, delicate, inconspicuous, arcuate-anastomosing at about 2—4
mm. from the margin; petiole about 4 mm. long. Racemes glabrous or minutely
puberulous, axillary, sometimes the flowers partly solitary, the fully developed ra-
cemes 7- to 11-flowered, the rhachis 2-3 cm. long, the pedicels filiform, 3-5 (-10)
mm. long, the uppermost flower usually sessile. Flowers white, small. Sepals up to
1.5 mm. long. Petals 3 mm. long, not covered by the calyx in the bud. Ovary
glabrous, 1 mm. high. Berry globose, 1-seeded, crowned by the small calyx.

Mexico, Costa Rica and Panama, in shady forests. Vernacular name: sequarra.

CANAL ZONE: in forest near the northerly arm of Quebrada Salamanca, Steyermark 8
Allen 17149; Rio Paraíso, Standley 29853; Darién station, Standley 31 596. CHIRIQUÍ:
Progreso, Cooper & Slater 158. PANAMÁ: in woods near the Hacienda de Juan Lanas,
Seemann 476; Bojio station, Hayes 600; in woods near Manuel station, Hayes s. n.

Closely allied to the preceding species, but apparently more slender in all parts
and with smaller flowers and leaves.

12. EUGENIA sp.

Shrubs or small trees about 4 m. high. Leaves oblong, rounded at base, obtuse
at apex, 20.0-26.5 cm. long and about 10 cm. wide, coriaceous, glabrous, dis-
colorous, the midrib flat above, prominent beneath, the lateral nerves numerous,
principal ones in about 20 pairs, prominulous, the lesser ones prominulous above
and obsolete beneath; petiole about 5 mm. long. Flowers apparently lateral,
fasciculate, possibly sessile. Berry oblong, tomentose, about 2 cm. long and 1 cm.
broad, crowned by 4 rounded sepals up to 3 mm. long.

DARIÉN: Paca, just below Cana, Williams 701.

(187)

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

188

f
|

i
J

U

3
N

A

L)

Z

Fig. 57. Eugenia acapulcensis

(188)

1958]
FLORA OF PANAMA (Myrtaceae) 189

Although quite distinct in appearance, the material presently available is
insufficient for the purpose of formal designation.

13. EUGENIA ACAPULCENSIS Steud. Nomencl. Bot. 1:601. 1840.

Myrtus maritima HBK. Nov. Gen. & Sp. 6:146. 1823.
Eugenia maritima (HBK.) DC. Prodr. 3:282. 1828, non E. maritima DC. loc. cit, 3:271.

1828.

Eugenia colipensis Berg, in Linnaea 29:245. 1858.
(?) Eugenia cartagenensis Berg, loc. cit. 1858.
Eugenia antiquae Riley, in Kew Bull. 1927:121. 1927.

Shrubs or trees 3-12 m. high, with glabrous twigs. Leaves elliptic, rarely
oblong or lanceolate, acute or attenuate at the base, shortly and obtusely acuminate
at the apex, 5-12 cm. long and 2-6 cm. wide, glabrous but densely glandular-
punctate, the midrib slightly impressed above, prominent beneath, the lateral
nerves distant, in 6—9 pairs, prominulous on both surfaces, the lower ones curved
at their base, passing into a marginal nerve at about 1—3 mm. distance from the
margin; petiole up to 1 cm. long. Flowers green or white, very small, in numerous
axillary abbreviated racemes, the rachis up to 1 cm. long though often much shorter
(in the latter case perhaps accrescent during fruiting), the pedicels filiform,
glabrous or puberulous. Ovary glabrous. Sepals 4, very short and scarcely 1 mm.
long, semi-orbiculate. Petals densely glandular-punctate, about 3 mm. long.
Berry globose, dark red to black (or sometimes yellow?), about 8 mm. in diameter,
crowned with the very small connivent sepals. Embryo homogeneous.

A polymorphic species distributed throughout Central America. In Panama
it seems to be the most common Myrtacea, on the Pacific coast especially frequent.
Possibly extending to Colombia and scarcely distinguishable from some forms of
the West Indian E. axillaris (Sw.) Willd. Eugenia cartagenensis Berg, described
from Costa Rica, is apparently nothing more than a small-leaved E. acapulcensis,
although with ovoid fruit (always?). The latter form is represented in Panama
by Williams 58 and Allen 2516 from Coclć, but with globose fruits. Eugenia
antiquae Riley was described from a Panama specimen, and that name must be
used in Panama if several species are recognized. Vernacular name in Panama:
baico.

CANAL ZONE: vicinity of Miraflores Lake, P. White 108, 118, G. Wbite 147, Allen
1712; Chagres, Fendler 183. cHIRIQUI: Finca Lérida, Woodson 3 Schery 210. PANAMA:
savana east of Pacora, Woodson, Allen & Seibert 726; Isla Taboga, Woodson, Allen 8
Seibert 1483; hills above Campana, Allen 1324; Old Panamá, Riley 142; Sabanas, Panama,
Riley 110; El Jagua Hunting Club on Río Jagua a or Hi ;
Trapiche Island, Allen 2631; Saboga Island, Miller 1955; San José Island, Jobnston 727,
734, 736, 748.

14. EUGENIA HIRAEIFOLIA Standl. in Field Mus. Publ. Bot. 17:202. 1937.
Shrubs or trees. Leaves obovate, cuneate toward the base, obtuse or subtrun-
cate and very shortly apiculate at the apex, 12-14 cm. long and 6-8 cm. wide,
glabrous, the midrib impressed above and prominent beneath, the lateral nerves
straight, in about 8—11 pairs, arcuate-anastomosing at some distance from the

5
a
5

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[Vol. 45
190 ANNALS OF THE MISSOURI BOTANICAL GARDEN

margin but for the greater part straight, ascending, nearly obsolete above, prom-
inent beneath; petiole robust, about 7 mm. long. Inflorescence (imperfectly
known) apparently cymose, 3-flowered; peduncle robust, about 2 cm. long.
Flowers not known. Berry large, obovoid, 1-seeded, about 2.5 cm. long and 1 cm.
broad, crowned by the persistent 4-merous calyx. Embryo homogeneous.

Endemic. A rather distinct but still imperfectly known species. The leaves,
by their venation, are strongly suggestive of a Psidium.

PANAMA: Monte Obscuro, near Panamá City, Zeżek 3550.

A deflorated specimen collected by Davidson (no. 439) in Chiriqui Province,
near Bajo Chorro, comes near E. hiraeifolia, but seems distinct by the definitely
acuminate leaves, the fasciculate inflorescences which are mixtures of 1-flowered
pedicels and one or two few-flowered racemes, and larger sepals up to 5 mm. long.

15. EUGENIA SALAMANCANA Standl. in Ann. Missouri Bot. Gard. 26:295. 1939.

Trees up to 6 m. high, the twigs terete, at first densely brown-pubescent, soon
glabrate. Leaves oblong or elliptical-oblong, rounded and shortly caudate-
acuminate at the apex, narrowly rounded toward the base, 7-9 cm. long and 2.5-
4.0 cm. wide, concolorous, glabrous with the exception of the midrib, the midrib
slightly impressed above, prominent beneath, the lateral nerves about 12 on each
side, obscure, the veins obsolete. Flowers probably sessile, solitary or in clusters of
few, lateral on the lower defoliated part of the twigs. Mature flowers and fruit
not known.

Endemic. An imperfectly known species.

CANAL ZONE: vicinity of Salamanca Hydrographic Station, Rio Pequeni, alt. 80 m.,
Woodson, Allen & Seibert 1570.

16. EUGENIA TAPACUMENSIS Berg, in Linnaea 27:222. 1856.
Eugenia ochra Berg, loc. cit. 216. 1856.
Eugenia roraimana Berg, loc. cit. 219. 1856.

Subglabrous shrubs or small trees up to 10 m. high. Leaves ovate- to elliptical-
oblong, obtuse or shortly and obtusely acuminate at the apex, obtuse or acute at
the base, 5-15 cm. long and 4-7 cm. wide, chartaceous to coriaceous, glabrous, the
midrib impressed or sulcate above, prominulous toward the base below, the lateral
nerves about 12-14 on each side, oblique, prominulous on both sides, arcuate-
anastomosing into a marginal nerve 2-3 mm. within the margin; petiole 4-10 mm.
long. Flowers mostly borne on the defoliated branchlets, pink or white, in short
racemes or fascicles of 2-15, subglabrous, the rachis if well developed up to 8 mm.
long, the pedicels 3-5 mm. long. Calyx waxen-white, the sepals rounded, glabrous,
ciliate, about 1 mm. long, even in the bud much shorter than the petals. Petals
chartaceous, orbiculate, about 5 mm. long. Ovary globose, glabrous, 2-celled;
placenta peltate; ovules about 15 in each cell. Berry subglobose, black, when dry
about 1 cm. in diameter; testa of seeds shiny; embryo undivide

Panama to the Guianas and Peru; West Indies (Martinique, St. Vincent).

(190)

1958]
FLORA OF PANAMA (Myrtaceae) 191

CANAL ZONE: Rio Grande railway station, Hayes 344. PANAMA: in woods near
Panamá, Hayes s. n.-

It is odd indeed that no more recent records of this species are available.

17. EUGENIA OCTOPLEURA Krug & Urban, in Engl. Bot. Jahrb. 19:653. 1895.

Shrubs 3-5 m. high, the young twigs very shortly pubescent at their tips with
dibrachiate hairs, soon glabrate. Leaves elliptical or elliptical-oblong, acute at
the base, obtusely acuminate at the apex, 7-13 cm. long and 4-5 cm. wide, cori-
aceous, glabrous, the midrib impressed above, prominent beneath, the lateral nerves
in about 12 pairs, obsolete above, prominulous beneath; petiole 8-10 mm. long.
Inflorescence shortly racemose, 2- to 8-flowered, as long as the petiole or somewhat
shorter, sparsely sericeous; pedicels 4-8 mm. long. Flowers 4-merous. Sepals un-
equal, the 2 outer orbiculate, 2.5-3.0 mm. long, the 2 inner 1.0-1.5 mm. long.
Petals obovate, 3-4 mm. long. Ovary obconic, pubescent outside but glabrescent,
2-celled; ovules 12—18 in each cell, inserted on a peltate placenta. Berries obovate-
oblong, distinctly 8-ribbed, up to 1.5 cm. long.

sser Antilles (Guadeloupe, Martinique, and Dominica) and Panama.

CANAL ZONE: Maumey station, Panamá Railroad, Hayes 577.

18. EUGENIA AUSTIN-SMITHII Standl in Field Mus. Publ. Bot. 18:1561. 1938.

Trees with slender glabrous twigs. Leaves elliptical-oblong to ovate-oblong,
acute at the base, acuminate at the apex, 6—7 cm. long an 2.5—3.5 cm. wide,
coriaceous, glabrous, the midrib impressed above, prominulous beneath, the lateral
nerves about 12 on each side, prominulous on both surfaces; petiole canaliculate,
up to 8 mm. long. Flowers mostly cauliflorous; pedicels robust, up to I cm. long.
Flower buds pear-shaped, the sepals enclosing the petals. Sepals greenish, rounded,
reflexed at anthesis, thick, unequal, the 2 inner up to 5 mm. long, the 2 outer nearly
3 mm. long. Petals white, about 7 mm. long. Ovary obconoid, glabrous or yim
a few hairs, 2-celled, about 3 mm. high, the cells about 12-ovulate. Berry said to
be oblong or globose, deep crimson, about 2.0—2.5 cm. in diameter.

Originally described from Costa Rica, in dry woods at medium altitudes. No
specimens from Panama have been seen by the writer, but it is to papę»
the highlands of western Panama.

19. EUGENIA CHRYSOPHYLLUM Poir. in Lam. Encycl. Suppl. 3:129. 1813.
Eugenia cricamolensis Standl. in Field Mus. Publ. Bot. 4:242. 1929.

Shrubs or trees; young twigs brown-tomentose, soon glabrate. Leaves oblong
to lanceolate-oblong, long acuminate at the apex, acute at the base, 7-15 cm. long
and 3-5 cm. wide, chartaceous to coriaceous, the young leaves densely golden
brown-sericeous on both sides, soon glabrescent above, with hairs closely appressed,
the adult leaves discolorous, glabrate above and paler and appressed-pubescent
beneath, the midrib flat, prominulous toward the apex above, prominent beneath,
the lateral nerves in about 12-16 pairs, prominulous on both sides; petiole 3-7 mm.

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[Vol. 45
192 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long. Flowers fasciculate or shortly racemose (the rachis up to 5 mm. long), in
clusters of 3-8, axillary; pedicels 2-7 mm. long, brown-tomentose; bractlets
minute, narrow. Sepals tomentose, very unequal, the 2 outer suborbiculate, about
1.5 mm. long, the 2 inner ovate, about 3 mm. long. Petals obovate or oblong,
about 5 mm. long, glandular. Ovary tomentose, 2-celled, many-ovulate. Berry
ellipsoid, about 15 mm. long and 9-12 mm. wide in our specimens (up to 2.5
cm. in diameter according to Cooper), grayish-puberulent, 1-seeded, the embryo
homogeneous.

Surinam and French Guiana, otherwise known only from the following speci-
men from Panama. The Panamanian specimen has fewer lateral leaf-nerves than
those of the Guianas (12 instead of 16) and also perhaps larger fruits, but other-
wise the populations appear to be conspecific. Vernacular name: cacique.

BOCAS DEL TORO: region of Almirante, Cricamola valley, Cooper 511.
20. EUGENIA OREINOMA Berg, in Linnaea 27:158. 1856.

Small trees 4-5 m. high, the trunk with grayish, exfoliating bark; young twigs
and petioles densely pubescent. Leaves ovate or broadly elliptical, rounded or ob-
tuse at the base, shortly acuminate at the apex, 9-12 cm. long and 4.5-6.0 cm.
wide, reticulate-veined, sparsely pubescent but glabrescent above, densely pubescent
with patent hairs beneath, especially on the nerves, the midrib flat above, prominent
beneath, the lateral nerves in about 8 pairs, arcuate-anastomosing at some distance
from the margin, prominulous above, prominent beneath; petiole robust, about 5
mm. long, pubescent. Flowers fasciculate but occasionally solitary, the pedicels
about 1 cm. long, pubescent. Flowers not known. Fruit globose, usually densely
pilose, 1-seeded, about 4 mm. long, black or purplish, crowned by 4 persistent,
striate sepals.

Costa Rica and Panama. Vernacular names: icaquillo, sortija.

CHIRIQUÍ: Sabana de la Tortuga, between El Boquete and Caldera, alt. 300-700 m»
Pittier 3297.

7. CALYCOLPUS Berg
CALYCOLPUS Berg, in Linnaea 27:378. 1856.

Trees or shrubs. Inflorescences axillary, usually 1-flowered, rarely 3-flowered,
singly or in fascicles of 2-5, at the tips of extremely abbreviate, minutely bracteate
shoots; bracteoles very small, paired at the base of the ovary; flowers 5-merous-
Sepals large and persistent, coriaceous-foliaceous, patent in the bud. Receptacle
not produced above the ovary. Petals obovate, large. Stamens numerous,
anthers linear-oblong, dorsifixed near the base, dehiscing longitudinally. Ovary p
to 6-celled, many-ovulate; placentas 2-lobed; ovules oblong. Berry subglobose,
many-seeded. Seeds hippocrepiform, yellowish white, one side with a blackish spo
shining, finely reticulate, hard. Embryo incurved, the radicle elongate, the Coty”
ledons very small.

Species about 7 in tropical America.

(192)

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(2023034) VINYNVA HO VUOTA

£61

[8561

[Vol. 45
194 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. CALYCOLPUS WARSZEWICZIANUS Berg, in Linnaea 27:382. 1856; Riley, in Kew
Bull. 1926:153. 1926

Glabrous shrubs or small trees up to 7 m. high, with exfoliating bark; twigs
subterete or faintly angled. Leaves ovate- or elliptical-oblong, obtuse or acute at
the base, acutely acuminate at the apex, 3-8 cm. long and 1.5-3.0 cm. wide,
firmly membranaceous to subcoriaceous, glabrous, the midrib impressed above and
prominent beneath, the lateral nerves delicate, numerous, prominulous on both
sides; petiole up to 3 mm. long. Peduncles glabrous, axillary, 1.5—3.0 cm. long,
mostly single; bractlets linear to oblong, 1-3 mm. long; flowers white to pinkish,
fragrant. Sepals oblong, obtuse, accrescent, 5-10 mm. long, with distinct mid-
nerve. Petals obovate-oblong, about 1.5 cm. long. Ovary subglobose, glabrous.
Berry subglobose, up to 1 cm. long, crowned with the large accrescent disc and
persistent sepals.

Panama and Costa Rica.

tias
Erlanson 60; in meadows near city of Panamá, Seeman

8. UGNI Turcz.
UGNI Turcz. in Bull. Soc. Nat. Moscow 21:579. 1848.

Low much branched and densely foliate shrubs. Leaves opposite or subopposite,
small. Peduncles axillary, 1-flowered. Flowers medium-sized, 4- or 5-merous.
Sepals patent in the bud, longer than the imbricated petals. Petals firm. Stamens
numerous, unequal; filaments straight or hardly curved even in the bud, short and
flat; anthers basifixed, 2-celled, dehiscing longitudinally, introrse. Ovary 3- to >
celled; cells several-ovulate; placenta central. Berry several-seeded; seeds hippo-
crepiform, with a hard and shining testa with a black spot at one side; embryo
incurved; radicle elongate; cotyledons very short.

ut 15 species of high altitudes, from southern Chile to Guatemala and
Venezuela.

1. UGNI WARSCEWICZI Berg, in Linnaea 27:390. 1856.
Eugenia warscewiczii (Berg) Hemsl. Biol. Centr. Amer. Bot. 1:411. 1880.

Low densely foliate shrubs 2-3 m. high, the branches glabrous, the internodes
on the defoliated parts of the twigs marked by the scars of the fallen leaves, the
innovation shoots more or less pubescent. Leaves oblong or elliptical-oblong, me
at both ends, up to 15 mm. long and 8 mm. wide, coriaceous, glabrous, more oF *
impressed-punctate, the midrib impressed above, prominent beneath, the vena T
obsolete. Pedicels recurved, up to 1 cm. long, pubescent; bractlets linear, ciliate,
persistent, longer than the ovary, about 6 mm. long. Sepals lanceolate, ob

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[8561

[Vol, 45
196 ANNALS OF THE MISSOURI BOTANICAL GARDEN

about 5 mm. long. Petals obovate, about as long as the sepals. Anthers of the
outer stamens lanceolate, acutish, crowned by an apical gland, the anthers of the
inner shorter stamens ovate, truncate, without glands; filaments flattened. Ovary
glabrous or sparsely pubescent, 3- to 4-celled; ovules inserted on a peltate central
placenta. Berry depressed-globose, 8 mm. in diameter, bluish black, several-seeded.

Endemic to Panama. U. oerstedii Berg is a closely allied species of Costa Rica.

CHIRIQUi: Volcán de Chiriqui, alt. 2500-4000 m., Woodson, Allen & Seibert 1066,
Woodson & Schery 439, Pittier 3075.

9. PSIDIUM L.
Psiprum L. Spec. Pl. 470. 1753.

Guiava [Tourn.] Adans. Fam. 2:88. 1763.
cca Berg, in Linnaea 27:138. 1856
Calyptropsidium Berg, loc. cit. 349. 1856.

Shrubs or trees. Leaves opposite or subalternate, the lateral nerves mostly
ascending, arcuate-anastomosing without forming a distinct marginal nerve.
Inflorescence cymose, 1- to 7-flowered; peduncles in the axils of the lower often
bract-like leaves of lateral branchlets. Flowers large or medium-sized. Receptacle
produced above the ovary. Calyx closed in the bud or open and then truncate or
5-lobed, at anthesis irregularly 3- to 5-fid, sometimes at length irregularly circum-
scissile. Petals 5, orbiculate or obovate, white. Stamens numerous, the filaments
incurved in the bud; anthers dorsifixed, dehiscing longitudinally. Ovary 2- to
7-celled, many-ovulate; placenta 2-lamellate; ovules in a single or triple row;
stigma capitate or peltate. Berry mostly globose, many-seeded; seeds embedded in
pulp, the testa dull, hard. Embryo incurved; radicle elongated; cotyledons very
short.

Species about 60 in tropical America. A few species are cultivated throughout
the tropics of the world for their edible fruits.

la. Calyx open in the bud; leaves small, subsessile........ ... 1. Ps. SALUTARE
b. Calyx closed in the bud; leaves shortly petiolate simu i slot ;
2a. Plants glabrous, ili LA IU UT NE :
b. Plants pubescent. Lucio ee NE ;
3a. Leaves 2.5—5.0 cm. long; young twigs subterete.......... 2. Ps. SARTORIANUM NUM
b. Leaves 8—12 cm. long; ng twigs narrowly 4-angled....................... 3. Ps, FRIEDRICHSTHALIA
4a. Leaves usually with 12 or mo rs of close parallel lateral nerves
impressed above; young twigs 4-angular................__ Ps. GUAJAVA

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5. Ps. GUINEENSE

1. PSIDIUM SALUTARE (HBK.) Berg, in Linnaea 27:356. 1856.
Myrtus salutaris HBK. Nov. Gen. & Sp. 6:105. 1823
Psidium cili

sidiu atum Benth. in Hook. Journ. Bot. 2:318, 1840.
0. 1856

Undershrub often less than 1 m. high, simple and sparingly branched or with
several stems; twigs compressed, puberulous or glabrous. Leaves subsessile, ova

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1958]

197

FLORA OF PANAMA (Myrtaceae)

Fig. 60. Psidium salutare

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[Vol. 45
198 ANNALS OF THE MISSOURI BOTANICAL GARDEN

to elliptical-oblong, rounded or obtuse at the base, acute at the apex, up to 6 cm.
ong and 2 cm. wide, coriaceous, at first more or less puberulous, later glabrate, the
midrib elevated on both sides or nearly flat above, the lateral nerves in about 6
pairs, arcuate-ascending, prominulous on both sides; petiole obsolete, often with
2 minute setaceous stipules. Peduncles 1-3 cm. long, solitary, axillary, 1-flowered.
Calyx open in the bud, later 5-fid to the base, the lobes rounded or obtusish. Petals
about 7 mm. long. Anthers suborbiculate, cordate. Ovary puberulous or glabrous,
obconoid, 2 mm. high; placenta 2-lamellate; ovules in a single or double row at
each side of the placenta. Berry subglobose, when dry about 1 cm. in diameter,
said to be about as large as a cherry and of good flavor.
Honduras and El Salvador to the Amazon basin. Vernacular names: guayabito
arrayán; guayabito del Perú.
ZONE: Ancón Hill, Piper 5535, Standley 25208, 26340. cHIRIQUI: Boquete
District, Davidson 835; Sabana de la Tortuga, between El Boquete and Caldera, Pittier
3300. COCLE: Penonomé and vicinity, William È NAMA: hills northeast of Haci-

e PA
enda La Gorga, 50-300 m., Dodge, Hunter & Steyermark 16899, Dodge, Hunter, Steyer-
mark & Allen 8751.

2. PSIDIUM SARTORIANUM (Berg) Ndz. in Engl. & Prantl, Nat. Pflanzenfam. 37:
69. 1893.

Mitranthes sartorianum Berg, in Linnaea 29:248. 1858.

Calyptropsidium sartorianum (Berg) Krug & Urb. in Engl. Bot. Jahrb. 19:581. 1895.
Calybtrantbes tonduzii Donn. Sm. in Bot. Gaz. 23:245. 1897.

Mitropsidium sartorianum (Berg) Burret, in Notizbl. 15:487. 1941.

Psidium solisii Standl. in Field Mus. Publ. Bot. 23 :133. 1944.

Shrubs or small trees, sometimes 15 m. high, the bark gray, smooth, in all parts
glabrous or nearly so; young twigs subterete. Leaves petiolate, ovate, rounded ot
obtuse to nearly acute at the base, acute or acutely acuminate at the apex, 2.5-5.0
cm. long and up to 2.5 cm. wide, coriaceous, glabrous, the midrib flat above,
prominent beneath, the lateral nerves in about 6 pairs, ascending, obsolete above,
prominulous beneath; petiole about 3 mm. long, slender. Peduncles 1-flowered,
1.0—1.5 cm. long, slender. Calyx closed in the bud, later irregularly splitting with
1 large more or less lid-like deciduous segment and 2—3 other minute persistent
segments.! Petals rounded, about 4 mm. long, ciliate, early-deciduous. Ant
ovate. Ovary glabrous, usually 3-celled; placenta 2-lamellate; ovules in a single
row at each side of the placenta. Fruit subglobose, 1.5-2.0 cm. in diameter,
greenish yellow, several-seeded; seeds angular, concolorous. Embryo coiled with
less than 2 spirals, embedded in a coarse-celled pulp.

Mexico to Panama, apparently becoming rarer toward the south; Cuba.
Arraydn, guayabillo.

PANAMA: San Josć Island, Jobnston 486.

_

e accompanying drawing, with neatly circumscissile calyx, is somewhat misleading; such

1 Th
abscission is exceptional

(198)

1958]
FLORA OF PANAMA (Myrtaceae) 199

Fig. 61. Psidium sartorianum

(199)

[Vol. 45
200 ANNALS OF THE MISSOURI BOTANICAL GARDEN

3. PSIDIUM FRIEDRICHSTHALIANUM (Berg) Ndz. in Engl. & Prantl, Nat. Pflanzen-

fam. 37:69. 1893
Calyptropsidium friedricbstbalianum Berg, in Linnaea 27:350. 1856.

Glabrous shrubs and small trees to 9 m. high, with spreading crown and crooked
bole, the bark smooth and fluted, brown, the wood fine-grained and hard; young
twigs 4-angular. Leaves shortly petiolate, elliptical or oblong, 8-12 cm. long and
4-5 cm. wide, membranaceous to chartaceous, glabrous, the midrib impressed above
and prominent beneath, the lateral nerves in about 8 pairs, prominulous on both
sides, especially beneath; petiole about 3 mm. long. Peduncles 1- to 3-flowered,

Fig. 62. Psidium friedrichsthalianum

1.0-2.5 cm. long. Calyx globose in the bud, closed and apiculate, later 2- to 3-fid,
the lobes circumscissile slightly above the ovary but remaining attached at one
point and at length deciduous, about 1 cm. long. Petals orbiculate, about 1.5 cm.
long or slightly less. Anthers oblong. Ovary glabrous, 3- to 5-celled; placenta
2-lamellate; ovules in 2-3 rows, involute; stigma peltate. Berry oval or globose,
yellow, edible.

Throughout Central America. Frequently cultivated for the fruits which are
smaller than those of the common guava, have a tart agreeable flavor and are some-
times used for jellies. Vernacular name: guayaba de agua.

BOCAS DEL TORO: Changuinola valley, Cooper & Slater 97, 97a. CANAL ZONE: around
Gamboa, Pittier 4809, 6521. cHIRIQUI: Progreso, Cooper & Slater 278.

4. PSIDIUM GUAJAVA L. Sp. PI. 470. 1753.
Psidium bomiferum L. loc. cit. 672. 1762.
Psidium pyriferum L. loc. cit. 1762.

Shrubs or small trees with peculiar pale brown bark that scales off in thin sheets;
young twigs 4-angular, pubescent. Leaves elliptical to oblong, rounded, obtuse oF
apiculate at the apex, rounded or obtuse at the base, 4—12 cm. long and A
cm. broad, chartaceous, glabrescent on both sides, the midrib impressed above 22
prominent beneath, the lateral nerves 12-16 on each side, impressed above a
prominent beneath, parallel and arcuate-anastomosing near the margin; petiole 3
mm. long. Peduncles axillary, 1- to 3-flowered, pubescent. Flower buds pubescent,

(200)

1958]
FLORA OF PANAMA (Myrtaceae) 201

constricted under the calyx, 1.0-1.5 cm. long. Calyx closed in the bud, ellipsoid,
irregularly 4- to 5-fid at anthesis, its segments about 1 cm. long, sericeous inside.
Petals 1.5-2.0 cm. long. Anthers linear-oblong. Ovary 3- to 5-celled; placenta
2-lamellate; ovules in a triple row at each side of the placenta, turned in all direc-
tions. Berry pear-shaped or globose, 3-6 cm. in diameter, glabrate.

Cultivated throughout the tropics.

Ancón, Pittier 2725. CHIRIQUÍ: Llanos del Volcán, Allen 1536. cocLé: hills south of El
Valle de Antón, Allen 2524 (in part: mixed with Ps. guineense) ; Penonomé and vicinity,
Williams 237. PANAMA: Juan Díaz region, near Tapia R., Maxon 6 Harvey 6738.

The common guava, native in tropical America and cultivated throughout the
tropics of the world, is abundant in pastures and rather dry thickets. On the
Pacific slope of Central America it often forms extensive thickets of characteristic
appearance known as guayabales. The wood is brownish or reddish gray, hard,
strong, elastic, close-grained and durable, but the trees are too small to be of much
use. The Spanish name for the fruit is guayaba, and for the tree guayabo. The
fruit varies greatly in size, shape, color and flavor, and has a musky odor that is
remarkably penetrating. Raw it is somewhat insipid, but it makes an excellent
jelly and is prepared in other ways. In general the fruit is little esteemed in Central
America, but slices of the stiff, dark colored jelly are a frequent dessert dish.

5. PSIDIUM GUINEENSE Sw. Prodr. 77. 1788.

Psidium polycarpon Anders. ex Lamb. in Trans. Linn. Soc. 11:231. £. 17. 1815.
Psidium araca Raddi, Mem. 5. £. I. 1821.

Psidium molle Bertol. Fl. Guat. 422. t. 9. 1840.

Guajava mollis (Bertol.) O. Ktze. Rev. Gen. 239. 1891.

Shrubs usually less than 1 m. high; young twigs compressed to subterete, pubes-
cent. Leaves elliptical to oblong, rounded, obtuse or acutish at the apex, obtuse
or acute at the base, up to 13 cm. long and 7 cm. broad, chartaceous, pubescent on
both sides when young, glabrate except the midrib and veins beneath when adult,
the lateral nerves about 7-10 on each side, prominulous to flat above, prominulous
beneath, arcuate-anastomosing near the margin; petiole 4-10 mm. long. Peduncles
l- to 3-flowered, 2-3 cm. long, pubescent. Flower buds pubescent, constricted
under the calyx, 1.0-1.5 cm. long. Calyx irregularly 4- to 5-fid at anthesis, the
segments about 7 mm. long, sericeous within. Petals 5, about 1.5 cm. long. An-
thers linear-oblong. Ovary about 5 mm. long, 3- to 5-celled; ovules in a triple
row at each side of the 2-lamellate placenta, turned in all directions. Berry sub-
globose, about 2 cm. in diameter, yellow.

Native of tropical America and cultivated in other tropical countries.
nacular names: guayabita, guayaba arraydn; guayaba de sabana.

R. Ser : Volcán and near Río
Chir ind Velo ale toe, T c pc wś pine ure de Chiriquí, W ood son
8 Schery 477, 505. cocLé: between Las Margaritas and El Valle, Allen ES Seibert 1295.
PANAMÁ: San José Island, Johnston 16, Erlanson 290; La Campana, Zetek 4228.

(201)

Ver-

MISSOURI BOTANICAL GARDEN . l 3 i: pa a
| | SQ ae um Acting Director ; DA x : 7 RE Ji |
EDGAR ANDERSON, ` = Oscar E. GLAESS v did y
Curator of Useful Plants — BIEGA ; A vinis. 2 i = JE Pipa Y

© Paleobotanist | —

a Fedde ES Cycadaccze to (RAR (June, 194 :
E | Fascicle 2 È frana to Cyclanthaceae _(September, 194).

Number 3

Annals
of the ^ 1

Missouri Botani

Annals
of the

Missouri Botanical Garden

A Quarterly Journal containing Scientific Contributions from the
Missouri Botanical Garden and the Henry Shaw School of Botany of
Washington University in affiliation with the Missouri Botanical
Garden.

i Information
The ANNALS OF THE Missouri BOTANICAL GARDEN appears four times
during the calendar year: February, May, September, and November. Four
numbers constitute a volume.
Beginning with Volume 45, 1958:
Subscription Price $12.00 per volume
Single Numbers __ | 3.00 each
of previous issues of the ANNALS OF THE MISSOURI BOTANICAL

Conten
GARDEN are re edi in the Agricultural Index, published by the H. W. Wilson
Company.

FLORA OF PANAMA

BY

ROBERT E. WOODSON, JR.
AND
ROBERT W. SCHERY

AND COLLABORATORS

PART VII

Fascicle 3

MELASTOMATACEAE (H. A. Gleason)

ANNALS
OF THE
MISSOURI BOTANICAL GARDEN

Vol. XLV SEPTEMBER, 1958

FLORA OF PANAMA

Part VII. Fascicle 3*

MELASTOMATACEAE

By H. A. GLEASON

Flowers regular, perigynous or epigynous, usually 4- to 6-merous. Hypan-
thium cylindric to saucer-shaped, bearing the sepals, petals and stamens on a
terminal torus. Calyx usually small, in most genera open in the bud, in some
genera calyptrate. Petals distinct, large or small, usually white to pink or purple,
rarely red or yellow, convolute in the bud. Stamens commonly twice as many
as the petals, rarely more numerous, rarely fewer by abortion, isomorphic or
dimorphic. Anthers basifixed, ovoid to subulate, 2-celled or 4-celled, opening by
one (rarely two) terminal pores or in a few genera by short sub-terminal slits;
connective often prolonged below the thecae or below the insertion of the filament
into one or more lobes, spurs or appendages of various types. Ovary 1, 2- to
several-celled, superior or inferior, commonly many-ovuled; style 1; stigma 1,
punctiform to capitate. Fruit a loculicidal capsule more or less enclosed by the
persistant hypanthium if from a superior ovary, or a berry if from an inferior ovary.
Trees, shrubs, herbs or vines; leaves simple, opposite, often unequal, occasionally
apparently alternate by abortion of one member of each pair, entire or toothed,
usually 3- to several-nerved (1-nerved in Mouriri and a few species of other
genera). Inflorescence always cymose, axillary or terminal, or flowers solitary.

About 200 genera and 4000 species, almost entirely tropical and most numer-
ous in South America.

a. ie a loculicidal capsule; ovary superior, free from the hypanthium.
- Connective of the anthers bearing one or more dorsal spurs or append-
ages

€. Flowers (in our species) 5- to 6-merous; seeds linear or cuneiform.
(Trib

d. Calyx open in the bud, exposing the corolla. BA
e. Climbing vines; leaves subrotund; pubescence of Malpighian
hairs; seeds linear, winged at both ends, the embryo near the

1. ADELOBOTRYS.

t
not Malpighian; seeds narrowly cuneiform,
embryo at the base.

> : . MERIANIA.
f. Basal spur of the connective sharp or conic.........———7 2. M

'_—@————
* : :
Assisted by a grant from The National Science Foundation.
n

Issued October 28, 1958. (203)

(203)

[Vol. 45
204

ANNALS OF THE MISSOURI BOTANICAL GARDEN

ff. Basal spur of the connective inflated, ri ed 3. AXINAEA.
dd. a calyptrate in the bud, at anthesis liże at or near the

e. "Pet em obovate or rounded, in our species about 18 mm. long. 4
ee. Petals lanceolate to ovate, acuminate, in our species up to
mm. long
cc. Flowers 4-merous; seeds short, cochleate. (Tribe R
bb. Connective of cale nther
orr

. CENTRONIA.

M
.

CALYPTRELLA.
RHEXIEAE.)........ 6. MONOCHAETUM.
s with ventral lobes, spurs, or appendages,

mple.
cross-section round, I or prismatic, or obscurely
angled; flower very rarely spica
d. > nin ot c ate; fertile bis 5 or leaves strongly di-
(Tribe MICROLICIEAE

e. Fertile ragà 5, each tipped by a sterile beak; leaves essentially

isomorphic........... 7. RHYNCHANTHERA.
. Fertile a athe s 8; leaves strongly dimorphic 8. CENTRADENIA.
ic Seeds cochleste; stamens all fertile; lari. essentially isomorphic.
(Tribe TIBOUCHINEA

e. Hairs of the upper dao (if present) not adnate to the
epidermis for a portion of de length; ovary not setose at
the su mmit Mo in Schwacka
f. Connective of the outer series ge stamens bearing at its >

a skile. “ui appendage which is 2- to 3-toothed o
3-setose at its apex.

. ARTHROSTEMA.
uter series of s tamens more o less evi-

lau]

s
a5
ma
A.

B od
0
S

et
>
o

nnective of the outer
e thecae and the summit of
lament, several times as long as that of the inner
h. Cosa wingless,
hh. Hypanthium narrowly
gg. sa wi
in size.
h. mon acute or acumina

mon: prolonged. at base into two conspicuous
ending s

campanulate to subglobose......... 10. HETEROCENTRUM.
8-winged, nearly cylindric ca 11. SCHWACKA
eakly : Siad usually differing somewhat

2. NEPSERA.
ii. God s at base, or with two very short
b

leaf-surface adnate to the

Acioris.
ACISANTHERA.
e upper

rmis for a portion of their length; ovary setose at the

it.

mm
hw»

£ Hypanthium e with stout hairs from a nag —

hairs of the s ae ner geniculate at base;

ff. — ium m. pubesee with simple or sda lale m or

cales or nearly deli flowers 4-merous or (in al
ies)

—
M
.

PTEROLEPIS.

TIBOUCHINA.

—
m

ongly 3-winged, its distal margin
ry concave summit; flowers sessile
es. (Tr she ae BERTOLONIEAE
e of the anther with a single ventral spur; herbs with
alla ROERO 7. MONOLENA.
dd. Connective ka 2 or 3 basal spurs; leaves cau
e. Basal sp
M

line.
of the connective 2; leaves pei distinctly

. DIoLE
leaves me isomorphic........... 19. PRAG
wholly or partly inferio
pening by a term inal or nearly terminal pore e or pores (or
per rap, longitudinal ventral cleft i in each theca), zara or
not na all glandular; leaves in all our r species with 3 or more primary
E m er subtended by two pairs of decussate bracts
inserted at th e of the hypanthium; anthers lola i in a ring,
each mes bol yes a separate pore. (Tribe BLAKEAE.)

(204)

_
oo

m în
sal spurs of the anther 3;
aa. Fruit a z bety ovary
b. Anthers
one

1958] |
FLORA OF PANAMA (Melastomataceae) 205

d. Anthers short, oval, oblong, or elliptic, blunt or broadly rounded
at the summit 20. BLAKEA.
dd. Anthers linear to lanceolate or subulate 21. TOPOBEA.
cc. Flowers not individually subtended by decussate bracts; anthers
separate at anthesis, in most species opening by a single pore. (Tribe
MICONIEAE.
d. Flowers lateral or axillary, solitary or cymose at the nodes of
the stem.
e. Petals obtuse or rounded.

f. Anthers linear to oblong or subulate, opening by a single
pore. (But see also Clidemia pittieri, with dimorphic, sessile,
cordate-clasping leaves.)

g. Flowers or flower-clusters from the axils of existing leaves. 22. CLIDEMIA.
gg. Flowers from leafless nodes below the existing leaves......... 23. HENRIETTEA.
ff. Anthers broadly oblong or dolabriform, more or less coherent
in a rin

g-
g. Anthers opening by two minute terminal pores................... 24. BELLUCIA.
gg. Anthers opening by a single pore 25. LOREYA.
ee. Petals acute or acuminate.
f. Flowers arising from leafless nodes below the existing leaves. 26. HENRIETTELLA.
ff. Flowers or flower-clusters arising from the axils of existing

eaves 27. OSSAEA.
dd. Flowers in terminal inflorescences.
e. Petals acute or acuminate 28. LEANDRA.
ee. Petals obtuse, rounded or retuse.
f. Calyx calyptrate in bud, deciduous at the torus at anthesis. 29. CONOSTEGIA.

ff. Calyx open in the bud, its lobes persistent at anthesis, but
often minute.
g. Formicaria present at the summit of the petioles of the
arger leaves 30. Tococa.
Bg. Formicaria none.
h. Younger stems and petioles (in the Panama species) at
once stellate-tomentose, glandular-hirsute, and simply
hirsute

w
N

. HETEROTRICHUM.
. MICONIA.

vw

hh. Pubescence never of three types at once à
bb. Anthers opening by two lateral slits, one on each side; connective
bearing a large gland on the back near the middle of the anther;
leaves (in our species) 1-nerved. (Tribe MEMECYLEAE.) 33. MOURIRI.

1. ADELOBOTRYS DC.
ADELOBOTRYS DC. Prod. 3:127. 1828.

Flowers 5-merous. Hypanthium campanulate to nearly tubular, becoming
strongly ribbed at maturity. Calyx-tube spreading, its lobes short or obsolete.
Petals obovate, conspicuous. Stamens isomorphic or dimorphic; filaments flat-
tened, adherent to the base of the petals; anthers linear or subulate, deflexed, often
arcuate; connective briefly prolonged below the thecae into a short, erect, conic
or 2-toothed, basal spur, bearing a long, antrorse, dorsal appendage which is cleft
at its apex and deflexed nearliy parallel to the thecae. Ovary free, 3- to 5-celled;
style elongate; stigma capitellate. Capsule many-seeded; seeds linear-cuneate,
winged at the apex, winged or caudate at base, bearing the embryo near the center.

oody lianas, climbing by adventitious roots, with oblong to rotund leaves and
Pink, white, or yellow flowers in panicled umbels or capitate clusters.
M 20 species of tropical America, extending from Mexico and Jamaica to

Via.

(205)

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 45

Fig. 63. Adelobotrys adscendens

(206)

1958]
FLORA OF PANAMA (Melastomataceae) 207

1. ADELOBOTRYS ADSCENDENS (Św.) Triana, in Journ. Bot. 5:210. 1867.
Melastoma adscendens Sw. Fl. Ind. Occ. 2:772. 1800.

Stems climbing several meters high, the younger parts, petioles, and inflores-
cence thinly brown-strigose; petioles 1-2 cm. long; leaf-blades firm or fleshy,
subrotund, 6-10 cm. long, short-acuminate, entire or serrulate, rounded or
truncate at base, 5-nerved, glabrous above; inflorescence a panicle 2-4 dm. long
from the upper axils or terminal, each component umbel 2- to 6-flowered;
pedicels 2-5 mm. long; hypanthium 5 mm. long; petals white, 10 mm. long;
anthers 6-10 mm. long, the dorsal spur 3-4 mm. long.

Mexico to Panama; Jamaica.

OCAS DE RO: Cricamola Valley, Almirante region, Cooper 201. cHIRIQUI: Río
Dai Viejo Valley, White 79. COLÓN: summit of Cerro Santa Rita, 360-450 m.,
len 5106.

2. MERIANIA Sw.
MERIANIA Sw. Fl. Ind. Occ. 2:823. 1800.

Flowers 5-merous or rarely 6-merous. Hypanthium campanulate to hemi-
spheric, pubescent or glabrous. Calyx truncate to conspicuously lobed; exterior
teeth often present and in some species exceeding the sepals. Petals large, purple
to white. Stamens isomorphic or dimorphic; filaments elongate; anthers subulate
to ovoid-lanceolate, declined in one direction transversely across the flower by
the twisting of the filaments; connective sometimes prolonged at base and ending
in a short, retrorse or divergent, basal spur, also in many species bearing a longer
dorsal spur directed forward. vary superior, 3- to 5-celled; style elongate;
stigma punctiform. Fruit a many-seeded capsule. Trees or shrubs, glabrous or
variously pubescent, the flowers solitary in the axils or paniculate.

ut 55 species; a few in the West Indies, others in southern Brazil, and
most numerous in the mountains from Venezuela and Columbia to Peru. The
following are the only known species in Central America.

» Leaves cuneate at base, glabrous beneath 1. M. PANAMENSIS.
aa. Leaves rounded or subcordate at base, canescent beneath 2. M. MACROPHYLLA.

l. MERIANIA PANAMENSIS Gl. ex Woods. & Schery, in Ann. Mo. Bot. Gard.
28:436. 1941.

Tree up to 20 m. tall, smooth throughout; petioles 2.5-4 cm. long; leaf-blades
ovate-elliptic, 8-13 cm. long, 4-7 cm. wide, shortly and obtusely cuspidate,
entire, broadly cuneate at base, 5-nerved, the secondary veins 1-2 mm. apart;
Panicle large with widely diverging branches; hypanthium cup-shaped, 6.5 mm.
wide; calyx spreading, 3-4 mm. wide, obscurely 5-lobed. Petals purple, 3 cm.
long; stamens dimorphic; filaments 17 or 12 mm. long; anthers subulate, arcuate,
12 or 7 mm. long; connectives in one series merely dorsally gibbous at base, in the
other series prolonged into a single spur 4 mm. long.

Endemic to Panama, so far as known.

(207)

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

208

Fig. 64. Meriania panamensis

(208)

1958]
FLORA OF PANAMA (Melastomataceae) 209

CHIRIQUÍ: Volcán de Chiriquí, 2300-3300 m., Woodson t$ Schery 361; vicinity of
Cerro Punta, 2000 m., Allen 3497

2. MERIANIA MACROPHYLLA (Benth.) Triana, in Trans. Linn. Soc. 28:66. 1871.
Davya macrophylla Benth. Pl. Hartweg. 75.
Conostegia excelsa Pittier, in Journ. Wash. Kad Sci. 14:450. 1924.

Tree to 25 m. tall, the younger parts and petioles densely furfuraceous,
glabrescent in age; leaves coriaceous, elliptic, up to 17 cm. long and 9 cm. wide,
rounded or subcordate at base, where a transverse ridge crosses the petiole, glabrous
above, canescent beneath; panicle widely branched; hypanthium and narrow
truncate calyx glabrous; petals magenta, 12-15 mm. long; stamens dimorphic,
the connective in the larger series prolonged into an erect bifurcate spur 7 mm.

Guatemala to Venezuela.

T cor Chiquero, Boquete district, 1650 m., Davidson 506; Las Siguas, Pittier
100.

3. AXINAFA Ruiz & Pav.

AXINAEA Ruiz & Pav. Fl. Peruv. & Chil. Prodr. 68. 1794.

Flowers 4- to 5-merous. Hypanthium campanulate to saucer-shaped, glabrous
or thinly pubescent. Calyx spreading, truncate or with short or depressed lobes;
exterior teeth none. Stamens 8 or 10, nearly or quite isomorphic; filaments flat-
tened, glabrous; anthers subulate, straight or somewhat arcuate; connective bearing
at base a large, inflated, usually rounded, dorsal appendage. Ovary free, 5-celled;
style usually slender and elongate; stigma truncate or punctiform. Fruit a many
seeded capsule. Trees or shrubs with broad, usually petiolate leaves and large,
white to purple flowers in terminal or axillary panicles; leaf-margins often some-
What dilated at base and more or less confluent over the petiole.

About 20 species of mountainous regions from Costa Rica to Venezuela and
Bolivia; the following is the only known species in Central America.

l. AXINAEA COSTARICENSIS Cogn. in DC. Monogr. Phan. 7:1182. 1891.

Good-sized tree, the younger parts thinly furfuraceous but soon glabrescent.
Petioles 1.5-3 cm. long. Leaf-blades oblong-elliptic, 10-15 cm. long, about

f as wide, very abruptly acuminate, entire, minutely expanded at the obtuse
base and bearing a median dorsal tubercle on the summit of the petiole, 5- nerved;
Panicle many-flowered, divergently branched, 1-3 dm. long, the flowers in ter-
minal umbels on pedicels 5-10 mm. long; hypanthium and calyx together about
8 mm. wide; petals blood-red, 10-15 mm. long; anthers about 6 mm. long, the
dorsal appendage white, rounded, nearly 4 mm. high.

ta Rica and Panama.

erą DEL TORO: Robalo Trail, northern slopes of Cerro Horqueta, 1800-2100 m.,

(209)

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

210

Axinaea costaricensis

Fig. 65.

(210)

1958]
FLORA OF PANAMA (Melastomataceae) 211

A —
rete ES

L

Fig. 66. Centronia tomentosa

(211)

[Vol. 45
212 ANNALS OF THE MISSOURI BOTANICAL GARDEN

4. CENTRONIA D. Don
CENTRONIA D. Don, in Mem. Wern. Soc. 4:314. 1823.

Flowers 5- to 7-merous. Hypanthium conic to hemispheric, pubescent or
hirsute. Calyx calyptrate in bud, at anthesis breaking irregularly and soon decid-
uous. Petals large, red or purple. Stamens isomorphic or weakly dimorphic;
filaments flat, more or less twisted at base; anthers subulate to lanceolate; con-
nective often elevated along the thecae, at base bearing a conspicuous dorsal spur.
Ovary free, 3- to 7-celled, often crowned with gibbosities or projecting append-
ages; style elongate; stigma truncate. Fruit a many-seeded capsule. Trees or
shrubs with large leaves and large red or purple flowers usually disposed in terminal
3-flowered cymules in a large terminal panicle.

About 15 species in the mountains from Central America to Peru and eastward
into Venezuela and British Guiana. C. phlomoides Triana, known from Costa
Rica and Colombia, may be expected in Panama; it is at once distinguished from
the following species by its strongly 5-pli-nerved leaves.

l. CENTRONIA TOMENTOSA Cogn. in Bull. Acad. Belg. III. 14:943. 1887.
Centronia grandiflora Standley, in Field Mus. Publ. Bot. 22:95. 1940.

Tree to 15 m. tall, the younger stems, petioles, and inflorescence rufous-
tomentose; petioles 2—4 cm. long; leaf-blades ovate-lanceolate to elliptic, up to
15 cm. long and about half as wide, acuminate, denticulate, rounded or obtuse at
base, 5-nerved or barely 5-pli-nerved, glabrous above, canescent on the surface
beneath with minute stellate hairs ending in a long bristle; panicle long-peduncled,
few-flowered, buds about 18 mm. long, short-rostrate; hypanthium about 1 cm.
long, densely pubescent; petals white or greenish white, about 18 mm. long.

Panama to Ecuador.

BOCAS DEL TORO: Robalo Trail, northern slopes of Cerro Horqueta, 1800-2100 m.,
Allen 4992. cHIRIQUI: vicinity of Casita Alta, Volcin de Chiriqui, 1500-2000 m.
Woodson, Allen & Seibert 842; vicinity of Bajo Chorro, 1900-2100 m., Davidson sa
360, Woodson t$ Schery 657; Boquete region, von Hagen & von Hagen 2102; vicinity 0
Cerro Punta, 2000 m., Allen 3492.

5. CALYPTRELLA Naud.
CALYPTRELLA Naud. in Ann. Sci. Nat. Bot. III. 18:115. 1852.

Flowers 4- to 9-merous. Hypanthium cup-shaped to campanulate. Calyx
calyptrate in the bud, at anthesis irregularly or regularly ruptured and soon
deciduous at the torus. Petals small or medium-sized, lanceolate to obovate,
commonly acute or acuminate. Stamens isomorphic or somewhat dimorphic;
anthers linear; connective more or less prolonged below the thecae and extended
beyond the apex of the filament into a single short dorsal spur. Ovary superior,
4- to 6-celled; stigma punctiform. Fruit a capsule; seeds linear with a central
embryo. Trees or shrubs, with petiolate leaves and terminal paniculiform cymes
the white, yellow, or purple flowers often in small glomerules at the ends of the
cyme-branches.

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1958]

213

FLORA OF PANAMA (Melastomataceae)

Fig. 67. Calyptrella micrantba

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[Vol. 45
214 ANNALS OF THE MISSOURI BOTANICAL GARDEN

About 10 species of mountainous regions or adjacent lowlands from southern
Mexico to Peru.

l. CALYPTRELLA MICRANTHA Gl. in Phytologia 3:346. 1950.

Shrub or tree 5-22 m. tall, essentially glabrous, but the hypanthium and
lower leaf-surface thickly punctate with hyaline atoms; petioles 3-5 cm. long;
leaf-blades thin, elliptic, up to 20 cm. long, two-fifths to half as wide, abruptly
caudate-acuminate, entire, bearing a small retrorse appendage at each side of the
obtuse base, 3-nerved; panicle divergently branched, 3-9 cm. long; flowers 5-
merous, on pedicels 3-5 mm. long; hypanthium cup-shaped, 2 mm. long; calyx
in bud slenderly conic, 3 mm. long, often falcate; petals white, narrowly lance-
olate, long-acuminate, 3 mm. long.

Endemic to Panama.

BOCAS DEL TORO: Fish Creek Mountains, von Wedel 2249. cocLé: Cerro Pajita,
north of El Valle de Antón, 1000-1200 m., Allen 4175. VERAGUAS: forested slopes o
Cerro Tuté, vicinity of Santa Fé, 900 m., Allen 4355.

6. MONOCHAETUM Naud.

MONOCHAETUM Naud. in Ann. Sci. Nat. Bot. III. 4:48. 1. 2. 1945.

Flowers 4-merous. Hypanthium tubular to narrowly campanulate. Calyx
divided nearly or quite to the torus; sepals lanceolate to ovate, commonly x
ate, persistent (in our species) or deciduous after anthesis. Petals medium-sized,
usually obovate. Stamens 8 and dimorphic, or rarely only 4 by abortion; filaments
slender, smooth; anthers slender, linear or subulate, those of the larger series
arcuate, bearing at base a single elongate, erect or ascending, dorsal appendage
those of the smaller series straight, the appendage commonly proportionately
longer. Ovary superior, 4-celled, pubescent to setose at the summit; stigma
punctiform. Fruit a 4-valved capsule; seeds cochleate. Small shrubs with com-
paratively small, linear to ovate, pli-nerved leaves and terminal cymes or cymules
of pink, purple, or rarely white flowers.

About 35 species of the mountains from southern and western Mexico to Peru.
a. Pubescence of the upper part of the stem distinctly spreading; leaves

ovate or ovate-lanceolate, commonly two-fifths to two-thirds as wide as

long; bracteal leaves of the inflorescence scarcely smaller than the

liage leaves Ld o oa

aa. Pubescence of the upper part of the stem distinctly strigose, the hairs
appr g at a narrow angle; leaves commonly a fourth to

1. M. RIVULARE.

b. Flowers in a freely branched inflorescence; bracts ovate or oblong,
much smaller than the leaves; large anthers 5-6 mm. long (excluding
the spur).
c. Leaves lanceolate, broadest well below the middle, firm, softly
villous bebeath ai. 2. M. CYMOSUM

rai 3. M. BRACTEOLATUM

i 1 uppermost leaves scarcely
reduced in size; large anthers 9-12 mm. ong

4. M. MACRANTHERUM

A

1958]
FLORA OF PANAMA (Melastomataceae) 215

Fig. 68. Monochaetum cymosum

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[Vol. 45
216 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. MONOCHAETUM RIVULARE Naud. in Ann. Sci. Nat. Bot. III. 4:50. 1845.

Shrub to 2 m. tall, laxly branched, the younger stems hirsute or hirsutulous
with spreading, usually smooth hairs; leaves ovate, two-fifths to two-thirds as
wide as long, hirsute above in poorly differentiated longitudinal strips, hirsute to
villous beneath; hypanthium strigose.

Forma RIVULARE. Pubescence of simple hairs.

Guatemala to Panama.

CHIRIQUf: Bajo Chorro, Boquete district, 1800 m., Davidson 88.

Forma GLANDULOSUM Gl. in Am. Journ. Bot. 16:590. 1929.

Pubescence of the inflorescence and hypanthium very dense and partly
glandular.

Costa Rica and Panama.

CHIRIQUf: Volcán de Chiriqui, 1500-2100 m., Davidson 884, Woodson, Allen 8
Seibert 906 (in bart).

2. MONOCHAETUM CYMOSUM Gl. in Am. Journ. Bot. 16:588. 1929.

Shrub to 3 m. tall; younger stems strigose with barbellate, appressed or some-
what ascending, eglandular hairs; leaves firm, lanceolate, 2-3 cm. long, a fourth
to a third as wide, often acuminate, strigose above in four well-marked longi-
tudinal strips each about as wide as the glabrous strip which parallels the con-
spicuously depressed primary nerves, hirsute or villous beneath, especially on
primary nerves; hypanthium strigose, eglandular; bracts ovate, 2-5 mm. long.

Endemic to Panama, so far as known.

_ CHIRIQUÍ: Volcán de Chiriquí, 1500-2000 m., Woodson, Allen & Seibert 906 (part);
Sierra del Boquete, Maurice 746, Pittier 2861; vicinity of Cerro Punta, 2000 m., Allen
1513.

3. MONOCHAETUM BRACTEOLATUM Triana, in Trans. Linn. Soc. 28:64. 1871.

Stem strigose with minutely barbellate eglandular hairs; leaves thin, narrowly
elliptic or oblong, 3-5 cm. long, merely acute, sparsely strigose above in four
strips between the nearly plane primary nerves, very sparsely strigose beneath;
inflorescence and hypanthium strigose, eglandular; bracts ovate, 2-4 mm. long.

Endemic to Panama.

CHIRIQUI: vicinity of El Boquete, Seemann 1668.

4. MONOCHAETUM MACRANTHERUM Gl. in Am. Journ. Bot. 16:591. 1929.

Stem closely strigose with barbellate eglandular hairs, densely pilose at the
nodes; leaves oblong-lanceolate, 2-3 cm. long, up to 1 cm. wide, subacute, Very
sparsely strigose on the upper side, often only toward the tip, sparsely strigose 0%
the primary nerves beneath; inflorescence of mostly 3-flowered terminal cymules;
hypanthium thinly strigose; large anthers 9-12 mm. long.

Panama and Costa Rica.

CHIRIQUi: Volcin de Chiriqui, Maxon 5305.

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1958]
FLORA OF PANAMA (Melastomataceae) 217

7. RHYNCHANTHERA DC.

RHYNCHANTHERA DC. Prod. 3:106. 1828.

Flowers 5-merous. Hypanthium ovoid to subglobose or campanulate, pubes-
cent. Sepals erect or spreading, lanceolate to linear, often exceeding the hypan-
ium. Petals conspicuous, obovate. Fertile stamens 5, episepalous; filaments
glabrous, usually stout, somewhat flattened; anthers subulate, rounded at base,
prolonged distally into a slender one-pored beak; connective slender, greatly
prolonged below the thecae, straight or arcuate, bituberculate, bicalcarate, or rarely
unappendaged at base. Sterile stamens 5, opposite the petals, reduced to slender
filaments, or lacking. Ovary free, 3- to 5-celled, glabrous, or distally setose or
glandular; style elongate, filiform, declined or somewhat sigmoid; stigma puncti-
form. Capsule thickly ovoid or subglobose; seeds ovoid to oblong, straight or nearly
so, minutely foveolate. Herbaceous or partly woody plants, usually densely
pubescent or pilose and often glandular; flowers large, in numerous few-flowered
cymes forming a terminal leafy panicle.
About 50 species, the others all in tropical South America, extending as far
as Paraguay.

Fig. 69. Rbynchanthera mexicana

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[Vol. 45

218 ANNALS OF THE MISSOURI BOTANICAL GARDEN

a: Perea > ae or CA equal in length; hairs of the hypan-
thiu ee 1. R. PALUDICOLA.
eros ns Padi in length, the connective of one greatly prolonged;
hairs of de ‘rale ben 0.5 mm. long or shorter 2. R. MEXICANA.

1. RHYNCHANTHERA PALUDICOLA (Donn. Sm.) Gl. in Phytologia. 1:136. 1935.
Tibouchina paludicola Donn. Sm. in Bot. Gaz. 42:293. 1906.

Stems, petioles, pedicels, and hypanthia sparsely glandular-pilose with hairs 1.5-
2 mm. long; stem erect, angular, freely branched; petioles slender, 10-15 mm.
long; leaf-blades ovate or ovate-oblong, 4-8 cm. long, 2-4 cm. wide, acuminate,
minutely denticulate, rounded to subcordate at base, 7-nerved, pilose on both sides;
inflorescence large, freely branched, the pedicels 2 mm. long; hypanthium cam-
panulate, 6 mm. long, 10-ribbed; sepals linear-subulate, 6 mm. long; petals purple,
broadly obovate, setose at the tip, 14 mm. long; sterile stamens 4.5 mm. long;
filaments 5.5 mm. long; anthers 6.5 mm. long, tipped with a bent beak 2 mm.
long, the connective prolonged 3 mm. at base; ovary free, 3-celled, minutely
glandular-pubescent at the summit; style 19 mm. long.

Costa Rica and Panama.

HIRIQUÍ: llanos del Volcán, Allen 1549; vicinity of ora ei 1200-1500 m., Woodson
8 debora 739b, Davidson 588. PROVINCE UNKNOWN: Pił 42

2. RHYNCHANTHERA MEXICANA DC. Prod. 3:108. 1828.

Semi-frutescent, freely branched above, the stem, petioles, inflorescence, and
hypanthium densely glandular-pubescent; petioles 8—12 mm. long; leaf-blades
lanceolate to ovate-lanceolate, 5-8 cm. long, 2-3 cm. wide, slenderly acuminate,
entire, rounded or broadly obtuse at base, 5- to 9- nerved, freely pubescent on both
surfaces; bracteal leaves gradually reduced in size and proportionately narrower;
o ee ample, many-flowered; pedicels mostly 2-5 mm. long; hypanthium
narrowly campanulate, 4-5 mm. long; sepals linear-subulate, exceeding the hypan-
thium, pubescent; petals 25-30 mm. long, lavender or white to deep rose-purple;
one connective much elongate; anthers, including the beak, 8-10 mm. long; capsule
subglobose, glabrous, 5 mm. in diameter, 3-valved.

Southern Mexico and Panama.

COCLE: El Valle de Antón, 600 m., Allen 28 Tarn of El Valle, VER & White
65, 66; Penonomé and vicinity, Williams 216. ERA: vicinity of "i ., Allen 4044
VERAGUAS: hills west of Soná, 020; vicin Ys ‘of Santiago, abiat 0 m., Allen Tones
between Cafiazas and the Cordillera Coi 300-600 m., Allen 73.

8. CENTRADENIA G. Don
CENTRADENIA G. Don, Gen. Syst. 2:755. 1832.

Flowers 4-merous. Hypanthium broadly campanulate, 8-ribbed, thin-walled,
somewhat 4-angled. Calyx-tube briefly or not at all prolonged; sepals erect, 0 ovate-
triangular, acute or obtuse; exterior teeth none. Petals obovate or broadly elliptic:
obtuse, entire or ciliate. Stamens dimorphic or isomorphic; filaments delicate, flat;
glabrous; anthers ovoid to oblong, linear-oblong, or subulate, obtuse or ac cuminate,

(218)

1958]
FLORA OF PANAMA (Melastomataceae) 219

Fig. 70. Centradenia maxoniana

opening by a minute terminal pore; connective prolonged below the thecae, bearing
at its base an anterior tubercle or a 2- to 3-lobed, truncate or rounded appendage.
Ovary free, 4-celled, often glandular or setose; style slender, glabrous; stigma
truncate. Capsule 4-valved; seeds clavate-ellipsoid. Herbs with terete, angled, or
winged stems and lanceolate to narrowly oblong dimorphic leaves, the smaller
member of each pair early deciduous; flowers few to many, in small cymes termi-
nating the stem and branches; petals white to pink-purple, 5-10 mm. long.
ven species in Mexico and Central America, one of them extending into

Colombia.

a. Stamens strongly dimorphic; principal leaves 3-8 mm. wide........-.----------- 1. C. INAEQUILATERALIS.
aa. Stamens weakly dimorphic, differing chiefly in size; leaves 1.5-3.5 cm.
Widk ooo 2392. 0000 A ADAK ERE e Aa la minele ac ac RD 2. C. MAXONIANA.

l. CENTRADENIA INAEQUILATERALIS (Cham. & Schlecht.) G. Don. Gen. Hist.

2:765. 1832.

Rbexia inaequilateralis Cham. & Schlecht. in Linnaea 5:567. 1830.

Stems slender, subterete or obscurely 4-angled, pilosulous with spreading hairs;
Principal leaves straight or slightly falcate, thin, lanceolate, 25—50 mm. long, 3-8
mm. wide, acuminate, cuneate at base into a pilose petiole 1-3 mm. long, 3-nerved,
minutely puberulous to almost completely glabrous; cymes laxly branched, often
shorter than the upper leaves, thinly pilosulous; hypanthium broadly campanulate,
2.5 mm. long, sparsely pilose; sepals broadly ovate-triangular, obtuse, 0.8—1 mm.
long; petals 6 mm. long, conspicuously ciliate; episepalous anthers 1.6 mm. long,

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[Vol. 45
220 ANNALS OF THE MISSOURI BOTANICAL GARDEN

the connective prolonged 1.6 mm. to the filament and 1.6 mm. below the filament
into a narrowly obtriangular, flat, truncate appendage; epipetalous anthers shorter,
the connective prolonged 1 mm. to the filament, bearing on its base above the
filament a flattened, ellipsoid, semi-adnate appendage.

Oaxaca and Veracruz to Panama.

CHIRIQUI: Mawi of El Boquete, 1000-1300 m., Pittier 3040. cocLE: vicinity of
El Valle, 500-700 m., Allen 766, Seibert 468; hills north of FI Valle, Allen 2907; El
Valle de Antón, Mieć 774.

2. CENTRADENIA MAXONIANA Gl. in Bull. Torr. Bot. Club 65:571. 1941.

Stems slender, terete, sparsely branched, densely strigose; petioles 5 mm. long,
strigose; leaf-blades lanceolate, straight or falcate, 8—15 cm. long, 1.5—3.5 cm.
wide, inequilateral, acuminate, entire, at base cuneate on one side, acute to rounded
on the other, 3-nerved, pubescent above, beneath strigose on the veins, thinly
pilose to glabrous on the surface; smaller leaves linear-lanceolate, soon deciduous;
panicle 2-3 cm. long, densely strigose; hypanthium 3.5 mm. long, glandular-
pubescent; sepals triangular, 1 mm. long; petals oblong, 5 mm. long; anthers nearly
isomorphic, 2.8 or 4 mm. long, linear-subulate; connective short, obliquely dilated
at base, the anterior lobe ascending, truncate, the posterior descending, ovate.

POS Costa Rica and western Colombi

RIQUÍ: between Hato del Jobo and Cerro Vaca, 700-1000 m., Pittier 5414. COCLÉ:
vicinity of El Valle, 600-1000 m., Allen 112, 1175. Bismark, above Penonomé, Williams
321

9. ARTHROSTEMA Ruiz 8 Pav.

ARTHROSTEMA Ruiz & Pav. Fl. Peruv. & Chil. 4: ź. 326. 1802.

Flowers 4-merous. Hypanthium obconic to narrowly campanulate, elongate,
ribless or obscurely ribbed, glabrous or sparsely glandular-setose. Calyx-tube erect
or spreading, persistent; sepals broader than long, rounded, obtuse, or apiculate.
Petals obovate, apiculate, caducous. Stamens more or less dimorphic; filaments
flat; episepalous anthers linear or oblong, straight or curved, the connective pro-
longed below the thecae and bearing a short or elongate, truncate, 2- to 3-t00
or 2-setose, anterior appendage; epipetalous anthers shorter, the connective shorter
or not prolonged, bearing a relatively short, 2-toothed or 2-setose, anterior append-
age. Ovary 4-celled, usually glabrous at the summit; style elongate, somewhat
sigmoid; stigma punctiform. Capsule 4-valved. Seeds flattened, semi-ovoid,
cochleate, marked with about 8 longitudinal, sometimes tuberculate ridges, the flat
sides smooth above the hilum. Erect or scandent, slender, sparsely branched herbs
or shrubs, with 4-angled to 4-winged stems, small, petiolate, lanceolate to ovate,
3- to 5-nerved leaves, and sessile or pediceled, pink to purple flowers in loosely
branched cymes.

About 8 species of tropical America.

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1958]

FLORA OF PANAMA (Melastomataceae)

Fig. 71. Artbrostema macrodesmum

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[Vol. 45

222 ANNALS OF THE MISSOURI BOTANICAL GARDEN
a. Flowers and ipee ce or nearly so; anthers nearly isomorphic, the
connective very briefly prolonged 1. A. ALATUM.
aa. rann bet Bis connective of the outer series of stamens prolonged
mm. below the anther . A. MACRODESMUM.

—

. ARTHROSTEMA ALATUM Triana, in Trans. Linn. Soc. 28:35. 1871.

Stems erect, freely branched, narrowly 4-winged, 4-6 dm. tall, very sparsely
glandular-setose; petioles 1-2 cm. long, winged distally; leaf-blades thin, bright
green, ovate-oblong to broadly ovate, 3-7 cm. long, 2—4 cm. wide, acute, minutely
ciliate-serrulate, rounded at base, 5-nerved, sparsely setose above, glabrous beneath;
inflorescence many-flowered, divided into elongate spiciform branches; flowers
sessile or nearly so; hypanthium narrowly campanulate, 4 mm. long, the erect
calyx-tube obscurely lobed into depressed-triangular, minutely apiculate sepals 1
mm. long or shorter; petals pink, about 3 mm. long; stamens almost isomorphic;
filaments 2 mm. long; anthers ovate, 0.8 mm. long; connective very briefly
prolonged, bearing a single short, subulate, anterior spur; fruiting hypanthium
long persistent, 10 mm. long, the flat sides of the seeds confluent over the end.

Southern Mexico, Salvador, Costa Rica, Panama, and Venezuela.

PANAMA: Punta Paitilla, Heriberto 234.

N

ARTHROSTEMMA MACRODESMUM Gl. in Williams, Fl. Trin. & Tob. 356, 357.
Artbrostema campanulare Triana, in Trans. Linn. Soc. Bot. 28:35. 1871. Not A. cam-
panulare DC. 1828.

Stems climbing, 1-3 m. tall, sharply 4-angled or narrowly 4-winged, glabrous
below, very sparsely glandular-setose above and in the inflorescence; leaves thin,
bright green, ovate or ovate-lanceolate, 3-7 cm. long, 1.5-3 cm. wide, acuminate,
truncate or subcordate at base, finely ciliate-serrulate, 5-nerved, glabrous or very
sparsely setose; inflorescence loosely branched, few-flowered; pedicels about 5 mm.
long, elongating in fruit; hypanthium glabrous, narrowly obconic, 8 mm. long;
sepals triangular-subulate, about 1 mm. long, sometimes glandular-setose at the
apex; petals pink to red or purple, 15-20 mm. long; episepalous anthers px
nearly straight, 4—4.5 mm. long, the connective prolonged 5—7 mm. to the
ment, its appendage 2.5—3 mm. long, 2- to 3-toothed at the apex; cipue
anthers curved, 2.5—3 mm. long; connective not prolonged, bearing at its base two
erect subulate appendages about 1 mm. long; fruiting hypanthium 15 mm. long.

From Chiapas southward through Central America and across northern South
America to Peru and Trinidad.

BOCAS DEL TORO: vicinity of Chiriqui Lagoon, von Wedel 1220, 1248, 1368, 1500,
2614; Teu Valley, von Wedel 831, 928, 2746; Shepherd vq von Wede | 2716; between
Finca Louis and Konkintóe, Woodson, Allen & Seibert 1907; Potrero, f: 8
Valley, Dunlop 72. CANAL ZONE: Ciara. poderi 112; Cola: Lione Crawford 51 :
CHIRIQUI: vicinity of gc — 699, W Woodson 8 Sober 7394, Davidson ei
Finca Lérida to Boquete, 700 m., Woodson, Allen & Seibert 1142. COCLÉ:
Valle de Antón, 600—1000 m., pre 1638.

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1958]
FLORA OF PANAMA (Melastomataceae) 223

10. HETEROCENTRON Hook. & Arn.
HETEROCENTRON Hook. & Arn. in Bot. Beech. Voy. 290. 1840.
Heeria Schlecht. in Linnaea 13:432. 1839.
Schizocentron Meissn. Gen. Comm. 355. 1843.

Flowers 4-merous. Hypanthium broadly campanulate, rarely subglobose,
setose at least at the summit. Sepals erect or spreading, separate to the base, ovate
to linear-lanceolate, acute or acuminate, often involute distally, few-nerved. Petals
rotund to obovate or broadly cuneate, commonly glandular-ciliate, somewhat in-
equilateral, flabellately veined. Stamens 8, dimorphic. Filaments slender, flat,
glabrous, longer in the outer series. Anthers linear or linear-subulate, straight or
nearly so, 2-celled, opening by a minute terminal pore; connective in the epi-
sepalous stamens prolonged conspicuously below the thecae to the insertion of the
filament, there continued in the same direction into two slender, parallel, anterior
spurs; connective of the epipetalous stamens very briefly prolonged below the
thecae, slightly swollen and sometimes minutely tuberculate on the anterior side.

Fig. 72. Heterocentron glandulosum

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[Vol. 45
224 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ovary free or adnate to the middle, 4-celled, its summit rounded or usually bearing
4 fleshy scales. Style slightly sigmoid, elongate, slender; stigma punctiform.
Capsule about equaling the hypanthium, 4-valved. Seeds minute, obovoid, cochle-
ate. Shrubs or half-shrubs, erect or rarely trailing, with 4-angled stems; leaves
(in our species) multipli-nerved or apparently pinnately veined, the usual secondary
veins lacking; inflorescence a terminal panicle or flowers rarely solitary; petals
purple, rose-purple, or pink, 5-12 mm. long.
About 15 species of Mexico and Central America.

1. HETEROCENTRON GLANDULOSUM Schenck. in Regel, Gartenfl. 227. £. 169.
1856.

Stem erect, rather stout, obtusely 4-angled, densely strigose; leaf-blades oblong
to obovate, 3-7 cm. long, 1.5-4 cm. wide, obtuse or acute, cuneate at base into a
slender petiole, multipli-nerved, densely strigose above, subsericeous beneath;
inflorescence ample, freely branched, rather crowded, strigose, becoming glandular
above, the short pedicels and hypanthium densely glandular with spreading hairs;
hypanthium broadly campanulate, 4 mm. long; sepals triangular-lanceolate, 4.5
mm. long, glabrous and entire; petals rotund, 7-8 mm. long; connective of the
larger stamens 2.4 mm. long, tangentially flattened; ovary crowned with four
quadrate scales nearly 1 mm. high and densely ciliate above.

Southern Mexico, Guatemala, Costa Rica, and Panama.

; PUE near E] Volcán, Wbite 233, 2334; Boquete district, 1140 m., Terry 6 Terry
1638.

11. SCHWACKAEA Cogn.

SCHWACKAEA Cogn. ex Durand Index Gen. Phan. 132. 1888.-Nomen; DC. Mon.
Phan. 7:182. 1891.

Flowers 4-merous. Hypanthium cylindric, prominently 8-ribbed, the ribs
somewhat tuberculate or papillose-strigose. Sepals linear-lanceolate, erect, pectinate-
ciliate. Petals small, obovate, setose at the apex. Stamens 8, dimorphic; filaments
glabrous, very slender; anthers of the outer series oblong-ovoid, somewhat curved,
the connective horizontal or declined, arcuate, as long as or longer than the anther,
bearing 2 short, blunt, basal, anterior spurs; anthers of the inner series ovoid,
straight, truncate, the connective very short, bearing 2 short, blunt, anterior spurs.
Ovary free, 4-celled, oblong, glabrous below, tipped with about 8 short ...
Style short, clavate, strongly incurved above to the broad stigma. Hypanthium
enlarged and thickened in fruit, with 8 rounded, corky, tuberculate ribs. Capsule
oblong, free. Seeds minute, cochleate. Annual, freely branched herb, the leaves
soon deciduous, with numerous small pink flowers in leafy cymes.

Only the following species.

l. SCHWACKAEA CUPHEOIDES (Benth.) Cogn. ex Durand, Index Gen. Phan. 132.
1888.

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1958]
FLORA OF PANAMA (Melastomataceae) 225

Fig. 73. Schwackaea cupbeoides

Heeria cupbeoides Benth. Bot. T salgo 93. 1.33.
Pterogastra cupheoides (Benth.) Seem. Bot. Voy. Heald i 1853.

Stem repeatedly branched, 2-10 dm. tall, reddish, 4- angled, setose on the nodes
and angles to nearly glabrous; petioles slender, 2-8 mm. long, leaf-blades oval or
oblong, 1-6 cm. long, 0.5—3 cm. wide, acute, entire, acute or obtuse at the base,
3-nerved, minutely pubescent, especially on the veins, to nearly glabrous; flowers
sessile; hypanthium 4 mm. long; sepals 2.5-3 mm. long; petals 5 mm. long; anthers
0.7 or 1.2 mm. long, the basal spurs of the connective 0.5 mm. long; fruiting
hypanthium 1 cm. long; seeds 0.4-0.5 mm. long.

Southern Mexico to Panama and Colombia.

ONE: Barro Colorado Island, Killip 40033, sati 299, 485;
Macbride 2 3702, polpa Hayes 151; Paraíso, Hayes 25; Chagres, Fendler 113; Río rir
near Culebra, Pittier 2108. PANAMA: near Alhajuela, Ca Valley, Pittier 2395; San
José Island, Jobnston 176, 219, 936A.

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226 ANNALS OF THE MISSOURI BOTANICAL GARDEN

12. NEPSERA Naud.
NEPSERA Naud. in Ann. Sci. Nat. Bot. III. 13:28. 1850.

Flowers 4-merous. Hypanthium campanulate to subglobose. Sepals triangular-
ovate or triangular-lanceolate, about equaling the hypanthium, acuminate. Petals
ovate to ovate-lanceolate, acute and apiculate. Stamens isomorphic, differing
slightly in size; anthers erect, linear or linear-subulate; connective prolonged briefly
below the thecae, somewhat expanded at the insertion of the lament, and bearing
2 erect slender anterior appendages. Ovary free, sub-globose, glabrous, 3-celled;
style straight, slender; stigma punctiform. Capsule 3-valved, closely invested by
the hypanthium; seeds stoutly semiovoid, cochleate, finely tuberculate. Slender,
freely branched, erect herbs, with very numerous, small, white to pink flowers in
a large, diffusely branched, terminal panicle.

Only the following species.

Fig. 74. Nepsera aquatica

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1958]
FLORA OF PANAMA (Melastomataceae) 227

1. NEPSERA AQUATICA (Aubl.) Naud. in Ann. Sci. Nat. Bot. III. 13:28. 1850.
Melastoma aquaticum Aubl. Pl. Guian. 1:430. £. 169. 1775.

Stems 3-10 dm. tall, 4-angled, glabrous to minutely pubescent; petioles slender,
5-10 mm. long; leaf-blades ovate or ovate-lanceolate, 2—6 cm. long, acute or
acuminate, finely serrulate, rounded or subcordate at base, 5- to 7-nerved, glabrous
or sparsely pubescent; panicle 1—4 dm. long, the pedicels slender and elongate;
hypanthium about 3 mm. long; petals 5-7 mm. long.

British Honduras to Panama; Jamaica, Hispaniola, Puerto Rico, and the Lesser
Antilles; widely distributed in tropical South America. e

S DEL TORO: Water Valley, von Wedel 825; vicinity of Chiriquí Lagoon, vom
Wedel 1133, 2485, 2551; Columbus Island, von Wedel 33. PROVINCE UNKNOWN: Moore
er.

13. AGIOTIS.D. Don

Acioris D. Don, in Mem. Wern. Soc. 4:300. 1823.

Flowers 4-merous. Hypanthium thin-walled, narrowly campanulate at anthesis.
Calyx-lobes minute, triangular, often glandular-setose at the apex. Petals lance-
olate to obovate, acute or acuminate, commonly glandular-setose at the apex.
Stamens 8, isomorphic or weakly dimorphic; anthers narrowly oblong to elliptic
or nearly globose, blunt; connective somewhat prolonged below the thecae, articu-
late to the apex of the filament, either simple or with two minute ventral lobes.
Ovary superior, 2-celled; stigma truncate or capitellate. Fruit a globose thin-
walled capsule, rupturing irregularly at maturity; seeds rugose, cochleate. Erect
or spreading herbs with lanceolate to ovate leaves and terminal, divaricately
branched, paniculiform cymes of small, ephemeral, white or pink flowers.

About 30 species, widely distributed in tropical America but most numerous in
lowland forests of high rainfall.
a. Leaves pubescent beneath.

b. Stems winged, pubescent with mostly simple hairs 1-1.5 mm. long..... 1. A. LEVYANA.

bb. Stems merely angled, pubescent with mostly glandular hairs 0.2-0.8
monelli er 2. A. ROSTELLATA.
aa. Leaves glabrous beneath...................22 ie 3. A. PURPURASCENS.

l. ACIOTIS LEVYANA Cogn. in Mart. Fl. Bras. 14?:460. 1885.

Stems herbaceous, branched, up to 1 m. tall, prominently and often broadly
winged, each wing ciliate and more or less hirsutulous on the surface with widely
spreading hairs 1-1.5 mm. long, commonly simple but occasionally glandular;
Petioles winged, pubescent like the stem; leaf-blades oblong to oblong-lanceolate,
to 1 dm. long, a third to half as wide, pubescent on both sides; fourth and fifth
Yeins arising from the second and third just above the base; panicle-branches
winged, hirsutulous, especially on the wings, with simple hairs commonly 1-1.5
mm. long.

British Honduras to Panama, chiefly at low elevations.

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[Vol. 45
228 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 75. Aciotis levyana

BOCAS DEL TORO: Water Valley, von Wedel 782, 1450. CANAL ZONE: Barro Colorado
Island, Bailey & Bailey 581; Gatun, Hayes 165; vicinity of Porto Bello, Pittier 2440; Santa
Rita trail, Cowell 139. cocrt: vicinity of El Valle, Allen 218.

2. ACIOTIS ROSTELLATA (Naud.) Triana, in Trans. Linn. Soc. 28:51. 1871.
Spennera rostellata Naud. in Ann. Sci. Nat. Bot. III. 14:143. 1850.

Similar to the preceding species in general habit and size; stem sharply 4-angled
but not winged, densely and softly pubescent with spreading, mostly gland
hairs commonly about 0.5 mm., rarely as much as 1 mm. long; petioles wingless,
pubescent like the stem; leaf-blades ovate or oblong, to 1 dm. long, about half as
wide, pubescent on both sides, the five nerves all arising together at the base;
panicle-branches merely angled, glandular-pubescent with hairs 0.2—0.5 mm. long.

Southern Mexico to Panama.

BOCAS DEL TORO: Nances Cay, von Wedel 582, 2863. cocLt: Bismarck, above
Penonomć, Williams 289. DARIEN; Cana and vicinity, Williams 735.

3. ACIOTIS PURPURASCENS (Aubl.) Triana, in Trans. Linn. Soc. 28:52. £. 3. 1871.

Melastoma purpurascens Aubl. PI. Guian. 1:402. 1775.
Glandular pubescence lacking. Stems herbaceous, to 1 m. tall, simple w
branched above, 4-winged, glabrous or ciliate on the wings with minute incurv

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19581
FLORA OF PANAMA (Melastomataceae) 229

hairs; petioles winged; leaf-blades lanceolate to oblong, to 15 cm. long, a fourth
to a third or even half as wide, glabrous beneath, glabrous or very sparsely
pilosulous above, often ciliate with incurved hairs; panicle-branches winged,
commonly glabrous; hypanthium glabrous.

Widely distributed in northern South America, northward to Trinidad and
Panama. The Panama collection cited below is the type of variety alata Cogn.,
and has leaves half as wide as long, pilosulous above, and distinctly ciliate.

CANAL ZONE: Chagres, Fendler 149.
14. ACISANTHERA P. Br.

ACISANTHERA P. Br. Hist. Jamaic. 217. f. 22. 1756.

Flowers 4-merous or 5-merous. Hypanthium subglobose to broadly campan-
ulate. Calyx-tube none or very short; sepals triangular to ovate or oblong, equaling
or exceeding the hypanthium. Petals usually small, mostly obovate. Stamens di-
morphic, the epipetalous series smaller or in some species imperfect. Anthers
obovoid to subulate; connective in the episepalous series prolonged below the thecae,
bearing at base two rounded to linear, anterior appendages; that of the epipetalous
stamens shorter or not prolonged, with two short or tuberculiform appendages.
Ovary free, usually subglobose, 2- to 4-celled; style slender; stigma punctiform or
truncate. Capsule 2- to 4-valved; seeds ovoid or oblong, more or less cochleate,
minutely foveolate. Herbaceous or suffrutescent, simple or freely branched plants,
with small, dentate to entire leaves and small, axillary or paniculate, white to pink
or purple flowers.

About 20 species, most numerous at low altitudes in South America; two others
occur elsewhere in Central America and one in Cuba.

a. Leaves ovate to ovate-oblong, conspicuously denticulate or crenate,
eeiy an

cuneate to the petiole; stems freely and diffusely branched.
b. Appendages of the episepalous stamens lanceolate to linear, acute or
subacute; connective of the epipetalous stamens straight to the
filament, about 0.2 mm. long 1. A. QUADRATA.
bb. Appendages of the episepalous stamens clavate or narrowly obovoid,
blunt or rounded; connective of the epipetalous stamens at right
to t

angles to the anther, prolonged about 0.5 mm. he filament............. 2. A. UNIFLORA.
22. Leaves ovate to su rotund, entire or essentially so, strictly sessile at the
broad ase; stems erect, simple or sparingly branched 3. A. LIMNOBIOS.

l. ACISANTHERA QUADRATA Pers. Syn. Pl. 1:477. 1805.

Stem freely branched, 2-4 dm. tall, sharply angled or narrowly 4-winged,
Setose at the nodes, on the internodes glabrous, pilosulous, or glandular-pubescent;
leaf-blades ovate, ovate-oblong, or oblong, 1-2 cm. long by half as wide, acute or
subacute, cuneate at base into a short petiole, 3-nerved, glabrous, pubescent, or
glandular; flowers solitary in the axils, on pedicels 1-4 mm. long; hypanthium

roadly campanulate, 2 mm. long, usually glabrous, sometimes sparsely pubescent
or glandular; sepals triangular, equaling the hypanthium; petals 5-6 mm. long;
episepalous anthers truncate, about equaling the connective, this bearing two

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[Vol. 45
230 ANNALS OF THE MISSOURI BOTANICAL GARDEN

nur

È DL
< RU.

Fig. 76. Acisanthera quadrata

slender spurs about 1 mm. long; epipetalous anthers smaller, the connective barely
prolonged and bearing two minute, rounded, basal tubercles.
Throughout Central America, north to Veracruz and Acapulco; Greater
Antilles; South America, where it is less abundant than the next species; reported
rom Dominica.

CHIRIQUÍ: l 0 m kes
1989; vicinity of El Valle, Allen Ir 74; vicinity of Penonomé, Williams 163. dg ss
hills west of Soná, about 500 m , Allen 1071. PANAMA: vicinity of Pacora, about it us
Allen 987; vicinity of Juan Diaz, Killip 3262; San José Island, Johnston 823. PRO
UNKNOWN: Pittier 4906.

2. ACISANTHERA UNIFLORA (Vahl) Gl. in Phytologia 3:346. 1950.

Rbexia uniflora Vahl, Symb. 2:48. 1791.
Rbexia recurva L. C. Rich. in Act. Soc. Nat. Hist. Paris 1792:108. 1792.
Acisanthera recurva of recent literature.

Much resembling the preceding in habit and foliage, but the leaves m
proportionately narrower and the leaves, pedicels, and hypanthium more

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1958]
FLORA OF PANAMA (Melastomataceae) 231

glandular-pubescent; sepals linear-oblong, obtuse, 3 mm. long; anthers of the
episepalous stamens linear-subulate, 2 mm. long, the erect appendages clavate or
narrowly obovoid; anthers of the epipetalous stamens similar, 1.5 mm. long, the
connective prolonged 0.5 mm. at base and bearing two erect, rounded appendages
0.2 mm. long.

Widely distributed in South America; reported from Panama and Costa Rica.

3. ACISANTHERA LIMNOBIOS (DC.) Triana, in Trans. Linn. Soc. 28:33. f. II.
1871.

Rbexia limnobios DC. Prodr. 3:117. 1828.

Stems stout, often inflated at base, 5-20 cm. tall, simple or sparingly branched,
sharply 4-angled or narrowly 4-winged, increasingly glandular-pubescent above;
leaves sessile, broadly ovate to subrotund, 5-10 mm. long, subcordate and clasping
at base, entire to remotely denticulate or the uppermost glandular-ciliate, obscurely
5- to 7-nerved; pedicels 2-3 mm. long, glandular-pubescent; hypanthium cam-
panulate, 2-2.5 mm. long, glandular-pubescent; sepals triangular-lingulate, 2.5-3
mm. long, obtuse, glandular-pubescent; petals obovate, 4 mm. long; filaments
2.6 or 2.2 mm. long; anthers narrowly oblong, 1.8 or 1.2 mm. long, the larger
declined, the smaller erect; connective of the larger anthers horizontal, nearly 1
mm. long, with two erect, flattened, completely separate, obtuse appendages 0.8 mm.
long; connective of the smaller anthers short, straight, with two rounded basal
tubercles

Panama and Colombia to Brazil; also known from Honduras.

cocLE: El Valle de Antón, about 600 m., Allen 1990.

15. PTEROLEPIS (DC.) Miq.

PTEROLEPIS Miq. Comm. Phyt. 72. 1840.

Flowers 4-merous. Hypanthium broadly or narrowly campanulate, sometimes
ribbed, pubescent with simple, branched, or glandular hairs, persistent and accres-
cent in fruit. Calyx-tube not prolonged; sepals erect, triangular, acute, ciliate.
Petals small, obovate, often ciliate. Stamens dimorphic. Filaments slender, flat,
straight, glabrous. Anthers subulate to ovoid or obovoid; connective more or less
Prolonged below the thecae, often horizontal or arcuate, terminating in two small
anterior appendages or lobes. Ovary free, oblong to cylindric, 4-celled, glabrous
below, crowned with stiff erect setae surrounding the base of the straight slender
style; stigma small, capitate. Capsule 4-valved, included by the hypanthium;
seeds minute, cochleate, finely tuberculate. Erect, freely branched herbs, with
Ovate to linear leaves and cymose or glomerate, sessile or subsessile, pink or white
flowers; hairs of the stem often geniculate or prolonged backward at base, those
of the hypanthium often branched from the end of a conspicuous stipe.

About 30 species, chiefly in tropical South America; two others have been
reported from Central America and a fourth from the Lesser Antilles.

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[Vol. 45
232 ANNALS OF THE MISSOURI BOTANICAL GARDEN

7 Te
y ES a
SAM 3 BTW mt
A jt r
"A
IN WYŻ
S |
X JE Le
> ¿dl

Fig. 77. Pterolepis pumila

l. PTEROLEPIS PUMILA (Bonpl.) Cogn. in Mart. Fl. Bras. 143:263. 1885.
Rhexia pumila Humb. & Bonpl. Rhex. £. 35. 1823.

Erect, freely branched, 1-6 dm. tall; stem strigose, the hairs geniculate or
prolonged at base; petioles densely strigose, 1-5 mm. long; leaf-blades firm, lance-
olate to elliptic or ovate, 20-45 mm. long, 6-17 mm. wide, acute or acuminate,
rounded or obtuse at base, 3- to 5-nerved, pilose on both sides; flowers very
numerous, terminating the stem and all its branches, soon appearing lateral from
proliferation, on pedicels 1-3 mm. long; hypanthium narrowly campanulate, 3-35
mm. long at anthesis, pilose, some hairs usually with a stout stalk 0.5-1 mm. long
and 3—6 slender terminal branches; sepals erect, persistent, ovate-lanceolate, long-
ciliate, acute; petals pink, obovate, 4-5 mm. long, glandular-ciliate; filaments =
2.5 mm. long; episepalous anthers 1-2 mm. long, narrowly oblong, truncate, the
connective prolonged 1 mm. or less and bearing two minute lobed or short anterior
spurs at its base; fruiting hypanthium 5 mm. long.

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1958]
FLORA OF PANAMA (Melastomataceae) 233

Southern Mexico southward through Central America and widely distributed
in northern South America.

cocLE: El Valle de Antón, 600-800 m., Allen 2860. PANAMA: vicinity of Pacora,
pe 35 m., Allen 988; San José Island, Johnston 824, 889. PROVINCE UNKNOWN:
Halsted.

16. TIBOUCHINA Aubl.

TIBOUCHINA Aubl. Pl. Guian. 1:445. £. 177. 1775.
Flowers 4- to 8-merous. Hypanthium tubular to broadly campanulate, thin-
walled, often costate at maturity. Calyx-tube briefly prolonged, usually erect;
sepals erect, spreading, or recurved, ovate to linear, small or large; exterior teeth
none. Petals usually ciliate. Stamens isomorphic to strongly dimorphic. Filaments
terete or flattened, in some species pubescent or glandular. Anthers subulate, linear,
or narrowly ellipsoid or oblong, 2-celled; connective more or less prolonged at base
aring two anterior lobes or short appendages. Ovary free, its locules
isomerous with the petals, its summit pubescent or setose and often glandular.
Style elongate, straight, sigmoid, or bent at the summit; stigma punctiform.
Capsule 4- to 8-valved, enclosed by the persistent hypanthium. Seeds minute,
cochleate, tuberculate. Herbs or shrubs, almost all pubescent or lepidote; leaves
isomorphic; flowers medium-sized to large, sometimes enclosed by bracts, solitary
or in terminal or axillary cymes; hairs of the upper leaf-surface adnate to the
epidermis for a part of their length.

At least 250 species, most numerous in southern Brazil and in the Andes of
Bolivia and Peru; about thirty species occur in other parts of Central America and
Mexico and four in the West Indies.

a. Flowers 4-merous.

b. — triangular-lanceolate, 4.5—5 mm. long; connective of the epi-

pa

ous stamens prolonged about 2 mm., much longer than that of
i 1. T. OERSTEDII.

the inner se
bb. Sepals triangular-ovate, 2—3 mm. long; connectives of both series of
equal 2. T. NANA.

Stamens about

aa. Flowers 5-merous.
b. Stem and hypanthium lepidote; connective pilose 3. T. BIPENICILLATA.
bb. Stem and hypanthium pubescent; connective glabrous 4. I. LONGIFOLIA.

l. TIBOUCHINA OERSTEDII (Triana) Cogn. in Mart. Fl. Bras. 14*:297. 1885.
Pterolepis oerstedii Triana, in Trans. Linn. Soc. 28:40. 1871.
Stems loosely spreading, freely branched, decumbent or prostrate, sharply 4-
angled, thinly strigose; petioles 3-6 mm. long; leaf-blades ovate, 1.5-3 cm. long
Y half as wide, acute, entire, acute or broadly cuneate at base, 5- to 7-pli-nerved,
densely and softly pubescent above, more thinly so beneath; flowers usually few, on
slender peduncles 2-3 cm. long; hypanthium narrowly campanulate-obconic, 4-4.5
mm. long, narrowly 4-winged, with 4 intermediate ribs, glabrous or setose on the
sides, strigose on the wings, more or less setose at the summit; sepals triangular-
ceolate, 4.5-5 mm. long, spreading, acuminate, 1-nerved, densely ciliate, tipped
With a stout seta 1-2 mm. long; petals narrowly obovate or almost spatulate, 11-

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[Vol. 45
234 ANNALS OF THE MISSOURI BOTANICAL GARDEN

12 mm. long; filaments 5 or 4.3 mm. long; thecae linear-subulate, 5—5.5 or 3-3.5
mm. long; connective 2—2.5 or 0.7 mm. long, with two small, rounded, anterior
lobes; ovary 4-celled, setose at the summit; style 8-10 mm. long.

Costa Rica, Panama, and northern Colombia.

CHIRIQUÍ: vicinity of Boquete, 1200-1500 m., Woodson & Schery 760. cocLÉ:
vicinity of El Valle, Allen 757. PROVINCE UNKNOWN: Pittier 5355.

2. TIBOUCHINA NANA (Standley) Gl. in Phytologia 1:133. 1935.
Chaetolepis nana Standley, in Field Mus. Publ. Bot. 4:247. 1929.

Stems herbaceous, ascending, 2-3 dm. long, forming dense mats, the branches
sparsely villous; petioles flattened, appressed-setose, 4-8 mm. long; leaf-blades thin,
ovate to broadly elliptic, 10-25 mm. long, 6-15 mm. wide, acute, entire, broadly
cuneate at base, 5- to 7-pli-nerved, above shortly soft-villous and sparsely setose,
beneath subsericeous; flowers solitary, terminating the branchlets, on slender
pedicels 5-8 mm. long; hypanthium obconic, 4-6 mm. long, obscurely 4-angled,
sparsely pubescent; sepals ovate, 2-3 mm. long, ciliate; petals purple, 5-6 mm.
long; stamens dimorphic; filaments 3.7 or 2.4 mm. long; thecae straight, 3.2 or
2.1 mm. long; connectives 0.6 or 0.3 mm. long, including the rounded basal
appendages.

Costa Rica and Panama.

BOCAS DEL TORO: Almirante region, Cooper 232, 233.

3. TIBOUCHINA BIPENICILLATA (Naud.) Cogn. in Mart. FI. Bras. 145:385. 1885.

Lasiandra bipenicellata Naud. in Ann. Sci. Nat. Bot. III 13:156. 1849.

Stems erect, woody, sparsely branched, obscurely 4-angled, scabrous with
brown, appressed, lanceolate scales; petioles 6-12 mm. long; leaf-blades sub-
coriaceous, oblong-lanceolate to ovate-oblong, 6-15 cm. long, 2-7 cm. wide,
acute, rounded at base, 5-nerved, the lateral nerves separate, strigose above, beneath
coarsely strigose on the nerves, softly pubescent on the surface; inflorescence many-
flowered, 5-10 cm. long, becoming lepidote above; bracts broadly ovate, 1-3 mm.
long; pedicels 1-3 mm. long; hypanthium narrowly campanulate, 8 mm. long,
silvery or cinereous with appressed lanceolate scales; sepals erect or nearly 0;
triangular, about 3 mm. long, acute; petals triangular-obovate, purple, about Lem
long; stamens dimorphic; filaments about 1 cm. long, often pilose distally, thecae
subulate, arcuate, 9-10 or 7 mm. long; connectives slender, prolonged 3.5 or 2 mm.
to the filament, their appendages slender, 1-2 mm. long, densely long-pilose; style
about 16 mm. long.

Costa Rica, Panama, Colombia, and Venezuela.

Allen

CHIRIQUÍ: Cerro Vaca, Pittier 5304. VERAGUAS: vicinity of Santiago, 50 m»
13.

4. TIBOUCHINA LONGIFOLIA (Vahl) Baill. in Adansonia. 12:74. 1877.
Rbexia longifolia Vahl Eclog. Am. 1:39. 1796.

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1958]
FLORA OF PANAMA (Melastomataceae) 235

Fig. 78. Tibouchina longifolia

Stems woody, erect, freely branched, densely pilose with stout, appressed,
ascending, or spreading hairs mostly 1.5-3 mm. long; petioles slender, 5-20 mm.
long; leaf-blades thin to firm, ovate-oblong to narrowly lanceolate, 3-10 cm. long,
1-3 cm. wide, acute or acuminate, essentially entire, at base narrowly to broadly
cuneate or rarely obtuse, 5-nerved, the lateral nerves confluent toward the base,
densely or sparsely, usually stiffly, pilose above, beneath softly villous on the surface,
more or less strigose on the veins; flowers in small cymes terminating the stem and
branches, often very numerous, forming large panicles; hypanthium broadly cam-
panulate, 3—3.5 mm. long, 10-ribbed, densely setose, the hairs mostly ascending;
sepals triangular-subulate to linear, setose and ciliate, shorter than, equaling, or
slightly exceeding the hypanthium, usually recurved; petals white or purplish, 5—7
mm. long; stamens nearly isomorphic; thecae straight, 2-3 or 1.5 to 2.5 mm. long;
connectives straight or curved, 0.3-0.7 mm. long, including the small or minute
basal lobes; fruiting hypanthia nodding.

Southern Mexico to Panama and southward to Brazil and Bolivia; Cuba, His-
Paniola, and occasional, possibly introduced in other parts of the West Indies; the
most abundant and most widely distributed species of the genus.

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[Vol. 45
236 ANNALS OF THE MISSOURI BOTANICAL GARDEN

BOCAS DEL TORO: Carleton 127; Changuinola Valley, Potrero Island, Dunlap 73.
CANAL ZONE: Barro Colorado Island, Aviles 105, Woodwortb & Vestal 672; Gatún, Hayes
3; Chagres, Fendler 129. cHIRIQUI: Boquete, Davidson 626, Terry © Terry 1050. cocLÉ:
Valle de Antón, 600-800 m., Allen 2855. DARIÉN: Chepigana district, Terry & Terry
1392; Cana and vicinity, Williams 733. PANAMA: vicinity of San Carlos, 0-10 m., Allen
1147.

17. MONOLENA Triana
MONOLENA Triana, in Benth. & Hook. Gen. Pl. 1:756. 1867.

Flowers 5-merous. Hypanthium obconic, glabrous, 3-angled. Calyx-lobes short,
obtuse. Petals obovate, ephemeral. Stamens nearly isomorphic, somewhat unequal
in size; anthers linear or narrowly oblong, obtuse; connective prolonged below the
thecae, minutely tuberculate dorsally at base, extended below the summit of the
filament into a single ventral spur. Capsule trigonous, expanded laterally into a
triangular summit depressed at the center; seeds obpyramidal. Rhizomatous herbs,
often epiphytic, the leaves apparently all basal, one member of each pair minute
and rarely persistent at anthesis; flowers few, pink or white, sessile in a secund
spike on an elongate scape.

Possibly 6 or 8 species, all poorly known and inadequately represented in
herbaria, ranging from Guatemala to Peru.

1. MONOLENA OVATA Cogn. in Bull. Acad. Belg. III. 14:945. 1887.

Plant glabrous; petioles 1-1.5 dm. long; leaf-blades ovate, 1.5-2.5 dm. long,
about three-fifths as wide, abruptly short-acuminate, broadly rounded at the base,
7-pli-nerved; scapes 2-4 dm. tall; spikes 4-6 cm. long, 3- to 5-flowered; petals
white, bordered with pink, about 2.5 cm. long; capsule about 1 cm. wide.

Panama and Colombia.

DARIÉN: crest of the Cana-Cuasi trail, 1650 m., Terry & Terry 1543.

18. DIOLENA Naud.

DioLENA Naud. in Ann. Sci. Nat. Bot. III. 15:329. 1851.

Closely resembling Triolena in the structure of the inflorescence, fruit, and
flowers, differing in the anthers which bear only two ventral spurs. S bs or
herbs, often of considerable size, in most species with strongly dimorphic leaves-

About 10 species of Andean South America and Amazonia, extending no
in Central America as far as Guatemala.

1. DIOLENA SPICATA Triana, in Trans. Linn. Soc. 28:81. 1871.

Shrub to 2.5 m. tall with densely strigose stems; larger leaves oblanceolate,
acuminate, ciliate, denticulate above the middle, at base oblique and cordulate,
tending to be convex on one margin toward the base and concave on the othets
sparsely setose on both sides, 5-pli-nerved, the inner pair of nerves alternate; petiole
2-4 mm. long; smaller leaves deflexed, reniform, 4-8 mm. wide; spikes up to 10
cm. long.

(236)

1958]

FLORA

OF PANAMA (Melastomataceae)

Fig. 79. Monolena ovata

(237)

237

[Vol. 45
238 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Western Colombia and Panama.

BOCAS DEL TORO: Water Valley, von Wedel 1510.
19. TRIOLENA Naud.

TRIOLENA Naud. in Ann. Sci. Nat. Bot. III. 15:328. 1851.

Flowers 5-merous. Hypanthium thin-walled, cup-shaped, usually strigose.
Sepals small; exterior teeth present and projecting. Petals small, ephemeral. Sta-
mens isomorphic, but some frequently partly abortive; anthers linear; connective
prolonged briefly at base and bearing three filiform or clavellate, ventral spurs,
each of them about as long as the anther and two of them usually curved-ascending
past the anther. Ovary 3-celled, superior. Fruit a 3-celled capsule of peculiar
structure; outer walls of the ovary growing laterally into three prominent longi-
tudinal wings and distally much beyond the actual summit of the ovary; upper
part of the ovary expanding into three slender lobes extending into the wings;
dehiscence apparently circumscissile between the summit of the ovary and the
lateral walls; capsule invested to its margin by the accrescent calyx. Seeds minute,
pyramidal, not cochleate. Caulescent shrubs with small flowers in dense secund

Fig. 80. Triolena hirsuta

(238)

1958]
FLORA OF PANAMA (Melastomataceae) 239

spikes or spike-like racemes; leaves isomorphic but often unequal, the smaller one
of each pair often deciduous.

About 8 species, chiefly in the mountains from southern Mexico to Peru.
Flowers are rarely collected and the species are not well understood.

1, TRIOLENA HIRSUTA (Benth.) Triana, in Trans. Linn. Soc. 28:81. 1871.
Bertolonia birsuta Benth. Bot. Voy. Sulphur 94. 1844.

Shrub up to 5 dm. tall; leaves essentially isomorphic; petioles strigose, 1-2 cm.
long; leaf-blades elliptic or oblong, 5-15 cm. long, half to two-thirds as wide,
abruptly acuminate, entire, rounded or broadly obtuse at base, 5- to 7-nerved,
hirsutulous above, beneath strigose on the principal veins, sparsely pilose on the
veinlets, glabrous on the surface; spike terminal, short-peduncled, 3—6 cm. long,
densely flowered, secund; hypanthium strigose, about 2.5 mm. long; petals white,
about 3 mm. long; capsule about 7 mm. wide.

Guatemala to Panama.

Bocas DEL TORO: Water Valley, von Wedel 702, 922; Pumpkin River, von Wedel
2568; vicinity of Chiriqui Lagoon, von Wedel 1020, 2613; vicinity of Nievecita, 0-50 m
dson

Woodson, Allen & Seibert 1825; without locality, von Wedel Zaj, 291; Chagulani
am Dunlap 279, 558. cocLE: vicinity of El Valle de Antón, 500-1000 m., Allen
1204, Se. ÓI. DARIEN: Chepigana district, 600 m., Terry Di > 5, 163 3; Cana

and vi T "William 757. PROVINCE UNKNOWN: Rowlee & Stork I
20. BLAKEA P. Br.

BLAKEA P. Br. Hist. Jamaic. 323. 1756.

Flowers usually 6-merous, each closely subtended by two pairs of bracts.
Hypanthium campanulate or cup-shaped, usually thick-walled. Calyx truncate
to deeply lobed. Petals usually large, commonly unsymmetrical. Stamens iso-
morphic; filaments short; anthers coherent in a ring, short, oval or oblong, abtuse
or rounded at the summit, laterally flattened, opening by two minute pores; con-
nective usually slightly prolonged or dilated at base into a dorsal spur. Ovary
wholly inferior, usually 6-celled; stigma punctiform or capitate. Woody plants,
ne epiphytic or climbing by adventitious roots when young, sometimes reaching

rge size, with conspicuous flowers solitary or fascicled in the axils of the leaves;
dl with the secondary veins usually close together, straight, and parallel.

At least 50 species, most numerous in the northern Andes; several others occur
in other parts of Central America.

a. Bracts, both outer and inner Imost to the summit.
b. Bracts glabrous externally « or v senti s o.
bb. Bracts dense ly pubesc :
c. Petals de tuberculae sepals eie with a red foliaceous
appendage; younger stems glabrous or merely koń SON
cc. Petals not tu Pale e ak unappendaged; ide stems
rsute

1. B. WOODSONII.

2. B. CALYCOSA.

pubescent to hirs
d ripae sessile; iaf-esius bonesch coarsely hirsute..............———— 3. B. CRINITA.

dd. Flowers on pedicel 5-20 mm. long; leaf-veins merely pubescent
beneath

4. B. BRUNNEA.

[Vol. 45
240 ANNALS OF THE MISSOURI BOTANICAL GARDEN

== A
"ST
RIZZI
> U

a

Fig. 81. Blakea woodsonii

(240)

1958]
FLORA OF PANAMA (Melastomataceae) 241

aa. Outer bracts, and usually also the inner, separate to the base. (Since the
pedicel passes imperceptibly into the hypanthium, they may appear to be
pane or a small part of their length.
Out

b. r bracts conspicuously longer than the hypanthium and calyx
Sebi ed.
c. Calyx ibi the lobes represented by 6 minute tubercles; pedicels
glabrous; secondary veins about 0.5 mm. apart 5. B. GRACILIS.
ec. LE — eres triangular with concave des about 1.5 mm.
long; pedicels stout, pubescent; secondary v —1.5 mm. apart..... 6. B. FOLIACEA.
bb. Outer hess distinctly shorter than the h Lee
- Calyx truncate, the lobes sagre by minute tc a OM aig 7. B. PAUCIFLORA,
cc. Calyx-lobes triangular, ac 8. B. PARVIFOLIA.

—
.

BLAKEA WOODSONII Gl. ex Woods. & Seib. in Ann. Mo. Bot. Gard. 26:296.
1939

Shrub or small tree to 9 m. tall, the younger branches and petioles smooth to
sparsely furfuraceous-hispid; petioles 1-3 cm. long; leaf-blades oblong-obovate to
obovate, 7-12 cm. long, half to two-thirds as wide, rounded to a small triangular
apex, entire, broadly cuneate to obtuse at base, glabrous above, essentially glabrous

neath, 3-nerved or barely 3-pli-nerved; pedicels solitary or rarely paired, stout,
1-5 cm. long; bracts connate nearly to the summit, the outer 12 mm. long, the
inner about 15 mm., glabrous or nearly so at anthesis; petals white, 2.5-4 cm. long;
anthers flattened, rounded, 5-7 mm. long.

Endemic to Panama.

CHIRIQUÍ: cloud forest on Cerro Horqueta, 1950 m., von Hagen & von Hagen 2045,
2028, 2071; vicinity of Casita Alta, Volcán de Chiriquí, 1500-2000 m., Woodson, Allen
8 Seibert 951; Casita Alta to Cerro Copete, 2300-3300 m., Woodson & Schery 370;
of Cerro Punta, 2000 m., Allen 3483; summit and southwest face of Cerro Copete, 2700

» Allen 4907; Cerro Punta to headwaters of Río Caldera, 2250-2500 m., Allen 1454;
Bajo Chorro, 2100 m., Davidson 401, 401A, 401B; Bajo Chorro, 1800 m., Davidson 1 75.

2. BLAKEA CALYCOSA Gl. in Phytologia 1:342. 1939.

Epiphytic shrub, the younger stems, petioles and leaf-blades glabrous or thinly
furfuraceous; petioles rather slender, 2—4 cm. long; leaf-blades thin, elliptic, up
to 25 cm. long, about a third as wide, abruptly and shortly caudate, acuminate,
rounded at base, 3-nerved with an additional pair of marginal nerves; flowers few
in the axils, on pedicels about 5 mm. long; bracts connate to the middle, pubescent
on the outside and distally on the inside; calyx-tube widely spreading; sepals ter<
minated by a foliaceous, reflexed, folded, broadly ovate, pubescent appendage;
petals rose-red, 2—2.5 cm. long, tuberculate on both sides; anthers broadly elliptic,
about 5 mm. long, the connective bearing a short, obtuse, basal spur.

Known only from the type specimen.

COCLÉ: north rim of El Valle de Antón, 600—1000 m., Allen 1788.

3. BLAKEA CRINITA Gl. in Phytologia 3:356. 1950.

Shrubs 3-6 m. tall, the younger stems and petioles densely long-hirsute; petioles
Very stout, 1-3 cm. long; leaf-blades coriaceous, broadly elliptic, up to 14 cm.
long and 11 cm. wide, rounded at both ends, 5-nerved with an additional pair of

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[Vol. 45
242 ANNALS OF THE MISSOURI BOTANICAL GARDEN

marginal veins, glabrous above, beneath densely long-hirsute on the principal veins,
less so on the veinlets, and sparsely pilose on the surface; flowers solitary or paired,
sessile or subsessile; bracts connate, densely hirsute with hairs about 4 mm. long;
petals pink, about 2 cm. long; anthers broadly elliptic, about 6 mm. long, the
connective bearing a very minute dorsal spur.

Known only from the type specimen.

COCLE: crest of Cerro Pajita, north of El Valle, 1200 m., Allen 3956.

*

BLAKEA BRUNNEA Gl. ex Woods. & Schery, in Ann. Mo. Bot. Gard. 28:435.
1941.

Tree to 12 m. tall or perhaps taller, also reported as an epiphyte, the younger
stems, petioles, pedicels, and sepals coarsely pubescent; petioles stout, 3—6 cm. long;
leaf-blades subcoriaceous, elliptic or elliptic-ovate, up to 25 cm. long and 15 cm.
wide, acute or apiculate, broadly rounded at base, glabrous above, brown and
glabrous on the surface beneath at maturity, permanently pubescent on the veins,
5-nerved, the outer pair diverging at right angles from the midvein and together
forming the arc of a circle; pedicels solitary or rarely paired, stout, 1-2 cm. long;
bracts connate, about 25 mm. long; petals large, white; anthers broadly rounded,
about 7 mm. long.

Endemic to Panama.

CHIRIQUI: Bajo Chorro, Boquete District, 2100 m., Davidson 332; cloud forest on Cerro
Horqueta, 1950 m., von Hagen & von Hagen 2131; Robalo Trail, northern slopes of

erro Horqueta, 1800-2100 m., Allen 4988; vicinity of Bajo Chorro, 1900 m., Woodson 3
Schery 618.

5. BLAKEA GRACILIS Hemsl. Diagn. Plant. Nov. 13. 1878.

Epiphytic shrub or vine, also becoming an independent tree and reported 12
m. tall, glabrous or nearly so; petioles 5-12 mm. long; leaf-blades commonly
obovate-oblong, 5-10 cm. long, two-fifths to three-fifths as wide, rather gradually
acuminate to a slender tip, cuneate to the base, apparently 3-pli-nerved but with
another pair of submarginal veins; secondary veins about 0.5 mm. apart; pedicels
solitary, slender, glabrous, 3-6 cm. long; outer bracts ovate, foliaceous, 2-3 cm.
long, many-nerved, much exceeding the inner ones and the calyx; calyx truncate,
the lobes represented by minute tubercles only; petals pink, broadly and obliquely
obovate, 1.5-2 cm. long.

Costa Rica and Panama.

4; >
m., Allen 315; vicinity of New Switzerland, central valley of Rio Chiriqui Viejo, !
2000 m., Allen 1354.

6. BLAKEA FOLIACEA Gl. in Phytologia 3:355. 1950.
Tree up to 18 m. tall, the younger stems, petioles, and pedicels closely red-

brown-pubescent; petioles slender, 1-2 cm. long; leaf-blades oblong or oblong-

(242)

1958]
FLORA OF PANAMA (Melastomataceae) 243

elliptic, 8-14 cm. long, about half as wide, abruptly acuminate to a slender tip,
obtuse to rounded at base, glabrous, 5-pli-nerved, the secondary veins 1-1.5 mm.
apart; peduncles stout, solitary, 3-6 cm. long; bracts foliaceous, spreading, separate
to the base, the outer about 3 cm. long, 2 cm. wide, the inner somewhat smaller;
calyx-lobes manifest, depressed-triangular; petals very obliquely cuneate-obovate,
white, about 3 cm. long

Endemic.

BOCAS DEL TORO: Fish Creek, vicinity of Chiriquí Lagoon, von Wedel 2219; Fish

Creek Mountains, von Wedel 2277. cocLé: vicinity of La Mesa, altitude 1000 m., Allen
2312.

7. BLAKEA PAUCIFLORA Gl. in Phytologia 3:357. 1950.

Small tree 6-7 m. tall, glabrous throughout. Petioles 8-10 mm. long. Leaf-
blades broadly elliptic-obovate, up to 5 cm. long and 3 cm. wide, rounded above
to an apiculum 6-10 mm. long, broadly cuneate at base, 5-nerved. Flowers
solitary in the axils on pedicels about 2 cm. long. Bracts separate to the base,
broadly triangular, the outer 2.5 mm. long, the inner a little longer. Hypanthium
3 mm. long. Calyx widely flaring, almost truncate, its lobes reduced to minute
tubercles. Petals white, semi-obovate, 9 mm. long. Anthers broadly oval, 2.5-3
mm. long, the connective neither elevated nor spurred.

Known only from the type specimen.

CHIRIQUÍ: foot of Sierra del Boquete, Maurice 744.

oo

. BLAKEA PARVIFOLIA Gl. in Phytologia 3:357. 1950.

Small tree, up to 10 m. tall, glabrous throughout. Petioles 5-8 mm. long.
Leaf-blades cuneate-obovate, 2.5—4 cm. long, about half as wide, rounded above to
a short apiculum, long-cuneate at base, 3-nerved. Flowers solitary in the axils on
pedicels about 1 cm. long. Bracts separate to the base, oblong-ovate, very blunt,
much shorter than the hypanthium. Hypanthium almost spherical, 3 mm. long.
Calyx-lobes depressed triangular, about 1 mm. long, acute. Petals pink, oblong-
obovate, 8-9 mm. long. Anthers semi-ovate, about 3 mm. long, the connective
neither elevated nor spurred.

Known only from the type specimen.

COCLE: crest of Cerro Pajita, El Valle de Antón, 1100 m., Allen 3761.

21. TOPOBEA Aubl.
TOPOBEA Aubl. Pl. Guian. 1:476. t. 189. 1775.

Stamens isomorphic; filaments relatively short; anthers coherent in a ring, often
laterally flattened, subulate or lanceolate, often somewhat cleft at the summit,
opening by two minute pores; connective in most species prolonged straight back
into a short, slender, dorsal spur. Otherwise precisely as in Blakea. ;

Probably 50 species of northern South America and Central America, especially

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[Vol. 45
244 ANNALS OF THE MISSOURI BOTANICAL GARDEN

in regions of high rainfall. Every type of leaf, inflorescence, bract, and calyx
known in the genus occurs also in Blakea, and the two genera can be accurately
distinguished only by the anthers. Besides the species described below, T. superba
Naud. was reported from Panama by Cogniaux. No specimen has been seen,
during the preparation of this treatment, which can be referred to it.

a. Leaves sessile or subsessile, rounded to cordate at bas
> a vers or e obtuse, dr 3-nerved; younger

s pubescent; pedicels 2-4 cm. long 1. T. ELLIPTICA.
bb. Blades Soia ra: ed plant glabrous; flowers sessile or
subsessile 2. T. CORDATA.
aa. Leaves distinctly petioled.
b. Outer bracts acute to acuminate, distinctly surpassing the inner ones.
E febr parts pan with barbellate hairs up to 1 mm. long;
sepals 2-4 mm 3. T. UROPHYLLA.
cc. Younger parts E furfuraceou
. Leaves thin para = pli- "ac? secondary veins about 1 mm.
apart; calyx-lob 4. T. PLUVIALIS.
dd. is caves > shining T preci y secondary veins 0.3—0.5 mm. apart;
anthesis ruptured nearly to the torus . T. COOPERI.

bb. Outer brac date rounded at the summit, equaling or shorter than
the i

c. Mower appearing when the leaves are lacking or at least not fully
expanded; sepals triangular-o 6. T. PRAECOX.
e ae appearing wh ile the sese are fully expanded; sepals almost

d. Loves Barsa coriaceous; flowers 1-2 per leaf axil on pedicels
long

Hd

. REGELIANA.

dd. dms p nene flowers 4—6 per leaf axil on pedicels
10-15 mm. long 8. T. MEMBRANACEA.

1. TOPOBEA ELLIPTICA Gl. in Phytologia 3:353. 1950.

Epiphytic shrub about 2 m. tall, the younger parts sparsely pubescent with
both simple and glandular hairs; leaves sessile, oblong or obovate-oblong, up to 7
cm. long and 4.5 cm. wide, rounded at the summit, subcordate and clasping at
base, 3-nerved; pedicels solitary in the axils, up to 4 cm. long; flowers small; bracts
foliaceous, separate to the base, broadly rounded, about equaling the hypanthium;
sepals triangular, recurved after anthesis.

Endemic.

an eue DEL TORO: Robalo trail, northern slopes of Cerro Horqueta, 1800-2100 m»
en 500

2. TOPOBEA CORDATA Gl. in Phytologia 3:354. 1950.

Small epiphytic shrub, glabrous throughout; leaves sessile, ovate, up to 8 o™
long, about two-thirds as wide, acute, broadly rounded to the subcordate base,
5-nerved with an additional pair of marginal veins; flowers solitary in the axils, 0%
pedicels about 1 mm. long; bracts free to the base, broadly elliptic, rounded at
both ends, the outer subcoriaceous, about 11 by 8 mm., the inner thinner, some”
what smaller; hypanthium nearly 4 mm. long; sepals broadly elliptic-ovate
somewhat imbricate at base, about 4 mm. long and 3 mm. wide.

Endemic.

(244)

1958]
FLORA OF PANAMA (Melastomataceae) 245

cocLE: Cerro Pajita, hills north of El Valle de Antón, 1000—1200 m., Allen 6 Allen
4178.

3. TOPOBEA UROPHYLLA Standl. in Field Mus. Pub. Bot. 22:162. 1940.

Shrub to 3.5 m. tall, the younger parts, petioles, and peduncles pubescent with
short, stout, rough hairs; petioles 1-2 cm. long; leaf-blades oblong or elliptic, up
to 13 cm. long, about half as wide, abruptly acuminate to a slender tip, rounded
to broadly obtuse at base, 5-pli-nerved, the outer pair of nerves submarginal,
glabrous above, sparsely pubescent on the veins beneath; pedicels solitary or few in
each axil, 4-8 mm. long; bracts separate to the base, thinly stellate-pubescent, the
outer ovate, acuminate, about equaling the hypanthium, the inner slightly shorter;
hypanthium stellate-pubescent; calyx-lobes triangular-ovate, 1.5-2.5 mm. long;
Petals pale pink, about 1 cm. long, pubescent externally.

Nicaragua and Panama.

DARIEN: Rio Balsa, above Tucuti, 30 m., Terry & Terry I4II; Cana-Cuasi trail,
Chepigana district, 1200 m., Terry & Terry 1517.

4. TOPOBEA PLUVIALIS Standl. in Field Mus. Pub. Bot. 22:162. 1940.

Shrub up to 3.5 m. tall, the younger parts minutely furfuraceous, otherwise
glabrous throughout; internodes frequently clavate-thickened; petioles 1-2 cm.
long; leaf-blades elliptic to broadly oblanceolate, up to 17 cm. long, about two-

fths as wide, acuminate to a slender tip, cuneate to the base, 5-pli-nerved, the
outer pair submarginal; flowers solitary or few in the axils, on pedicels 1-1.5 cm.
long; bracts free to the base, the outer acuminate, about equaling the hypanthium,
the inner somewhat shorter; calyx nearly truncate at the summit, the lobes repre-
sented by six minute teeth; petals white, about 1 cm. long.

Endemic.

¿ros Cana-Cuasi trail, Chepigana District, 1200-1650 m., Terry 6 Terry 1557,
560.

5. TOPOBEA COOPERI Gl. in Phytologia 3:354. 1950.

Small epiphytic shrub, glabrous except on the youngest parts and the bracts;
stems thickened at the nodes; petioles stout, 1-2 cm. long; leaf-blades elliptic to
elliptic-obovate, up to 10 cm. long, half to two-thirds as wide, abruptly narrowed
to a short slender tip, very broadly cuneate at base, 3- to 5-nerved or barely 3-pli-
nerved, the secondary veins 0.6 mm. apart; pedicels fascicled in the axils, 2-11 mm.
long; bracts distinct, thinly furfuraceous, the outer oblong-obovate, acute, 8
mm. long, the inner broadly rounded at the summit, about two-thirds as long;
hypanthium glabrous, 6 mm. long; calyx nearly closed in bud, at anthesis irregu-

ly ruptured; petals pink, about 12 mm. long, unsymmetrically oblanceolate.

Endemic.

per 199. COCLÉ: vicinity
7

Bocas DEL TORO: Cricamola valley, region of Almirante, Coo
of La Mesa, north of El Valle de Antén, about 1000 m., Allen 2837.

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[Vol. 45
246 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fig. 82. Topobea praecox

6. TOPOBEA PRAECOX Gl. in Phytologia 3:355. 1950.

Shrub or tree 6—20 m. tall, with thick stems and on the flowering branches
very short internodes; leaves lacking or very immature at anthesis; petioles 12
cm. long; leaf-blades elliptic, certainly up to 10 cm. long and about three-fift
as wide, abruptly acuminate, broadly cuneate to subrotund at base, 5-nerved, o
secondary veins 1-2 mm. apart; flowers several in a fascicle, on slender pedicels
up to 1 cm. long; bracts distinct to the base, nearly equal, rotund or even wider
than long, 7-8 mm. long, glabrous or nearly so; hypanthium 4-5 mm. long; sepals
depressed-ovate, rounded at the summit, 3—3.5 mm. long from the torus; peta
rose-purple, narrowly and obliquely obovate, about 17 mm. long.

OCLE: vicinity of La Mesa, north of El Valle de Antón, 1000 m., Allen 2788; Mee

c
of El Valle de Antón, 600 m., Allen 2067, 1742. DARIEN: trail between Pinogana
Yavisa, about 15 m., Allen 294.

7. TOPOBEA REGELIANA Cogn. in DC. Monogr. Phan. 7:1085. 1891.

Tree 10 m. tall, the youngest parts coarsely furfuraceous, otherwise glabrous;
petioles slender, 2-4 cm. long; leaf-blades broadly elliptic to obovate-rotund, uP
to 11 cm. long, half to three-fourths as wide, apiculate, obtuse to subrotund "
base, 5-nerved, the outer pair submarginal, the secondary veins 2-3 mm. apart;
flowers solitary or paired, on pedicels 3-6 mm. long; bracts distinct to the =
the outer obovate-oblong, obtuse, 4-6 mm. long, the inner much broader an

(246)

1958]
FLORA OF PANAMA (Melastomataceae) 247

slightly longer; hypanthium about 7 mm. long; calyx nearly erect, about 2 mm.
long, its lobes obscure or obsolete.
Panama and probably also Costa Rica.

vicinity of Salamanca Hydrographic Station, Rio Pequeni, 80 m.,
Nx. ae. M Seibert 1589; Chagres, Fendler 295.

8. TOPOBEA MEMBRANACEA Wurdack, in Brittonia 9:108. 1957.

Tree or large vine, the young parts puberulous, the leaves as in T. regeliana
but membranous rather than thin-coriaceous at anthesis; flowers 4-6 per upper
leaf axil on slender pedicels about 1—1.5 cm. long, otherwise as in T. regeliana.

Eastern Panama to northwestern Colombia.

DARIÉN: Ensenada Guayabo, Stern $ Chambers 175.
22. CLIDEMIA D. Don

CLIDEMIA D. Don, in Mem. Wern. Soc. 4:306. 1823.

Flowers 4- to 7-merous. Hypanthium cup-shaped to tubular; sepals erect or
spreading, of various shapes and sizes, in some species obsolete; exterior teeth
present, in many species exceeding and more or less concealing the sepals; petals
small, obtuse, mostly white; stamens isomorphic; anthers subulate, linear, semi-
ovoid, or rarely obovoid; connective usually simple, or briefly prolonged below the
isse. or rarely prolonged into a short dorsal appendage; ovary partly or wholly
inferior, its cells commonly isomerous with the petals; style slender; stigma capitate
to punctiform; fruit a globose many-seeded berry. Herbs, shrubs, or rarely even
trees, with axillary flowers or flower-clusters of various types.

About 160 species, widely distributed throughout tropical America. Several
other species are known from Costa Rica and some of them may be expected in

anama.
a. Flowers 5- to 7-m

b. Leaves Pre. dimorphic, one member of each pair always 2-12
times as long as the o

d. Leaves rounded at the base 1. C. GRACILIS.
dd. Leaves cordate and clasping at base 2. C. PITTIERI.
cc. Las short-petioled, more or less pubescent beneath 3. C. OBLONGA.

bb. Leaves isomorphic (in lateral pouce the leaves of each alternate
Pair may be somew = unequal in
c. Torus bearing a me: s fiti or ciliate scales; hypanthium
e with simple e
- Lea să mearga the i inner > of rina veins usually alternate;
bada a ually acute or ol at base, averaging nearly three
times as long a as dn connective not pro! tai below the summit
ati the pres IS AI RI rne
Leaves 5- to >. nerved, usually rounded or subcordate at base,
płodna, less than twice as long as wide; connective minutely
onged into a short dorsal spur...
cc. Toral scales none.
+ Inflorescence distinctly pedunculate, elongate, with a well

defined central axis.

4, C. DENTATA.

a.
pe

5. C. HIRTA.

(247)

[Vol. 45
248 ANNALS OF THE MISSOURI BOTANICAL GARDEN

e. Principal leaves 7- to 9-nerved, cordate or rounded at base;
Me drsd subulate, not convolute; connective si

. C. STRIGILLOSA,
. C. NOVEMNERVIA.

"^
"^
m
M
et
o
je)
Q
R
et
c
©
et
Jg"
o
o
5
PSE
5 3.
CJE
o
n
To
a
E
<
a
g
M
"3
5
2
3
99
rà
= pal
4)
"d
a
Na

A Is
ee. Principal leaves 5-nerved, obtuse or rounded at base; anthers
ea de subulate with ben ud. thecae;
f. Leav ine tomentose on the actual surface beneath......... 8. C. NEGLECTA.
ff. Tian pubescent beneath with c si imple bristles which
obscure or conceal the underlying s sel hairs.
g. Inflorescence elongate, the flex internodes mostly >
cm. long, sparsely lic fort male hairs 1-3 mm. lon
and co pick iof glandular; hairs of the Liana
mostly 1-3 ong
i ee pe pei t and compact, the internodes rarely as
uch as 1 c "aes; * ey hirsute with simple hairs up
mm. long early or quite pit vp e few
gas ir rosa of the hypanthium up t E mm.

9. C. CAPITELLATA.

99
a

. C. DEPENDENS.

_
o

dd. tini not distinctly as cg or elongate, or rarely
with a short peduncle, more or less trichotomous from the base,
without a distinct central axis.

e. Messina none; pubescence of the stem tomentose, hirsute, or

substrig
f ile teeth equaling or bra than the PARA na Sage
hypanthium hirsute abovi nse furfuraceous tomentum..... 11. C. REITZIANA.
f. Exterior meri distinctly frese the sepals; Feud stems
densely subst 12. C. DEPPEANA.
ee. Formita. Bm resent on the petioles; pubescence of the stem
reflexed-set
f. Lower saio po setose = dy veins and veinlets;
setae of the hypanthium 8— 13. C. COLLINA.
ff. Lower edited hirsute tai dea setae of the hypan-
thium 14. C. TAURINA.

aa. Flowers 4-merous.
. Leaves or Lege bearing formicaria
. Formicaria pendent at the base Pri the petiole 15. C. TOCOCOIDEA.
Lr eem summit of the peti sole 16. C. SETOSA.
bk Formica lac skin

c. Exterior teeth Kinda, sug ni equaling or exceeding the
hypanthium; leaves strongly dimorphic

cc. Exterior uel c onic to subulate, much shorter dun DM = prez:
d. Flowers sessile or nearly so in dense cia È clus

. C. EPIPHYTICA»

-
N

e. Leaves densely hirsute above, 5-pli-nerv 18. C. RUBR
ee. Leaves glabrous on the surface above, 3 ui 19. C. DENSIFLORA.
dd. Sf wers in branched cymes
. Leaves broadly ovate, acuminate 20. C. PURPUREO-
VIOLACEA:
ee. Leaves parte sori contracted into a slender linear
apiculum 2-3 cm. 21. C. OMBROPHILA.

1. CLIDEMIA GRACILIS Pittier, in Journ. Wash. Acad. Sci. 13:390. 1923.

Shrub, up to 1.5 m. tall, the younger stems flattened and two-edged, furfur-
aceous when young, later glabrous; leaves dimorphic, the larger sessile, ovate-
lanceolate, 1-2 dm. long, 3-7 cm. wide, acuminate, entire, obtuse or roun =
base, 3-nerved, glabrous and shining above, puberulent on the veins beneath, che
smaller leaves lanceolate, 1-3 cm. long, soon deciduous; inflorescences long-
peduncled, much-branched, many-flowered, up to 18 cm. long; bp
campanulate, 3 mm. long; sepals scarcely differentiated, the exterior tee
stoutly subulate, scarcely surpassing the sepals; anthers linear, the connective

(248)

1958]

FLORA OF PANAMA (Melastomataceae)

249

Fig. 83. Clidemia pittieri

(249)

[Vol. 45
250 ANNALS OF THE MISSOURI BOTANICAL GARDEN

prolonged briefly below the filament into a minute subrotund dorsal appendage.
Panama and Colombia.
ocas DEL TORO: Water Valley, von Wedel 715, 962; Fish Creek Mountains, von
Wedel 2339. CANAL ZONE: lake shore along Gatun River valley, Pittier 6748. COCLE:
El Valle, 500-700 m., Seibert 467; western slope and summit of Cerro Valle Chiquito,
700-800 m., Seibert 408. DARIEN: Chepigana district, 600 m., Terry & Terry 1439.
2. CLIDEMIA PITTIERI Gl. in Bull. Torr. Bot. Club 68:252. 1941.

Shrub to 8 m. tall, glabrous throughout, the young stems terete; leaves sessile,
cordate-clasping, abruptly acuminate, entire, strongly dimorphic, the larger ovate-
lanceolate, 8-15 cm. long, 3-7 cm. wide, 5-nerved, the smaller broadly ovate to
rotund, 2-3 cm. long, nearly or quite as wide; cymes pedunculate, trichotomous,
3-6 cm. long, few-flowered; flowers 5-merous; hypanthium subglobose, 3.3 mm.
long; sepals about 0.6 mm. long, depressed-triangular; exterior teeth resembling
the sepals but slightly smaller; petals red, subrotund, 4 mm. long; anthers thickly
ovoid, the connective elevated into a prominent dorsal ridge.

Endemic to Panama, so far as known.

CHIRIQUÍ: southern slope of Cerro de la Horqueta, 1700 m., Pittier 3177; cloud forest,
Cerro Horqueta, 1950 m., von Hagen & von Hagen 2039; Finca Lérida to Pefia Blanca,
1750-2000 m., Woodson & Schery 289; vicinity of Bajo Mona and Quebrada Chiquero,
1500 m., Woodson & Schery 562; Bajo Chorro, Boquete district, 1800 m., Davidson 397.

3. CLIDEMIA OBLONGA Gl. in Brittonia 3:106. 1939.

Shrub climbing by roots; stems at first paleaceous, soon becoming glabrous;
petioles paleaceous, about 4 mm. long; leaf-blades narrowly obovate-oblong, up to
18 cm. long and 7.5 cm. wide, one member of each pair about half as long as the
other, short-acuminate, entire, narrowed to a rounded base, 3-nerved, glabrous
above, glabrous beneath on the surface, paleaceous on the nerves; flowers in clusters
1-3 cm. long; hypanthium tubular, about 2.5 mm. long, very minutely puberu-
lent; sepals very short, depressed semi-circular; anthers stoutly subulate, 2 mm.
long, the connective prolonged below the filament into an ovate dorsal lobe 0.4
mm. long; ovary wholly inferior.

Known only from the type collection.

SAN BLAS: forests around Puerto Obaldia, San Blas coast, Pittier 4304.
4. CLIDEMIA DENTATA D. Don, in Mem. Wern. Soc. 4:308. 1823.

Shrub 1-4 m. tall, densely hirsute throughout with simple hairs; petioles stout,
1-1.5 cm. long; leaf-blades thin, oblong-ovate to ovate-lanceolate, often unequa
in each pair, commonly 8—20 cm. long, 3-8 cm. wide, acuminate, entire or minutely
serrulate, acute to rounded and usually oblique at base, sparsely pilose on both
sides, 5-pli-nerved, the inner pair usually alternate; inflorescence 1-3 cm. long,
few-flowered, short-stalked; hypanthium campanulate, about 3 mm. long; calyx
truncate; exterior teeth subulate, erect, 2-5 mm. long, hirsute; petals white,
obovate, 6-7 mm. long; filaments arcuate near the summit; anthers subulate, the
connective not prolonged.

(250)

1958]
FLORA OF PANAMA (Melastomataceae) 251

Southern Mexico to Panama, Bolivia, and southern Brazil.

BOCAS DEL TORO: Water Valley, von Wedel 953; Nances Cay, von Wedel 582A; Old
Bank Island, von Wedel 1915; Sansan Swamp, Almirante region, Cooper 192; Changuinola
Valley, Dunlap 1024, 378. CANAL ZONE: Chagres, Fendler 37; old Fort Lorenzo, mout
of Rio Chagres, Piper 5956; Barro Colorado Island, Woodworth 8 Vestal 607, Wilson 50,
Aviles 912; Cerro Ancón, Heriberto 134; between Mt. Hope and Santa Rita trail, Cowell
75. COCLE: Bismarck, Williams 287.

5. CLIDEMIA HIRTA (L.) D. Don, in Mem. Wern. Soc. 4:309. 1823.
Melastoma hirtum Linn. Sp. PI. 390. (Partim). 1753.

Shrub 1-3 m. tall, hirsute throughout with simple hairs; petioles 0.5-3 cm.
long; leaf-blades ovate, 5-15 cm. long, often unequal in each pair, short-acuminate,
serrulate or entire, rounded to subcordate at base, 5- to 7-nerved; flowers numerous
in loosely branched cymes 3-5 cm. long, 5-merous or more commonly 6-merous;
hypanthium campanulate, about 5 mm. long; calyx about 1 mm. long, truncate or
obscurely lobed, the exterior teeth slender, projecting about 3 mm.; petals white,
oblong-obovate, 8-11 mm. long; anthers about 5 mm. long, the connective
minutely prolonged below the filament into a dorsal spur; ovary glabrous.

uthern Mexico, throughout the West Indies and Central America, southward
in South America to Bolivia and southern Brazil; introduced as a weed in the Old
World tropics.

BOCAS DEL TORO: vicinity of Chiriqui Lagoon, von Wedel 118. CANAL ZONE: Ancón
Hill, Woodson, Allen, & Seibert 1334, Seibert 401. PANAMA: near Arraijan, Woodson,
Allen & Seibert 1336; Isla Taboga, Woodson, Allen 6 Seibert 1546.

6. CLIDEMIA STRIGILLOSA (Sw.) DC. Prodr. 3:159. 1828.
Melastoma strigillosa Sw. Prodr. Veg. Ind. Occ. 71. 1788.

Shrub 1-2 m. tall; stems densely stellate-tomentose and sparsely glandular.
Petioles pubescent like the stem, about 1 cm. long. Leaf-blades firm, ovate-
lanceolate, 8—14 cm. long, 3-6 cm. wide, acuminate, minutely serrulate, cordate or
subcordate at base, 7-nerved, densely setose above, the hairs rising from conic bases,
beneath reticulate-veined, densely stellate-tomentose, and more or less setose on the
nerves; inflorescence with a distinct central axis 3-5 cm. long; flowers 5-merous;
hypanthium hemispheric, 3 mm. long, glandular-hirsute and densely stellate-
tomentose; sepals oblong, about 2 mm. long, surpassed by the subulate exterior
teeth; petals white, 4-5 mm. long; ovary glandular-setose at the summit.

Guatemala, British Honduras, West Indies, and northern South America;
reported from Panama.

7. CLIDEMIA NOVEMNERVIA (DC.) Triana, in Trans. Linn. Soc. 28:136. 1871.
Heterotrichum novemnervium DC. Prodr. 3:173. 1828.

Shrub, the stems and petioles densely stipitate-stellate-tomentose, also with
spreading glandular hairs; petioles 1.5—4 cm. long; leaf-blades ovate to ovate-
oblong, 10-15 cm. long, about half as wide, falcately acuminate, bullate-hirsute
and sparsely stellate-pubescent above, beneath foveolate, densely stellate-tomentose;

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[Vol. 45
292 ANNALS OF THE MISSOURI BOTANICAL GARDEN

inflorescence racemose, up to 10 cm. long, with a well developed central axis and
stalked lateral branches; flowers sessile; hypanthium 4-4.5 mm. long, densely
stellate-tomentose and glandular-hirsute; sepals round-ovate, 1-1.5 mm. long and
about as wide; exterior teeth triangular, stellate on the back, barely surpassing the
sepals; petals oblong-obovate, 4.5-5 mm. long; anthers subulate, 2.5-3 mm. long;
connective very briefly prolonged below the thecae; ovary villous and glandular
at the summit.
Costa Rica and Panama.

PANAMA: exact locality lacking, Piper 5568.
8. CLIDEMIA NEGLECTA D. Don, in Mem. Wern. Soc. 4:307. 1823.

Shrub 1-2 m. tall, densely glandular pubescent, stellate-tomentose, and also
setose on the stems, petioles, inflorescence, and hypanthia; petioles 5-15 mm. long;
leaf-blades oblong-ovate, rather thick, 7—15 cm. long, about half as wide,
acuminate, serrulate, broadly rounded at base, 5-nerved, more or less rugose or
bullate above, densely pubescent on both sides, more or less stellate beneath;
hypanthium broadly tubular, densely stellate beneath the longer simple hairs; sepals
oblong-ovate, 2.5-3 mm. long, slightly exceeded by the subulate exterior teeth;
petals white or greenish, oblong-obovate, 6-7 mm. long; anthers stoutly subulate,
3.5-4 mm. long; ovary glandular-pilose at the summit.

British Honduras to Panama; Cuba; northern South America to Brazil.

Bocas DEL TORO: Big Bight, von Wedel 2885. CANAL ZONE: Old Fort Lorenzo,
mouth of Rio Chagres, Piper 5968. cocLE: hills south of El Valle de Antón, 3 2509.

PANAMA: Cerro Campana, 800 m., Allen 3988; San José Island, Andersson s. n. VINCE
UNKNOWN: Fendler 34.

9. CLIDEMIA CAPITELLATA (Bonpl.) D. Don, in Mem. Wern. Soc. 4:310. 1823.
Melastoma cabitellata Humb. & Bonpl. Melast. f. 3. 1816.
hrub 1-3 m. tall; stems thinly stellate-tomentose and pilose with simple hairs

1-2 mm. long, often more or less glandular distally; petioles 1-2 cm. long; leaf-
blades firm, ovate-oblong to ovate-lanceolate, 1-2 dm. long, 4-9 cm. wide, acu-
minate, minutely serrulate, obtuse to rounded at base, 5- to 7-nerved, hirsute above
and below; inflorescence spicate with well defined central axis, up to 15 cm. long,
its internodes conspicuously glandular and hirsute; hypanthium 3.5-4 mm. long;
short-hirsute and also thinly stellate; sepals broadly quadrate, 1-1.5 mm. long;
exterior teeth about 1 mm. longer, conic or stoutly subulate; petals white, obovate-
oblong, 4-5 mm. long; anthers stoutly subulate.

Southern Mexico to Panama and Colombia.

CANAL ZONE: Gatün, Hayes 575. DARIÉN: Cana and vicinity, Williams 738, 7 69.
10. CLIDEMIA DEPENDENS D. Don, in Mem. Wern. Soc. 4:307. 1823.

Shrub 1-3 dm. tall; stems thinly stellate-tomentose and densely hirsute “i
non-glandular hairs 2-4 mm. long; petioles densely hirsute, 5-20 mm. long; leat-
blades ovate to ovate-oblong, 7-12 cm. long, 3-7 cm. wide, abruptly short-

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1958]
FLORA OF PANAMA (Melastomataceae) 253

acuminate, entire or minutely serrulate, broadly rounded at base, 5-nerved, hirsute
above, densely pubescent beneath with both stellate and simple hairs; inflorescence
2-6 cm. long, with a well defined central axis, densely villous with non-glandular
hairs 3-6 mm. long, the glomerules nearly or quite contiguous; hypanthium
campanulate, about 3 mm. long, densely long-hirsute; sepals depressed-ovate, about
1 mm. long from the sinus, exceeded by the exterior teeth; petals white, oblong-
obovate, about 6 mm. long; anthers stoutly subulate; connective simple.

Southern Mexico to Panama and southward to Bolivia and northern Argentina;
western Cuba.

L ZONE: west slope of Ancón Hill, near Balboa, Seibert 395. COCLE: between Las
Marcaricas ad El Valle, Woodson, Allen & Seibert 1237. PROVINCE UNKNOWN: Seemann
72.

11. CLIDEMIA REITZIANA Cogn. 8 Gl. ex Gl. in Brittonia 3:115. 1939.

Shrub about 1 m. tall, the younger stems densely or sparsely hirsute with simple
hairs projecting from a dense furfuraceous tomentum; petioles 1-3 cm. long,
pubescent like the stem; leaf-blades ovate, 8-17 cm. long, about half as wide,
short-acuminate, broadly obtuse to rounded at base, 5-nerved, sparsely hirsute
above, beneath densely pilose on the surface and also furfuraceous on the veins;
hypanthium densely hirsute and also minutely stellate; calyx spreading, the minute
sepals depressed-triangular, the exterior teeth slightly shorter; petals quadrate,
about 1.5 mm. long; anthers stoutly subulate, the connective prolonged into an
oblong dorsal spur about 0.4 mm. long.

Costa Rica and Panama.

AS DEL TORO: Fish Creek Mountains, von Wedel 2307. DARIEN: Cana-Cuasi trail,
Tys) & Terry 1460.

12. CLIDEMIA DEPPEANA Steud. Nom. Bot. ed. II. 1:384. 1841.

Shrub 1-2 m. tall; stems loosely hirsute with ascending or somewhat appressed
hairs. Petioles hirsute, about a third as long as the blade. Leaf-blades ovate-
oblong to obovate-oblong, 6-10 cm. long, about half as wide, acute or acuminate,
distinctly serrulate, ciliate, obtuse or rounded at base, 3-nerved, densely pilose on
both sides, often with purplish hairs; cymes short-peduncled, branched from near
the base, 2-4 cm. long, hirsute; hypanthium urceolate to tubular, 3-3.5 mm. long,
densely hirsute, some of the hairs occasionally gland-tipped; sepals ovate, obtuse,
about 1 mm. long; exterior teeth slenderly subulate, 1.5-2.5 mm. long; >
white, obovate-oblong, about 4 mm. long; anthers stoutly subulate, about 2 mm.
long, notched at the apex, the connective minutely prolonged at base.

uthern Mexico to Panama, at low or moderate altitudes.
OCAS DEL TORO: Isla Colón, tor Sa 2982; Water Valley, von Wedel 604, 947;
vicinity of Almirante, Cooper 108, I
13. CLIDEMIA COLLINA Gl. in Phytologia 3:359. 1950.
Shrub to 2 m. tall, the stem densely setose with stiff reflexed hairs up to 6 mm.

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[Vol. 45
254 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long; petioles 1-2 cm. long, almost completely occupied by a densely setose formi-
carium; leaf-blades obovate-oblong, up to 2 dm. long, sparsely setose above, densely
hirsute on the veins beneath, almost glabrous on the surface, 7-pli-nerved; flowers
5-merous, in a densely setose cluster; hypanthium campanulate, setose with spread-
ing hairs up to 10 mm. long; sepals oblong, obtuse, nearly 2 mm. long from the
torus; exterior teeth slender, sparsely long-setose, surpassing the sepals by 2 mm.;
petals white, 5-6 mm. long.

Endemic to Panama, so far as known.

cocLÉ: vicinity of El Valle, 800-1000 m., Allen 1820.
14. CLIDEMIA TAURINA Gl. in Phytologia 3:360. 1950.

Shrub to 3 m. tall, the stem densely setose with stiffish reflexed hairs 4-7 mm.
long; petioles similarly setose, 1-3 cm. long, the distal half occupied by a didymous
long-setose formicarium; leaf-blades oblong-ovate, up to 25 cm. long and three-
fifths as wide, 5- to 7-nerved, hirsutulous above, densely hirsute over the whole
lower side; inflorescence compact, few-flowered; flowers 5-merous; setae of the
hypanthium and exterior teeth rarely more than 3 mm. long; sepals oblong, obtuse,
much exceeded by the slender exterior teeth.

Endemic to Panama, so far as known. This and the preceding species are
closely related to C. reflexa Gl. and C. spectabilis Gl. of Costa Rica and to
myrmecina Gl. of Colombia. A re-alignment of species may be necessary when
more and better material is available.

BOCAS DEL TORO: Water Valley, von Wedel 703, 925; Old Bank Island, von Wedel
1917.

15. CLIDEMIA TOCOCOIDEA (DC.) Gl. in Bull. Torr. Bot. Club 58:81. 1931.
Calophysa tococoidea DC. Prodr. 3:166. 182
Maieta tococoidea Cogn. in Mart. Fl. Bras. ae :465. 1888.

Small shrub, long-hirsute throughout; petioles 1-4 cm. long, bearing formicaria
at the base; leaf-blades thin, ovate or ovate-oblong, 1-2 dm. long, 7-12 cm. wide,
more or less unequal in each pair, abruptly acuminate, finely serrulate, ciliate,
rounded to subcordate at base, 5- to 7-nerved, thinly pilose on both sides; cymes
small, compact, few-flowered, sessile or nearly so in the upper axils; hypanthium
cylindric, 3-3.5 mm. long, sparsely hirsute; sepals minute; exterior teeth widely
spreading, stout, about 1 mm. long; petals 4, white, about 2 mm. long; anthers
linear.

British Honduras to Panama and the upper Amazon Valley.

OCAS DEL TORO: vicinity of Almirante, Cooper 480. DARIEN: Cana-Cuasi rail
res 8 Terry 1514.
16. CLIDEMIA SETOSA (Triana) Gl. in Bull. Torr. Bot. Club 58:82. 1931.
Calophysa setosa Triana, in Journ. Bot. 5:2 867.
Maieta setosa (Triana) Cogn. in Mart. Fl. R, ią :462. 1888.
Low shrub, the stems and petioles densely covered with stiff retrorse setae 7-10

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1958]
FLORA OF PANAMA (Melastomataceae) 255

mm. long; petioles 2-5 cm. long, bearing the formicarium at the summit; leaf-
blades ovate or ovate-oblong, 1-2 dm. long, about half as wide, short-acuminate,
entire or serrulate, ciliate, rounded to subcordate at base, 5- to 7-nerved, long-
hirsute above, densely setose near the base beneath; inflorescence long-peduncled,
trichotomous, 3-8 cm. long; hypanthium 2-2.5 mm. long, 4-angled, minutely
puberulent, usually sparsely setose near the summit; exterior teeth subulate, about
1.5 mm. long; petals 4, about 5 mm. long; anthers linear.
Southern Mexico to Panama.

cocLE: El Valle de Antón, near Cerro Turrega, 700 m., Woodson ES Schery 200.

17. CLIDEMIA EPIPHYTICA (Triana) Cogn. in DC. Monogr. Phan. 7:1025. 1891.
Sagraea epiphytica Triana, in Trans. Linn. Soc. 28:139. 1871
Prosanerpis banamensis Blake & Standley, in Field Mus. Pub. Bot. 4:245. 1929.

A root-climbing or epiphytic shrub; stems nearly glabrous; petioles of the
larger leaves 4—10 cm. long; leaf-blades thin, ovate or ovate-oblong, 12-20 cm.
long, about two-thirds as wide, sharply acuminate, conspicuously serrate, cordate
at base, 7-nerved, essentially glabrous; smaller leaves 1-5 cm. long; cymes usually
3-flowered, branched at base; hypanthium urceolate, 2-2.5 mm. long, bristly;
sepals triangular-ovate about 1 mm. long from the torus; exterior teeth foliaceous,
oblong, up to 5 mm. long, long-ciliate.

Panama and at low elevations in northwestern South America.

DARIEN: Cerro de Garagará, 500—974 m., Pittier 5611.

18. CLIDEMIA RUBRA (Aubl) Mart. Nov. Gen. & Spec. 3:152. f. 287. 1829.
Melastoma rubra Aubl. Pl. Guian. 1:416. 1775.

Suffrutescent or woody, up to 2 m. tall, often blooming when young and
herbaceous, the stem and foliage densely hirsute or villous throughout; petioles
none, or up to 3 cm. long, usually manifest but short; leaf-blades thick, oblong-
lanceolate to broadly elliptic, commonly 6—12 cm. long and about half as wide,
Obtuse to short-acuminate, at base cuneate to subcordate, 5-pli-nerved; flowers
few, sessile in dense axillary glomerules, 4-merous; hypanthium cylindric, 3-4 mm.
long; exterior teeth triangular below, narrowed to a short, divergent, subulate tip,
equaling or slightly exceeding and more or less concealing the sepals; petals obovate-

oblong, 2-3 mm. long, white or pinkish; anthers subulate, 3-4 mm. long.

Tropical Mexico to Panama, chiefly at low altitude; very abundant in the
lowlands of South America as far south as southern Brazil and Bolivia.

ONE: Ancón Hill, Williams 1, ru d i Pe 5558. CHIRI 0: I uete,
Davidson “676, Woodson & Schery 784. cocr y of Natà, 50 m., Allen ru
Vicinity of Penonomé, Williems 92; between Oli "y dun, Pittier 5097.
San sand Island, Johnston 34, Erlanson 382, 428; Taboga Island, Macbride łaa:
VERAGUAS: between Cafiazas and the Cordillera Central, 300-600 m., Allen 165.

19. CLIDEMIA DENSIFLORA (Standley) Gl. in Phytologia 3:346. 1950.

Henriettella densiflora Standley, in Field Mus. Pub. Bot. 4:247. 1929.

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[Vol. 45
256 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Shrub 1-3 m. tall, the younger stems and petioles densely furfuraceous-
tomentose; petioles 1-3 mm. long; leaf-blades thin, narrowly elliptic to oblong,
12-20 cm. long, about a third as wide, acuminate, gradually narrowed to the base,
3-nerved, glabrous and shining above, beneath glabrous on the surface, densely
furfuraceous on the primary nerves; flowers few, crowded and sessile in small axil-
lary clusters; hypanthium cup-shaped 2—2.5 mm. long; sepals triangular; exterior
teeth minute; petals oblong, 2.8 mm. long; connective of the anthers prolonged
at base into a minute, glandular, dorsal appendage.

British Honduras and Panama.

AS DEL TORO: Almirante region, Cooper 575. DARIEN: Cana-Cuasi Trail, 600 m.,
dy. 3 Terry 1424.

20. CLIDEMIA PURPUREO-VIOLACEA Cogn. in Bull. Soc. Bot. Belg. 30':263. 1891;
in Dur. & Pitt. Fl. Cost. 166. 1891.

Shrub 1-5 m. tall, the stems, petioles, lower leaf-surface, and hypanthia appar-
ently glabrous, actually very minutely furfuraceous-puberulent; petioles 2-8 cm.
long; leaf-blades thin, bright green above, purple beneath, broadly ovate, 12-22
cm. long, 6-13 cm. wide, acuminate, minutely ciliate-denticulate, cordate or sub-
cordate at base, 5- to 7-nerved; inflorescences mostly below the leaves, about 1 cm.
long, few-flowered; hypanthium urceolate, 2-2.5 mm. long; sepals depressed-
triangular, about 0.5 mm. long; exterior teeth ending in a deflexed seta about 0.5
mm. long; petals oblong, about 2 mm. long; anthers obovate, 2 mm. long.

Costa Rica and Panama.

ANAL ZONE: Chagres, Fendler 38. RIEN: Cana and Maid about 750
Williams 956; Ensenada Gosia. Stern 8 Cheer 182. san BLas: San Blas district
Cooper 235.
21. CLIDEMIA OMBROPHILA Gl. in Brittonia 3:138. 1939.

Stems slender, essentially glabrous; petioles 8—12 mm. long; leaf-blades firm,
shining, elliptic, 6-9 cm. long, 2.5-4 cm. wide, abruptly contracted at the apex
into a linear tip 2-3 cm. long, entire, cuneate at base, 3-pli-nerved, essentially
glabrous; panicle sessile, trichotomous from the base, its divaricate branches 3-5
cm. long, each bearing 3—5 short-pediceled flowers; hypanthium subglobose, about
2 mm. long, very minutely lepidote; sepals triangular, subacute, 0.9 mm. tong;
exterior teeth adnate and exactly equaling the sepals.

Known only from the type collection.

DARIÉN: foothills of Garagará, Pittier 5610.
23. HENRIETTEA DC.

HENRIETTEA DC. Prodr. 3:178. 1828.

Flowers 5- to 6-merous. Hypanthium campanulate, thick-walled. a
tube distinctly prolonged; sepals small or large, ovate or triangular or pe
obsolete; exterior teeth minute or concealed. Stamens isomorphic; filamen

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1958]
FLORA OF PANAMA (Melastomataceae) z”

>> ud
AAU
NU 8
SS

Fig. 84. Henriettea succosa

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[Vol. 45
258 ANNALS OF THE MISSOURI BOTANICAL GARDEN

slender, glabrous; anthers subulate or linear, often beaked; connective neither
appendaged nor prolonged. Ovary inferior, 5- to 6-celled; style stout, enlarged
toward the summit; stigma punctiform or truncate. Fruit a many-seeded berry.
Trees or shrubs with large, usually pli-nerved leaves and medium-sized flowers borne
in small axillary clusters below the existing leaves.

About 16 species, mostly in Amazonia but extending through the West Indies
and Central America, usually at low altitudes.

1. HENRIETTEA succosa (Aubl.) DC. Prodr. 3:178. 1828.
Melastoma succosa Aubl. PI. Guian. 1:418. 1775.

Small tree to 10 m. tall, the younger stems and petioles densely strigose; petioles
stout, 5-15 mm. long; leaf-blades firm in texture, narrowly obovate, up to 2 dm.
long, a third or nearly half as wide, abruptly acuminate or apiculate, entire, cuneate
at base, 3-pli-nerved, scabrously pilose above, beneath strigose on the veins, stellate-
pubescent and rather canescent on the surface; flowers few, short-pediceled; hypan-
thium densely strigose, 8-9 mm. long; sepals ovate, 4-5 mm. long; petals white
or pink.

Central America, Trinidad, the Guianas, and eastern Brazil.

Bocas DEL TORO: Old Bank Island, vicinity of Chiriqui Lagoon, von Wedel 2113.

CANAL ZONE: Barro Colorado Island, Shattuck 1093. PANAMA: San José Island, Jobn-
ston 740, Erlanson 340, 502.

24. BELLUCIA Neck.
BELLUCIA Neck. Elem. Bot. 2:142. 1790.

Flowers 5- to 8-merous. Hypanthium thick-walled, hemispheric. Calyx large,
at anthesis divided into 2-8 regular or irregular lobes. Petals large, leathery,
oblong or obovate. Stamens isomorphic; filaments short and stout; anthers more
or less coherent in a ring, laterally compressed, broadly oblong, elliptic, or dolabri-
form, opening by two terminal pores; connective neither prolonged nor appendaged.
Ovary inferior, 5- to 15-celled; style stout; stigma large, capitate. Fruit a large
many-seeded berry. Trees or tall shrubs, with large ovate or oblong leaves and
large white or pink flowers on short pedicels solitary or few in the leaf-axils.

About 10 species of low altitudes, most numerous in northern South America.
B. grossularioides (L). Triana is widely distributed and has been reported from
Panama: its calyx exhibits two to five lobes of irregular shape and size at anthesis.

1. BELLUCIA COSTARICENSIS Cogn. in Bull. Soc. Bot. Belg. 301:264. 1821. in
Dur. & Pitt. Fl. Cost. 167. 1891.

Small tree, glabrous throughout; petioles stout, 2—4 cm. long; leaf-blades firm,
broadly elliptic, commonly 1.5—3 dm. long and about two-thirds as wide, obtuse
or short-acuminate, entire, obtuse or rounded at base, 5-pli-nerved, the outer pair
submarginal; cymes axillary, sessile, trichotomous from the base, 3- to 7-flowered;
pedicels about 1 cm. long; hypanthium hemispheric; calyx spreading, nearly 2 cm.
wide; petals about 15 mm. long.

(258)

19581

259

FLORA OF PANAMA (Melastomataceae)

Fig. 85. Bellucia costaricensis

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[Vol. 45
260 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Costa Rica and Panama.

CANAL ZONE: near Ferry along Thatcher Highway, White & White 140; Barro
Colorado Island, Shattuck III9. cHIRIQUI: vicinity of Remedios, 0-150 m., Allen 3669.
DARIEN: Cana and vicinity, 750 m., Williams 808. PROVINCE UNKNOWN: Pittier 5450.

25. LOREYA DCE.
LoREYA DC. 3:178. 1828.

Flowers 5- to 6-merous. Hypanthium cup-shaped to broadly campanulate.
Calyx more or less spreading, its lobes shorter than the tube or lacking.
obtuse, oblong-obovate or obovate. Stamens isomorphic; filaments short; anthers
often coherent in a ring, oblong or ovoid, blunt, opening by a minute terminal
pore; connective simple at base, often thickened or elevated along the back of the
anther. Ovary wholly inferior, 3- to 6-celled; style elongate; stigma capitate.
Trees, often of large size, with broad, usually pli-nerved leaves and small or
medium-sized flowers in axillary cymes or fascicles from the leafless nodes.

About 10 species, the others all in lowland forests of Amazonia.

1. LOREYA BRUNNESCENS (Standley) Gl. in Phytologia 3:346. 1950.
Henriettea brunnescens Standley, in Field Mus. Publ. Bot. 4:247. 1929.

Small tree to 8 m. tall, the younger stems, petioles, pedicels, and hypanthia
densely brown-hirsute; petioles 3-5 cm. long; leaf-blades rhombic-elliptic, up to
2.5 dm. long, about half as wide, abruptly acuminate, obtuse at base, 5-pli-nerved
with an additional pair of marginal nerves, the inner pair arising 3—5 cm. from the
base, above glabrous on the surface, strigose on the veins, beneath roughly pubes-
cent; flowers several in axillary clusters, on pedicels about 2 cm. long; hypanthium
deeply cup-shaped, about 8 mm. long; sepals triangular, about 2 mm. long; petals
and stamens unknown.

Known only from the type collection.

BOCAS DEL TORO: Buena Vista Camp, Chiriquí Trail, 375 m., Cooper 594.
26. HENRIETTELLA Naud.

HENRIETTELLA Naud. in Ann. Sci. Nat. Bot. III. 18:107. 1852.

Flowers 4- to 5-merous. Hypanthium small, globose to cup-shaped. Sepals
small, triangular or ovate to depressed or obsolete; exterior teeth usually present,
conic or subulate, small. Petals ovate to lanceolate or triangular, usually acute "
subacute, commonly cucullate at the summit and often exceeded by a minute
exterior tooth. Stamens isomorphic; filaments slender, glabrous; anthers oblong»
usually obtuse, opening by a large terminal pore; connective simply or b
prolonged below the thecae, rarely with minute basal lobes. Ovary inferior, uos
5-celled; style slender; stigma truncate to capitate. Fruit a many-seeded PE
Shrubs or trees with small cluster of flowers at the leafless nodes or rarely in che
axils of the leaves.

(260)

1958]
FLORA OF PANAMA (Melastomataceae) 261

o = br => Ex == ui
w A =
ei. S

Fig. 86. Henrietella seemanii

(261)

[Vol. 45
262 ANNALS OF THE MISSOURI BOTANICAL GARDEN

About 40 described species, chiefly in the West Indies and northern South

America.
a. Leaves long-tapering at base, strongly pli-nerved 1. H. TUBERCULOSA.
aa. Leaves rounded or obtuse at base, 3-nerved or obscurely pli-nerved.
b. Pedicels about 1 mm. long; hypanthium densely pubescent................... 2. H. SEEMANNII.
bb. Pedicels 4-8 mm. long; hypanthium essentially glabrous.....................-- 3. H. FLAVESCENS.

1. HENRIETTELLA TUBERCULOSA Donn. Sm. in Bot. Gaz. 27:335. 1899.

Shrub or tree up to 8 m. tall; younger stems and petioles densely strigose; leaf-
blades oblanceolate, up to 15 cm. long and about a third as wide, slenderly
acuminate, long-tapering at base, strongly 5-pli-nerved, scabrous above with short
stiff hairs from conic bases, densely pubescent beneath; flowers subsessile;
hypanthium densely strigose, about 2 mm. long; sepals minute, triangular; petals
white, about 2 mm. long, the blade ovate, subacute.

Costa Rica and Panama.

BOCAS DEL TORO: Old Bank Island, Isla Colón, 0-120 m., von Wedel 1933.
2. HENRIETTELLA SEEMANNII Naud. in Ann. Sci. Nat. Bot. III. 18:108. 1852.

Small tree, the younger stems and petioles very densely pubescent with ascend-
ing hairs 1-1.5 mm. long; petioles 8-15 mm. long; leaf-blades oblong-lanceolate,
up to 13 cm. long, about a third as wide, acute, ciliate, rounded at base, 3-nerved,
scabrous above, pilose beneath; flowers several in a cluster, 5-merous, on pe icels
about 1 mm. long; hypanthium about 3 mm. long, densely pubescent with ascend-
ing hairs from swollen bases; sepals depressed-triangular, about 0.4 mm. long; petals
lanceolate, nearly 5 mm. long, acuminate, irregular on the margins.

Costa Rica and Panama to northern Colombia.

CANAL ZONE: near Las Cruces, Seemann s. n.

3. HENRIETTELLA FASCICULARIS (Sw.) Triana, in Trans. Linn. Soc. 28:14).

1871.

Melastoma fascicularis Sw. Prodr. Veg. Ind. Occ. 71. 1788.

Small tree, up to 10 m. tall, with densely hirsute stems and petioles; petioles
stout, 6-12 mm. long; leaf-blades elliptic, oblong, or oblong-obovate, 8-15 cm.
long, about half as wide, acute, obtuse or subacute at base, weakly 5-pli-nerved
with an additional pair of marginal nerves, scabrously pubescent on the surface
and hirsute on the nerves above, densely pubescent beneath; pedicels slender,
mm. long; hypanthium campanulate, 3 mm. long, essentially glabrous; S€
minute, depressed-ovate; petals white, ovate, acute, about 4 mm. long.

British Honduras, Panama, Guatemala, and the West Indies.

DARIÉN: Cana and vicinity, Williams 753.

27. OSSAEA DC.
OssaEA DC. Prodr. 5:168. 1828.

Flowers 4- to 5-merous. Hypanthium urceolate or subglobose to campanulate

(262)

1958]
FLORA OF PANAMA (Melastomataceae) 263

Fig. 87. Ossaea micrantha

or nearly tubular. Calyx-tube often conspicuously prolonged; sepals minute or
obsolete; exterior teeth varying from minute tubercles to long subulate appendages
often simulating a sepal. Petals small, lanceolate to ovate, acute or acuminate,
usually with an infra-apical exterior tooth. Stamens isomorphic, erect; filaments
slender; anthers linear to oblong; connective simple or very briefly prolonged below
the thecae and in a few species continued below the filament into a minute dorsal
appendage. Ovary inferior, 3- to 5-celled; style filiform; stigma punctiform or
capitellate. Fruit a many-seeded berry, often terminated by the enlarged calyx-
tube. Small trees or shrubs with white flowers in fascicles or paniculiform cymes
in the axils of existing leaves.

About 90 species, distributed from southern Mexico to Bolivia and southern
Brazil but probably most numerous in the West Indies.

4-merous; fruit 8-ribbed; leaves acute to obtuse at base........... 1. O. MICRANTHA.
bb. Flowers 5-merous; fruit obscurely 10-ribbed; leaves typically cuneate
aa. FI at base and m decurrent along the petiole........------------------ 2 O. DIVERSIFOLIA.
a. Flowers subsessile, few, closely crowded in clusters up to 1.5 cm. in
ameter; fruiting hypanthium ribless...........— m0 3. O. TRICHOCALYX.

[Vol. 45
264 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1. OssAEA MICRANTHA (Sw.) Macfadyen, Fl. Jamaic. 2:49. 1850.
Melastoma micrantba Sw. Prodr. Veg. Ind. Occ. 71. 1788.

Shrub or small tree to 10 m. tall; young stems, petioles, and inflorescence nearly
or quite glabrous; petioles slender, 1.5-3 cm. long; leaf-blades thin, narrowly
oblong-elliptic, 10-17 cm. long, about a third as wide, long-acuminate, entire or
nearly so, broadly cuneate to the base but not decurrent, 3-pli-nerved with an
additional pair of marginal nerves, the lateral nerves often alternate, essentially
glabrous on both sides; panicles loosely branched, 2-5 cm. long; flowers 4-merous;
hypanthium subglobose, about 2 mm. long; exterior teeth minute tubercles or
lacking; petals narrowly lanceolate, 3-4 mm. long; fruiting hypanthium with 8
rounded elevated longitudinal ribs.

Southern Mexico to Panama and Peru; Jamaica.

BOCAS DEL TORO: Cricamola Valley, region of Almirante, Cooper 478; Fish Creek
Mountains, von Wedel 2354; locality not stated, von Wedel 192; Changuinola Valley,

Cooper & Slater 115. cHmiQui: Quebrada Velo, 1800 m., Woodson & Schery 267; Bajo
Chorro, Boquete District, 1800 m., Davidson 105.

2. OssaEA DIVERSIFOLIA (Bonpl.) Cogn. in Bull. Acad. Belg. III. 14:968. 1887.
Melastoma diversifolia Humb. & Bonpl. Melast. 138. f. 59. 1816.

Shrub to 1.5 m. tall, the younger stems, petioles, and inflorescence densely but
finely stellate-furfuraceous; petioles 1.5—5 cm. long; leaf-blades thin, ovate to
ovate-lanceolate, 8-20 cm. long, half to two-thirds as wide, abruptly acuminate or
apiculate, ciliate, often conspicuously dentate, abruptly narrowed at base and de-
current along the petiole almost to the stem, 5- to 7-pli-nerved, sparsely pilose
above, glabrous beneath or furfuraceous on the veins; inflorescence loosely branched,
2-5 cm. long; flowers 5-merous; hypanthium subglobose, about 2 mm. long, in
fruit shallowly 5-sulcate and obscurely ribbed; petals lanceolate.

Costa Rica, Panama, and northwestern South America.

Bocas DEL TORO: Lincoln Creek, Carleton 87; region of Almirante, Cooper an.
CANAL ZONE: Barro Colorado Island, Chrysler & Roever 4819, 4870; Gatun, Hayes 17 ;
between Juan Mina and El Vigía, Pittier 2401; vicinity of Madden Dam, Allen 2009;
Ahorca Lagarto, Cowell 250. cHIRIQUI: vicinity of Puerto Armuelles, Woodson 8 Sc

É Sabana, Leopold

$55. DARIÉN: headwaters of Río Chico, 150-225 m., Allen 4612; Río Sa
III 189. PROVINCE UNKNOWN: Kuntze 194.

3. OSSAEA TRICHOCALYX Pittier, in Journ. Wash. Acad. Sci. 13:391. 1923.

Shrub or small tree to 6 m. tall, the younger stamens and petioles very finely
furfuraceous; petioles 1-5 cm. long; leaf-blades thin, ovate-lanceolate to oblong,
10-22 cm. long, 4-10 cm. wide, long-acuminate, entire, abruptly narrowed below
to a long cuneate base, glabrous above, essentially glabrous beneath, 5- to 7-P li-
nerved, the inner pair arising 2-5 cm. from the base; flowers 4-merous, su
or very short-pediceled, densely crowded in clusters 10-15 mm. wide at a
nodes; hypanthium urceolate, 2.5 mm. long, essentially glabrous; exterior te
subulate, spreading, about 1.5 mm. long, glandular-setose; petals 1.6 mm. long.

British Honduras to Ecuador, chiefly at low altitudes.

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1958]
FLORA OF PANAMA (Melastomataceae) 265

BOCAS DEL TORO: vicinity of Chiriqui Lagoon, von Wedel 2197A, 2911; Water Valley,
von Wedel 1462. DARIEN: Cana-Cuasi Trail, 900 m., Terry & Terry 1450; above Paca,
Williams 742.

28. LEANDRA Raddi

LEANDRA Raddi, in Mem. Mod. 18 Fis.: 385. 1820.

Flowers 4- to 8-merous. Hypanthium globose to tubular, often constricted
at the mouth. Sepals small or obsolete; exterior teeth present, minute to elongate.
Petals ovate to narrowly lanceolate, rarely obovate, acute to acuminate, the covered
side often much the wider, usually erect at anthesis. Stamens nearly or quite
isomorphic; filaments slender, smooth; anthers linear to oblong or rarely subulate,
2-celled or 4-celled; connective simple or rarely slightly prolonged at base, never
appendaged, often gibbous on the back near the base. Ovary mostly or wholly
inferior, its cells often isomerous with the petals; style slender; stigma capitellate
to punctiform. Fruit a many-seeded berry. Shrubs or small trees, almost always
pubescent, often hirsute or hispid, with terminal panicles or bracteate heads of
small, usually white flowers.

More than 200 species, extending from Mexico to Bolivia and southern Brazil,
avoiding the West Indies.

a. Pubescence of younger parts of the stem and axis of the inflorescence of

short, spreading, Pun le hairs surpassed by projecting, stouter, glandular
1. L. MEXICANA.

hai rs; leaves 7-nerv
aa. Pubescence of en stem never pere
b. In rn of the younge of t and the axis of the
i e a dense mass of strongly baia Mas: commonly with
Polexciut porta bristles; leaves all or mostly 5-nerved.
c. Leaves sparsely hirsute on che surface Ate th with straight or
curved s simp pe bristles (commonly more or less stellate on the
principal veins)... I o Pia AREA eie © —
cc. OH ste ate pesce on the surface beneath L. SUBSE
bb. Indument of the stem and axis of the Acción composed entirely
of pers lites.
c. Hairs of the stem more or less retrorse, at least at their bases............. 4. L. DICHOTOMA.
cc. Hairs of the stem merda distinctly appressed or ascending.
d. Flowers 5-merous; exterior teeth none or very short and
InCcODSPICHOUS. 53, il; aliis .... 5, L. STRIGOSA.
Flowers Amero exterior ig subulate, 5-6 mm. long............. 6. L. SUBULATA.
bbb. dei of the younger parts of the stem and the axis of the
inflorescence edt uec rer leaves strongly 5-pli-nerved. 7. L. CONSIMILIS.
1886.

l. LEANDRA MEXICANA (Naud.) Cogn. in Mart. Fl. Bras. 14*:77.
Clidemiastrum mexicanum Naud. in Ann. Sci. Nat. Bot. III. 17:87. 1852.

Shrub 1—3 m. tall, the stems pubescent with slender simple hairs slightly sur-
Passed by glandular ones; petioles 2-6 cm. long, similarly pubescent; leaf-blades
thin, ovate, 1-2 dm. long, 4—15 cm. wide, acuminate, serrulate, broadly rounded
to subcordate at base, 7-nerved, hirsutulous above and also sparsely pilose with
longer stiff hairs; panicle 6-10 cm. long, its branches eventually greatly elongate,
divaricate, and more or less flexuous; flowers commonly 7-merous; hypanthium
broadly tubular, 3—3.5 mm. long, densely villosulous with simple hairs and sparsely

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[Vol. 45
266 ANNALS OF THE MISSOURI BOTANICAL GARDEN

glandular-hirsutulous, the hairs up to 0.5 mm. long; petals about 3 mm. long;
sepals depressed-ovate, 1—1.5 mm. long.
Southern Mexico to Panama, usually at low altitudes.

AS DEL TORO: Water Valley, von Wedel 1903, 1913; Little Bocas, von Wedel 2493;

o
Big Bight, von Wedel 2890. cHIRIQUI: San Félix, Pittier 52 Colón

i OI. CAN
Kuntze; Monkey Hill, Cowell 43. PANAMÁ: San José bias pa 461.
2. LEANDRA MELANODESMA (Naud.) Cogn. in Mart. Fl. Bras. 14*:115. 1886.
Clidemia melanodesma Naud. in Ann. Sci. Nat. Bot. III. 17:353. 1852.

Shrub or small tree up to 5 m. tall; younger stems, petioles, and inflorescence
densely covered with a mass of strongly barbellate or plumose hairs surpassed by
projecting simple bristles; petioles slender, up to 4 cm. long; leaf-blades ovate-
lanceolate or oblong-ovate, 8-15 cm. long, 2-7 cm. wide, acuminate, entire, obtuse
to rounded at base, 5-nerved, very sparsely and minutely hispidulous above, sparsely
hirsute beneath with simple bristles, the barbellate pubescence of the petiole con-
tinuing along the primary nerves and to some extent along the secondaries; panicle
slender with ascending branches, 5-15 cm. long; flowers 5-merous; hypanthium
subglobose, minutely stellate and hirsutulous; sepals depressed-triangular, very
short, the exterior teeth minute; petals white, about 3 mm. long.

In mountainous regions, southern Mexico to Colombia.

iqui: Finca Lérida to Peña Blanca, Woodson & Schery 309; vicinity of Finca
pinto 1750 m,, Woodson 6 Schery 213; vicinity of Casita Alta, 1500-2000 m., Woodson,

» Seibert 253, White 32; valley of the upper Rio Chiriqui Viejo, W bite 23; Boquete,
Davidson 857; Volcán de Chiriquí, Davidson 919.

3. LEANDRA SUBSERIATA (Naud.) Cogn. in Mart. Fl. Bras. 144:73. 1886.
Clidemia subseriata Naud. in Ann. Sci. Nat. Bot. III. 17:354. 1852.

Very like L. melanodesma in all features except the pubescence; lower surface
of the leaves sparsely pubescent with short-stipitate stellate hairs; hypanthium often
stellate only without simple bristles.

Southern Mexico to Panama; also in Venezuela, Colombia, and Ecuador.

CHIRIQUÍ: vicinity of New Switzerland, 1800-2000 m., Allen 1343.

4. LEANDRA DICHOTOMA (D. Don) Cogn. in Mart. Fl. Bras. 14%:200. 1886.
Clidemia dichotoma D. Don, in Mem. Wern. Soc. 4:307. 1823.

Shrub 1-3 m. tall; younger stems, petioles, and axis of the inflorescence densely
hirsute with straight or curved, distinctly retrorse or retrorsely outwardly curved,
simple bristles; petioles 2-5 cm. long; leaf-blades ovate-lanceolate to ovate-oblong
or oblong, 1-2 . long, 5-12 cm. wide, acuminate, denticulate and strongly
ciliate, broadly obtuse to rounded at base, commonly 7-nerved, more or less hirsute
on both sides, especially on the nerves beneath; panicles 1-2 dm. long, the branches
becoming widely divaricate and crooked, with secund 5-merous flowers; hypan-
thium subglobose, densely hirsute; sepals minute, depressed-triangular, the exterior
teeth subulate, about 1 mm. long; petals white, 2.5-3 mm. long.

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1958]
FLORA OF PANAMA (Melastomataceae) 267

V a =
DOM ` — Th
Birr ES

à ORNE
DPS
iv

SRA
A J AX u hy
x = ^
` y
i 4 €

Leandra melanodesma

Fig. 88.

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[Vol. 45
268 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Guatemala and British Honduras to Panama and Bolivia.

BOCAS DEL TORO: Water Valley, von Wedel 1419; Old Bank Island, von Wedel 1971;
vicinity of Chiriqui Lagoon, von Wedel 1075; Fish Creek Mountains, von Wedel 2370;
Cricamola, Almirante region, Cooper 514. CANAL ZONE: Barro Colorado Island, Aviles
913, Bailey & Bailey 470.

5. LEANDRA STRIGOSA Gl. in Phytologia 3:345. 1950.

Shrub up to 2 m. tall; younger stems, petioles, and axis of the inflorescence
densely strigose with appressed or closely ascending hairs 0.5—1.5 mm. long; petioles
1-3 cm. long; leaf-blades ovate-lanceolate or ovate, up to 15 cm. long and 10 cm.
wide, acuminate, denticulate, densely ciliate, rounded at base, 5-nerved, densely
pubescent on both sides; panicle 6-20 cm. long; flowers 5-merous, sessile, secund;
hypanthium cup-shaped, densely strigose, some of the hairs gland-tipped; sepals
triangular, about 0.5 mm. long; exterior teeth none; petals white, about 2.3 mm.
long.

Panama and Colombia.

cHIRIQUÍ: between Río Tinta and Río Tabasará, Allen & Seibert 415. COLÓN: vicin-
ity of Camp Pina, 25 m., Allen 3672.

6. LEANDRA SUBULATA Gl. in Phytologia 3:345. 1950.

Epiphytic shrub up to 2 m. tall; stem, petioles, inflorescence, and hypanthia
densely covered with straight, appressed, spinulose hairs 1—2 mm. long; leaves
long-petioled, the blades ovate, 2-3.5 cm. long, about half as wide, acute, spinulose-
ciliate, rounded at base, obscurely 5-nerved or weakly 7-pli-nerved, strigillose on
both sides; flowers 4-merous, in terminal cymules; hypanthium obconic; sepals
very short, acute, triangular, the exterior teeth subulate, erect, 5-6 mm. long;
petals ovate, pink, 4 mm. long.

Known only from the type specimen.

BOCAS DEL TORO: northern slopes of Cerro Horqueta, 1800-2100 m., Allen 4804.
7. LEANDRA consimilis Gleason, spec. nov.

Arbuscula, caulibus 4-sulcatis, petiolis, et ramis inflorescentiae minute pulveru-
lento; folia petiolata, anguste elliptica, utrinque acuminata, valde 5-plinervia,
supra glabra, subtus ad venas pulverulenta caeterum glabra.

Small tree 5 m. tall, the younger parts of the 4-sulcate stems, petioles,
branches of the inflorescence minutely brown-pulverulent; petioles about 1 =
long; leaf-blades thin, narrowly elliptic, up to 2 dm. long, about a third as wide,
equally and gradually tapering to both ends, strongly 5-pli-nerved, pulverulent on
the veins beneath, otherwise glabrous; panicle widely branched, about 1 dm. long;
flowers 5-merous; hypanthium 1.4 mm. long, nearly glabrous; calyx about 0.8 mm
long, the acute triangular lobes about equaling the tube, barely exceeded by the
minute, subulate, exterior teeth; petals oblong-lanceolate, about 3.5 mm. long;
anthers short, oblong; style about 4 mm. long; stigma rounded, not dilated.

Known only from the type specimen.

and

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1958]
FLORA OF PANAMA (Melastomataceae) 269

PANAMA: Las Minas, Allen 2702. (Herb. New York Bot. Gard., HOLOTYPE).
The plant exhibits a striking superficial similarity in leaves, pubescence, and
inflorescence to Conostegia micromeris Standl.

29. CONOSTEGIA D. Don.

CONOSTEGIA D. Don, in Mem. Wern. Soc. 4:316. 1823.

Flowers 5- to 10-merous. Calyx calyptrate, deciduous at or near the torus in
anthesis, the flower-buds globose to pyriform or obovoid. Petals white or pink,
often very inequilateral or conspicuously retuse. Stamens isomorphic, two to five
times as many as the petals; filaments slender, glabrous; anthers linear or oblong,
4-celled, more or less laterally compressed, opening by a terminal or ventro-terminal
pore; connective neither appendaged nor prolonged; thecae often prolonged a short
distance below the apex of the filament. Ovary wholly inferior, 5- to 12-celled;
style straight or bent; stigma punctiform, truncate, capitate, or broadly peltate
and retuse. Fruit a many-seeded berry. Trees or shrubs, with usually pli-nerved
leaves and terminal panicles of small or medium-sized white or pink flowers.

More than 50 species have been described, ranging from southern Mexico and
the West Indies to Ecuador and Brazil but most numerous in Central America.
The key which follows is “a artificial, since a reasonable classification of the
species has not yet been ac

a. Buds globose, apiculate or not, Ca thick-walled, with no distinc-

tion between the h ial se calycine portions; stigma (so far as
own) broad, retuse, radiali
b. silenzi about 7 m za (measured on the axis), commonly

verrucose; cad very emily presta anthers elliptic, about half
. C. MACRANTHA.

s wide a
bb. Hypanthium BL 4 mm. long, smooth; leaves strongly pli-nerved;
anthers about a third as wide as long 2. C. OERSTEDIANA.
aa. Buds for to obovoid, rar ody obtuse, commonly acute, acumin:
or apiculate, with obvious — distinction oig the Road
and calycine > ons; sind neither lobed n
b. Pubescence of the lower leaf-surface uere or c chiely of slender

simple se i È
c. Leaves pli-nerved, bearing petiolar formicaria; buds about 5-6 mm.
lon:

d. Petioles 1.5—5 cm. long; zag e basally rounded..................... 3. C. HIRSUTA.
ey Petioles at most 1 cm. long; leaf-blades basally pg nunt bodie 4. C. SETOSA.
- Leaves 3-nerved; formicaria is PORE; buds about 8 mm. long............... . C. BRACTEATA.
bb. Paetaa of | lower leaf-surface, petioles, stems, sa (TA of
lo; ati stellate hairs C. SPECIOSA.
bbb. Pubescence of lower leaf-surface să sessile stellate hairs, often minute
r — or wholly lacking at maturit
c. Lower lea Soahace completely lai by a fine, close, canescent
sociali tum . C. XALAPENSIS.
€c. Lower lea £ a Lal ber tha Indum
d. Style a distally to a truncate or sda stigma.
e. Leaves 3-nerved, short cip sen at the obtuse or rounded
pee. buds about 10 mm. scusate. uil . C. POLYANDRA.
-pli-nerved, Lu de w obtuse, 3.5 mm. long....... C. MICROMERIS.
dd. Style; of RZ diameter or widened distally; — RE
to
= the range 10. C. PUBERULA.

e. Leaves of an oblanceolate type, broadest well above
ee. Leaves of an ovate or oblong type, broadest near or below t
middle.

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[Vol. 45
270 ANNALS OF THE MISSOURI BOTANICAL GARDEN

f. Leaves hirsutulous above, conspicuously dentate and ciliate. 11. C. SUBCRUSTULATA.
ff. Leaves glabrous above, entire or nearly so, never ciliate
z SA y = i
: rostr

g trate, 7—13 mm. long.
h. Petals 4-6 mm. long; buds 7-8 mm. long; leaves
stellate-tomentulose on the veins beneath 12. C. MONTANA.
hh. Petals 10-15 mm. long; buds 10-13 mm. long; leaves
glabrous beneath 13. C. CHIRIQUENSIS.
gg. Buds rounded at the summit, 3.5-4 mm. long.............-------- 14. C. MICRANTHA.

1. CONOSTEGIA MACRANTHA Berg, ex Triana, in Trans. Linn. Soc. 28:97. 1871.

Tree to 10 m. tall, usually more or less furfuraceous, especially about the
inflorescence; petioles 2-4 cm. long; leaf-blades elliptic, 10-15 cm. long, half to
two-thirds as wide, obtuse or apiculate, at base obtuse or rounded, 5-nerved or
barely 5-pli-nerved, glabrous on both sides or minutely stellate beneath; panicle
widely branched, few-flowered; buds globose, 10—15 mm. long; hypanthium at
anthesis about 7 mm. long, usually verrucose; flowers white, usually 7-merous;
anthers oval, about half as wide as long.

Costa Rica and Panama.

cHIRIQUI: Rio Chiriquí Viejo Valley, near El Volcán, White & White 194.

y.

CONOSTEGIA OERSTEDIANA Berg, ex Triana, in Trans. Linn. Soc. 28:98. 1871.

Tree to 15 m. tall, glabrous throughout; petioles 1-3 cm. long; leaf-blades
varying oblong-ovate to oblanceolate or obovate, 6-15 cm. long, usually less than
half as wide, apiculate or acuminate, at base cuneate, acute, or rarely obtuse, 3- to
5-pli-nerved; panicle small, widely branched; buds globose, 7-10 mm. long,
usually apiculate; hypanthium at anthesis about 4 mm. long; flowers white, com-
monly 7-merous; anthers elliptic, about a third as wide as long.

Costa Rica and Panama.

CHIRIQUÍ: Valley of the upper Rio Chiriquí Viejo, White & White 98.
3. CONOSTEGIA hirsuta Gl. nom. nov.

ae setosa Standley & Macbride, in Field Mus. Pub. Bot. 4:244. 1929. Not
7 1871.

ostegia setosa Triana

Shrub, the stem, leaves, panicle, and hypanthium setose with simple bristles;
petioles up to 5 cm. long, the upper half and a portion of the midnerve occupied
by a slender formicarium; leaf-blades obovate-oblong, up to 2 dm. long and hal
as wide, abruptly acuminate, shallowly undulate-dentate, obtuse or rounded at base;
panicle small, many-flowered; buds 4-4.5 mm. long, the pyriform body tipped by
a slender beak; petals white, about 5 mm. long; style straight; stigma capitate.

Endemic to Panama, so far as known.

BOCAS DEL TORO: Buena Vista, Cooper 219.

*

CONOSTEGIA SETOSA Triana, in Trans. Linn. Soc. 28:99. 1871.

Shrub, similar to C. birsuta but the petioles only 0.5—1 cm. long, the formicaria
short and broad, the leaf-blades attenuate basally, the buds 5—6 mm. long.
Eastern Panama to northwestern Colombia.

(270)

1958]
FLORA OF PANAMA (Melastomataceae) 271

DARIÉN: Ensenada Guayabo, Stern & Chambers 178.

5. CONOSTEGIA BRACTEATA Triana, in Journ. Bot. 4:209. 1867.

Shrub to 3 m. tall, the younger stems hirsute; petioles 5-12 mm. long; leaf-
blades narrowly elliptic to oblanceolate, 8-15 cm. long, a fourth to a third as wide,
acuminate, obscurely serrulate, hirsutulous on both sides, 3-nerved; panicle small,
the glomerulate flowers subtended by oblong-obovate bracts 8 mm. long and by
smaller bractlets; buds pyriform, about 8 mm. long, abruptly acuminate, hirsute;
hypanthium at anthesis about 3 mm. long; petals white, 7-8 mm. long; style
straight; stigma ovoid, scarcely wider than the style.

Nicaragua to Panama. The species exists in two forms: plants of the lowlands
have leaves distinctly acuminate to a narrow base; plants of the highlands have
leaves convexly narrowed to an obtuse base.

ZONE: mea Mw pe Monte Lirio, 30 m., Killip 12158; Barro Colorado
Island, pos 876. co garto region, Gatün Take, Seibert 1530. DARIÉN:

ana-Cuasi trail, bin ndo mb m., Terry 6 Terry 1462. PROVINCE UNKNOWN:
Pittier 5619.

6. CONOSTEGIA SPECIOSA Naud. in Ann. Sci. Nat. Bot. III. 16:109. 1851.

Shrub to 7 m. tall, the stems, petioles, lower leaf-surface, panicles, and hypan-
thia softly pubescent with stellate hairs on stipes 1-2 mm. long; petioles stout, 1-2
cm. long; leaf-blades ovate, oblong-ovate, or oblong-obovate, up to 25 cm. long,
half to two thirds as wide, acuminate, usually serrate, obtuse or rounded at base,
5- to 7-pli-nerved, hirsute above with simple bristles; panicle many-flowered,
divaricately branched, up to 15 cm. long; buds pyriform, subacute, about 7 mm.
long; hypanthium at anthesis 3-4 mm. long; petals pink, 5-7 mm. long; style 3-4
mm. ES abruptly bent near the summit beneath the expanded flat stigma.

BOC TORO: Bastimentos, Mariano Creek, von Wedel 2904. CANAL ZONE:
Ancón Hill, ‘Killip 3011, Williams 16, Woodson, Allen & Seibert 1332; Barro Colorado

Island, Aviles 892, Bailey & Bailey 6, 232, Bangham 301, 412, Brown 134, Fairchild,
Shattu ck 707, Starry 123, Wetmore & Abbe 9, Wheeler 83, Woodson & Seibert 972,
d V al die a EA

; bet
P ANE r 6709; Santa Rita trail, La ell 152. cocLE: Llano Bonito, north of Las Margaritas,
400—500 m., Seibert 532; between Las Margaritas and ELY alle, Woodson, Allen & Seibert
1742. DARIEN: Macbride 2710. PANAMA: forests near Arraij án, Woodson, Allen 8
Seibert 1361; Isla Taboga, 0-186 m., Woodson, Allen 8 "Seibert 1524; vicinity of Pacora,
35 m., Allen 1118; San José Island, Johnston 446, 1166, Erlanson 567, Harlow 69. Prov-
INCE UNKNOWN: Ducbasssink s. n., Halsted s. n., Hayes 833.

7. CONOSTEGIA XALAPENSIS (Bonpl) D. Don, in Mem. Wern. Soc. 4:317. 1823.

Melastoma xalabense Humb. Eo heus 126. £. 54.
Conostegia lanceolata Cogn. in DC. Monogr. Phan. 7:708. "1891.

b or small tree to 10 m. ill the younger stems thinly but closely tomen-
tulose; petioles 1—1.5 cm. long; leaf-blades variable in shape, commonly narrowly
lanceolate, up to 15 cm. long and 4 cm. wide, occasionally oblanceolate or ovate-

(271)

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

272

dY
:
MS

eciosa

Conostegia sp

Fig. 89.

(272)

1958]
FLORA OF PANAMA (Melastomataceae) 273

lanceolate, acuminate, commonly dentate at least beyond the middle, obtuse to
cuneate at base, glabrous above when mature, permanently canescent beneath with
a fine close tomentum, 3- to 5-pli-nerved; panicle up to 1 dm. long; buds pyriform,
acute, 5-6 mm. long, tomentose; petals pink or white, 5-8 mm. long; style 4-5
mm. long, abruptly bent and slightly dilated below the capitate stigma.

Abundant and widely distributed from southern Mexico into northern South
America. Narrow-leaved plants, distinguished by Cogniaux as C. lanceolata, are
the most abundant in Panama

OCAS DEL TORO: southwest of Bocas at Macaw Hills, von Wedel 530; without locality,
von Wedel 334; Changuinola Valley, Dunlab 143. CANAL ZONE: Barro Colorado Island,
Aviles 99, Sbattuck 857, 1062; Chagres, Fendler 30; Gatún, Hayes 252; between Mt. Ho
and Santa Rita trail, Cowell 93. cHiRiQui: Bajo Mono, Boquete district, 1350 m., David-
son 517; vu of El Boquete, 1000-1300 m., Maxon 4930; Finca Lórida to Boquete,
1300—1700 +, Woodson, Allen 6 Seibert 1122; Rio Chiriqui Viejo Valley, near El Volcin
Wbite 210. COCLE: vicinity of El Valle de Antón, Allen 1771, 2502, 3546, Maurice ra
Seibert 404. COLÓN: along Río Fató, 10—100 m., Pittier 3883. PANAMA: hills
Campana, 600-800 m., Allen 1693.

8. CONOSTEGIA POLYANDRA Benth. Bot. Voy. Sulphur 96. t. 35. 1844.

Shrub or small tree to 6 m. tall, the younger parts minutely stellate-puberulent,
soon glabrescent; petioles slender, 1—2.5 cm. long; leaf-blades elliptic or oblong,
5-9 cm. long, about half as wide, obtuse or short-apiculate, entire, obtuse to
rounded at base, 3-nerved, essentially glabrous at maturity; panicle widely branched,
5-10 cm. long; buds pyriform, acuminate, glabrous, about 10 mm. long; hypan-
thium about 4 mm. long at anthesis; petals white, about 10 mm. long; style
straight, 8-10 mm. long, tapering distally to a punctiform stigma.

Costa Rica, Panama, and Colombia.

OCAs DEL TORO: Old Bank Island, von Wedel 1876, 1920; Bastimentos, Mariano
Crock, von Wedel 2895; Isla de Colón, von Wedel 75, 102, 531; Fish Creek Hills, von
Wedel 2416; Nances Cay, von Wedel 2935; without locality, von Wedel 341.

9. CONOSTEGIA MICROMERIs Standley, in Contr. Arn. Arb. 5:117. pl. 15. 1933.

“Tall tree,” the younger parts minutely furfuraceous, soon glabrescent; petioles
2-5 mm. long; leaf-blades thin, oblanceolate, up to 22 cm. long, about a third as
wide, acuminate, entire, long-cuneate to the base, 5-pli-nerved, glabrous above at
maturity, minutely furfuraceous on the veins beneath; panicle 3-6 cm. long,
divaricately branched; buds 3—3.5 mm. long, minutely furfuraceous; hypanthium
-2.5 mm. long at anthesis; petals white, 3—3.5 mm. long; style about 5 mm. long,
narrowed above to a small truncate or punctiform stigma.

Panama and western Colombia.

CANAL ZONE: Barro Colorado Island, Killip 39988, Woodworth 6 Vestal 602.

10. CONOSTEGIA PUBERULA Cogn. in DC. Monogr. Phan. 7:703. 1891.
Conostegia sororia Standley, in Field Mus. Pub. Bot. 22:161. 1940.
Tree to 12 m. tall, the younger parts finely but completely brown-stellate-

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[Vol. 45
274 ANNALS OF THE MISSOURI BOTANICAL GARDEN

tomentulose; petioles slender, 2-3 cm. long, tomentulose; leaf-blades oblanceolate,
up to 2 dm. long, a third to two-fifths as wide, caudate-apiculate or abruptly
short-acuminate, entire, gradually narrowed to an acute or cuneate base, glabrous
above, thinly stellate beneath and along the veins and veinlets; panicle 1-1.5 dm.
long; buds pyriform, obtuse or subacute, never apiculate, about 8 mm. long,
completely and permanently brown-stellate-tomentulose; hypanthium at anthesis
about 4 mm. long; petals white, 6-8 mm. long; style stout, columnar, 4-5 mm.
long; stigma hemispheric-capitate.

Costa Rica and Panama.

BOCAS DEL TORO: Almirante region, Cooper 412, 487, 581; Cricamola Valley, Cooper
486; Changuinola Valley, Cooper & Slater 91, Dunlap 412, 508. COLÓN: aroun
Bocas, Rio Fató Valley, 40-80 m., Pittier 4218. DARIÉN: Chepigana district, 240 m.,
Terry 6 Terry 1414.

11. CONOSTEGIA SUBCRUSTULATA (Beurl.) Triana, in Trans. Linn. Soc. 28:98.

1871.

Miconia subcrustulata Beurl. Prim. Fl. Portob. 130. 1854.

Shrub or small tree, up to 8 m. tall, the younger parts of the stem, petioles,
panicle, and lower leaf-surface floccose-tomentulose with stellate hairs; petioles 1-4
cm. long; leaf-blades usually ovate, varying to ovate-oblong, up to 2 dm. long,
half to two-thirds as wide, acute or short-acuminate, conspicuously serrate and
ciliate, rounded to subcordate at base, 7- to 9-pli-nerved, sparsely hirsutulous above
with simple hairs; panicle many-flowered, 1-2 dm. long; buds pyriform, 4-5 mm.
long, acuminate and often falcate, the hypanthial portion stellate-tomentulose, the
calycine portion nearly glabrous; hypanthium 2.5 mm. long at anthesis; petals pink,
4-5 mm. long; style about 4 mm. long, abruptly dilated and bent just below the
capitate stigma.

Nicaragua to Colombia.

ocas DEL TORO: Almirante region, Cooper 341; Fish Creek lowlands, vos Wedel
2386; Little Bocas, von Wedel 2548; vicinity of Nievecita, Woodson, Allen & Seibert 1874;

12. CONOSTEGIA MONTANA (Sw.) D. Don, in Mem. Wern. Soc. 4:317. 1823.
Melastoma montana Sw. Prodr. Veg. Ind. Occ. 69. 1788.

Small tree to 10 m. tall, the younger parts of the stem, petioles, and veins of
the lower leaf-surface thinly but distinctly stellate-tomentulose; petioles slender,
2-4 cm. long; leaf-blades thin, up to 15 cm. long, about a third as wide, acumina
entire, obtuse at the base, glabrous above, 5-pli-nerved; panicle 10-15 cm. long,
widely branched, many-flowered; buds pyriform, 7-8 mm. long, acute or short-
apiculate, glabrous; hypanthium at anthesis 2.5-3 mm. long; petals white, 4-6 mm.

(274)

1958]
FLORA OF PANAMA (Melastomataceae) 275

long; style 4-5 mm. long; stigma capitate, 1—1.3 mm. in diameter.
Costa Rica, Panama, and the West Indies.
OCAS DEL TORO: Robalo trail, area slopes of Cerro Horqueta, 1800-2100 M.,

Allen 4042, 5006. CHIRIQUÍ: Quebrada Velo, vicinity of Finca Lérida, 1500 m., Allen
4; between Alto de las Palmas ay top of Cerro de la Horqueta, 2100-2268 m., Pittier
E

13. CONOSTEGIA CHIRIQUENSIS Gl. ex Woods. & Schery, in Ann. Mo. Bot. Gard.
28:436. 1941.

Tree to 20 m. tall, glabrous except the very youngest stems and unfolding
leaves; petioles slender, 1-3 cm. long; leaf-blades firm, elliptic or obovate-oblong,
up to 15 cm. long, a third to half as wide, apiculate or abruptly short-acuminate,
entire, obtuse at the base, 5-pli-nerved; panicle 5-15 cm. long, few-flowered, the
branches ascending; buds pyriform, apiculate, 10-13 mm. long; hypanthium at
anthesis 3-4 mm. long; petals white, more or less unguiculate, 10-15 mm. long;
style stout, 4-5 mm. long; stigma hemispheric, capitate, about 1.5 mm. in diameter.

Endemic to Panama, so far as known.

RIQUf: Río Chiriquí Viejo valley, White 47, 324; trail from Paso Ancho to Mon
Lirio, valley of Rio Chiriqui Viejo, 1500-2000 m., Allen 1480; vicinity of Callejón ha
n de Chiriquí, 1700 m., Woodson & Schery 479; forests around El Boquete, 1000—
1300. m., Pittier 3142.

14. CONOSTEGIA MICRANTHA Standley, in Trop. Woods 16:18. 1928. Nomen;
Field Mus. Pub. Bot. 4:246. 1929.

Tree to 10 m. tall, the younger parts of the stem, petioles, and panicle loosely
stellate-pubescent; petioles 2.5-3.5 cm. long; leaf-blades elliptic, varying to
somewhat ovate or obovate, up to 18 cm. long, about half as wide, abruptly
caudate-acuminate, entire or undulate, broadly obtuse at base, barely to distinctly

5-pli-nerved, glabrous above, stellate-pubescent beneath; panicle pyramidal, many-
flowered, 3-12 cm. long; buds obovoid, 3.5-4 mm. long, rounded at the summit,
truncate or retuse at the base, glabrous or nearly so; hypanthium 2 mm. long at
anthesis; petals white, about 4 mm. long; style columnar, 3-4 mm. long; stigma
Capitate.

Costa Rica and Panama.

BOCAS DEL TORO: Buena Vista Camp on the Chiriqui trail, 375-540 m., Cooper 578,

619
30. TOCOCA Aubl.

Tococa Aubl. Pl. Guian. 1:437. £. 174. 1775.

Flowers usually 5-merous. Hypanthium obconic to campanulate. Calyx-tube
more or less prolonged, erect to flaring; sepals usually short or obsolete; exterior
teeth often conspicuous. Petals obovate to oblong, inequilateral, usually retuse.
Stamens isomorphic or nearly so; filaments stout; anthers erect, linear or subulate,
always more or less incurved at the summit; connective elevated into a low ridge

(275)

[Vol. 45
276 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ji

Fig. 90. Tococa guyanensis

(276)

19581
FLORA OF PANAMA (Melastomataceae) 277

along the thecae and often gibbous near their base, below the thecae neither pro-
longed nor appendaged. Ovary half-inferior, 3- to 5-celled; style stout, glabrous
or pubescent; stigma capitate, often large. Fruit a many-seeded berry. Shrubs or
even small trees with broad leaves often unequal in each pair and in many species
bearing formicaria at the summit of the petiole or base of the blade; flowers
medium-sized, white to pink, in terminal panicles.

About 60 species, most numerous in Amazonia.

1. TOCOCA GUYANENSIS Aubl. Pl. Guian. 1:438. £. 174. 1775.

Shrub 1-3 m. tall, the younger stems strongly flattened, glabrous to setose;
petioles stout, 1-6 cm. long, pubescent on the back, many of them bearing an
inflated formicarium at the summit; leaf-blades ovate-lanceolate to oblong or
elliptic, 10-25 cm. long, half to two-thirds as wide, usually ciliate, 5-nerved,
sparsely setose above, thinly pubescent beneath; panicle 1-2 dm. long; flowers
sessile; hypanthium about 5 mm. long; calyx almost truncate, with broadly
triangular glandular-setose exterior teeth; petals about 8 mm. long.

British Honduras and Panama; widely distributed in South America from
Columbia to the Guianas and Bolivia.

BOCAS DEL TORO: vicinity of Chiriquí Lagoon, von Wedel 1139. cocLE: Bismarck,
above Penonomé, Williams 586; locality not stated, Cooper & Slater 151.

31. HETEROTRICHUM DC.
HETEROTRICHUM DC. Prodr. 3:173. 1828.

Flowers 5- to 9-merous. Hypanthium campanulate to cup-shaped, always
pubescent. Sepals usually short, conspicuously exceeded by the exterior teeth.
Petals obovate. Stamens isomorphic; filaments smooth, slender; anthers subulate
or linear, 2-celled; connective more or less elevated along the thecae, unappendaged,
not prolonged at base or very briefly so. Ovary inferior only at base, 5- to 9-celled;
style slender, elongate; stigma capitate to punctiform. Fruit a berry with numer-
ous minute seeds. Shrubs or small trees of various aspect, with petiolate leaves and
terminal panicles of large or medium-sized flowers.

Twelve species of tropical America, each of them, except the following, re-
stricted in distribution.

1. HETEROTRICHUM OCTONUM (Bonpl.) DC. Prodr. 173. 1828.

Melastoma octonum Humb. & Bonpl. Melast. 7. t. 4. 1816.
Shrub 1-5 m. tall; stem, petioles, panicle, and hypanthium densely covered with
three types of pubescence, short, nearly sessile, stellate hairs, somewhat longer,
Projecting, glandular bristles, and elongate (4-8 mm.) simple bristles; petioles ies
cm. long; leaf-blades thin, ovate, 12-18 cm. long, half to two-thirds as wide,
acuminate, denticulate, cordate at base, 7-nerved, setose above with long hairs,
conescent-stellate beneath; panicle many-flowered, 5-10 cm. long; flowers 5- to
8-merous; sepals triangular-ovate, very short; exterior teeth subulate, 3-4 mm.

(277)

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

278

Heterotrichum octonum

Fig. 91.

(278)

1958]
FLORA OF PANAMA (Melastomataceae) 279

long; petals white, 8-10 mm. long; anthers subulate, 5-6 mm. long
Southern Mexico to Peru and Brazil, at low or moderate elevations; also in
Cuba.

OCAS DEL TORO: Little Bocas, vicinity of Chiriqui Lagoon, von Wedel 2402. CANAL

ZONE: Ancón Hill, vicinity of Balboa, Seibert 400, Woodson, Allen & Seibert 1321; Barro
gp Island, Aviles 2 22, ris abi 48 5. CHIRIQUÍ: San Félix to Cerro Pa" 100-850
AN an

9. ntí and vicinity, Williams 664. San José
ed: bai ES sputi I 0 470% 262. PROVINCE UNKNOWN: Hon. 22 f

32. MICONIA Ruiz & Pav.
MICONIA Ruiz & Pav. Fl. Peruv. & Chil. Prodr. 60. 1794.

Flowers 4- to 7- (commonly 5-)merous, in terminal paniculiform, spiciform,
or rarely corymbiform inflorescences. Calyx open in the bud, usually with small
but evident lobes and short, more or less adnate exterior teeth. Petals commonly
white, mostly small, often inequilateral or retuse. Stamens isomorphic or di-
morphic; anthers of various shapes, subulate, linear, oblong, or obovate, 2-celled
or 4-celled, opening by 1, 2, or 4 terminal pores or the pore extended into 1 or 2
longitudinal slits; connective simple or variously lobed or appendaged. Ovary
partly to wholly inferior; stigma truncate, capitate, or peltate. Fruit a many-
seeded berry. Trees or shrubs of diverse size and habi

At least 900 valid species, distributed throughout wici America. The genus
is ordinarily divided into twelve sections characterized by the shape and mode of
dehiscence of the anthers and by the nature of the calyx; three of these are not
represented in Panama. Miconia is closely related to several other genera, especially
Conostegia, Heterotrichum, and Tococa. The characters by which these are dis-
tinguished from Miconia are far weaker than those which separate certain sections
within the genus. Species of Miconia can seldom be identified satisfactorily with-
out recourse to the structure of the anthers, although many of them may be
recognized by various intangible characters not easily expressed in words. Two
keys are provided below, the first almost wholly artificial, the second more tech-
nical; the two should be used together. Besides the species described here, a few
hes may be represented by flowerless material now in herbaria. No attempt has
been made to include them in the keys. The flowers of one admitted species, M.
shattuckii, are unknown and it is placed at the end without assignment to a section.

ARTIFICIAL KEY
a. ee fac surface of the leaves distinctly and permanently pubescent or
epido
b. Pubescence of stellate hairs or scales.
e ssile.
i panim and calyx together 2-3 mm. long; petals about 2.5

11. M. IMPETIOLARIS.

5. M. AMPLEXANS.

d

I inflorescence.
e. Younger branches s scored tate and vier two-edged.....
a Younger branches terete or angled.

(279)

ce. Leaves ane
d.

10. M. ARGENTEA.

[Vol. 45

280 ANNALS OF THE MISSOURI BOTANICAL GARDEN

f. Petals entire
ff. Petals ią endless

dd. Flowers not at jka
. Petals 8—

dace flowers commonly 6-merous; leaves

— > i
8. ubt truncate or undulate; anthers oblong, blunt...
gg- = 5 lobes about 3 mm. long, deciduous at anthesis;

ff. I 2—3.5 mm. long; flowers 5-mer
ves 3 nerved, creed E ago ‘glabrous above.........

i Leaves ka -nerved, short-acuminate.
h. r stellate above when young, soon glabrescent,
nently and densely ferruginous-stellate beneath.
hh. Leaves enro setose “ba very sparsely stellate on

the actual surface beneath

bb. Pubescence entirely o or chiefly of simple bristles es by stellate
hairs in certain species; frayed or split at the apex in M. barbinervis).
c. Petals deep " 6-8 mm. long; filaments PETI mm. lear style

-25 mm. long
cc. Fatale phe or pale, less than 5 mm. long; filaments and style
uch shor
d. “Flowers S and wee along the branches of the inflorescence;
irs on es stem 8 mm. long; hypanthium glabrous; style

t to
dd. Flowers al in sessile, lateral, nearly or quite contiguous
glom ise along the few bran ches o£. the info orescence, forming
pound spiciform inflorescence; hypanthium pubescent;

E aig
ddd. Flowers in paniculiform inflorescences, although often glomeru-
late; hypanthium opere or pubescen nt; “ale strai

e. Pubescence of the stem, petioles, and pri veins distinctly
ascending-strigose; leaves long-tapering to aha base, the lateral
veins arising 3—8 cm. from bas
ee. Pubescence of the pt mone ci leaves 5-nerved...........
eee, Pubescence of the stem more EN ess spreading, frana never

ons dion

. ase.
ff. Leaves 3- to 5-nerved or 3- to 5-pli-nerved, obtuse or
t the base

escence of both stellate and simple hairs; leaves
inctly 5-pli-nerved

disti
gg- Pubescence of elongate matted hairs frayed or divided at
he apex

ggg. Pubescence entirely of simple bristles.
h. Plants densely hirsute > ROD hairs up to 3
ute, ru

mm. piar stigma min
hh. Plants hirsutulous with TAC irs up to 1 mm. long; stigma
arge, peltate, 4- to 5-ang obed A

ge, peltate led or |
aa. Lower surface of mature leaves glabrous or essentially so. ost
every species is somewhat stellate ia young and a few minute bairi
rs

the base.
d. Calyx-lo! € triangular
Eu. Calyx- lobes iet 1

pa

the base.
ul: Petals 5-10 m m. long; an thers 4—6 mm. lon
e. ‘a Plans glabrous chedeghiuts duce iol
Plants

ie or less stellat i erat la on the younger stems,
es, yy o: - thers blun
dd. mah! 2-3 mm. long; anthers 2 mm. ts or shorter.

* See footnote on p. 296.

(280)

19. M. ALBIC
9, M. Cruel E

28. M. SCHLIMII.
2. M. MUCRONATA.

3. M. CAUDATA.
22. M. RUBIGINOSA.

23. M. RUFOSTELLULATA.

55. M. MELANOTRICHA.

27. M. LACERA.

7. M. AERUGINOSA.
19. M. NERVOSA.
32. M. PITTIERI.
36. M. SHATTUCKII.
12. M. IBAGUENSIS.

13. M. BARBINERVIS.

29. M. COSTARICENSIS.

30. M. GONIOSTIGMA.

15. M. PRASINA.
16. M. PTEROPODA-

*
4. M. HONDURENSIS-

28. M. SCHLIMII.

1958]
FLORA OF PANAMA (Melastomataceae) 281

. Anthers less than 1 mm. long 34. M.
ee. i dute about 2 mm. long 33. M. EL A

1. M. LATERI

d. Stem Mes Tisk glabrous RIFLO
23. M. arti o

dd. Stem and lomis side of the leaf stellate
cc. Blowers 5-m
d. Flowers secund along the branches of the inflorescence; style bent
near me su
e. m grid, two-fifths to half as wide as long . M. CILIATA
ves purple-red beneath, a fourth to a third as wide as long. ic M. OINOCHROPHYLLA.
dd. “Flower not at all se und

and elon
e Flower ears along our go” axis of the spike-like
e
££ Piles = at least the lower) sessile on short Wn branches,
uppermost often solitary on a lateral bra
e. Inflorescence paniculiform
eaf-blades comparatively large, commonly half to two-

6. M. TRIPLINERVIS.

8. M. GRACILIS.

da as wide as

g. Flowers plico deve caudate; anthers subulate........... 3. M. CAUDAT
. Flowers sessile; leaves short-acuminate; a a nega AGE 14. M. CALVESCENS.
ff. PARRA “comparatively small, rarely up t wide,
arely as much as half as wide as lon

g. Leaves ge an pes ng or o blong-lanceolate type, commonly
widest lens below the

h. = and fifth veins abetted and — d
. M. BOREAL
hh. (pe es per ind veins cin id marginal and obscure..... t M. usum
gg. Leaves of an oblong to obovate type, commonly broadest

h. iere X A the calyx. sali. but distinctly developed.
of t inflorescence marked align
Ama sera anthers linear
ii. Nodes of the severa lacking setae; n
oblong to obovate.
j. Serale branches of the inflorescence angular and
bi isul cate.
jj. Smaller branches of the inflorescence terete or

M. HYPERPRASINA.

31. M. THEAEZANS.

3. M. AM
18. M. INSUL

nearly so
hh. Lobes of the calyx obsolete
KEY TO THE SECTIONS

a. Anthers €— OMS to tai quam, opening by a minute

ter ris po e commonly
b. L pelado connective Viduae or glabr
c. Sepals or — eeth or both well d jeg i in intra
with t aier Po I. JUCUNDA (spp. 1-2).
cc. Sepals de exterior teeth poorly developed in comparison
. with the hypanthium . II. TAMONEA (spp. 3-4).
bb. Leaves sess sile; connective din mpm - vnius RSA III. ADENODESMA (sp. 5).
aa. Anthers linear or broader.
b. Anhana pening by a a erminal por ^
Anthers s slender, Í linear or slightly widened distally.
Pore minute, rarely more than half the width of the
antha n (nf, Ę MIcoNIA (spp. 6-23).
dd. Pore conspicuous, about as na as ii o . GLOSSOCENTRUM (sp. 24).
ce. Anchen oblong to obovate or
d. minute, much less than half as ; wide as the anther.
Eyck zę 2-celled; style abruptly bent near the summit. Ai HARTIGIA (spp. 25-27).
ee. Anthers 4-ce zp: style up LAUR HB AL ET. iio . AMBLYARRHENA (spp. 28-30).
di Pote at least half as wide as B anther, the A E
tween the thecae commonly protruding.....-------------------- un CREMANIUM (spp. dgr
bb. Anthers opening by two longitudinal età M - CHAENOPLEURA (sp

(281)

[Vol. 45
282 ANNALS OF THE MISSOURI BOTANICAL GARDEN

I. Section JUCUNDA (Cham.) Tr. ex Benth. & Hook. f. Gen. Pl. 1:764. 1867

Anthers subulate, usually with convolute thecae, opening by a minute pore;
connective simple or variously lobed or prolonged; either the sepals or the exterior
teeth greatly developed and the calyx therefore apparently deeply lobed; exterior
teeth often much exceeding the sepals and mistaken for them; leaves petioled (in
our species).

Twenty known species, most numerous in Amazonia.

a. Sepals proper nearly obsolete, much surpassed by the narrowly triangular

exterior teeth; flowers 4-merous; leaves 3-nerved 1. M. LATERIFLORA,
aa. Sepals 3-4 mm. long from the torus; exterior teeth minute; flowers
mostly 6-merous; leaves 3- to 5-pli-nerved 2. M. MUCRONATA.

1. MICONIA LATERIFLORA Cogn. in Bol. Mus. Goeldi 5:255. 1909.

Ossaea disparilis Standley, in Contr. Arn. Arb. 5:120. pl. 17. 1933.
Miconia disparilis (Standley) R. O. Williams, in Dept. Agric. Trinidad & Tobago 1:388.
1934.

Shrub to 3 m. tall, the stems and leaves glabrous; petioles slender, 5-25 mm.
long; leaf-blades shining, thin, elliptic, 10-25 cm. long, two-fifths to half as
wide, slenderly acuminate or caudate, obscurely crenulate and sparsely ciliate,
obtuse to broadly cuneate at base, 3-nerved; panicle loose and open, sparsely
branched; flowers predominantly 4-merous, sessile in small terminal glomerules;
hypanthium tubular, becoming ovoid in fruit, glabrous or minutely punctate, 3.354
mm. long; calyx-lobes nearly obsolete; exterior teeth narrowly triangular, 1.5-2
mm. long; petals obovate, 1.5-2 mm. long; stamens isomorphic, the anthers sub-
ulate, 3—3.5 mm. long; connective minutely 4-lobed at base; ovary nearly free,
4-celled; stigma truncate; fruit strongly 8-costate, sometimes glandular-pilose.

British Honduras to Panama, Colombia, Trinidad, Surinam, and northern
Brazil.

Bocas DEL TORO: Fish Creek, von Wedel 2230; Water Valley, von Wedel 1470; Buena
Vista Camp on the Chiriqui trail, 375 m., Cooper 593. CANAL ZONE: Barro Colorado
Island, Aviles 21, Bailey & Bailey 201, 506; Frijoles, Standley 27553. DARIEN: Cana-
Cuasi trail, Chepigana district, 600 m., Terry & Terry 1455.

2. MICONIA MUCRONATA (Desr.) Naud. in Ann. Sci. Nat. Bot. III. 16:120. 1851.

Melastoma bolosericea L. Sp. Pl. 390. 1753.

Melastoma mucronata Desr.; Lam. Encyc. 4:46. 1796. ia DE

Miconia bolosericea Triana, Trans. Linn. Soc. 28:101. 1871. Not M. holosericea ^^^
1828.

Shrub or small tree to 5 m. tall, the smaller twigs somewhat flattened, very
finely tomentulose, as are also the petioles, lower leaf-surface, panicle, and hypan-
thia; petioles 1.5-2.5 cm. long; leaf-blades broadly elliptic, varying to ovate oF
obovate, up to 4 dm. long, a third to half as wide, abruptly short-acuminate, m”
obtuse to rounded at base, glabrous above, 3- to 5-nerved or 3- to 5-pli-nerved;
panicle to 15 cm. long, rather few-flowered; flowers commonly 6-merous, sessile
hypanthium tubular, about 6 mm. long; sepals ovate to lanceolate, 2,555 E

long, deciduous at anthesis; petals obovate-oblong, 8-10 mm. long; stamens 1507

(282)

1958]
| FLORA OF PANAMA (Melastomataceae) 283

morphic; anthers subulate, 6-8 mm. long, the connective prolonged into two obtuse
basal lobes and a short erect dorsal spur; ovary almost free; style about 18 mm.
long; stigma truncate.
British Honduras and Panama; in South America south to southern Brazil and
Bolivia.
CHIRIQUÍ: vicinity of San Félix, 0-120 m., Pittier 5261.
II. Section TAMONEA Cogn. in Mart. Fl. Bras. 14*:238. 1887.

Anthers subulate, usually with convolute thecae, opening by a minute terminal
pore; connective usually simple, occasionally very briefly prolonged and lobed,
rarely glandular; calyx-tube well developed but scarcely lobed, the exterior teeth
none or minute.

About 50 species, most numerous in Amazonia.

a. Leaves ovate, usually (not always) cinereous beneath, glabrous above;

hypanthium cinereous 3. M.
aa. Leaves oblong or oblong-obovate; plant glabrous throughout..........-...------- 4. M.

CAUDATA.
HONDURENSIS.
3. MICONIA CAUDATA (Bonpl.) DC. Prodr. 3:187. 1828.

Melastoma caudata Humb. & Bonpl. Melast. 13. £. 7. 1807.

Shrub or small tree to 12 m. tall, the younger stems ferruginous-stellate-
tomentose, glabrescent in age; petioles slender, 2-5 cm. long; leaf-blades firm,
ovate, 1-2 dm. long, about half as wide, caudate-acuminate, entire, broadly rounded
at base, 5-nerved, glabrous and dull green above, beneath varying from cinereous
with a close tomentum of stellate hairs to completely glabrous; panicle 5-10 cm.
long, compact, much branched, the flowers in small terminal glomerules; hypan-
thium campanulate, about 2.5 mm. long to the torus, cinereous-stellate, calyx-tube
about 0.5 mm. long, obscurely 5-lobed; petals white, obovate, 3-3.5 mm. long,
stellulate externally; stamens isomorphic or somewhat dimorphic; anthers stoutly
subulate, 4-4.5 mm. long, somewhat incurved, with convolute thecae; connective
simple or elevated into a dorsal ridge near the base, smooth or beset with sessile
glands; filaments smooth or villosulous; style often villosulous.

Southern Mexico to Ecuador.

CHIRIQUi: Boquete, 1350 m., Maxon 5145, Terry 1278. COCLÉ: region of El Valle de
Antón, 1000 m., Allen 3734.

4. MICONIA HONDURENSIS Donn. Sm. in Bot. Gaz. 40:3. 1905.
Miconia gatunensis Pittier, in Journ. Wash. Acad. Sci. 14:447. 1924.

Shrub to 4 m. tall, glabrous throughout; petioles 1-2 cm. long; leaf-blades
oblong to oblong-obovate, up to 20 cm. long and 8 cm. wide, abruptly caudate-
acuminate, entire, narrowed to an obtuse base, minutely but densely verruculose-
puncticulate above, slightly 3-pli-nerved; panicle pyramidal, 5-15 cm. long, pe
flowers mostly in clusters of three at the end of the branches, the terminal on a
pedicel about 2 mm. long, the lateral sessile at the end of a very short en
hypanthium narrowly campanulate, 3.5—4 mm. long; calyx-tube slightly flaring,

(283)

[Vol. 45
284 ANNALS OF THE MISSOURI BOTANICAL GARDEN

about 1 mm. long, the lobes nearly obsolete, the exterior teeth lacking; petals white,

oblong-elliptic, retuse, about 5 mm. long; stamens dimorphic; filaments about 6 or

about 3.5 mm. long; anthers subulate, with obscurely convolute thecae, the larger

nearly straight, about 6 mm. long, the smaller incurved, about 5 mm. long;

connective thickened toward the base and curved around the base of the thecae

as two small lateral lobes; style slender, about 12 mm. long; stigma capitellate.
British Honduras to Panama.

CANAL ZONE: along the Rio Indio de Gatún, near sea-level, Pittier 2786. DARIEN:
Tucuti, Chepigana district, about 15 m., Terry 6 Terry 1373.

III. Section ADENODESMA Naud. ex Cogn. in Mart. Fl. Bras. 14*:253. 1887.

Anthers subulate, usually strongly arcuate, opening by a small terminal pore;
thecae convolute; connective expanded at the base, prolonged and variously lobed
below the filament, glandular; leaves pli-nerved, usually sessile.

Eleven known species, all but two Amazonian. Only the following species in
Panama.

5. MICONIA AMPLEXANS (Crueg.) Cogn. in Mart. Fl. Bras. 14:256. 1887.
Pogonorbynchus amplexans Crueg. in Linnaea 20:107. 1847.

Shrub or small tree to 8 m. tall, the younger stems closely brown-tomentose
with stellate hairs; leaves sessile, firm in texture, obovate, up to 4 dm. long and
about half as wide, abruptly acuminate, narrowed from above the middle to 2
rounded or auriculate base, 3-pli-nerved, the lateral nerves arising 5-10 cm. above
the base, soon glabrescent above, beneath permanently thinly stellate-pubescent;
inflorescence narrow, 1-2 dm. long, stellate-tomentose; hypanthium tubular-
campanulate, 5.5-6 mm. long to the torus, finely stellate-tomentose; calyx-tube
prolonged 1-1.5 mm.; sepals 0.5-1 mm. long from the sinus, semicircular; petals
white, inequilateral, oblong-obovate, about 7 mm. long; stamens only slightly
dimorphic; anthers slenderly subulate, about 6 or about 8 mm. long, the connec-
tive at base glandular-pubescent and curved into a half-circle below the thecae;
filaments sparsely glandular; ovary glandular; stigma truncate. :

British Honduras, Trinidad, and Panama, and southward at low elevations in
South America to Bolivia.

PROVINCE UNKNOWN: Hayes 972.

IV. Section MICONIA

Anthers elongate, linear, opening by a terminal pore seldom half as wide as the
anther, 2-celled; connective simple in a few species, usually more or less prolonged
below the thecae into various types of lobes or appendages.

About 250 species, distributed over the entire range of the gen :
numerous in Amazonia. The nature of the connective, upon which the major
divisions of the key are based, can not be determined from dried anthers; they
must first be restored to natural form by brief boiling.

us but most

(284)

19581 3
FLORA OF PANAMA (Melastomataceae)

al. Connective of tt episepalous stamens simple at the base, not prolonged

below the thec 6.

a2. og dd of the episepalous stamens prolonged below the thecae into
wo deflexed lateral lobe es.
tig Lateral branches from the central axis of the inflorescence themselves
unbranche
c. Lateral branches of the rn? bearing aue A contiguous)

285

M. TRIPLINERVIS.

lomerules of flowers; pubescence of simple bristles . M. AERUGINOSA.

ce: arii rane hes of the panicle bearing a fex sessile solitary flowers;
plan early or qui te glabro throug out
bb. Lateral “notti es from the fatt axis of the inflorescence also

c. Flowers secund along the ultimate branches of the panicle; leaves
$ Ai 1

encath,

d. Petals glandular-ciliate; upper internodes strongly angled but not

o-edge
dd. stia not glandular; upper internodes strongly flattened and
arply two-edge 10.
cc. Flowers not secund; leaves om ter stellate, or pubescent beneath,
erruginous in one speci
d. Leaves du sessile rion E broad base, permanently stellate
eneat

dd. Leaves petioled, or the blade long-cuneate at base and decurrent
along the —

e. Younger s and lower le gti hirsute (the bristles may
be BA per or rashod at the summ:
f. Leaves > -nerved, glabrous or > 30 ADOVE ede 12;
ff. Leaves 3-nerved, hirsute abov 15.
ee. Younger stems and lower ca surface glabrous or obscurely
pubescent sk minute stellate
f. sta es broadly rounded to bede? at base, 3- to 5-nerved;
capitate
ff. Leaves gradually ponad at base and cuneate into a ca
nt petiole; stigma tru
z Cal pie triangular, wad or quite equaled by the
triangular external tee 15.
gg- mendo -lobes depressed semi cular; exterior teeth obsolete
y ute thickenings
33. Connective of i episepalous stamens dilated at base to the width of
the thecae, decurved around the filament and prolonged below the thecae

more or less t rgan.
b. Leaves sessile by a broad base, permanently stellate beneath 11.

bb. Leaves petio oled, glabrous or obscurely and minutely stellate beneath.
c. Inflorescence branches an ne. br anchlet usually opposite; calyx lobes

distinct, the external t projecting 17.

ob; — branches d ranches | whorled, 3-6 per node; calyx
truncate, the external ii arly 18.

a4, ler t dilat t base, zar er d deus back above the
filament as a single rue 1 doba 19.

a5. Connective of the episepalous stamens prolonged below the thecae and

"Ww ated into a broad, oblique, cordate structure with the apex of the
een ini its in
amens morphic or nearly so; flowers secund on the ultim
is of the panicle; > ves — cinereous-tomentose diras SUE 20.
bb. Stamens dimor orphic; flow:
€. Panicle large ini di pina great the main axis of the
eafy branches.
d. Xe iuf stems and lower leaf-surfaces thinly and sparsely pubes-
with stellate hairs.
dd. Stems and lower poż: ży ori danny and permanently pubescent

21.

22.

erruginous stellate hai
ce. Pe ti and several, Ama short, divaricate, fno levi,
lat ater. ral branches

e).

(285)

M. GRACILIS.

9. M. STENOSTACHYA.

M. ARGENTEA.

11. M. IMPETIOLARIS.

M. IBAGUENSIS.
M. BARBINERVIS.

. M. CALVESCENS.

M. PRASINA.

16. M. PTEROPODA.

M. IMPETIOLARIS.

M. HYPERPRASINA.
M. INSULARIS.

M. NERVOSA.

M. ALBICANS.

M. PANAMENSIS,
M. RUBIGINOSA.

M. RUFOSTELLULATA.

[Vol. 45
286 ANNALS OF THE MISSOURI BOTANICAL GARDEN

6. MICONIA TRIPLINERVIS Ruiz & Pav. Syst. Veg. 1:105. 1798.
Miconia angustispica Blake, in Contr. U. S. Natl. Herb. 24:15. pl. 5. 1922.

Shrub or small tree to 10 m. tall, the younger parts, including the leaves, more
or less stellate-tomentose when young, glabrescent in age; petioles slender, 5-15
mm. long; leaf-blades thin, oblanceolate to obovate, up to 25 cm. long, a third to
half as wide, abruptly but conspicuously acuminate, entire, gradually narrowed
below to an obtuse or acute base, 3-nerved; inflorescence normally a single, elongate,
spiciform cluster 1-2 dm. long, floriferous nearly to the base, the flowers sessile in
lateral glomerules of 2—4; hypanthium campanulate, 2.5 mm. long to the torus,
thinly stellate-pubescent; calyx about 0.5 mm. long, nearly truncate; exterior
teeth none; petals white, 1.5-2 mm. long, thinly stellulate on both sides; stamens
isomorphic; anthers 2.5-3 mm. long, somewhat incurved, with convolute thecae
and simple connective.

Southern Mexico and Jamaica to Brazil. Panama plants, segregated as a distinct
species by Blake, have in general larger, proportionately broader leaves and a denser
tomentum than typical plants of Amazonia.

BOCAS DEL TORO: Fish Creek, von Wedel 2197; Isla Colón, von Wedel 1929; region
of Almirante, Cooper I6I.

7. MICONIA AERUGINOSA Naud. in Ann. Sci. Nat. Bot. III. 16:135. 1851

Shrub or small tree to 6 m. tall, the stems, petioles, both sides of the leaves,
panicle, and hypanthia more or less hirsute with straight simple hairs, also (except
the upper side of the leaf) more or less finely stellate-tomentulose; petioles stout,
1-3 cm. long, more densely hirsute on the upper side; leaf-blades ovate or oblong-
ovate, up to 20 cm. long, half to two-thirds as wide, acuminate, commonly dentate,
broadly rounded or subcordate at base, 7-nerved, the outer pair nearly marginal,
sparsely hirsute above; panicle with a central axis up to 15 cm. long, with several
simple branches bearing small sessile glomerules, often contiguous, of sessile flowers;
hypanthium obconic, about 2 mm. long to the torus; calyx-tube prolonged about
0.5 mm., its lobes depressed-triangular, minute; exterior teeth subulate, barely
surpassing the lobes; petals white, obovate-oblong, 3.5—4 mm. long; stamens di-
morphic; anthers linear, about 2 or about 2.5 mm. long, connective of the smal
ones simple, that of the larger prolonged at base into two small lateral lobes; stigma
truncate.

Southern Mexico to Colombia and Venezuela.

CHIRIQUi: vicinity of El Boquete, 1140-1300 m., Davidson 604, 724, Pittier

o

2907;
Finca Lérida to Boquete, 1300-1700 m., Woodson, Allen & Seibert 1143. COLÓN:
Monk[ey?] Hill, Kuntze s. n.

8. MICONIA GRACILIS Triana, in Trans. Linn. Soc. 28:107. 1871.

Shrub to 5 m. tall, glabrous or essentially so throughout; petiole 5-10 (rarely
25) mm. long; leaf-blades thin, bright green up to 10 cm. long, oblong or elliptic,
a third to nearly half as wide, the members of a pair often unequal, acuminate 10 3
blunt tip, entire, acute at the base, 3-nerved; panicles slender, up to 15 cm. long

(286)

1958]
FLORA OF PANAMA (Melastomataceae) 287

and 2 cm. in diameter, the numerous short branches seldom again branched and
bearing 1—5 sessile flowers; hypanthium campanulate, 3 mm. long, costate in fruit;
calyx closed in bud, at anthesis ruptured nearly to the torus into 3-5 broadly
triangular lobes; exterior teeth none; petals white, obovate-oblong, 3 mm. long;
stamens nearly isomorphic, differing in size; anthers linear, 2.5-3 or 3.5-4 mm.
long, the connective very briefly prolonged below the thecae into two lateral lobes,
stigma capitate.
Nicaragua to Colombia.

BOCAS DEL TORO: Fish Creek, von Wedel 2190, 2196, 2234; Old Bank Island, von
Wedel 1922, 2166. cocLÉ: vicinity of El Valle, Allen 237.

9. MICONIA STENOSTACHYA DC. Prodr. 3:181. 1828.

Shrub or small tree to 6 m. tall, the younger branches, petioles, lower leaf-
surface, panicle, and hypanthia thinly cinereous-tomentulose; branches strongly
angled; petioles 1-4 cm. long; leaf-blades thin, oblong or ovate-oblong, 8-18 cm.
long, about two-fifths as wide, obtuse to abruptly short-acuminate, entire, rounded
at the base, glabrous above, 3- to 5-nerved; panicle usually dense, up to 15 cm.
long; flowers sessile and secund; hypanthium campanulate, about 3 mm. long;
calyx-tube scarcely prolonged, the calyx-lobes broadly triangular; petals 3-5 mm.
long, ovate, minutely glandular-ciliate; stamens isomorphic; anthers linear, about
4 mm. long, the connective barely prolonged below the thecae into two deflexed
lateral lobes and one minute dorsal lobe; stigma conic or punctiform.

Southern Mexico and Trinidad to Bolivia and southern Brazil.

CANAL ZONE: vicinity of Summit, Allen 4556; road to Corozal, Gervais 142.

10. MICONIA ARGENTEA (Sw.) DC. Prodr. 3:182. 1828.
Melastoma argentea Sw. Prodr. Veg. Ind. Occ. 70. 1788.

Tall shrub or tree to 15 m. tall, the younger stems, petioles, lower leaf-surface,
panicle, and hypanthia thinly but completely canescent with stellate hairs; younger
stems strongly flattened and two-edged; petioles stout, 2-6 cm. long; leaf-blades
firm, broadly elliptic, up to 25 cm. long, about half as wide, abruptly short-
acuminate, entire or denticulate, obtuse to rounded at base, 5-nerved, glabrous
above; panicle widely branched, 1-2 dm. long; flowers sessile, 5-merous; hypan-
thium obconic, about 2 mm. long; calyx almost truncate, the obscure lobes 0.2
mm. long; petals inequilateral, obovate-oblong, 3 mm. long; connective of the
larger stamens prolonged at base into two small lateral lobes, that of the smaller
stamens unlobed at base or with a minute dorsal lobe only; terminal pore large, as
wide as the anther; stigma capitate.

Southern Mexico to Panama.

CANAL ZONE: Barro Colorado Island, Aviles 31, Brown 133, 150, Wilson 104, towa?
— & Vestal 315, Zetek 4636; Río Agua Salud, = PA portó 58 54 Gatún, Cowe
district, ee s i : El Pedregal de David, near sea-level, Pittier 511 6. cocLE:
vicinity of El Valle de Antón, 700-1000 m., Allen 102, 2480; Penonomć and vicinity,

(287)

[Vol. 45
288 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Williams 215. COLÓN: Juan Mina plantation, Rio Chagres sone Ga mboa, 25 m., Allen
4154. DARIEN: vicinity of La Palma, 0-50 m., Pittier 5492. PANAMA: Trapi cha Perlas

I75, 218. san BLAS: Permé, Cooper 657. VERAGUAS: Santa Fé to Rio Santa Marta, 300
m., Allen 4418.

11. MICONIA IMPETIOLARIS (Sw.) D. Don, in Mem. Wern. Soc. 4:516. 1823.
Melastoma impetiolaris Sw. Prodr. Veg. Ind. Occ. 70. 1788.

Shrub or small tree, the younger stems, petioles, lower leaf-surface, inflores-
cence, and hypanthia cinereous or ferruginous with stellate hairs; leaves sessile,
oblanceolate to obovate, up to 4 dm. long, a third to half as wide, abruptly short-
acuminate, entire or denticulate, rounded to a cordate-clasping base, glabrous above
at maturity; panicle large, divaricately branched; flowers 5-merous, sessile in small
glomerules; hypanthium cup-shaped to obconic, 1.5-2.5 mm. long; calyx-tube
prolonged about 0.5 mm., nearly truncate, often erose; exterior teeth broadly
triangular, adnate, projecting about 0.2 mm.; petals white, about 2.5 mm. long;
stamens weakly dimorphic; ovary 3-celled, half-inferior; style slender, 8-10 mm.
long; stigma truncate.

Southern Mexico to Panama; West Indies and (in a variety) northwestern
South America. The two varieties described below are clearly distinct in Panama
and other parts of Central America but intergrade almost completely in the West
ndies.

Var. IMPETIOLARIS.

Leaves 3-nerved or 3-pli-nerved with an additional marginal pair; hairs of the
lower leaf-surface not contiguous, exposing the surface; secondary veins com-
monly 35 or fewer on each side of the midvein, the middle ones diverging at an
angle of about 75°; upper leaf-surface not punctate; connective dilated at base
into two small lateral lobes.

S DEL TORO: banks of the Chaguinola River, Dunlap 522. CANAL ZO

: Barro
Colo ide Island, Wilson 62, wie 6G Vestal 1597; between Mt. Hope and "de Sants
Rita Trail, Cowell 66; Santa Rita Trail, Cowell 11

Var. PANDURIFOLIA Naud. in Ann. Sci. Nat. ser. 3. 16:137. 1851.

Leaves 5-nerved or 5-pli-nerved with an additional marginal pair; hairs of the
lower leaf-surface with interlaced arms, forming a continuous indument; secondary
veins commonly 40 or more on each side of the midvein, the middle ones diverging
almost at right angles; upper leaf-surface minutely verruculose-punctate; i
tive dilated into a large, more or less erose-margined lobe extending over the ba
and deflexed over the sides of the base of the anther.

OCAS DEL TORO: Quebra Nigua, von Wedel 2740; Snapper Point, vom Wedel 2643;
Water Valley, von Wedel 816, 890, 1412, 1734, 1742. CANAL ZONE: Chagres Fendler
29; vicinity of Sum ummit, 150 m., Allen 4518; Naval Amm. Depot, Co 300 Road, z:
& Chambers 37. DARIÉN: headwaters of Rio Chico, 150-225 m en 4599. PA
vicinity of Bella Vista, Macbride 2747, Piper 5332; Šan riki Island, peri 690,
72, 239. PROVINCE UNKNOWN: Kuntze s. n., von Wedel 8.

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FLORA OF PANAMA (Melastomataceae) 289

12. MICONIA IBAGUENSIS (Bonpl.) Triana, in Trans. Linn. Soc. 28:110. 1871.

Melastoma ibaguense Humb. & Bonpl. Melast. 105. /. 45. 1816.

Shrub to 6 m. tall, the younger stems and petioles stellate-tomentose and also
more or less densely hirsute; petioles 3-10 mm. long; leaf-blades firm, oblong to
oblong-lanceolate, up to 15 cm. long, a third to nearly half as wide, acuminate,
entire to minutely serrulate, obtuse or rounded at the base, 5- pli-nerved, the veins
usually conspicuously impressed, sparsely stellate above when young, later nearly
or quite glabrous, below hirsutulous with simple hairs; panicle compact, up to 1

m. long, more or less hirsute, the sessile flowers commonly crowded toward the
ends of the branches; hypanthium campanulate, 2.5—3 mm. long, finely stellate and
sparsely hirsute; calyx-tube about 0.7 mm. long, its lobes minute, triangular, about
equaled by the exterior teeth; petals white, obovate, about 4 mm. long; stamens
somewhat dimorphic; anthers linear 3—3.5 or 4—4.5 mm. long, the connective
expanded below the thecae into two lateral lobes, somewhat deflexed on the smaller
anthers, curved forward on the larger ones; summit of the ovary commonly setose;
stigma capitate.

Widely distributed from southern Mexico and Cuba to southern Brazil and

ivia.

savannas, Soquete district, 1200 m., Davidson 687; rocky plains near
TES Alles 4688. PANAMA: San José Island, Erlanson 281. VERAGUAS: trail between
Cañazas and the foot of dei Cordillera P nd 300—600 m., Allen 170.

13. MICONIA BARBINERVIS (Benth.) Triana, in Trans. Linn. Soc. 28:113. 1871.
Clidemia barbinervis Benth. Bot. Voy. Sulphur 95. 1844.

Shrub to 4 m. tall, the stems, petioles, lower leaf-surface, panicle, and hypan-
thia densely hirsute or tomentose with straight or matted bristles 0.7-4 mm. long
and stellate or frayed at the summit; petioles stout, 5-20 mm. long; leaf-blades
ovate-lanceolate, oblong, or somewhat obovate-oblong, a third to two-fifths as
Wide, acuminate, more or less saliently dentate, 3-nerved, hirsute above with
usually simple, rarely stipitate-stellate hairs; panicle pyramidal, up to 1 dm. long;
flowers 5- to 6-merous, sessile in terminal few-flowered glomerules, each subtended
by a small obovate-oblong bract; hypanthium nearly cylindric, 3.7 mm. long;
calyx-tube prolonged about 0.5 mm.; calyx-lobes triangular, blunt, considerably
surpassed by the subulate exterior teeth; petals oblong-obovate, about 4 mm. long,
unsymmetrical; stamens essentially isomorphic but unequal, the anthers 4-5 mm.
9r 3—4 mm. long; connective continued at base into two rounded lateral lobes;
Stigma capitate, about 1 mm. wide

icaragua to Panama; at low elevations in northern South America to Peru
á

BOCAS DEL TORO: Water Valley, von Wedel 905, 1581.

14. MICONIA CALVESCENS DC. Prodr. 3:185. 1828.
Small tree to 10 m. tall, the younger branches and base of the panicle some-

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290 ANNALS OF THE MISSOURI BOTANICAL GARDEN

what flattened, thinly cinereous with minute stellate hairs; petioles 3-6 cm. long;
leaf-blades elliptic to obovate-oblong, 15-35 cm. long, half to two-thirds as wide,
abruptly short-acuminate, entire or undulate, obtuse to broadly rounded at base,
3-nerved with an additional pair of marginal veins, pubescent when young, soon
glabrescent, or with a few stellate hairs persistent along the veins beneath; panicles
1-3 dm. long, divaricately branched, thinly cinereous-tomentulose; flowers sessile
in small lateral and terminal glomerules, 5-merous; hypanthium about 2 mm. long,
thinly tomentulose; calyx-tube prolonged about 0.5 mm., sepals broadly triangular,
about 0.3 mm. long; petals obovate, white, 2.5-3 mm. long; stamens nearly iso-
morphic; anthers linear, 2-2.5 or 3-3.5 mm. long; connective very briefly pro-
longed below the thecae into two deflexed lateral lobes; stigma capitate.

British Honduas to Panama, and southward to Bolivia and southern Brazil.

DARIÉN: Cana-Cuasi trail, Chepigana district, 600 m., Terry & Terry 1492.

15. MICONIA PRASINA (Sw.) DC. Prodr. 3:188. 1828.
Melastoma prasina Sw. Prodr. Veg. Ind. Occ. 69. 1788.
Miconia darienensis Pittier in Contr. U. S. Natl. Herb. 18:248. 1917.

Shrub or small tree, reported to 10 m. tall, the younger branches, lower leaf-
surface, panicle, and hypanthia very minutely and sparsely stellate-furfuraceous;
petioles 5-20 mm. long; leaf-blades firm in texture, oblong or lanceolate, com-
monly 1-2 dm. long and a third to two-fifths as wide, acuminate, entire or
somewhat undulate, acuminate or acute at base and often decurrent on the petiole,
usually completely glabrous above, 3- to 5-pli-nerved; panicle pyramidal, up to
15 cm. long; flowers sessile or nearly so, tending to be crowded toward the ends of
the branches; hypanthium narrowly campanulate, 2—2.5 mm. long; calyx-tube
prolonged about 0.5 mm.; calyx-lobes triangular, acute or obtuse, 0.2-0.4 mm.
long; petals white, obovate, 2.5-3 mm. long; stamens somewhat dimorphic; anthers
linear, 3—3.5 or 2.5—3 mm. long, the connective prolonged at base below the thecae
into two deflexed lateral lobes; style slightly dilated upward to the truncate stigma.

Abundant, widely distributed, and highly variable throughout tropical America
from southern Mexico and the West Indies to Paraguay and Bolivia.

B L TORO: Fish Creek Mountains, von Wedel 2293. CANAL ZONE: Chagres,
Fendler 32; Rio Gatún, Hayes 390. DARIÉN: Garachiné, Pittier 5478.

16. MICONIA PTEROPODA Benth. in Hook. Journ. Bot. 2:314. 1840.

Shrub to 9 m. tall, the younger parts of the stem, panicle, and hypanthia
tomentulose with minute brown stellate hairs; leaves firm in texture, narrowly
oblong-elliptic, up to 25 cm. long, a fifth to a third as wide, acuminate to 2 blunt
tip, entire or somewhat undulate, abruptly cuneate below into a narrow wing
extending to the stem, strongly 5-pli-nerved, at maturity glabrous above and on
the surface beneath, minutely furfuraceous along the veins beneath; panicle 1-2
dm. long; flowers sessile; hypanthium cup-shaped, 2.5-3 mm. long to the torus;
calyx-tube prolonged about 0.5 mm., its lobes depressed-semicircular, about 02
mm. long from the sinuses; petals obovate, about 3 mm. long, densely furfura-

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FLORA OF PANAMA (Melastomataceae) 291

ceous; stamens slightly dimorphic; anthers 3.2—3.7 mm. or 2.6-3 mm. long, linear;
connective prolonged into two lateral lobes at base, slender and rounded on the
smaller anthers, on the larger flat and truncate; stigma truncate. Perhaps not
distinct from the preceding species.

British Honduras to Panama; widely distributed in northern South America.

CANAL ZONE: vicinity of Summit, 150 m., Allen 4519. cHIRIQUI: Boquete, Davidson
679, 8 38; San Félix, Pittier 5258. COCLE: south of El Valle de Antón n, 600-800 m., Allen
2852; E Las Margaritas and El Valle, Woodson, Allen & Seibert 1307; Binari
above é, 600-900 m., Williams 552. PANAMA: Bella Vista, Macbride 2742; San
José land, „Jobnizow 760, 770, IIII, Prieto 89, 265.

17. MICONIA HYPERPRASINA Naud. in Ann. Ści. Nat. Bot. III. 16:186. 1851.

Shrub or small tree to 8 m. tall, the younger stems, petioles, inflorescence, and
hypanthia thinly but closely stellate with minute brownish hairs; petioles 1-2 cm.
long; leaf-blades commonly oblong-lanceolate, varying to oblong or somewhat
oblanceolate, 1-2 dm. long, a fourth to a third or rarely half as wide, acuminate,
entire or undulate, acute or somewhat obtuse at base, glabrous above, glabrous or
very sparsely furfuraceous along the veins beneath, 3-nerved, often also wit
conspicuous marginal veins, the veins beneath often pale; panicle widely branched,
up to 10 cm. long, always minutely setulose at the nodes, bractless; flowers sessile;
hypanthium'cup-shaped, about 2 mm. long to the torus; calyx-lobes nearly semi-
circular, 0.5—0.8 mm. long; exterior teeth small, conic; petals white, obovate, 2—2.5
mm. long; anthers dimorphic; anthers about 3 or about 2.5 mm. long, the con-
nective of the larger expanded at base into a large lobe with its sides curved around
the filament and its margins meeting beneath, that of the smaller prolonged at base
into a narrow short dorsal lobe and two minute lateral lobes; stigma truncate.

Southern Mexico to Panama.

wg > Neun 434; Frijoles, Piper 5 © enono: ‘
: Cana and vicinity, ya m., Williams 98I. PANAMA: San José Island,
Joon 668, 762, 1133, Erlanson 88, 96, 138, 229, 206. SAN BLAS: Permć, Cooper 635,

18. MICONIA INSULARIS Gl. in Sargentia 8:234. 1949.

Shrub to 4 m. tall, the younger branches minutely and sparsely stellate-canes-
cent; petioles 1-2 cm. long; leaf-blades elliptic or oblong, 10-16 cm. long, about
two-fifths as wide, acuminate to a blunt tip, entire or somewhat sinuate, acute at
the base, glabrous and puncticulate above, glabrous beneath, 3-nerved; panicle
about 1 dm. long and usually somewhat wider, branched from the base; flowers
sessile or very shortly pediceled; hypanthium 1.3 mm. long to the torus, sparsely
and minutely stellate; calyx-tube prolonged 0.4 mm., truncate, the minute exterior
teeth scarcely projecting; petals white, oblong, 3-3.5 mm. long; anthers dimorphic,
about 3 or about 2.5 mm. long, the connective of the larger expanded at base into
a large dorsal lobe with its sides deflexed around the filament, that of the smaller

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292 ANNALS OF THE MISSOURI BOTANICAL GARDEN

prolonged at base into one dorsal and two lateral lobes about equal in size; stigma
truncate.

Endemic to Panama.

PANAMA: San José Island, Jobnston 448, Erlanson 403, 573.

19. MICONIA NERVOSA Ta Triana, in Trans. Linn. Soc. 28:111. 1871.
Melastoma nervosum Sm. in Rees, Cycl. 23: Melastoma n. 31. 1819.
Miconia septuplinervia Pittier, in Bol. Soc. Venez. Cienc. Nat. 11:26. 1947.

Shrub to 6 m. tall, the younger stems, petioles, panicle, and primary nerves of
the leaves densely strigose; petioles 1-2 cm. long; leaf-blades thin, elliptic or
oblong, up to 25 cm. long, a third to half as wide, often unequal in each pair,
acuminate or rarely acute, entire or nearly so, long-cuneate at the base, 5- to 7-pli-
nerved, the inner pair of lateral nerves arising 2—8 cm. from the base, sparsely
pubescent on the upper surface, softly pubescent or subsericeous on the lower;
panicle commonly elongate, 5-15 cm. long, the flowers mostly sessile in lateral
glomerules, occasionally on short lateral branches; hypanthium narrowly cam-
panulate, 3.5—4 mm. long, densely strigose; calyx-tube about 0.5 mm. long; sepals
minute, triangular; petals white, oblong-obovate, 3—3.5 mm. long; stamens
isomorphic; anthers linear, straight, 3.5—4 mm. long, the connective prolonged
straight back into a single dorsal appendage 0.2—0.3 mm. long; ovary pubescent;
style about 6 mm. long; stigma truncate.

In regions of heavy rainfall, southern Mexico to Panama; southward to Peru
and eastward through the Amazon valley.

AS DEL TORO: Water Valley, von Wedel 804, 1425, 1467; Changuinola Valley,
Dunlap, I 131; Talamanca Valley, Carleton 131. CANAL ZONE: Chagres, Fendler 35; Gatún,

CHIRIQUi: San Félix, Pittier 5175. DARIÉN: Cana-Cuasi trail, Chepigana
dirci 660 m., Terry 6 Terry 1521. PROVINCE UNKNOWN: Moore 23.

20. MICONIA ALBICANS (Sw.) Triana. Trans. Linn. Soc. 28:116. 1871.
Melastoma albicans Sw. Prodr. Veg. Ind. Occ. 70. 1788.

Shrub or small tree to 6 m. tall; younger branches finely cinereous-tome entulose
soon glabrescent; petioles stout, tomentulose, 5-15 mm. long; leaf- blades ub
coriaceous, oblong, ovate-oblong, or elliptic, 7-15 cm. long, about two-fifths as
wide, obtuse to short-acuminate, entire, emarginate or subcordate at base, 5- nerved,
the outer pair commonly submarginal, above when mature glabrous, dark green,
and shining, beneath closely and finely cinereous-tomentulose; panicle 8-20 cm.
long, freely branched, cinereous or ferruginous; flowers 5-merous, sessile, secund
and alternate on the ultimate branches of the panicle; hypanthium about 2 MM:
long, densely tomentose; calyx-tube prolonged about 0.5 mm., its lobes very short,
triangular; petals obovate, about 3 mm. long; stamens nearly isomorphic; COP-
nective briefly prolonged below the thecae and dilated into an oblique cordate
organ; stigma capitate.

Widely distributed from southern Mexico and the West Indies to Bolivia and
southern Br

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FLORA OF PANAMA (Melastomataceae) 293

CHIRIQUÍ: Boquete, 1200 m., Davidson 678. cocLE: between Las Margaritas and El
Valle, Woodson, Allen & Seibert 1250. PANAMA: in llanos at Chorrera, Wbite 178; Isla
Taboga, Woodson, Allen 6 Seibert 1445; San José Island, Erlanson 230, 283. VERAGUAS:
trail between Cafiazas and the Cordillera Central, 300-600 m., Allen 204.

21. MICONIA PANAMENSIS Gl. in Phytologia 1:341. 1939.

Shrub, the younger stems and petioles very thinly and sparsely stellate-furfura-
ceous, eventually glabrescent; petioles stout, 3-5 mm. long; leaf-blades firm in
texture, ovate-lanceolate or oblong-lanceolate, up to 16 cm. long, a third to two-
fifths as wide, long-acuminate, entire, rounded to subcordate at the base, 3-nerved
with an additional pair of submarginal nerves, glabrous above, very sparsely stellu-
late on the primaries beneath; panicle many-flowered with ascending branches, to
15 cm. long; flowers sessile; hypanthium campanulate, 2 mm. long to the torus,
very thinly and minutely stellulate; calyx-tube prolonged 0.4 mm., sepals tri-
angular, 0.2 mm. long from the sinuses; exterior teeth minute conic thickenings;
petals elliptic, about 2.5 mm. long, white; stamens dimorphic; anthers linear, about
3.5 mm. or 2.7 mm. long, the connective of the larger prolonged below the thecae
into a relatively large, cordate organ, the sides deflexed and antrorse, connective of
the smaller anthers smaller, prolonged into a narrow dorsal lobe and two small
deflexed lateral lobes; stigma capitellate.

Endemic to Panama. Perhaps to be combined with Miconia cbamissois Naud.
Which scarcely differs except in its 3-pli-nerved leaves.

BOCAS DEL TORO: vicinity of Macaw Hills, von Wedel 6, 528.

22. MICONIA RUBIGINOSA (Bonpl) DC. Prodr. 3:183. 1828.
Melastoma rubiginosa Humb. & Bonpl. Melast. 109. £. 47. 1816.

Shrub to 5 m. tall, the stems, petioles, lower leaf-surface, panicle, and hypanthia
tomentose with ferruginous stellate hairs; petioles stout, 5—10 mm. long; leaf-
blades firm in texture, oblong, 8-15 cm. long, two-fifths to half as wide, acuminate,
entire, broadly rounded to subcordate at base, above sparsely stellate when young,
soon glabrescent and somewhat shining, 3-nerved, often with an additional con-
spicuous pair of marginal nerves; panicle up to 15 cm. long, commonly with super-
posed branches; hypanthium campanulate, 2-2.5 mm. long to the torus; calyx-tube
slightly flaring, prolonged about 0.7 mm.; calyx-lobes obscure, depressed-
semicircular; petals obovate, white, about 3 mm. long; stamens dimorphic; anthers
inear, about 3 or about 2.5 mm. long, the connective of the larger prolonged
dorsally and laterally into a heart-shaped organ oblique to the axis, the lateral lobes
antrorse and bearing the filament in the sinus, that of the smaller anthers prolonged
into a minute dorsal lobe and two minute deflexed lateral lobes; stigma truncate.

Puerto Rico and Panama to southern Brazil.

COCLÉ: between Agua Dulce and the Río Chico, near sea level, Pittier 5001; between
Las Margaritas and El Valle, Woodson, Allen & Seibert 1285. panamá: Isla Toboga,
0-186 m., Woodson, Allen & Seibert 1442.

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294 ANNALS OF THE MISSOURI BOTANICAL GARDEN

23. MICONIA RUFOSTELLULATA Pittier, in Journ. Wash. Acad. Sci. 13:390. 1923.

Shrub 1.5 m. tall, the younger stems, petioles, panicle, and hypanthia rufous-
tomentose with stellate, often stipitate hairs; stems later glabrescent; petioles 5-8
mm. long; leaf-blades lanceolate, up to 8 cm. long, a third to two-fifths as wide,
acuminate, entire, rounded at base, 3-nerved, minutely setose above, beneath
stellate densely on the primary veins, very sparsely so on the surface; panicles 2-3
cm. long, terminating short bifoliate divaricate branches; flowers 4-merous, sub-
sessile, each (and each node of the panicle) subtended by a short subulate bract;
hypanthium cup-shaped, about 1.5 mm. long to the torus; calyx-tube very short,
its lobes triangular, acute, 0.7 mm. long from the sinus; exterior teeth adnate
almost to the summit of the sepals, thence divergent, subulate, 0.5 mm. long;
petals white, obovate-oblong, about 2.5 mm. long; stamens dimorphic; anthers
linear, about 2.5 or about 2 mm. long, the connective of the larger broadly dilated
below the thecae into a cordate organ, its sides folded around the sides and meeting
at the apex of the filament; connective of the small anthers scarcely dilated,
prolonged into one minute dorsal and two small lateral lobes; stigma capitate.

Known only from the type collection.

DARIÉN: in clearings around Pinogana, Pittier 6535.

V. Section GLOSSOCENTRUM Triana, ex Benth. & Hook. f. in
Benth. & Hook. Gen. 1:764. 1867.

Anthers linear or slightly dilated distally, opening by a terminal pore about as
wide as the anther; connective variously prolonged or lobed; exterior teeth small
or obsolete; flowers usually small.

About 80 species, chiefly in southern Brazil. Only the following species in
Panama.

24. MICONIA BOREALIS GI. in Bull. Torr. Bot. Club 55:118. 1928.

Shrub or small tree, reported to 15 m. tall, the young stems finely stellate-
tomentulose, soon glabrescent; petioles 5-15 mm. long; leaf-blades lanceolate to
oblong, 8-15 cm. long, a fourth to a third or rarely two-fifths as wide, long-
acuminate, entire, rounded at the base, sparsely stellate on both sides when young;
soon glabrescent, 3- (rarely 5-) nerved; panicle much branched, often as wide as
long, its axes thinly stellate; hypanthium 1-1.5 mm. long, cup-shaped, glabrous
or nearly so; calyx-lobes broadly triangular, only 0.2 mm. long; petals obovate-
oblong, 2-2.5 mm. long; stamens dimorphic; anthers opening by a wide te i
pore, 2-2.5 or 1.5-2 mm. long, the connective of the larger prolonged into a single
flat dorsal lobe wider than the filament, that of the smaller scarcely expanded;
style slightly dilated distally to a truncate stigma.

Southern Mexico to Panama; western Cuba.

CANAL ZONE: Barro Colorado Island, Bangham 486, Shattuck 701; east of Fort Clay-
ton, Harvey 5123; near Miraflores Lake, Harvey 5132, White 251. cHrriqui: San Félix,
Pittier 5138. COCLE: vicinity of El Valle, Allen 776. panamá: west of Chorrera,

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FLORA OF PANAMA (Melastomataceae)

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296 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Woodson, Allen & Seibert 1677, Paul 499; Isla Taboga, Woodson, Allen & Seibert 1462;
Bella Vista, Macbride 2744; San José Island, Jobnston 32, 1097, 1210, 1304. VERAGUAS:
wes 4, Woodson, Allen & Seibert 513; trail between Cańazas and the foot of the
Cordillera Central, 300—600 m., Allen 161. PROVINCE UNKNOWN: Allen 2676.

VI. Section HARTIGIA (Miq.) Griseb. Fl. Br. W. Ind. 258. 1860.
Anthers short, oblong, about as thick as wide, 2-celled, opening by a minute
terminal pore; connective simple; style abruptly bent near the summit.
About 30 species, a few widely distributed, the others chiefly in northwestern
South America. The species have been distributed by earlier students in sections
Tamonea, Miconia, and Amblyarrhena.*

a. Calyx truncate or merely undulate at the summit, glabrous.

b. Leaves green beneath, two-fifths to half as long as wide 25. M. CILIATA.
bb. Leaves red-purple beneath, a fourth to a third as long as wide............. 26. M. OINOCHROPHYLLA.
aa. Calyx conspicuously lobed and ciliate. 27. M. LACERA.

25. MICONIA CILIATA (Rich.) DC. Prodr. 3:179. 1828.
Melastoma ciliata Rich. in Act. Soc. Hist. Nat. Par. 1:109. 1792.

Shrub to 3 m. tall, the younger branches glabrous, or rarely with a few setae
at the nodes; petioles slender, 1-3 cm. long, usually setose near the summit on the
upper side; leaf-blades firm in texture, oblong, varying toward ovate or obovate,
up to 15 cm. long, two-fifths to half as wide, acute or abruptly acuminate, con-
spicuously ciliate and commonly serrulate, glabrous, 3-nerved; panicle pyramidal,
divaricately branched; flowers sessile and secund; hypanthium globose or urceolate,
2-2.5 mm. long; calyx-tube short, erect, truncate; petals about 3 mm. long;
stamens isomorphic; anthers stoutly oblong, 1.5 mm. long, 2-celled, the connective
simple.

Abundant and widely distributed from southern Mexico and the West Indies
to central Brazil and Peru.

CANAL ZONE: Ancón Hill, Killip 3007.

26. MICONIA OINOCHROPHYLLA Donn. Sm. in Bot. Gaz. 40:4. 1905.

Glabrous shrub 1-3 m. tall; petioles stout, 1-3 cm. long; leaf-blades red-purple
beneath, narrowly oblong to lanceolate, 12—25 cm. long, a fourth to a thi
wide, acuminate, entire but ciliate, acute at the base, 3-nerved with a conspicuous
pair of submarginal nerves; panicle small, 5-10 cm. long, sparingly branched;
flowers sessile, secund, subtended by triangular bracts about 1 mm. long; hypan-
thium subglobose, 2—2.5 mm. long at anthesis; calyx-tube very short, truncate or
nearly so around the summit; petals white to pink, oblong-obovate, 2.5-3 mm.

* Stern 8 ine 19 187, both from Pifias Bay, Darién, have recently been referred by
e i e G

3 and
to M. robinsoniana Cogn., a species otherwise en
artificial key to the genus Miconia, this species would fall next to M. bondurensis, differing however
in the quadrate — denser leaf blade venation, secund flowers, and eglandular stamen connectiV
ns i :

n obvious superfluity of epithets, no phytogeographic speculations concerning the apparent distribu-
tional disjunction of M. robinsoniana would be valid at present.

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FLORA OF PANAMA (Melastomataceae) 297

long, oblique; stamens isomorphic; anthers slenderly oblong, 1.7 mm. long, 2-celled;
connective simple; ovary almost completely inferior, 5-celled; style stout, 2.5 mm.
long, abruptly bent near the summit; stigma punctiform.

British Honduras to western Colombia

COCLE: north rim of El Valle de Antón, near Cerro Turrega, 650-700 m., Allen 17.
OLÓN: summit of Cerro Santa Rita, 360-500 m., Allen 5008. PANAMA: Cerro Campana,
600—800 m., Allen 26

Allen 5098 is referred here with some doubt, since it was described by the
collector as a vine and the panicle is more widely branched than in typical plants.

27. MICONIA LACERA (Bonpl.) Naud. in Ann. Sci. Nat. Bot. III. 16:152. 1851.
Melastoma lacera Humb. & Bonpl. Melast. 9. t. 5. 1816.

Shrub to 4 m. tall, the stems, petioles, and panicle more or less hirsute with
purple hairs as much as 10 mm. long; petioles 1-3.5 cm. long; leaf-blades thin,
lanceolate to oblong or ovate-lanceolate, up to 15 cm. long and 6 cm. wide, entire
or serrulate, acuminate, conspicuously ciliate, obtuse or rounded at base, more or
less pilose on both sides, especially on the veins, 3- to 5-nerved; panicle small and
compact, commonly 4-8 cm. long, the sessile flowers secund along the branches;
hypanthium cup-shaped, about 2 mm. long to the torus, glabrous; calyx-tube
prolonged nearly 1 mm.; sepals broadly ovate or nearly semicircular, about 0.5
mm. long from the sinus, conspicuously ciliate with long hairs; exterior teeth
triangular, almost completely adnate, the slender free tip about 0.4 mm. long;
petals white to pink, oblong-obovate, about 4 mm. long, distinctly retuse; stamens
isomorphic; anthers oblong, about 1.5 mm. long, 2-celled, the connective simple;
style slender, bent or hooked at the summit, about 3 mm. long.

Abundant in Central America from southern Mexico to Panama; also in the
West Indies and Trinidad, and southward to Peru

A Hill, Killip Taro; Piper sod, apri 393, Williams ró Barto
Colorado Island Bailey 8 Bai , 376, Shattuck 317, 1072; sag Fendler 33
vicinity of Gatün Lake, Chrysler s Roever 4874, tima 251, 560; Jp orenzo,
Piper 5992; Monk[ey?] Hill, Kun .". CHIR between
tai Woodson, Allen 6 Seibert at "icini inity of El Galle Ales 755% berrien
as Margaritas and El Valle, Wobdps, Allen ie uberi 12733 Bismarck, above Penonomć,
Williams 290, 1159; La Chorrera, Paul 507. NAMA: thickets and forests near Arraijan,

Woodson, Allen 6 Seibert I 352; Cerro Ca węża vicinity of Chica, 600-800 m., Allen
2651 i Trapiche, Perlas pese Allen 2617; Isla Taboga, 0-186 m ., Woodson, Allen

VII. Section AMBLYARRHENA Triana, ex Benth. & Hook. f. Gen. Pl. 1:763. 1867.
Anthers oblong or ovoid, thick and stout, as thick as wide, 4- celled, opening
by a minute terminal pore; style straight; connective in most species not prolonged
below the thecae.
About 130 species, most numerous in the northern Andes mountains from
Colombia to Peru.

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[Vol. 45
298 ANNALS OF THE MISSOURI BOTANICAL GARDEN

a. Connective simple, without lobes or appendages; filaments glabrous;
stigma small, capitate or truncate

ate.
b. Pubescence thin and sparse, of minute stellate hairs 28. M. SCHLIMII.
bb. Pubescence dense, of long simple bristles 29. M. COSTARICENSIS.
aa. Connective prolonged as a short truncate lobe over the summit of the
glandular filament; stigma peltate, conspicuously lobed 30. M. GONIOSTIGMA.

28. MICONIA SCHLIMII Triana, in Trans. Linn. Soc. 28:102. 1871.

Small tree, up to 10 m. tall, the younger stems, petioles, lower leaf-surface,
panicle, and hypanthia thinly stellate-tomentose; petioles about 1 cm. long; leaf-
blades lanceolate, as much as 17 cm. long but usually much shorter, about two-

ths as wide, acuminate, remotely denticulate, more or less ciliate, acute at the
base, 5-pli-nerved, glabrous above; panicles relatively few-flowered, 3-8 cm. long,
divaricately branched; flowers 6-merous, sessile at the end of the branchlets;
hypanthium broadly campanulate, about 3.5 mm. long (measured on the axis);
calyx-tube flaring, about 2 mm. long, truncate or undulate, stellate within and
without; exterior teeth minute, conic; petals white, about 10 mm. long, broadly
obovate; stamens isomorphic; anthers oblong, blunt, 4-celled, 4.5-5 mm. long,
opening by one minute terminal pore; connective simple; ovary 5-celled; style
elongate; stigma capitate.

British Honduras to Colombia.

ocas DEL TORO: Changuinola Valley, Dunlap 103. cHIRIQUI: between Concepción

z

B
d El Volcán, White 312; Progreso, Cooper & Slater 297. cocLé: vicinity of El Valle
de Antón, Allen 1774, 2496; between Las Margaritas and El Valle, Woodson, Allen 8
9

29. MICONIA COSTARICENSIS Cogn. in DC. Monogr. Phan. 7:887. 1891.

Shrub to 4 m. tall, the stems, petioles, leaves, inflorescence, and hypanthia
densely hirsute with simple red-purple bristles up to 3 mm. long; petioles 5-20
mm. long; leaf-blades ovate-lanceolate to oblong-elliptic, up to 15 cm. long, two-
fifths to half as wide, acuminate, obscurely denticulate or crenulate, obtuse to
rounded at base, 5-pli-nerved or sometimes merely 5-nerved; panicles 2-5 cm.
long, congested; hypanthium cup-shaped, about 2.5 mm. long to the torus; calyx-
tube prolonged about 0.5 mm.; calyx-lobes depressed-triangular, about 0.5 mm.
long from the sinus; exterior teeth stoutly subulate, hirsute, about 2 mm. long;
petals pink, about 3 mm. long; stamens isomorphic; anthers oblong, blunt, 4-celled;
stigma truncate.

Costa Rica and Panama.

cHIRIQUI: Bajo Chorro, Boquete district, Davidson 239; Bajo Mono-Robalo trail,
1500-2100 m., Allen 4823.

30. MICONIA GONIOSTIGMA Triana, in Trans. Linn. Soc. 28:124. 1871.

Shrub 1-2 m. tall; all parts of the plant to the hypanthia more or less hirsute
or hirsutulous with slender simple hairs seldom more than 1 mm. long; petioles 3-8
mm. long; leaf-blades oblong or elliptic, up to 10 cm. long, two-fifths to half as
wide, short-acuminate, entire or denticulate, rounded at the base, 5-nerved or

(298)

1958]
FLORA OF PANAMA (Melastomataceae) 299

somewhat 5-pli-nerved; panicle small, sparingly branched and few-flowered;
flowers 4-merous or 5-merous, sessile; hypanthium broadly cup-shaped, 2-2.5 mm.
long to the torus; calyx glabrous, the tube prolonged 0.4 mm., the lobes semi-
circular, 0.7 mm. long from the sinus; exterior teeth reduced to minute thicken-
ings; petals subrotund, 3.5-4 mm. long; stamens isomorphic; filaments very broad,
glandular on the outer face at the summit; anthers oblong, 4-celled, nearly 2 mm.
long, opening by a dorso-terminal pore; connective widened toward the base and
projecting as a short truncate body over the summit of the filament; ovary com-
pletely inferior; stigma peltate, 2.5 mm. wide, 4 to 5-angled or lobed.
Panama to Ecuador.

DARIÉN: Cana-Cuasi trail, Chepigana district, 600 m., Terry & Terry 1516.

VIII. Section CREMANIUM (Don) Triana, ex Benth. & Hook. f. Gen. 1:764. 1867

Anthers oblong to obovate or obovoid, often flattened, commonly widest above
the middle, 2-celled or 4-celled, opening by a wide terminal pore through which
the septum separating the thecae commonly protrudes; filaments usually perman-
ently geniculate above the middle.

ut 150 species, throughout the range of the genus but most numerous in
the Andes mountains from Colombia to Bolivia.
a. Actes the two thecae separated by a radial septum, each theca
al sept

also divided by a tangential septum; septa extending into the terminal
= m 31. M. THEAEZANS.

pore and divid g
aa. Anthers 2-celled or 4-celled only below the middle, only the radial
septum extending into the terminal pore and dividing it into two
segments.
b. Connective at base elevated into a dorsal protuberance and prolonged
around the base of the thecae into two lateral lobes.
Pubescence of simple unbranched hairs 32. M. PITTIERI.
: 33. M. CUSPIDATISSIMA.

a

. rs
bb. Connective at base widened and thickened but not lobed, prolonged

34. M. RUBENS.

31. MICONIA THEAEZANS (Bonpl.) Cogn. in Mart. Fl. Bras. 14*:419. 1888.

Melastoma theaezans Humb. & Bonpl. Melast. 17. f. 9. 1816.

Younger stems more or less sparsely stellate-pubescent, commonly glabrous at
maturity; petioles slender, 1-2 cm. long; leaves varying from lanceolate to obovate-
oblong, as much as 15 cm. long, commonly smaller, always acuminate and entire,
commonly acute but varying to obtuse or even subrotund at base, 3-nerved, rarely
3-pli-nerved, glabrous above, glabrous beneath or retaining more or less stellate
Pubescence along the veins; panicle 5-15 cm. long, glabrous to stellate-pubescent,
widely branched; flowers mostly short-pediceled; hypanthium cup-shaped, glabrous
or sparsely stellate, 2-3 mm. long; calyx-tube briefly prolonged, its lobes depressed-
semicircular to triangular, 0.3-0.5 mm. long from the sinus; exterior teeth minute
conic thickenings; petals white, broadly elliptic to subrotund, commonly somewhat
inequilateral and retuse; stamens isomorphic; filaments flat, gradually narrowed to
the summit, geniculate above the middle; anthers obovate or obovate-oblong,

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[Vol. 45
300 ANNALS OF THE MISSOURI BOTANICAL GARDEN

flattened tangentially, opening by a wide terminal pore, 4-celled, the septa at right
angles to each other and commonly visible through the pore; connective prolonged
0.3—0.5 mm. below the thecae, elevated into a prominent dorsal protuberance and
extending below the thecae into two lateral lobes; stigma capitate.

Southern Mexico and the West Indies to southern Brazil and Bolivia. A highly
polymorphic species, variable in size and shape of leaves and somewhat in size of
flowers but constant in the structure of its anthers; Cogniaux distinguished
seventeen varieties.

BOCAS
Bajo Chorro, 1800 m., Davidson 96, 97, 344; Chiquero, Boquete district, Davidson 507;
Cerro Horqueta, 1950 m., von Hagen 2014; valley of the upper Rio Chiriqui Viejo, White
& White 117. COCLE: crest of Cerro Pajita, 1200 m., Allen 3955.

32. MICONIA PITTIERI Cogn. in DC. Monogr. Phan. 7:1191. 1891.

Tall shrub, the stems densely retrorse-strigose with brownish hairs; petioles 1-3
cm. long, densely hirsute; leaf-blades ovate-oblong, up to 13 cm. long and about
half as wide, short-acuminate, obscurely denticulate, obtuse to subrotund at base,
5-nerved, densely pubescent with simple hairs on both sides; panicle up to 17 cm.
long, densely pubescent; pedicels about 1 mm. long; hypanthium cup-shaped, about
2 mm. long to the torus, densely hirsute, the hairs 1—1.5 mm. long; calyx-tube
very short, its lobes semicircular, about 0.5 mm. long from the sinus, barely
surpassed by the triangular-acuminate exterior teeth; petals white, suborbicular,
inequilateral, retuse, 1.6 mm. long; stamens isomorphic; anthers obovoid-oblong,
1.6 mm. long, opening by a wide terminal pore, the connective elevated at base
dorsally and prolonged 0.3 mm. below the thecae into two lateral lobes; stigma
capitate.

Costa Rica and Panama.

CHIRIQUÍ: vicinity of Bajo Chorro, 1900 m., Woodson & Schery 630.

33. MICONIA CUSPIDATISSIMA Pittier, in Journ. Wash. Acad. Sci. 14:449. 1924.

Small tree, reported up to 7 m. tall, the younger parts, petioles, panicle, and
hypanthium thinly tomentose with stellate hairs; petioles 1-2 cm. long; eaf-blades
oblong to oblong-elliptic, up to 15 cm. long, two-fifths to half as wide, abruptly
narrowed above into a caudate tip 1-2 cm. long, entire, obtuse to rounded at base,
3-nerved with an additional pair of marginal nerves (in one specimen 5-pli-
nerved!), glabrous above, stellate-tomentose on the primary nerves beneath, thinly
so on the veinlets, glabrous or nearly so on the surface; panicle to 15 cm. long,
widely branched; actual pedicels about 1 mm. long; hypanthium cup-shaped, 2-2.5
mm. long to the torus; calyx-tube prolonged about 0.4 mm., the lobes semicircular,
about 0.5 mm. long from the sinus, surpassing the triangular adnate exterior teeth;
petals white, broadly obovate, 2—3 mm. long; stamens isomorphic; anthers oblong
or oblong-obovate, about 2 mm. long; connective dilated toward the base, elevated
into a conspicuous dorsal protuberance, and prolonged below the thecae into two
deflexed lateral lobes; stigma capitate.

Endemic to Panama, so far as known.

(300)

1958]
FLORA OF PANAMA (Melastomataceae) 301

pumau; Bajo Chorro, Boquete district, 1800 m., Davidson 387, 715; between Alto
almas and Cerro de la Horqueta, 2100—2268 m., Pittier 3224; upper Caldera
Micerhed, above El Boquete, 1650-1925 m., Maxon 5654.
34. MICONIA RUBENS (Sw.) Naud. in Ann. Sci. Nat. Bot. III. 16:169. 1851.
Melastoma rubens Sw. Prodr. Veg. Ind. Occ. 71.
Topobea micrantba Pittier, in Journ. Wash. Acad. "er 14: 451. 1924.

Shrub or tree, reported to 18 m. tall, glabrous or essentially so throughout;
petioles slender, 1-2 cm. long; leaf-blades thin, elliptic or oblong, varying to
obovate-oblong, 3-10 cm. long, a third to half as wide, conspicuously acuminate
or caudate to a blunt point, entire, acute, obtuse, or cuneate at base, 3-pli-nerved;
panicle small, 3—6 or rarely 10 cm. long, many-flowered; hypanthium cup-shaped,
1.5-2 mm. long to the torus; calyx-tube scarcely prolonged, its lobes semicircular
or depressed-triangular, about 0.5 mm. long; exterior teeth varying from a minute
thickening to conic, subulate, and 0.5 mm. long; petals white, suborbicular, about
2 mm. long; stamens isomorphic; anthers oblong, about 0.7 mm. long; thecae about
0.4 mm. long, opening by a large ventro-terminal pore; connective conspicuously
prolonged below the thecae, more or less dilated, not lobed.

Jamaica, Costa Rica, and Panama.

Qui: vicinity of Bajo Chorro, 1800-1900 m., Davidson 181, 334 390, Woodson
E sebery 665; Bajo Mono-Robalo trail, 1500-2100 m., Allen 4843; La Peńa Blanca, 1650
; between Alto de las Palmas and Carro de la torna. 2100-2268 m.,

Pittier - dU; Cerro Horqueta, 1950 m., von Hagen 2042.

IX. Section CHAENOPLEURA (Rich. ex DC.) Triana, ex
th. & Hook. f. Gen. 1:764. 1867.

Anthers oblong to obovate or obovoid, the terminal pore tending to elongate
toward the base into clefts.

A section of doubtful validity. Most species may possibly be regarded as
members of section CREMANIUM with large ventro-terminal pore. About 50
species have been referred here, chiefly from the West Indies and the Andes moun-
tains. Only the following species in Panama.

35. MICONIA MELANOTRICHA dics Gl. in Brittonia 3:101. 1939.
Clidemia melanotricba nes in Trans ;28:136. 1871.
Miconia atrosanguinea Cogn. in Engl. a yc 8: 23. 1886.

Shrub to 2 m. tall, the stem densely pubescent or hirsute; petioles 1-3 cm.
long, pubescent like the stem; leaf-blades dark green, ovate or ovate-lanceolate, 5-2
cm. long, two-fifths to half as wide, acuminate, rounded or subcordate at base,
pubescent on both sides, 5-nerved; panicle trichotomous from the base, broader
than long, 1-2 dm. wide, with linear bracts at each node; hypanthium cup-shaped,
3.5-4 mm. long to the torus; sepals oblong, rounded at the summit, about 1.5 mm.
long; petals deep red, 6-8 mm. long; stamens isomorphic; filaments 13-18 mm.
long; anthers oblong, 3-3.5 mm. long, each theca dehiscing by a longitudinal slit;
style 15-25 mm. long; stigma capitate.

(301)

[Vol. 45
302 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Costa Rica, Panama, and western Colombia. The species exists in Panama in
two well-marked varieties.

Var. MELANOTRICHA. Stem densely hirsute, the hairs 3-4 mm. long; hairs of
the upper leaf-surface 3-4 mm. long, those of the lower surface 1-1.5 mm. long;
panicle and hypanthium long-hirsute like the stem.

CHIRIQUÍ: Bajo Chorro, Boquete district, 1800 m., Davidson 75; vicinity of Cerro
Punta, 1800 m., Allen 3489; Bajo Mono-Robalo trail, 1500-2100 m., Allen 4819.

Var. panamensis Gl. var. nov. Caules densissime pubescentes pilis vix 0.5
mm. longis eis superficie superioris foliorum usque 1 mm. longis inter se vix 0.2
mm. distantibus eis superficie inferioris similiter densis sed tantummodo 0.2 mm.
longis.

CHIRIQUi: above El Boquete, 1650-1925 m., Maxon 5653 (Herb. N. Y. Bot. Gard.
HOLOTYPE) ; Bajo Mono-Robalo trail, 1500-2100 m., Allen 4817.

SECTION UNKNOWN
36. MICONIA SHATTUCKII Standley, in Contr. Arn. Arb. 5:119. pl. 16. 1933.

A shrub or small tree; younger stems, petioles, and panicle thinly pubescent
with very short simple hairs; petioles stout, 1-2 cm. long; leaf-blades broadly
elliptic, up to 3 dm. long, half to two-thirds as wide, short-acuminate, entire,
rounded below to a shallowly cordate base, 7-nerved, glabrous above, sparsely and
shortly pubescent beneath; mature inflorescence paniculiform, about 1 dm. long;
flowers not seen, but, fide Standley, on pedicels 1-2 mm. long; petals white, 4 mm.
long; anthers linear-attenuate, biporose; berries depressed-globose 4.5 mm. diam.

Apparently endemic to Panama.

CANAL ZONE: Barro Colorado Island, Shattuck 335.

Terry 6 Terry 1444, an immature plant from rain-forest on the Cana-Cuasi
trail, Darién Province, altitude 600 m., is questionably referred here also.

33. MOURIRI Aubl.
MOURIRI Aubl. Pl. Guian. 1:452. 1775.

Flowers 5-merous. Hypanthium obconic to pyriform or cup-shaped. Sepals
commonly coherent in the bud, at anthesis separating nearly or quite to the torus.
Petals mostly yellow, ovate to lanceolate, acute or acuminate, often very unsym-
metrical. Stamens isomorphic; anthers stout, blunt; thecae much shorter than the
anther, opening by a lateral longitudinal cleft on both sides of the anther; con-
nective thickened toward the base, bearing a conspicuous dorsal gland near
middle, often prolonged at base into a short, stout, dorsal spur. Ovary wholly
inferior, 1- to 5- celled; stigma punctiform; ovules few in each locule, basal or
nearly so. Fruit a berry. Trees or shrubs, usually glabrous, with 1-nerved (in our
species) leaves and small axillary cymes.

About 65 species, distributed throughout tropical America, mostly at low
elevations and most numerous in Amazonia.

(302)

1958]

FLORA OF PANAMA (Melastomataceae) 303

a

i
|
/
|
\
|
y LI

3 7

21

N

N

= ANI
1

Fig. 93. Mouriri parvifolia

(303)

[Vol. 45
304 ANNALS OF THE MISSOURI BOTANICAL GARDEN

a. Leaves sessile, up to 7 cm. long; hypanthium and sepals each 2.5-3 mm.
ong 1. M. PARVIFOLIA.
aa. tame = petioled, 10-25 cm. long; hypanthium and sepals each 5—6

b. gą ‘blades papyranenss, the tertiary veins evident; hypanthium

rown-t tulose 2. M. BRUNNEICALYX.
bb. Leaf Turis NE the tertiary veins not visible; hypanthium
(fide Pittier) glabro 3. M. COMPLETENS.

1. MOURIRI PARVIFOLIA Benth. Bot. Voy. Sulphur 97. £. 36. 1844.

Shrub 2-6 m. tall with divaricately branched twigs; leaves sessile, lanceolate
or ovate-lanceolate, 3-7 cm. long, a third to half as wide, broadest muc ow
middle, acute or acuminate, rounded at base; secondary veins obscure or obsolete;
flowers few in a fascicle; pedicels 2—4 mm. long, bracteolate near the summit;
hypanthium glabrous, 2.5-3 mm. long; sepals narrowly triangular, to 3 mm. long;
petals 5-6 mm. long.

Southern Mexico to Bolivia.

CHIRIQUÍ: Cierito, gei & Slater 248. coc mountains beyond Pintada, 400-
600 m., Hunter & Allen ; Penonomé and vic midi > Williams 84, 579. PANAMA: Rio
Las Ln. 20 m., Allen n lower Rio Cabra, 30 m., Paul 794.

2. MOURIRI BRUNNEICALYX Standley, ex Woods. & Seibert, in Ann. Mo. Bot. Gard.
24:193.

Small tree to 7 m. tall; petioles 2-3 mm. long; leaf-blades thin, oblong or barely
ovate-oblong, 1-2 dm. long, a third to two-fifths as wide, rounded at the summit
to a triangular cusp 5-10 mm. long, broadly rounded at the base; secondary veins
prominulous; flowers apparently solitary, on a glabrous peduncle 8 mm. long ‘and
a shorter tomentulose pedicel; hypanthium brown-tomentulose, 6 mm. long; sepals
triangular-ovate, 5 mm. long; petals at least 8 mm. long.

Endemic to Panama.

PANAMÁ: vicinity of Arenoso, 26-50 m., Seibert 609.

3. MOURIRI COMPLETENS (Pittier) Burret, in Notizbl. 11:151. 1931.
Aulacocarpus completens Pittier, in Smiths. Misc. Coll. 63, No. 4:2. 1914.

Tree to 18 m. tall, glabrous; petioles 4-5 mm. long; leaf-blades coriaceous,
oblong to ovate-oblong, 14-25 cm. long, 5-11 cm. wide, abruptly short-acuminate
at the apex, shortly cordate at the base, the secondary veins subprominulous, the
tertiaries not visible; flowers solitary or aggregate on bibracteolate pedicels 12-15
mm. long; hypanthium and sepals each about 5-6 mm. long; petals pink, about
10 mm. long.

Panama and (fide Morley) Colombia.

El Real de Sta. Maria, Pittier 6074; Pinogana, Pittier 6573. SAN n
Pana OR. Pittier 4310.

(304)

MISSOURI BOTANICAL GARDEN

STAFF
Director
Frits W. WENT
Oscar E. GLAESSNER,
ller

EDGAR ANDERSON,
Curator of Useful Plants Contro.

HENRY N. ANDREWS, Norton H. NICKERSON,
Paleobotanist Pte st

HucH C. CUTLER, TRIFON VON SCHREN
Executive Director Assistant Curator via of

Economic Plants
CARROLL A Dopce,
dda JULIAN A. STEYERMARK,
ROBERT L. DRESSLER, Honorary Research Associate

sheer nas and È Editor iran B. VAN SCHAA
f the ANNALS urator of Grasses and Pia
q... D. Dwre : ROBERT E. ROSA: Jr.
earch Ad pa of Herbari

BOARD OF TRUSTEES

President
ROBERT BROOKINGS SMITH
Vice-President
Leicester B. FAUST
Second Vice-President
y B. PFLAGER
Jonn S. LEHMANN
J. Lock woobp
Warren McK. SHAPLEIGH

DANIEL K. CATLIN
“DUDLEY FRENCH
HENRY HITCHCOCK

EX.OFFICIO MEMBERS
STRATFORD LEE MORTON;

THUR C. LICHTENBERGER,
Bishop of the Diocese of President of the ao» of
Missouri Science of St. Loui
JAMES J. MCCAFFERY, a ETHAN A. H. SHEPLEY,
: ington

President of the Board of
Education of St. Louis

Chancellor of Washin
University
RAYMOND R. TUCKER, .
Mayor of the City of St. Louis

Oscar E. GLAESSNER, Secretary

FLORA OF PANAMA

Part II
Fascicle 1.
Fascicle 2.
Fascicle 3.

Part III
Fascicle 1.
Fascicle 2.
Fascicle 3.
Fascicle 4.
Fascicle 5.

Fascicle 3.
Part VII
Fascicle 1.
Fascicle 2.
Fascicle 3.

Fascicles Published to Date

Cycadaceae to Gramineae (June, 1943)
Cyperaceae to Cyclanthaceae (September, 1943)
Araceae to Pontederiaceae (March, 1944)

Juncaceae to Marantaceae (February, 1945) Out of Print
Orchidaceae, first part (March, 1946) Available only in sets. —
Orchidaceae, second part (December, 1946) Out of Print
Orchidaceae, third part (March, 1949)

Orchidaceae, conclusion (June, 1949) Ont of Print

Piperaceae (March, 1950)

Lauraceae to Cracifers: (March, 1948)
Resedaceae to Leguminosae, first part (June, 1950)
Leguminosae, second part (March, 1951)

Passifloraceae to Cactaceae (March, 1958)
Thymelaeaceae to Myrtaceae (May, 1958)
Melastomataceae (September, 1958)

Volume XLV Number 4

Annals
of the

Missouri Botanical
Garden

Annals
of the

Missouri Botanical Garden

A Quarterly Journal containing Scientific Contributions from the
Missouri Botanical Garden and the Henry Shaw School of Botany of
Washington University in affiliation with the Missouri Botanical
Garden.

Information

The THE URI BOTANICAL GARDEN appears four times
during the calendar year: February, May, September, and November. Four
numbers constitute 2 volume

Beginning with Volume 45, 1958:
__. $12.00 per volume

Single Numbers ________ 3.00 each

Contents of previous issues of the ANNALS OF THE MISSOURI BOTANICAL

GARDEN are listed in the Agricultural Index, published by the H. W. Wilson
Company.

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Issued January 6, 1959.

Annals
of the

Missouri Botanical Garden

Vol. 45 NOVEMBER, 1958 No. 4

SYNTHYRIS RANUNCULINA
F. W. WENT

On recent trips to the Charleston Mountains in Southern Nevada I found a
plant which I could not possibly identify nor even tentatively assign to a family.

The plant is an herb, growing at about 3,000 m. altitude in moist places; it
has a short perennial stem with leaves placed in a rosette. The leaves are round in
outline, reniform, with lobed margin, and the whole habit very much resembles
Heuchera rubescens, which is a common plant at the same altitude in the Charleston
area. There are one or two inflorescences per rosette, shorter or slightly longer
than the leaves; the peduncle is scapose and carries a short ear-like inflorescence.
The flower buds are enclosed between the green bracts, which close tightly into an
almost globose structure. Each bract carries an almost sessile flower, which has 2
sepals which are slightly united at the base. The 3 light-blue strap-shaped petals
are entirely separate; between the single wider abaxial and the two adaxial petals
2 stamens are inserted, and the ovary is two-celled with one style. The fruit is
two-valved, with several seeds per locule, and resembles that of Veronica.

This latter fact should have made me look among the Schrophulariaceae, but
neither the habit, nor the structure of the inflorescence, nor the flower structure,
made this a likely family. Looking through the figures in Abrams Illustrated
Flora of the Pacific States, several species of Synthyris were found to have the
habit, inflorescence and fruit characteristics of the Charleston Mountains plant,
which, it turned out, had been described by Pennell (1933) as Syntbyris ranun-
culina. He had seen only fruiting material (the plant occurs in areas covered
until June or July with snow, apparently flowers immediately after the snow
melts, and toward the end of July or in August, when most collecting is done, has
only fruits) and thus had not noticed that the flower structure of this plant
differs radically from the other described Synthyris species. They are supposed to
have 4 calyx lobes, a corolla tube with 4 lobes, 2 stamens and 2 carpels.

Since the Charleston Mountain Synthyris differed so extensively in flower
characters from the description of this genus, I inspected specimens of all Synthyris
species described, which I found in the herbarium of the Philadelphia Academy of

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306 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Sciences (my thanks are due to Dr. Smith who helped me and who had photo-
graphs prepared of all species). It became immediately clear that although
Synthyris ranunculina differs radically in its flower morphology from the better-
known species, there are species with intermediate flower characters, in that S.
stellata and S. laciniata both have only 3 corolla lobes. This poses interesting
evolutionary problems, since S. ranunculina occurs at the extreme edge of the area
of distribution of Synthyris, and the nearest species in habit, in flower characters,
and in geographical distribution is S. laciniata from Southwestern Utah.

To see the evolutionary history of Synthyris ranunculina in perspective, I shall
briefly summarize Pennell’s revision (1933) of the genera Synthyris and Besseya.

Syntbyris and Besseya are certainly closely related to Veronica, a Scrophulari-
acea. Typically they have, like Veronica, only two stamens, inserted at the base
of a short tubular or rotate blue corolla, with a flattened two-valved capsule. The
main difference from Veronica lies in their vegetative development: on a short
upright rootstock the long-petioled leaves are inserted close together, and the
flowers are placed in axillary scapose inflorescences. Mostly they occupy moist
places in sub-alpine, montane or arctic localities on the North American continent,
resembling Saxifragaceae such as Heuchera and Mirella in habit and habitat.

Syntbyris and Besseya differ mainly in vegetative characters. The leaves of
Besseya are cordate and ovate or oblong with palmate venation. They are usually
serrate, and the peduncles are more or less densely covered with sessile bract-like
leaves. In Synthyris the leaves are cordate-reniform with palmate venation, and
they are doubly crenate and more or less deeply cleft; the inflorescences are scapose,
and the flowers in the ear-like spikes are subtended by bracts exceeding the flower
in length.

Besseya occurs in the Rocky Mountain region and further eastward, with one
species (B. rubra) extending from Montana west to Eastern Washington and
Oregon, and two (B. arizonica and B. gooddingii) extending into N.E. Arizona.
It seems doubtful that the latter two species are distinct, and also B. plantaginea
does not seem specifically separated from these two. In the genus Besseya, whi
forms a natural group of species, an interesting reduction in flower parts occurs:
Whereas 7 of the 9 species have a white, or light or dark purple corolla (presum-
ably all with 4 lobes, of which the posterior one is enlarged to a lip), in the two
species which extend most northwesterly (B. rubra and B. cinerea) the corolla is
lacking. In the former the calyx is 4-lobed, but in B. cinerea this is reduced to
2 lobes.

As far as the flower characters is concerned, Besseya alpina, through its purple
flower color, and the relatively few bracts on the inflorescence (4-6), seems to
be nearest the genus Syntbyris, and occupies a central position in the distribution
pattern of the genus, predominantly in Colorado in the alpine zone of the Central
Rocky Mountains, with occurrences in S. Wyoming, N. New Mexico and E. Utah.
A group of 5 species, which seem to differ only slightly from each other, mainly
in not well-defined quantitative characters (apparent thickness of leaf blade, which
in the specimens I have seen seems to be a function of the method of pressing
specimens; more or less elongated leaves; size of flowers, with overlapping ranges)»

1958]
WENT—SYNTHYRIS RANUNCULINA 307

occurs to the South-West of the range of Besseya alpina. Three of these are also
alpine, occurring between 3000 and 4000 m. (B. ritteriana, B. oblongifolia, B.
gooddingii), the other 2 species (B. plantaginea and B. arizonica) are montane
(2000-3000 m.). In flower characters they seem to be very much alike.

From this core of Besseya species the other three are clearly separated, both
morphologically and in their distribution. Besseya bullii has flowers like the pre-
ceding species, but has cordate-ovate, palmately veined leaves, which are densely
hirsute. It extends from Minnesota and Iowa eastward to Ohio and Michigan, far
outside the range of the other species of Besseya and Syntbyris, at low altitudes.

The leaves of Besseya cinerea resemble most those of B. bullii in hairyness and
shape, but it lacks a corolla and its calyx is 2-lobed. This species occurs in montane
or alpine localities north of the range of B. alpina, from Nebraska and South
Dakota to Idaho. The other corolla-less species, Besseya rubra, occurs at low
altitudes west of the range of B. cinerea.

The genus Syn£byris is more varied and gives us more indications of evolution.
In contrast to Besseya, which has a very uniform leaf form in all species, leaves in
Syntbyris are of different shape, and there are more variations in flower structure,
although reduction does not proceed as far as in Besseya where the corolla is lacking
in 2 species.

Syntbyris reniformis occurs in forests at low altitude in western Oregon, mainly
in the coastal ranges, extends into Washington and California; it flowers very
early in the year with bright blue campanulate, short 4-lobed corollas. Its leaves
are shaped like those of Viola, ovate or orbicular-cordate. This species stands apart
from the others in the genus, also as regards seed characters (only 2 per locule,
brown instead of yellowish in the other species).

The next two species are vegetatively very much alike, with large reniform
leaves, both occurring on bluffs or cliffs on the border between Washington and
Oregon, S. schizantha occurring in the coastal mountains, S. stellata along the
Columbia river gorge. Their corollas are very different, however. In both the
corolla tube is very short, but S. schizantha has 4 laciniate lobes like S. platycar pa
from Idaho, with the lobes slit over more than half their length, and S. stellata has
3 entire lobes, the larger abaxial one obovate, the two lateral lobes strap-like. These
three species seem to be most closely related to S. missurica, which has the widest
range of distribution of any Synthyris species, occurring at high altitudes through-
out Idaho, in adjacent areas in Washington and Oregon, as far as N.E. California.
The flowers of this species are typical of Syntbyris: a short wide corolla tube, with
4 lobes, of which the abaxial one is widest, with the two stamens inserted at its base.

The other species of Synthyris are all smaller, and grow in high montane or
alpine localities. They can be divided into two distinct groups, one with laciniately
once or twice divided leaves, the other with entire reniform leaves, more (S.
canbyj) or less deeply crenate. The former group consists of a number of very
closely related species, each with a very limited distribution range. It seems
doubtful to me that some of the species which Pennell distinguishes are really
distinct. Thus S. lanuginosa from alpine slopes in the Olympic Mountains of
Washington and S. bendersoni from Central Idaho, both tomentose, seem to be

5 [Vol. 45
308 ANNALS OF THE MISSOURI BOTANICAL GARDEN

conspecific, whereas S. dissecta and S. cymopteroides, both alpine in Montana, both
with long corolla tubes and 4 corolla lobes, also seem the same species. The two other
species with bipinnate leaves are S. paysoni from Wyoming and S. pinnatifida from
the Wasatch Mountains in Utah, both occurring in alpine habitats. The latter is
small, and in size, habit, and shape of inflorescence, very much resembles S. ranun-
culina, but it has a short 4-lobed corolla tu

The remaining 3 species of Syntbyris are ‘all small, with reniform leaves. S.
borealis, villose, occurs in Alaska at timberline. S. laciniata is found in Utah near
snowbanks in the high country of the Wasatch Mountains. It has a wide, almost
campanulate corolla, halfway slit into 3 oblong lobes, sky blue, and has 4 calyx
segments. In very similar localities S. ranunculina grows in the Charleston
Mountains at 3000-3200 m. altitude, mear snowbanks. It has lost its corolla tube,
but still has 3 narrow strap-like light blue petals, the two stamens inserted next
to the sides of the wider abaxial petal. There are only 2 calyx segments.

In figure 1 all the described species of Synthyris are depicted through actual
photographs of herbarium specimens, all at the same magnification. These photo-
graphs are placed on the map of the Western United States in the approximate
place of their occurrence, with only the Alaskan S. borealis out of place. This
figure shows the central position of S. missurica and the vegetatively closely
similar S. schizantha, S. stellata and S. platycarpa towards the North and North-
West. S. reniformis is off to the West. The group of species with bipinnate
leaves is closely spaced in the North-Eastern and Eastern part of the distribution
range, with only S. lanuginosa outside this range. And the small species with
reniform leaves extend toward the NW, NE and South outside the general range
of distribution of the genus. In figure 1 sketches of the flower or of the corolla
of several species are also included, with the number of corolla lobes as far as I
have determined them.

Synthyris and Besseya provide excellent material for an evolutionary analysis.
Since most of the species are montane or alpine, occurring in an area with an
abundance of extensive mountain ranges and isolated mountains, individual popula-
tions have usually been separated for long periods of time, and had time to evolve
as separate entities.

Flower structure places these genera close to Veronica. ‘This genus of the
Scrophulariaceae has about 200 species, and has evolved especially in its vegetative
characters. With the exception of 2 (V. ciliolata and V. pulvinaris from New
Zealand with 5 lobes), all species have 4 corolla lobes and 2 stamens, im implanted
next to the widest lobe. Most species are herbs, but more than one-third are
shrubby or almost tree-like (Section Hebe from New Zealand). In several species
the corolla is tubiform, but in most it is rotate. All have upright stems with
opposite leaves, at least on the lower part of the stem. All species grow in tem-
perate regions or are montane, and practically all have the same sky-blue flower
color.

The genera most closely related to Veronica as indicated by the 2 stamens, the
typical capsule, the sky-blue flower color, and the generally rotate corolla, have a
very different habit: they all have short perennial caudices with rosettes of

1958]
WENT—SYNTHYRIS RANUNCULINA 309

S, lanuginosa
Washington

WASHINGTON

d
N 1
Rie
KŻ S. canbyi MONTANA
>
a A S3

e pe a id
iium A e RY —
Idaho ES ri » È z=

¥

Vu

5 S. paysoni

S. cymopteroides * Wyom. , Idaho
Montana, Idaho
Ne
WYOMING
a :

IDAHO $. hendersoni

4 lobes

Idaho

S. laciniata
3 lobes
` Utah

S. ranunculina
3 free petals
Nevada

Fig. 1. Distribution and Variation in Syn£byris

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310 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long-petioled leaves. In the Old World the genus Wulfenia represents these
Veronica-derivatives, all at high altitudes in Carinthia and in the Himalaya; in the
New World Besseya and Synthyris seem to be direct derivatives of Veronica. As
mentioned earlier, the latter two genera are separated only by vegetative characters:
leaf form and bracts on inflorescence. Even if these differences are not admitted
for generic differentiation, we still have to recognize that there are two or three
lines of differentiation: one of oblong dentate leaves, one of reniform crenate
leaves, and one of pinnatifid leaves. In each line there is a tendency for reduction
in the flowers: in calyx lobes, corolla lobes and petals in general. In addition,
there is a reduction in the number of calyx lobes or sepals. This is of special
interest since generic differentiation usually follows flower characters, while specific
differences often reside in vegetative characters. But in the case of Veronica,
Wulfenia, Besseya, and Syntbyris generic differences are based on vegetative char-
acters, whereas the most significant subsequent evolutionary differentiation of the
species of the latter two genera resides in basic flower characters.

It is usually believed that flower characters are only slightly influenced by
environment, whereas vegetative characters have a strongly adaptive background.
In Syntbyris and Besseya, however, reductions in the flower structure are of specific
significance. But in Besseya the reductions occur only in the species in the north-
western part of the generic range, whereas in Syn£byris these reductions are most
pronounced in the southern limits of the genus, where Besseya does not show flower
reductions. Therefore, both arguments tend to strengthen the conclusion that
the evolution of these genera in the direction of reductions in the flower structure
was not a response to the environment nor due to an environmentally controlled
selection process, but rather that it was in the nature of an orthogenetic process.
In another respect, however, evolution in these genera may have proceeded along
lines of selection by the environment. For in both Besseya and Syntbyris the
major specific development occurs in montane or alpine habitats, but in both
genera species at the northern limit of the genus (with the exception of Syntbyris
borealis) have found habitats near sea level. In both cases the lowland forms are
considerably larger than the alpine or montane species.

Culture of Syntbyris ranunculina in the laboratory has raised the question of
how basic are the differences between the genera Synthyris and Besseya. Normally
Synthyris has a short ear-like inflorescence, with not more than 2 bracts lower

own on the peduncle. But by growing S. ranunculina at very low temperatures
(14° C in light and 4°C in darkness) the peduncles are very short, and the
ear-like part elongates, with bracts over its whole length subtending rudimentary
flower buds. "Therefore, at the lowest temperature where growth is possible these
Syntbyris inflorescences look superficially as if they belong to Besseya.

The condition of evolutionary flux which exists in the flower development of
Syntbyris is reflected in the instability of flower structure in plants grown in
Earhart Laboratory. A plant kept in the shade at 20? day — 14? night temperature

19581

WENT—SYNTHYRIS RANUNCULINA 311
mber Green Petaloid
Observed Sepals Sepals Petals Stamens Carpels

Flowers on Plants
Growing Wild in

Charleston Mountains... Many 2 0 3 2 2
Flowers Developed Two Lower
in Greenhouse... Flowers 2 1 3 3 1?
2 2 1 3 2 2
1 2 2 3 3 Li
1 2 2 + 2 2
1 2 2 455, 2 2

Table I. Number of Flower Parts in Flowers of Syntbyris ranunculina

had flowers of many different structures. In the sepal series there were only one
or two additional ones which were petaloid by being narrow and blue towards the
tip. When there were two extra sepals they were adjacent and abaxial. In one
case a petal was partially laterally adnate to a petaloid sepal, with a small adventi-
tious petal wedged between them at the point of adnation. The number of petals
was unstable, but they were always choripetalous.

Perhaps the most amazing abnormality was the occurrence, in 3 out of 7
flowers, of a third stamen, in the gap between the 2nd and 3rd petal (the 2
normally occurring stamens are implanted at the sides of the adaxial 1st petal).
This instability in stamen number is all the more amazing, since in Veronica and
the other species of Synthyris and Besseya, the stamen number is completely fixed
to 2 and is not known to differ. In 3 of the flowers the ovary seemed reduced to
only a simple carpel, but this could not be established. Table I summarizes the
flower characteristics, showing that there are at least 6 different flower types in
Syntbyris ranunculina, and that probably all conceivable combinations of 2-4
sepals, 3-4 petals, 2-3 stamens and 1-2 carpels can ound.

Summary. A reexamination of the species of Synthyris shows that they can
be placed in definite evolutionary lines, which show both morphological and geo-
graphic progression. Of special interest are the progressive reduction in flower
structure, from 4 corolla lobes towards 3 lobes towards 3 free petals, and from
4 sepals to only 2, with the most reduced forms occurring at the greatest distance
from the geographical center of the genus. In Besseya, a closely related genus,
the reduction of petals is similar, and in two species the corolla has completely
disappeared. Again these reduced forms occupy the geographically remotest
localities,

[Vol. 45
312 ANNALS OF THE MISSOURI BOTANICAL GARDEN
LITERATURE CITED

Abrams, LeRoy. Illustrated Flora of the Pacific States. Vol. III. Stanford Univ. Press.
Gail, F. W. and F. W. Pennell, 1937. A new species of Synthyris from Idaho. Amer. Jour. Bot.
24:39—40.

Pennell, F. W. 1933 A revision of Synthyris and Besseya. Proc. Acad. Nat. Sc. Philadelphia.
85:77—106.

THE NEW WORLD SPECIES OF CYNOMETRA

JOHN D. DWYER!

Cynometra, a Linnean genus of approximately 75 species is widely distributed
throughout the tropics. The New World species have not been studied in detail
since Bentham's limited treatment in Martius Flora Brasiliensis in 1876. The
present study seems particularly timely in view of the current intensive studies
on the African species.

HISTORY OF THE GENUS

Linnaeus established the genus Cynometra in 1741 (Act. Soc. Upsal. 79) and
included it in the first edition of Species Plantarum (p. 382. 1753), describing
two species from India: C. cauliflora and C. ramiflora. Vogel described the first
New World species: C. americana in 1836 (in Linnaea 10:602). In 1840 Bentham
(Jour. Bot. 2:74) established the tribes Cynometreae and Amherstieae, including
three species of Cynometra from tropical America: C. baubiniaefolia, C. marginata,
and C. crassifolia; his subsequent treatment of the genus in Martius Flora Brasili-
ensis (1876) is classical. The well-known Cynometra spruceana was described
therein. Four years before Bentham's initial work, Tulasne described two species
from French Guiana: C. hostmanniana and C. parvifolia (Arch. Mus. Nat. Paris
4:181. 1844). With the exception of C. cubensis described by de la Sagra (Hist.
Cub. 10:233. 1845), all the remaining New World species appeared after 1895.

Recently Léonard (1951) presented an excellent history of the African species.
In this same paper he made an important contribution to the taxonomy of the
African cynometras when he distributed some 60 species, classically assigned to
the genus, among 11 genera. Those species with the stipe of the pistil free in the
receptacle-cup were placed in Cynometra in limited sense and in Gilletiodendron,

INTERGENERIC REATIONSHIP

To establish relationships between Cynometra and other genera is a challenging
problem. If one considers the question on a regional basis, e.g. New World species
only, and accepts the tribe Cynometreae as defined by Bentham as being clearly
distinguishable from the allied tribe Amherstieae, the problem seems narrowed
considerably. It has been traditional to follow this course in relating the New

orld cynometras. The Mexican, tropical American, and West Indian species
have been related to the New World species of Copaifera and to the pantropical
genus, Crudia; the leaves of these genera are multijugate and the flowers are
apetalous, in contrast to Cynometra (New World) with unijugate leaves and 5
petals. While such characters are useful in distinguishing the genera, nevertheless,
additional significant characters tend to make the problem of relationships of
Cynometra on a world-wide scale a formidable task. È

1 Missouri Botanical Garden and St. Louis University, St. Louis, Missouri.

(313)

[Vol. 45
314 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Initially the validity of the tribe Cynometreae (as defined by Bentham) is
jeopardized by the fact that the most fundamental character utilized by its author:
the stipe of the pistil free in the receptacle cup, is not constant for all species of
several genera included in the tribe: Crudia, Cynometra (in wide sense), Schotia
etc.! Léonard, in his recent monograph (1951) of the African cynometras, recog-
nized the inconsistency and divided some sixty African species previously accepted
in Cynometra among 11 genera. Those species with the stipe of the pistil free in
the receptacle-cup were placed in the emended genus Cynometra and in Gilletio-
dendron (5 species). In his later treatment (1957) of the African genera of the
Cynometreae-Amherstieae complex, he studiously avoided stressing this same
character in emending the two tribes under discussion. He elected as his principal
distinguishing character the non-envelopment (Cynometreae) or envelopment
(Amherstieae) of the floral buds by bracteoles. Unquestionably this was an ex-
cellent choice for the African genera; whether it can be applied with success to the
non-African Old World genera is an interesting problem. Any discussion of this
here would be out of place.

A noteworthy point in considering broad intergeneric relationships involving
Cynometra is the lack of attention paid to the tropical American tribe Sclero-
lobieae (nine genera and approximately 60 species). In my opinion it is impossible
to define the limits of the tribes Cynometreae and Amherstieae without giving
careful consideration to this group. Among its nine genera we find some genera
with free central attachment of the stipe (Sclerolobium, Diptychandra etc.),
some with eccentric attachment (Dicymbe); two have the bracteoles enveloping
the floral buds (Dicymbe and Tbylacantbus), while the remaining seven have
bracteoles not enveloping the buds.

A third factor in establishing intergeneric relationships is the obvious relation-
ship of Cynometra with the Papuasian genus Maniltoa. A superficial glance at
herbarium material of Maniltoa (unfortunately poorly represented in herbaria)
suggests their proximity. The foliage, young strobiliform racemes, and legumes
are strikingly similar. The flowers of Maniltoa differ in several respects from those
of Cynometra in the narrow sense: the receptacle-cup is usually more elongate,
the stipe is often eccentric in position, and the stamens are numerous. The occa-
sional fusion of the stipe to the wall of the hypanthium suggests affinity with one
of the African segregates of Cynometra, the genus Schotia, also known in the
Papuasian-Melanesian region. The need for a critical study of the relationships
between Cynometra and Maniltoa is suggested by A. C. Smith in two recent papers
(Sargentia 1:36—38. 1942; and Jour. Arnold Arb. 21:166—171. 1950.).

1 The stipe is constantly free in all New World cynometras, and likewise in all African spe qee
according to Léonard's concept. While the same is probably true for the Indian and Papetti
Melanesian species, a thorough study of Cynometra and it ally Manilfoa in these areas is much n ;

2 . "v. i

2 It is my opinion that the tribes Cynometreae, Sclerolobieae, and Amherstieae should be united to
form one large tribe Amherstieae. This will be the subject of a separate paper.

1958]
DWYER—NEW WORLD CYNOMETRA 315

In relating the New World cynometras therefore, we must seek affinities with
genera traditionally placed in three tribes: Cynometreae, Amherstieae, and Sclero-
lobieae. The remarkable floral stability of the New World cynometras suggests
that this group diverged from the main line evolving in Africa and were able to
maintain themselves in a stable fashion in the New World. With the exception
of the anomalous C. fissicuspis all the New World cynometras seem to resemble
more closely (in a limited number of characters) Léonard's "groupe 2." Except
for the fact that the extra-African Old World cynometras frequently have multi-
jugate leaves, or occasionally simple leaves, the New World cynometras are
structurally quite similar. Bentham in 1840 (in Hook. Lond. Jour. Bot. 100)
remarked that "These American Cynometrae are similar to the Asiatic . . . decan-
drous ones, in the flower and inflorescence."

In viewing the problem of intergeneric relationships on a strictly regional basis,
ie. in the New World, Cynometra seems to be more closely related to the genus
Peltogyne than to Copaifera and Crudia. The presence of petals, the subimbricate
nature of the sepals, as well as the unijugate character of the leaves, suggest this.
Their ranges are similar, extending from Mexico to south-eastern Brazil, although
Peltogyne is absent from the West Indies. Pelżogyne is readily distinguished from
Cynometra by its pellucid-punctate leaflets, usually larger flowers, subclavate buds,
glandular-punctate and usually densely pubescent sepals, more dilated stigmas,
several to numerous ovules, paniculate inflorescences, and the often dehiscent
legumes. That some of the species of Peltogyne have the stipe of the pistil
eccentric is worthy of note.

MORPHOLOGICAL CHARACTERISTICS

STIPULES—these were not found on herbarium specimens, except a few of
seedlings deposited in Utrecht. As Britton and Rose (1930) and Amshoff (1939)
have indicated, the stipules are filiform and setaceous and measure up to 1 cm.
in length.

LEAVEs—the leaves of the New World cynometras are consistently unijugate,
in this respect differing from several Old World species, e.g. C. megalocepbala and
C. mannii. Petioles are obvious in the New World species while petiolules are
occasionally measurable, e.g. C. bostmanniana, C. trinitensis, and C. crassifolia, or
are represented by a black glandular area at the base of the lamina. Occasionally
a weak filiform extension terminates the petiole proper. While most cynometras
have leaflets which are oblong or lanceolate, several divergent forms exist. Of the
New World species C. bostmanniana alone has consistently equilateral laminas,
while those of C. marginata var. acuminata are occasionally equilateral. Strongly
inequilateral leaflets, on the other hand, characterize several species: C. cubensis,
C. oaxacana, and C. duckei. In most species the blades are obviously acuminate,
with a few: C. spruceana var. pbaselocarpa, C. microfłora, and C. duckei, with
very obtuse leaflets. In two collections (Purpus 7383, the type of C. oaxacana,
and Spruce s. n., K. type of C. spruceana var. procera) the leaves were found to

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316 ANNALS OF THE MISSOURI BOTANICAL GARDEN

vary widely in texture and venation, some being thin and obviously reticulate,
while others of the same collection were found to be thicker and smoother, with
the principal veins immersed or evanescent. This is disconcerting inasmuch as
considerable emphasis is placed on these characters in segregating the species.

The leaflets of the New World cynometras are not glandular-punctate although
Pittier, the author of the confusing C. fissicuspis, assigns this character to the
leaflets (cf. my discussion p. 342, this paper) ; Tulasne, too, in his original descrip-
tion of C. parvifolia, characterizes the blades as "interdum pellucido-punctatis.”
In the present study considerable attention was paid to the rugulose nature of the
blade (under mag.) of some species as well as the papillate nature of the leaflets
of other species. These are not completely reliable diagnostic characters at the
species level inasmuch as the blades of some collections of C. marginata may be
rugulose while others on the same twig are papillate. In general, however, those
leaves with obviously reticulate leaflets are invariably papillate, and conversely
those with immersed or plane secondary veins are rugulose. Among the New
World cynometras only the new species, C. duckei, has the leaflets neither papillate
nor rugulose above.

While petioles are invariably pubescent, the leaflets are rarely so (C. baubiniae-
folia var. macrophylla, some collections of C. cubensis, C. crassifolia, C. longi-
cuspis). In most cases the hairs are confined to the mid-veins or may extend to
the secondary veins, although the type of C. baubiniaefolia var. macrophylla has
completely pubescent blades.

INFLORESCENCES—two features of the inflorescence are useful for diagnostic
purposes: the relative length of the rachis, and secondly the numbers of flowers
present. Scarcely measurable rachises are occasionally densely flowered, e.g. C.
schottiana. The usually elongate pedicels tend to persist on the rachis until the
petals fall away. Initially the rachis emerges from a gemmiform cluster of concave
and imbricate bracts, which are often persistent. Such collections of bracts, fre-
quently described as strobiliform, are characteristic of other genera of the tri
Cynometreae: Guibourtia, Copaifera, etc. Unlike the bracteoles the bracts are
stiffer, more crassate, and more uniformly pubescent (except in C. bostmanniana).

The bracteoles are oblong, carinate (doubtfully so in section NUMMUS), and
paired, being attached at the base of the pedicels in most species. In three species:
C. spruceana, C. longifolia, and C. stenopetala these appear to be located along the
middle third of the pedicel. Bracteoles were not observed in C. americana, C.
cubensis, C. portoricensis, C. schottiana, and C. crassifolia. Since bracteoles are
soon deciduous it is possible that they may be present on immature inflorescences
of these species.

The pedicels of the flowers are slender in all species, being better described as
filiform in C. spruceana, C. cuneata, and C. longiflora. The reduction or absence
of hairs on the floriferous pedicels of C. bostmanniana and C. microflora 3s
noteworthy.

1958]
DWYER—NEW WORLD CYNOMETRA 317

FLORAL STRUCTURE—the receptacle-cup or hypanthium, while small, is obvious
as in most genera of the Amherstieae-Cynometreae complex, with the perianth
and the androecium borne on the margin of same. ‘The external pubescence of the
hypanthium and the thickness of the wall are of little importance as interspecific
characters in the New World cynometras.

The sepals are always somewhat unequal and reflexed at anthesis though not
markedly so in C. fissicuspis. Although varying little in puberulence, venation
etc., they are uniformly scarious in texture except in C. americana. These
number 4 or 5, although reduction to three per flower is not uncommon. The
flowers of C. oaxacana and C. marginata are occasionally bisepalous. The sepals
are puberulent on the outside, often sparsely so; occasionally sepals of C. oaxacana
and C. spruceana may be glabrous.

The petals of the New World cynometras show little variation in shape and
venation, and are regularly thin except in C. americana. They are glabrous, except
that those of C. baubiniaefolia var. macrophylla and C. parvifolia may occasionally
have a few apical or marginal hairs. In C. portoricensis and C. stenopetala the
petals are of about the same length as the sepals.

The stamens are constantly 10 and glabrous. No staminodia were found in
the numerous flowers dissected; the filaments are consistently slender, unequal in
length, and united very briefly at the base. The anthers are always small, not
exceeding 1.5 mm. in length, and versatile. In general the androecium furnishes
no diagnostic characters of importance.

Equally disappointing in furnishing diagnostic characters is the pistil. In the
key to species only three ovarian characters are utilized: the few hairs in C.
trinitensis, the appressed sericeous condition of C. cuneata, and the glabrous
condition of C. fissicuspis.

FRUITS—the fact that only about 10% of the herbarium material examined
bore fruit presented considerable difficulty in interpreting the material Despite
this the legumes provide several important characters: size, thickness of the wall
of the valves, and surface modification of the valves. Most ripe fruits are glabrous
although the valves of C. baubiniaefolia are velutinous while those of C. spruceana,
and occasionally C. porforicensis, are puberulent. Whether this is a consistent
character in the first two species is questionable. The pods are usually oblong, and
turgid, and only occasionally are compressed, e.g. in C. portoricemsis and C. bon-
durensis. They are regularly indehiscent. Amshoff states that C. hostmanniana
has “infertile pods . . . at last splitting into 2, showing no cavity at al" A
collection of C. bostmanniana (Froes 25771) has pods which seem to have split
open irregularly on drying. This same irregular dehiscence was noted on a collec-
tion (Elmer 12954) of C. bijuga from the Philippine Islands.

The pericarp in dried fruit is usually very spongy in appearance on the broken
surface and light in weight. The wall may range from 0.5 mm. (C. parvif olia)
to 3 mm. (C. retusa) in thickness. This is probably an excellent diagnostic char-
acter, the limited fruiting material however, makes it difficult to evaluate. The pods

[Vol. 45
318 ANNALS OF THE MISSOURI BOTANICAL GARDEN

are of 4 types with reference to the character of the surface: relatively smooth
with minute granulations or excresences (C. bemitomophylla, C. portoricensis);
obviously rugate and bearing some resemblance to the convolutions of a mam-
malian cerebrum (e.g. C. baubiniaefolia); tuberculate (e.g. C. duckei) ; smooth,
lustrous and vaguely reticulate (one species, C. fissicuspis). Rugate and tuber-
culate types are themselves often minutely granulose.

e fruits of C. fissicuspis are noteworthy inasmuch as they bear some resem-
blance to the pods of several African species placed by Léonard in his "Groupe 3."
On the other hand these African species diverge markedly in having several pairs
of leaflets, five sepals, bracteoles often articulate at the summit of the pedicels, and
dehiscent fruits. e fruits of the taxa of "Groupe 3" are apparently more
strongly reticulate and are not round but oblong in shape

n examination of the legumes of the non- African Old World species indicates
that they are regularly indehiscent and in general are either distinctly rugate or
tuberculate, some being deeply incised on the surface.

INTERSPECIFIC RELATIONSHIPS AND GEOGRAPHIC DISTRIBUTION

As suggested previously the New World cynometras are closely knit morpho-
logically with surprisingly few characters of diagnostic importance. In view o
the paucity of useful floral characters and the few legumes available for study, the
foliage assumes added importance. Among the numerous species placed in section
COMATOVARIA, those confined to Mexico and Central America tend to have con-
siderably larger leaflets and to a lesser degree more compressed racemes than their
South American allies. The fact that approximately one-half of the New World
cynometras are found in Amazonia, with four species of the group: C. longifolia,
C. longicuspis, C. stenopetala and C. duckei, restricted to the Brazilian states of
Pará and Amazonas, suggests that Amazonian Brazil is the center of geographical
distribution for the New World species.

ree new sections are proposed herein for the New World cynometras:
COMATOVARIA, GUIANA and NUMMUS. A view of Léonard’s 3 "groupes", pro-
posed for the African species of the genus, indicates that the 3 sections propo
for the New World species bear little relationship with those tentatively proposed
for Africa.

ECONOMIC IMPORTANCE

Although the cynometras are trees, the New World species are apparently of
little use commercially. In discussing C. sphaerocarpa, Pittier, the author of the
species (in Enum. Plantas Usuales 184. 1926) states that the wood is excellent for
construction. In the same discussion he adds that the seeds "son estomáquicas Y

i : ;cuspis to 2

s (L. = coin) refers to the fancied resemblance of the legumes of C. fair ca
coin. COMATOVARIA (comatus L. — densely hairy) refers to the fact that the ovaries of all spec
LI 1 | ls
y

P .

1958]
DWYER—NEW WORLD CYNOMETRA 319

antipalúdicas.” No oleoresins of commercial value have been reported for the
New World cynometras.*

Standley in his Flora of Lancetilla Valley (Honduras), in Field Mus. Nat. Hist.
10:219. 1931, states that the wood of C. re£usa is used for charcoal.

Record and Hess (Timbers of tbe New World, p. 252, 1943) list several
common names which I have not seen on the labels of herbarium material: "cour-
baril (Haiti); bruta de danto (Hond.); . . . herairo (Peru), guarabu amarello,
iauaranami, jutahy-rana, pororoca (Brazil), ingá-puita (Arg.)."

ACKNOWLEDGEMENTS

I wish to express my appreciation to the Missouri Botanical Garden for the use
of its herbarium and library facilities. I wish also to thank the directors of the
institutions listed below who were kind enough to allow me to examine herbarium
material of Cynometra. For purposes of citation a letter designating the particular
institution is used:

Arnold Arboretum, Harvard University, Cambridge, Mass. (A)
Jardin Botanique de l'Etat, Bruxelles, Belgium. (
Chicago Natural History Museum, Chicago, Ill. )
Conservatoire et Jardins Botaniques, Geneva, Switzerland. (G).
Department of Agriculture, Georgetown, British Guiana. (BRG).
Royal — Garden, Kew, wa eu
Herbariu epartment of Botany, University of iena Los Angeles, Calif. (LA).
Missoni” Bon cal Garden KA tas s, Mo. an
New York Botanical node. New York, N. Y
Museum National d'Histoire Naturelle, Laborataice de Dhan£togstuie, Paris, France. (P).
Seccao de Botanica Sistematica, Jardin Botanico do Rio de Janeiro, Rio de Janeiro,
Brazil.
Botanische Museum en Herbarium van de Rijksuniversiteit te Utrecht, Utrecht, Nether-

División de = vd "> rio de Agricultura y Cria, Caracas, Venezuela. (V).
Naturhistor n um, Botanische Abteilung, Wien, Austria. )
Yale Valve School of Forestry, New Haven, Conn. (Y).

TAXONOMY
CYNOMETRA L. Act. Soc. Upsal. 79. 1741 and Sp. PI. ed. 1. 382. 1753.

Trees or shrubs. Stipules filiform, soon deciduous. Leaves simple or pinnately
compound (exclusively unijugate in New World), the blades equilateral to in-
equilateral, usually acuminate, emarginate, the venation pinnately reticulate, the
margin entire. Racemes or panicles. Racemes usually strobiliform in young state,
usually one to several per axil, the bracts scarious or membranaceous, the bracteoles
paired, not enveloping the young bud, deciduous. Flowers obviously pedicellate,
occasionally articulate. Receptacle cup short, usually without a thickened disc.
Sepals usually 4-5, but occasionally 2 or 3, free, imbricate, unequal, usually
reflexed. Petals 5, usually equal in length; stamens 10 (8-12 by exception)

ue copal is obtained from a number of trees of the Leguminosae. C. sessiliflora Harms of the
Old endi is the only member of the genus known to yield copal.

[Vol, 45
320 ANNALS OF THE MISSOURI BOTANICAL GARDEN

glabrous, the anthers small, versatile, the filaments slender, usually very briefly
united at base; stipe of pistil free in receptacle cup, the ovary usually hairy, the
style slender, the stigma capitate, the ovules 1-2 (3 or 4 by exception). Legumes
dehiscent or indehiscent, usually rugate or tuberculate on surface, occasionally
smooth, the seeds 1-2.

Type species: C. cauliflora L.

KEY TO SECTIONS or New WORLD CYNOMETRAS*

a. Ovaries hairy; fruit not reticula
b. Leaflets obviously lae feni pedic els of flowers ques e: largar bracts 2-5 mm.

long; bracteoles present (some species uncertain) but soon decidu
Sectio COMATOVARIA Dwyer
bb. een regularly equilateral; pedicels of fora glabrous to scarcely ‘pubescent bracts
ab mm. long, glabrous; bracteoles absent.........................-.....---- n 2. IANA Dwyer

pean 21: G: boria

aa. Ovary glabrous; fruit reticulate Section 3. MMUS Dwyer
(Species 2. "c. fissicuspis)

Section 1—COMATOVARIA Dwyer

a. Perianth obviously carnose Subsection A—COMATOVARIA
. C. americana.
aa. Perianth non-carnose Subsection B—TENUIFLORA Dwyer
b. Leaflets more than 2 cm. long exa some leaflets of C. per C. microflora and
C. sini aż an t narrow lances lat
c. Lowerm eins (if seen) llamaba along costa and not spreading flabellately;
labia aia velu tinous e gems moz C. spruceana).

d. Secondary veins v:
e. Leaflets (1—)2—4 zy obtuse or only vaguely acuminate. = Indies.
f. Leaflets obliquely Sblong-rotund or obliquely oblong, (1- —)2-2.5 cm. long,
tals long, obviously longer than Rede “legumes

plump, p a pil ate s ina rea cubensis.
ff. Leaflets narrow-oblong, 5.5 cm. long, obtuse or only vaguely acuminate at
spesi 2. up to 3.5 mm. long, as - or shorter da ues legumes
essed to turgid st with — Own excrescences C. portoricensis.
ee. Lees iti m. long, acuminate. Mexico s tropical Americ
rgest leoni uide cm. long; legumes 2-4 cm. long, am N Central
erica. (C. trinitensis?).
h. Leaflets coriaceous; legumes turgid.
i We densely disposed on valves of o
eaflets narrow-oblong or siate ia 4-11 cm. long, 2-3.5 c
ra the seco veins asseruit s or immersed; fruit Zini
like in shape, ras ied up t thick. Mexico, re
Honduras, and British dr. „4. C. refuse.

* In arder to publish validly the new section and subsections the following latin descriptions are

COMATOVARIA Dwyer, sect. nov.: des e ecc qp Pedicelli lorum dense puberuli. Legumina
non petto Species typica: C. ameri Sepala
Subsect. COMATOVARIA. Sepala eae Pa TENUIFLORA Dwyer, subsect. nov.
petalaque tenuia. — Se C. baubiniaefolia. A
GUIANA Dwyer, : Folia aequilateralia. Pedicelli lorum vix puberuli vel glabri. Leg:
non reticulata. siet dla C. bostmanniana. Species
Dwyer, sect. nov. Ovaria glabra. Legumina reticulata laevia compressa.
typica: C. fissicuspis.

1958]

DWYER—NEW WORLD CYNOMETRA 321
jj. Leaflets elliptic, 9-14 cm. long, 3-5 cm. wide, the secondary veins

prominulous; Ele tu raid. but pe ete the pericarp up t

i ta R 5 C. bemitomopbyla.

ii. Hairs s pary sid on valves of ovary C. trinitensis.

hh. Leaflets stiff -papyraceous to stiff-chartaceous.
k. Rachise de 8-2 cm. long; pedicels 0.5-0.6 cm. long................- 7. C. hondurensis.
kk. Rachises 0. 253-5 cm. ne pedicels up to 1.3 cm. long............. 8. C. schottiana.

gg. pera leaflets 4-9 cm. long (except C. cuneata and C. longifolia often longer) ;
legumes 1-2 cm. long (ex cept G. es lye are longer), crustulose or rugu-

l. Leaflets minutely papi illat ak mag.) (except C. oaxacana);
prominulous; rachises of a toan 0.2-2 c patie on pedicels of flowers
ses to 1.2 cm. long (except C. marginata var. up to 1.8 cm. long

ta up to 2 cm. long), the Lula pues basally.

m. Parola 0.2-0.8 cm. long; rachises of inflorescences 3-10 flowered,

asi cm. long; petals 2.8—5.2 cm. long; hairs of ovary not uniformly
S
Leas ae to thin-coriaceous; sepals 2.5—4.5 mm. long;

9
T
să!

tals 3-5 m
^ “Leafs oblong, ‘sublanceolate or narrow-elliptic, 4-7 cm. long; 1. k
$ € wide, epals 3—4; wall of pencep up S 1m
thi ka Sur "i: Venezuelas Brazil, British Guiana............. cC. marginata
oo. Leaflets par die to ovate-sublanceolate, neges cm. e 1.5-
cm. wide, obtuse; sepals Pamela only 2; wall of paent up to
Mexico. 0. C. oaxacana.
nn. Leaflets sarite sepals 2.0-3.5 mm. long; petals 2.3—5.2 mm. long.
p. Leaflets petiolulate for 1 mm.; lowermost srai i ann shsrpiy
spesi y sepals 2. 5-3.5 mm. Tong; petals 4.5— ime
wide

"©. ala
n dee not obv Pu. onmes: lowermost veins ses mol X
about the same ang zi upper veins; sepals 2-3 mm. long; peta
2.3—2.8 mm. long, 0 mm. wide o pna
mm. sej 0.5—1.8 c sp poo f inflorescences 10-20 icd up
appressed . cuneata.
Ak aa rugulose (sub. mag.); costa usually plane ibo oud. veins
plane or immersed above; rachises of inflorescences 0.2—6 m. long; pedicels
of flowers 1-2 cm. long, = Laica (or scars) baos prr but not
e. basal.
. Rachises of inflorescences 0.2-2 cm. long; majority of bracts deciduous
at anthesis; flo wers up to 15 per recht:

"3
Mz]

> C. spruceana.
. C. longicuspis.

but not
q a of — E to M cm. long; majority of iu. persistent
is . C. longifolia.

mehess flowers 30—50 per
dd. Leaflets e seco veins si i visible da (sub. mag.), son papillate or
rugulose deka Sura tuberculate C. duckei.
cc. Lowermost pair of veins on Son side costone disposed and usually of same ee

asally as costa; 2d egumes rug velutin
s. pra obliquely mor rw to poni pape 1-4 cm. long, 0. ifa cm. wid
pedicels pi $6 bawbinicefolia.
SS. Lese ora run or oval-lanceolate, up to 2 cm. long, up to 13 cm. wide;
s glabrous.............. ernst retener 19. C microflora

bb. Leaflets ems than 2 cm. long, narrow-lanceolate......-.------- 220. C. servil.

l. CYNOMETRA AMERICANA Vogel, in Linnaea 10:602. 1836.

Small tree. Branchlets glabrous, ultimately lustrous and rubescent, the lenticels
pustulate. Petioles 0.4-1 cm. long, glabrous. Leaflets oblique-lanceolate,
oblique-oblong or ovate-oblong, occasionally subtrapeziform, 5-14 cm. long,
2.5-5 cm. wide, tapering cuneately into an acumen, ultimately obtuse and

retuse, the wider side tapering narrowly or frequently subauriculate at base,

the costa prominulous above, the secondary veins prominulous above, more prom-
inulous beneath, the lamina thin-coriaceous, rugulose above, papillate beneath,

[Vol. 45
322 ANNALS OF THE MISSOURI BOTANICAL GARDEN

lustrous, inequilateral, glabrous. Bracts oblong or oblong-rotund, up to 2.5 mm.
long, up to 2 mm. wide, puberulent dorsally and marginally. Bracteoles somewhat
concave, oblong, up to 2 mm. long. Racemes 1-3 per axil, the rachises 0.1—0.5 cm.
long, puberulent, the flowers about 10 per rachis; sepals 3—4, oblong-rotund, ovate-
rotund to rectangular, 3.5-4.5 mm. long, 2—3.5 mm. wide, carnose, glabrous; petals
obovate-oblong to oblong, 4.2—5 mm. long, 2—3.5 mm. wide, spatulate or obtuse
at base, carnose; filaments 4.5—7 mm. long, carnose; anthers subrotund, 0.8-1 mm.
long; ovaries oblong to subrotund, 2—3.5 mm. long, carnose, densely pubescent, the
style 2.5-3.5 mm. long, carnose; legumes (here immature?) oblique-oblong, up to
1.5 cm. long, compressed, marcescent, finely velutinous.

Harti: Les Atricots, Eyerdam 310 (A, NY, US); Petionville, Ekman H2343 (A, US);

Port de Paix, sw R (A, US); vicinity of Jean Rabel, River Cote de Fer, Leonard

& Leonard 12748 (U

C. americana, dia to Haiti has several obvious distinguishing character-
istics: the carnose perianth parts and the relatively large leaflets which are strik-
ingly bicolor and which usually taper cuneately into an acumen. The carnose
character of the perianth is not encountered in any other New World species, and
is the chief reason for placing C. americana in a separate subsection.

2. CYNOMETRA CUBENSIS Rich. in de la Sagra Hist. Cuba 10:233. 1845. (T.:

de la Sagra s. n.!).

Shrub or small tree. Branchlets glabrous, stiff, the lenticels obvious. Petioles
0.2-0.4 cm. long, glabrous to puberulent. Leaflets obliquely oblong-rotund or
oblique-oblong, occasionally subtrapeziform, 0.8—2.5 cm. long, almost as wide as
long, very inequilateral, the narrow side often 1/6th the width of the wide side,
reticulose especially below, obtuse at apex, ultimately retuse, obtuse at base on wide
side, vaguely papillate (under mag.), the costa prominulous above but evanescing
apically, occasionally pubescent, the secondary veins about 7, subprominulous to
subimmersed, the lamina coriaceous to thin-coriaceous, smooth and lustrous above,
obviously inequilateral, glabrous. Bracts oblong-rotund to narrow-oblong up to
3 mm. long, ferrugineo-puberulent dorsally; bracteoles not seen. Racemes ap-
parently one per axil, the rachises 0.5—1 cm. long, ferrugineo-puberulent, the
flowers 6—10 per rachis, the pedicels 0.6—1.2 cm. long, about 0.6 mm. wi
ferrugineo-puberulent. Sepals 4-5, oblong to oblong-rotund, 4-6 mm. eee
1.5—3.2 mm. wide, pubescent basally within; petals lanceolate, 6-8 mm. long,
2—2.6 mm. wide, acute at apex, spatulate at base; filaments 7-11 mm. long, the
anthers oblong, about 0.8 mm. long; ovary obliquely-oblong, about 3 mm. long,

8 mm. wide, densely pubescent, the style about 5 mm. long, subcrassate prox-
imally, obviously eccentric. Legume obliquely elliptic, up to 3 cm. long, plump,
granulose or crustulose, the wall of the pericarp up to 0.8 mm. thick.

UBA: without locality, Mr 2386 (G, US); Pinar del Rio: Pe de los O rganos,
Ekman 12684 (A, G), 12978 (A, US); Marl. Ekman 17375 (A, G, NY); Buenavista,
yajabos, Léon 13552 (NY), 13805 (NY); Rio Francisco, Ekman 18204 (A, NY);

S ea (?), de la Sagra s. n. (P); Havana, de la Sagra 307 (A, G).

1958]
DWYER—NEW WORLD CYNOMETRA 323

C. cubensis is closely allied to the only other West Indian species, C. porto-
ricensis.? Both have unusually small obtuse to vaguely acuminate leaflets. The
Cuban species has more or less orbicular leaflets in contrast to the narrow-oblong
blades of C. portoricensis. The flowers of C. cubensis are much larger and the
rachis of the racemes more elongate. The fruits are more turgid with the crustu-
lose excrescences less prominent than in its ally.

e de la Sagra collection in Paris is probably the type despite the fact that
the locality on the label (Valenzuela) does not agree with the cited type locality:
"Vuelta de Abajo." It is an excellent match for the illustration (plate 41) in
de la Sagra's Hist. Cuba 1845

In several dissections the sepals proved to be puberulent at the base within.
This was not observed in the calyx of any other New World Cynometra. The
type description indicates that "filaments (stamens) . . . subvillosa". This was
not observed and is probably an error inasmuch as the filaments of all cynometras
are glabrous. The common name of C. cubensis is “Pico de Gallo.”

3. CYNOMETRA PORTORICENSIS Krug and Urban, in Symb. Ant. 1:312. 1899.

(T.: Sintenis 5600!).

Small tree. Branchlets somewhat smooth, rimose, głabrous, the lenticels often
papillate. Petioles 0.2—0.3 cm. long, puberulent. Leaflets narrow-oblong to oblong
or lanceolate, 1.6 cm. long, 0.5—2.6 cm. wide, obtuse to vaguely obtuse-acuminate
at apex, obtuse to subauriculate at base on wide side, the costa prominulous above
but evanescing apically, the secondary veins + 10, often plane or subimmersed
above, the lamina reticulate especially below, not papillate above, vaguely papillate
below, coriaceous, lustrous, inequilateral to equilateral, glabrous. Bracts ovate-
rotund, about 2 mm. long; bracteoles not seen. Racemes apparently solitary in
the axils, the rachises up to 0.5 cm. long, puberulent, the flowers few per rachis,
the pedicels 0.4—0.6 cm. long, densely pubescent; sepals 4—5, oblong, about 5 mm.
long; petals oblong, 2.5-3 mm. long, obtuse to subspatulate at base, white in vivo;
filaments up to 5 mm. long, the anthers oblong, 0.8-1 mm. long; ovaries densely
pubescent, the style about as long as ovary. Legume oblong, 2.5-3 cm. long,
1.5—1.8 cm. wide, compressed or turgid, the surface with crustaceous excrescences,
velutinous or glabrous, the wall of pericarp about 1 mm. thick.

” 1 Salto”, — 600 (A G,

v Yao ka m i die ghee az (US); near „p Lo a
RO =, (NY); Rosario, San German (NY); Veri Aur Gleason & Cook A-2I,
“Dow NICAN REPUBLIC: zadanie Fuertes 1452 (NY, P); Santo Domingo City, Rio
Oxum man 12511 (A, US); Com. Palmar, J. Jiménez 2495 (US); Lopez, Prov.
Sung; Jiménez 458 (US).

The description of the flowers is of particular interest inasmuch as Krug &
Urban did not include this in their original diagnosis. Noteworthy is the fact that

5 This statement is based on the assumption that C. frinitensis is not a West Indian species.

[Vol. 45
324 ANNALS OF THE MISSOURI BOTANICAL GARDEN

the petals are shorter or barely as long as the sepals. Krug and Urban originally
described the legumes as "postremo dehiscentia.” This seems questionable especially
in view of the fact that the herbarium material lacks dehiscent fruits. As indicated
in the introduction, legumes with regular dehiscence are not found in the New
World cynometras.

4. CYNOMETRA RETUSA Britton and Rose, in Trop. Woods 7:5. 1926. (T.:

Record 8832!).

Small tree. Branchlets smooth, glabrous to puberulent, the lenticels often
scattered-punctate. Petioles 0.4-0.7 cm. long, glabrous to puberulent. Leaflets
narrow-oblong to lance-oblong, 4—11 cm. long, 1.2—4 cm. wide, usually attenuate-
acuminate at apex, often falcately so, obtuse or cuneate at base on wide side, the
costa prominulous above, the secondary veins + 10, prominulous or immersed, the
lamina vaguely to obviously rugulose or papillate above, papillate beneath, thin-
coriaceous, lustrous above, inequilateral, glabrous. Bracts disposed as a lanceolate
cluster in bud, up to 2 cm. long, ovate to ovate-lanceolate, 3.5-5 mm. long,
densely puberulent dorsally; bracteoles oblong, about 2 mm. long, ciliate on back,
soon deciduous. Racemes apparently solitary in the axils, the rachises 0.4—0.6
cm. long, puberulent, the flowers 10—20 per rachis, the pedicels slender, 0.4—
1.5 cm. long, about 0.35 mm. wide, ferrugineo-villose. Sepals 4—5, oblong, 2.8-5.2
mm. long, 1.5—2 mm. wide, sparsely ciliate at apex and on margins; petals obovate-
oblong to oblong, 5—6.5 mm. long, 0.8-2 mm. wide, obtuse to subacuminate at
apex, subspatulate at base; filaments of stamens up to 7 mm. long, the anthers
oblong, 0.7—1.2 mm. long; ovaries obliquely oblong, densely lanate, the styles
2—3.2 mm. long. Legumes oblong-rotund to inequilaterally oblong, up to 1.8 cm.
long, 1.8 cm. wide, nuciform, minutely apiculate, the surface minutely crustose,
the pericarp wall about 3 mm. thick.

MEXICO: TABASCO: Nacajuca, Rovirosa 748 (US).

GUATEMALA: Los Andes, Record 8832 (NY, Y); ALTA VERAPAZ: Chinajá, Steyermark
45505 (F, US); Cubilguitz, Steyermark 44557 (F); zagar: Rio Chaciün, Jocoló, H.
prese cai (F, US); Quiriguá, Standley 24254 (NY, US); near Cristina, Steyermark
30071 (F).

ONDURAs: YORO: Progreso, Hottle 23 (F); ATLANTIDA: San Alejo (?), Standley
7873 (F); Tela, Standley 52915 (A, F, NY, US), 56686 (A, F, G, NY, US); La Ceiba,
Yuncker, Koepper & Wagner 8007 (F, G, MO, NY, US).

BRITISH HONDURAS: British Honduras Forest Dept. 5 (Y).

This relatively well collected species is, according to S. J. Record (cf. Britton &
Rose's original description), * . . . one of the commonest trees of the forest"
(Guatemala). The narrowly oblong to oblong-lanceolate leaflets, lustrous above,
are striking. The densely flowered and compressed racemes resemble those of C.
trinitensis but their rachises are shorter. The thickness of the wall (3 mm.) of the
small nutmeg-like fruits is noteworthy. According to Record and Hess (Timbers
of New World p. 252, 1943) the vernacular name is “pata de cabra” ( goat's foot)
because of the fancied resemblance of the paired leaflets to the foot of a goat.

1958]
DWYER—NEW WORLD CYNOMETRA 325

5. CYNOMETRA HEMITOMOPHYLLA (Donn. Sm.) Donn. Sm. in N. Am. Fl. 23:
220. 1930. (T.: Tonduz 7012!).

Copaifera bemitomopbylla Donn. Sm. in Bot. Gaz. 27:332. 1899.

Tree. Branchlets zig-zag, nodose or gnarled, glabrous, the lenticels numerous,
small, and verrucose. Petioles 0.6-0.8 cm. long, glabrous. Leaflets elliptic, 9-14
cm. long, 3-5 cm. wide, obviously acuminate at apex, the wide side obtuse at base,
the lamina reticulate, papillate (under mag.) especially above, the costa sub-
prominent above, the secondary veins 15—20, curving arcuately near margin to
join veins above, thin-coriaceous, inequilateral, curved falcately, glabrous. Bracts,
bracteoles, and flowers not known. Rachises of racemes (in fruit) 0.5—1 cm. long,
apparently solitary in the axil, glabrous. Pedicels 1-1.5 cm. long, 0.3-0.4 cm.
wide, glabrous. Legumes suborbicular, 3-4 cm. long, up to 3.5 cm. wide, nuci-
form, the surface crustose-granular, the excrescences deciduous, the wall of the :
pericarp up to 2.8 mm. thic

CosrA Rica: LIMON: Río Reventazón below Cairo, Standley 6 Valerio 48799 (US);
Santo Domingo de Osa (Golfo Dulce), Tonduz 9972 (MO, NY); Palmar Norte Río
Terraba, Paul & Dorothy Allen 5264 (MO).

C. bemitomopbylla, with perhaps the largest leaflets of the New World cyno-
metras (C. bostmanniana excepted), is unfortunately known only from fruiting
material? The principal secondary veins of the leaflets are more numerous (15—
20) than in any other species. Another character of especial interest is the
glabrous pedicel of the fruit.

The local name of C. bemitomopbylla is “Guapinol Negro” and the common
name is "Cativo" (fide P. Allen, loc. cit.).

6. CYNOMETRA TRINITENSIS Oliver, in Hook. Ic. 2443. 1896.

Small tree. Branchlets smooth, glabrous or puberulent, the lenticels inconspic-
uous. Petioles 0.6-1 cm. long, puberulent; petiolules 1-2 mm. long. Leaflets
obliquely ovate or oblong, 5-11 cm. long, 1.5—4.5 cm. wide, vaguely to obviously
acuminate at apex, tapering acutely or cuneately on wide side, the costa prominent,
the secondary veins 10—12, prominulous, the lamina inequilateral, coriaceous,
smooth or rugulose above and below, lustrous, glabrous, obviously inequilateral, the
intervenal areas rather indistinct above. Bracts rotund to ovate-lanceolate, 3—6
mm. long, 1.5—3.5 mm. wide, densely puberulent dorsally; bracteoles oblong, up to
4.2 mm. long, 0.5-0.8 mm. wide, carinate and densely puberulent dorsally.
Racemes 1-3 per axil, the rachises 0.8—1 cm. long, puberulent, the flowers about
15 per rachis, the pedicels up to 0.8 cm. long. Sepals usually 4, oblong to sub-
rotund, 3—3.5 mm. long, 1.5-3 mm. wide, puberulent dorsally at apex and base;
petals lanceolate to oblong-lanceolate, 4-5.8 mm. long, 1.3-2 mm. wide, tapering
toward base; filaments 4.5-6 mm. long, the anthers 0.6-0.2 mm. long; ovary

W

6 Paul Allen writes (The Rain Forest of Golfo Dulce Costa Rica p. 191. 1956), that the flowers
of C. Bei itomopbsilo = “pale-brown [and] are produced from late August to nearly November.

[Vol. 45
326 ANNALS OF THE MISSOURI BOTANICAL GARDEN

oblong, about 2.5 mm. long, about 1.5 mm. wide, villose especially basally and on
sutures, the hairs scattered on valves, the style 4-5 mm. long. Legume rotund,
2-4 cm. long, 2-4 cm. wide, the pericarp smooth, minutely granulose or crustulose,
2—2.5 mm. thick.

JAMAICA: Castelton Garden, collector ?, collection no. II (NY).

TRINIDAD: Botanical Garden, collector ? (G); Government House grounds, Broadway
s.n. (NY, US, P); Botanical Garden, Britton 1183 (NY, US).

BRITISH GUIANA: Botanical Garden, Georgetown, collector ? (BRG).

C. trinitensis bears a striking resemblance to the Costa Rican C. hemitomophylla
which unfortunately is known only from fruiting material. The principal sec-
ondary veins of the leaflets of C. bemitomopbylla, however, are more numerous
and proximate. The densely flowered and compressed racemes resemble those of
C. refusa. The latter species, however, has more narrow leaflets, a shorter rachis,
apparently longer sepals, and a densely hirsute ovary. The reduction in the number
of hairs on the ovary is the most conspicuous floral character of C. frinifensis and
is unique for the New World cynometras, C. fissicuspis and possibly C. stenopetala
excepted. In the plate accompanying the original description this character was
not accurately depicted despite the reference in the text to “ovario parce piluloso.”

The type collection (obviously from Trinidad) was not located. Oliver, fol-
lowing the original diagnosis, states: “Trinidad, Crueger, Prestoe etc." Whether
the species is native to Trinidad is not certain.

7. CYNOMETRA hondurensis Dwyer, sp. nov. (T.: Schipp 1192 (A)!).

Arbores parvae. Ramuli laeves glabri saepe lenticellis inconspicuis. Folia
glabra. Petioli 0.6-0.8 cm. longi. Foliola sessilia anguste oblongo-lanceolata, 6-12
cm. longa, 1.8—4.5 cm. lata, apice plerumque attenuato-acuminata, acumine circ.
1 cm. longo saepe subfalcate disposito, ultime emarginato, subequilateralia vel
evidenter inequilateralia latere lato basi cuneata, latere angusto attenuato, Costa
supra prominula infra subprominente, venis secundariis principalibus © 15 pro-
minulis, laminis reticulosis tenui-coriaceis vel rigide chartaceis supra in siccitate
obscure olivaceis. Racemi compressi, rhachidibus (hic alibus in fructu) 0.8-2 cm.
longis, circ. 0.12 cm. latis puberulis. Bracteae bracteolaeque non visae. Sepala 4,
oblonga, 2.5—3.8 mm. longa, 1—1.8 mm. lata, acuta vel acuminata (rare obtusa)
omnino glabra praeter dorso basim puberulo; petala alba in vivo, anguste ovato-
lanceolata vel angusto-oblonga, 3.2—4.6 mm. longa, 0.8—1 mm. lata, acuminata vel
vix obtusa; antherae non visae; ovaria omnino pubescentia comis aliis appressis,
valvis oblongis, 2.3—2.5 mm. longis, 1.5—1.8 mm. latis, stylo circ. 2.5 mm. long.
Legumina (hic juvenilia) impare obovato-oblonga, 2.3 cm. longa, 1.5 cm. lata,
apiculata compressa minute crustulosa.

BRITISH HONDURAS: Rio Grande River, Schipp 1192 (A, G, MO, NY).

The foliage of the new species closely resembles that of C. retusa and C. scbotti-
ana, differing from the former in having only thin-coriaceous or sia a
leaflets, much longer rachises of the inflorescences, longer petals, and larger an

1958]
DWYER—NEW WORLD CYNOMETRA 327

more compressed fruits. C. schottiana on the other hand, has much larger and
more numerous flowers, shorter rachises of the racemes, more elongate pedicels, and
sepals with hairs restricted marginally. Like C. retusa, the new species bears
apiculate fruits.

8. CYNOMETRA SCHOTTIANA Hochr. in Bull. N. Y. Bot. Gard. 6:275. 1910.

(T.: Schott XII 857!).

Tree. Branchlets glabrous, the lenticels obvious and numerous. Petioles 0.6—1
cm. long, glabrous. Leaflets oblong to oblong-lanceolate, 8-13 cm. long, 3-5.5 cm.
wide, the acumen up to 1.5 cm. long, obtuse to subauriculate on wide side at base,
the costa subprominent, the secondary veins up to 20, prominulous, uniting near
margin to form an undulating vein subparallel to margin, the lamina scarcely stiff-
chartaceous, obviously inequilateral, glabrous. Bracts ovate or oblong, puberulent
dorsally; bracteoles not seen. Racemes apparently one per axil, the rachises 0.2—0.3
cm. long, puberulent, about 3-flowered. Sepals 4-5 (often two almost completely
united marginally), oblong-lanceolate to oblong, 4—5.3 mm. long, 1.8-2.8 mm.
wide, glabrous except puberulent marginally; petals lanceolate, 6-7 mm. long,
about 2 mm. wide, acute to acuminate at apex; filaments up to 9 mm. long, the
anthers about 1 mm. long; ovaries densely pubescent, the style 2—3.8 mm. long.
Legume oblong (here very immature).

COLOMBIA: EL CHOCÓ: Atrato and Truando Rivers, Schott XII 857 (NY).

The few-flowered racemes and the elongate petals exceeded in length only by
those of C. cubensis and C. cuneata, are noteworthy. Hochreutiner in his original
description erred in describing the sepals as glabrous. The common name is
“Tremantino”.

The type collection in New York bears the collection number “XII 857” and
not simply “7” as listed by Britton & Killip (Ann. N. Y. Acad. Sciences 35:193.
1936).

9. CYNOMETRA MARGINATA Benth. in Hook. Lond. Jour. Bot. 2:100. 1840.

Tree. Branchlets rough, usually conspicuously lenticellate, glabrous to puberu-
lent. Petioles up to 3.5 mm. long, glabrous to puberulent. Leaflets sessile, oblong,
sublanceolate or narrow elliptic, 4-7 cm. long, 1.5-3 cm. wide, scarcely attenuate
to acuminate, emarginate, the wider side subauriculate to scarcely obtuse at base,
the costa prominulous above and below, the principal secondary veins 10-15, im-
mersed to prominulous above, prominulous to subevanescent below, the lamina
thin-coriaceous, inequilateral to subequilateral, minutely papillate to rugulose above
and below, reticulate, glabrous, the margin often thin-callose, occasionally crisp.
Bracts oblong, up to 2 mm. long, densely pubescent on back. Bracteoles oblong, up
to 1.5 mm. long, obviously carinate, the hairs elongate along carina. Racemes 1-2
per axil, the rachises 0.2-1.8 cm. long, 8- to 15-flowered, densely aureo-puberulent,
the pedicels 0.5-1 cm. long, about 0.35 mm. wide (at anthesis), puberulent. Sepals
3-4, often 2 united marginally, almost completely or only to middle, ovate-oblong,

: [Vol. 45
328 ANNALS OF THE MISSOURI BOTANICAL GARDEN

oblong-rotund to narrowly ovate-oblong, 2.5—6.5 mm. long, 2-5 mm. wide, obtuse
to subacute at apex, puberulent to scattered-villose on back, the hairs often con-
fined below middle; petals narrow-lanceolate to narrow-oblong, 3.5-7 mm. long,
1-3.5 mm. wide, acuminate to obtuse at apex, obtuse to somewhat clawed at base;
filaments of stamens up to 13 mm. long; the anthers 0.6-1.1 mm. long; ovary
oblong, up to 3 mm. long, densely hirsute, most hairs elongate and ascending, some
shorter and more flaccid, the style 2.8-4 mm. long. Legume oblong, up to 2 cm.
long, vaguely rugulose to somewhat smooth, crustulose, the pericarp about 1 mm.
thick.

KEY TO THE VARIETIES

Leaflets obviously reticula
Leaflets oblong, błocie or narrow-elliptic, scarcely acuminate, coriace0us.......------------
9a marginata var. marginata,
Leaflets elliptic, obviously acuminate, the acumen 1-1.5 cm. long, Char ei cd
9b. C. marginata var. guianens
Leaflets smooth and not reticulate 9c. C. marginata var. [Pe

9a. CYNOMETRA MARGINATA Var. MARGINATA.
ANAMA: Savanna north of Panama City, Brotber Paul 463 (F); Paraíso Station,

SURINAM: without specific pel B. W. 658 (U); Forest vue Section, B. W.
1257, W); „aa B. W. 4885 (K, NY, U); Sectie O, B. W. 5868 (MO, U); Sectie O,
6 (U Aż Tibiens River, sad 300 (U); Coppename River near Raleighfalls,
pia "E. 58 (MO, U); Surinam River near Kabel, famio I2I (K, NY); He
River, Lanjouw $ ‘Lindemans 2964 (U); Saramacca River Headwaters, Maguire 2
pięt “giga Saramacca River, village Jacob Kondre, oa & Maguire 23856 (F,
„NY, V); Posoegronoe, Saramacca River, Maguire & Maguire 24040 (MO, NY, U);
"BEŻ Pana 24040a (U, V).

BRITISH GUIANA: British Guiana Forest Dept. 6217 (NY, U).

VENEZUELA: Rio > Between mous Río fade and Cano Masagua, Level 89
(NY); BOLIVAR: Cerro Guaiq mile above Raudal Aquacanta, Maguire 32705
(NY, V); Laguna de = Cha (Manicino Upata), Saldo 216 (V

BRAZIL: PARA: without specific locality, Schomburgk 197 ( 169) (F, K, V, W); H
Remansao, Tocantins, Froes 23600 Qo: Rio Capim between Aproagu and iudi
Canduru; Rio Tanti, Ducke 16898 (R

C. marginata is apparently the most difficult of all the New World pue
to circumscribe because of the variability of its leaflets, which are typically
coriaceous, narrowly lanceolate, and scarcely attenuate apically. While Pn
nutely papillate laminae furnish a character permitting one to segregate it fom
its ally, C. spruceana, nevertheless, other foliage characters are not stable, viz. the
reticulate surface and the prominulous character of the secondary veins.
rachises of the racemes are strikingly variable in length, apparently an un
feature for the New World cynometras. Other characters, perhaps of secondary
importance, are the occasional reduction of sepals to 2, occasionally glabrous sepals,
and the variability in the length and width of the petals. The reduction in num
of sepals to 2 is otherwise found only in the Mexican C. oaxacana.

1958]
DWYER—NEW WORLD CYNOMETRA 329

9b. CYNOMETRA MARGINATA var. guianensis Dwyer, var. nov. (T.: Schom-
burgk 777 (NY)!).

Foliola elliptica, 3-8 cm. longa, 1.2-3 cm. lata, conspicue attenuato-acuminata
acumine 1-1.5 cm. longo, costa supra plerumque pubescente, chartacea, plerumque
vix inaequilateralia.

SURINAM: ioni Gongyryp 8 Stabel roro (U).

BRITISH GUIANA: without specific locality, Richard Schomburgk s.m. (G, U W);
Richard Schomburgk I 533 (U); Robert Schomburgk (Herb. Hance 7667) (W); Schom-
burgk 777 (NY, V

VENEZUELA: it Alto Rio Paragua, Cardona 884 (US).

The Schomburgk collections of the new variety are recognized with ease
inasmuch as the leaflets are strongly attenuate-acuminate and chartaceous. The
material from Surinam and Venezuela was difficult to assign to this taxon.

9c. CYNOMETRA MARGINATA var. laevis Dwyer, var. nov. (T.: Froes 6 Murça

24014 (NY!)).

Foliola laevia nitida venis secundariis supra evanescentibus. Fructus globosi
suturis conspicuis.

RAZIL: PARA: Rio Capini between Aproaga and Igarapé Candiri, Froes $ Murça
dor, 4 (NY).

The glossy leaflets with the secondary veins scarcely visible mark the new
variety. The fruit is more rotund than in C. marginata var. marginata. It is difficult
to say whether the conspicuous sutures of the valves represent a substantial dis-
tinguishing character inasmuch as fruiting material of C. marginata is so poorly
represented in herbaria.

10. CYNOMETRA OAXACANA Brandgee, in Univ. Cal. Publ. Bot. 6:180. 1915.
(T.: Purpus 73831).
Cynometra colimensis Britton and Rose, N. Am. Fl. 23: 220. 1930. (T.: Ferris 62371).
Shrub to large tree. Branchlets glabrous or puberulent, the lenticels prom-
inent. Petioles about 0.5 cm. long, glabrous. Leaflets obliquely ovate-oblong,
ovate-lanceolate, rarely oblong-subrotund or narrow-oblong, 3.5-8 cm. long, 1.5-5
cm. wide, obtuse on wide side at base, the costa prominent or prominulous, the
secondary veins + 10, prominulous, the lamina vaguely to obviously papillate
above (sub mag.), reticulate above and below in thinner more mature leaves, thin-
coriaceous, very inequilateral, glabrous. Bracts oblong to ovate-oblong, 2.5-3 mm.
long, up to 3 mm. wide, puberulent dorsally and marginally; bracteoles located
about 1 mm. from base of pedicel, oblong, up to 2 mm. long, carinate, scattered
pubescent on back. Racemes one to 3 per axil, the rachises 0.5-2 cm. long,
= 8-flowered; pedicels 0.6-1.2 cm. long, densely aureo-pilose; sepals 2-5, often a
pair fused below middle, oblong, ovate-oblong to ovate, 2.5-4 mm. long, 1.2-3
mm. wide (or up to 4 mm. wide if 2 fused), glabrous or puberulent dorsally; petals
oblong to obovate-oblong, 3.5-5 mm. long, 1-1.6 mm. wide, often subfalcately

[Vol. 45
330 ANNALS OF THE MISSOURI BOTANICAL GARDEN

disposed, obtuse to vaguely clawed at base; filaments up to 7 mm. long, the anthers
subrotund, 0.7-1 mm. long, 0.6-8 mm. wide; ovaries obliquely oblong to sub-
rotund, 3—4.5 mm. long, 2—2.5 mm. wide, densely pilose, style up to 3.5 mm. long,
subcrassate, glabrous to pilose above middle. Legume oblong, up to 3 cm. long,
up to 1.6 cm. wide, minutely granulose or crustulose, the wall of the pericarp up
to 0.5 mm. thick.

Mexico: without locality, Hinton 10893 (U); corma: Manzanillo, Ferris 6188
(US), 6237 (A, NY); GUERRERO: La Lagunilla, Nelson 7004 (US); Zihuatanejo, Lan-
glassé s. n. (P), Langlassé 441 (F, G, US); oaxaca: Cerro de Picacho, Purpus 7383 (F,
LA, MO, NY); CHIAPAS: Aguacate Palenque, Matuda 3796 (F, NY).

Two characters mark C. oaxacana: the oval-oblong or ovate-oblong leaflets
scarcely acuminate at the apex and the reduction in the number of the sepals (2-3).
Some of the smaller leaflets of Purpus 7383 (MO) have a thick lamina which is
not obviously papillate and reticulate below, thus differing markedly from the
larger leaflets. A similar variation in the thickness, surface character, and venation
of the leaflets on a single collection (Spruce s. n. (K), type of C. spruceana var.
spruceana) was encountered.

11. CYNOMETRA CRASSIFOLIA Benth. in Hook. Lond. Jour. Bot. 2:100. 1840.

Small tree. Branchlets often wiry and zig-zag, angular in cross-section, rough
or smooth, glabrous or puberulent. Petioles 0.4-0.8 cm. long, slender, glabrous or
puberulent. Petiolules up to 0.1 cm. long. Leaflets oblong-lanceolate, ovate-
lanceolate, rarely subtrapezoid, 3.6-9 cm. long, 1.5—4 cm. wide, short-acuminate
at apex, the wide side obtuse or tapering narrowly at base, the costa prominulous,
glabrous to puberulent, the secondary veins prominulous, the most basal 1 or 2
strict and spreading flabellately, the upper arcuate-ascending, plane or prominulous
above, prominulous and conspicuous beneath, the lamina minutely papillate above
and below, reticulate, subcoriaceous or stif-chartaceous, inequilateral, the wide side
usually thrice the diameter of the narrow side, glabrous above, glabrous or puberu- -
lent below. Bracts ovate to subrotund, up to 2 mm. long; bracteoles not seen.
Racemes solitary in the axils, the rachises up to 0.5 cm. long 8- to 10-flowered.
Pedicels 0.6—1.2 cm. long, about 0.7 mm. wide in middle. Sepals 4, ovate to oblong,
2.5-3.5 mm. long, 1.3-2.8 mm. wide, puberulent dorsally; petals narrow-oblong,
narrow-lanceolate to obovate-oblong, 4.5—5.2 mm. long, 1.2-1.6 mm. wide, obtuse
to somewhat clawed at base; filaments up to 8 mm. long, the anthers oblong,
about 0.6 mm. long; ovaries obliquely oblong, completely pubescent. Mature
legumes not seen.

È Pen locality and collector unknown (P, probable type); without specific locality,
eoyre $) S. 7. .

Ate sk MANABI: Recreo, Eggers 15752 (F, US); oro: Piedras, Little 6622 (US).

In the herbarium in Paris I located a specimen labeled Cynometra crassifolia
Benth. ex ipso Brésil. This is probably the type inasmuch as Bentham, in descri"
ing C. crassifolia, stated that “I have seen the three following American species m
the Paris Herbarium” (including C. crassifolia).

1958]
DWYER—NEW WORLD CYNOMETRA 331

12. CYNOMETRA stenopetala Dwyer sp. nov. (T.: Krukoff 6631 (NY)!).

Arbores 18-30 m. altae. Ramuli laeves glabri. Petioli 2-5 mm. longi, mar-
cescentes glabri. Foliola 2 sessilia inaequilateralia vix falcata oblique oblongo-
lanceolata vel oblonga, 3-8 cm. longa, 1-3.3 cm. lata, breviter acuminata
retusaque, latere angusto gradatim attenuato, latere lato obtuse disposito, costa
utrimque subprominente glabra, venis secundariis principalibus 10—12 prominulis
vix ascendentibus ultime venam marginalem conspicuam dispositis, coriacea laevia
brunnea glabra?. Racemi fasciculati rhachidibus plerumque pluribus per axillam,
1-7 (-10) mm. longis. Bractae crebrae mox deciduae ovatae vel ovato-rotundae
intus glabrae extus puberulentes. Bracteae oblongae, circ. 2.5 mm. longae, 0.5 mm.
latae, carinatae dorso ciliatae infra medium pedicellorum affixae. Flores 4—10
per rachidem. Pedicelli (hic in fructu) circ. 6.5 mm. longi, circ. 0.8 mm. lati,
lignosi. Sepala 4, oblonga vel ovata-oblonga, 2-3 mm. longa, 0.9-2 mm. lata,
obtusa glabra praeter dorso marginibusque; petala lanceolata, circ. 2.8 mm. longa,
circ. 1 mm. lata, acuta basi subspathulata glabra; stamina 10, filamentibus vix basi
cohaerentibus, ad 6.5 mm. longis, circ. 0.1 mm. latis, antheris non visis; legumina
turgida subrotunda, ad 1.3 cm. longa, in forma nuculae constructa suturis copiose
aliter sparse pubescentibus, crustulosa vel granulosa pericarpio spongioso, ad 3 mm
crasso, seminibus solitariis loculum explentibus.

BRAZIL: AMAZONAS: Humayatá near Livramento on Rio Livramento, Krukoff 6631
(A, NY, U, US) ; Humayatá near Tres co Krukoff 6439 (A, F, NY, US); Sao Paulo
de Olivenca near Palmares, Krukoff 8418 (U

All of the specimens are in the late flowering or juvenile fruit stage. For-
tunately sepals and to a less extent, petals may be found after a careful search.
The petals, not exceeding 2.8 mm. in length are smaller (2-3 mm.) than those of
any of the other New World cynometras except C. baubiniaefolia and C. micro-
flora. On the basis of leaflet venation, however, the species appear unrelated,
inasmuch as C. stenopetala has the principal secondary veins on the wider side of
the lamina regularly spaced, while C. baubiniaefolia and C. microflora have the
lowermost 2 or 3 secondary veins flabellately disposed. Vegetatively the new
species appears to be closely related to C. spruceana. The short-pedicelled flowers
borne on short rachises scarcely resemble the long-pedicelled flowers of C. spruce-
ana which are disposed in lax racemes on relatively elongate rachises. C. stemo-
betala seems to be more closely related to C. marginata and to C. crassifolia; it is
readily distinguished from these by its more coriaceous leaflets, smaller sepals and
petals, and the ovary (judging from the condition of the juvenile fruit) presum-
ably bearing only few hairs on the valves. Reduction in the number of ovarian
hairs is elsewhere encountered only in C. trinitensis.

13. CYNOMETRA CUNEATA Tul. in Arch. Mus. Paris 4:179. 1844. (T.: Richard
$. 2.1).
Tree. Branchlets smooth, drying silver-brown, puberulent, the lenticels in-
conspicuous. Petioles 0.5-1.8 cm. long, crassate, glabrous. Leaflets rhomboid-

[Vol. 45
332 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ovate to elliptic-lanceolate, 6-11 cm. long, 2.3-4.3 cm. wide, the acumen up to
1.2 cm. long, or short-acuminate to almost obtuse, obviously obtuse at base on
wider side, the costa prominulous above, usually rather prominent beneath,
glabrous, the principal secondary veins + 15, very prominulous but rather evanes-
cent above, prominulous beneath, the margin thin-callose, the lamina thin-
coriaceous, minutely papillate above and below, scarcely to obviously inequilateral,
obviously reticulate below, glabrous. Bracteoles narrow-oblong, up to 3 mm. long,
puberulent on back, attached basally. Racemes prominent, usually one per axil,
the rachises slender, 2—3.5 cm. long, 10- to 20-flowered, puberulent, the pedicels
0.8-2 cm. long, about 0.5 mm. wide (up to 1.5 mm. wide in fruit), scarcely
pubescent, smooth, rubescent. Sepals oblong, 4—5.5 mm. long, 2—2.8 mm. wide,
obtuse, glabrous except for few scattered hairs at apex; petals lanceolate to obovate-
oblong, 7—7.5 mm. long, 1.6—2.2 mm. wide, acute to obtuse at apex; filaments up
to 9 mm. long, the anthers 0.8—1 mm. long; ovary oblong, about 3.5 mm. long,
about 2.2 mm. wide, the hairs dense, appressed, sericeous. Legumes oblong, up to
3.3 cm. long, up to 1.5 cm. wide, plump, finely rugulose, crustulose or minutely
papillate, velutinous at first, then glabrate or glabrous.

VENEZUELA: BOLIVAR: Raudal Arutani, Alto Paragua, Guayana, Cardona 865 (V).

RAZIL: PARÁ: without specific locality, Ricbard s.n. (P); Juruá, Ducke 16894 (K,
US); Belém do Pará, Ducke 20208 (US); Caracari, Rio Branco, Kuhlmann 1067 (R);
RIO DE JANEIRO: without specific locality, Glaziou 13756 (G).

The relationship of C. cuneata and C. spruceana is suggested by their similar
foliage. C. cuneata, however, has longer petioles, often rhomboid-ovate leaflets
which are papillate and bear prominulous secondary veins. Likewise the rachises
and pedicels are longer, the flowers more numerous and the petals longer, thus
suggesting closer relationship with C. longifolia. Unlike C. spruceana the bracte-
oles are attached basally. The uniformly appressed hairs of the ovary are unique
among the New World cynometras.

14. CYNOMETRA SPRUCEANA Benth. in Mart. Fl. Bras. 15(2):247. 1876. (T.:
Spruce s. n. ?)

Small trees; branchlets smooth, glabrous to puberulent; petioles 2-7 mm.
long, usually puberulent; petioles absent or up to 1 mm.; leaflets narrow-oblong,
subovate-oblong to obovate oblong, 2-8.5 (10) cm. long, 1-4 cm. wide, vaguely
to obviously acuminate at apex, ultimately emarginate, oblique at base, the wider
side tapering obtusely, or subtruncate at base, the narrow side acute (tapering
usually beginning near middle part of blade), the costa plane to prominulous
above, often with groove on each side, prominulous below, the secondary veins
scarcely prominulous to plane or immersed (?) above, the margin regular, occa-
sionally revolute, thin to moderately thick-coriaceous, obviously inequilateral,
rugulose, often obviously reticulate below; racemes usually 1-2 per axil, the rachis
slender, puberulent, 0.7—2 cm. long, (6-) 10- to 15-flowered, the pedicels filiform
(about 0.5 mm. wide), 0.5-2 cm. long, densely puberulent; bracts soon deciduous,
oblong-rotund to rotund, up to 5 mm. long and 5 mm. wide, densely or moderately

1958]
DWYER—NEW WORLD CYNOMETRA 533

puberulent on back and margins; bracteoles deciduous, up to 3 mm. long, densely
puberulent on back, carinate, thickened apically, the bracteole scars 2—5 mm. above
proximal end of pedicel; sepals 4 (or 3 by fusion of 2), oblong, ovate-oblong to
subrectangular, 3-4.8 mm. long, 1-2.3(—3) mm. wide, obtuse to subacuminate at
apex, usually obviously puberulent on back, often ciliate on margins; petals oblong
to obovate-oblong, 5—7.5 mm. long, 1.8—2.3 (—3.2) mm. wide, obtuse to acuminate
at apex, vaguely to obviously clawed at base; filaments 5-10 mm. long, the anthers
0.9-1.3 mm. long, 0.5-0.7 mm. wide; ovaries oblique-oblong, 2-3.8 mm. long, 1-2
mm. wide, densely pubescent, the style up to 5 mm. long; legume oblong to sub-
rotund, 2.5-3 cm. long, 1-1.5 cm. wide, the suture of valves distinct, plump,
obviously rugulose, minutely granulate and puberulent.

KEY TO THE VARIETIES

Leaflets conspicuously or only vaguely acuminate at apex 14a. C. spruceana var. spruceana.
Leaflets obviously obtuse at apex 14b. C. spruceana var. phaselocarpa.

14a. CYNOMETRA SPRUCEANA Var. SPRUCEANA.

Trachylobium martianum Hayne, in Flora 10(11):744. 1827
Cynometra spruceana var. B procera Benth. in Mart. Fl. Bras. 152; :248. 1876. (T.: Spruce

34 >
Cyn onera ec var. y macrophylla Benth. in Mart. Fl. Bras. 152:248. 1876. (T.:

Spruc
Cy sii La (Hayne) Macbr. in Contr. Gray Herb. 59:19. 1919.

rea GUIANA: without specific sin IE 197 (168) (W).

VENEZUELA: BOLÍVAR: Laguna de los Chivos (Municipio Upata), Delgado 216 (G,
US); Pittie er 216 (G); AMAZONAS: Canis Api Capihuara, Alto Casiquiare, Llewelyn
Williams 115593 (US, V).

MBIA: Río Apaporis between Rio Pacoa and Río Kananari, Schultes & Cabrera
Pa (US 7;

BRAZIL: peep, Barra, Rio Negro, Spruce 1513 (K, W); San Carlos, Rio Do»
Spruce 3479 (NY, P, V, W). MARANHAO: Pedreiras, Ducke 2320 "n US);
ini 358 di Igarapé de Cachoeira Grande, Ule 8864 (G, NY, U , US), Veche 719

US), Ducke (U, US); Igarapé de vecia Ducke 24247 PARA
rada specific as Spruce s.n. (NY, P, W); Lago de Faro, Ducke 8412 (G, US);
Esposente municipio Almeirim, Ducke 3 549 wę: SRR De do Chao, Ducke 10820;

as Xingu, Victoria, Snethlage 10408 (G); Santarem, Spruce 872 (K, P); Belém, 2
ec of LA.M., Archer 7906 (F, K), he (E), wę (US); Conceicas perto da Fóz do
Rio Juruena, Pires 3912 (NY); Eu uos do Rio Arua, Pires & Silva 4210 (NY); Rio
= near Rio Xingu, Kablm nn 17655 (G, P, U, U
ORETO: Mishuyaca, prać, Klug 1448 (F, "NY, US); Maraüón, Iquitos,
iens Peer (F, G).

In 1919 Macbride effected the combination Cynometra martiana (Hayne)
Macbr. maintaining that Baillon’s earlier combination Cynometra martiana
(Hayne) Baillon is untenable inasmuch as "it is a valid species of Hymenaea (H.
martiana Hayne).” According to Macbride, Hayne’s specific name martiana may
be used for the “species of Cynometra which has heretofore borne the later pub-
lished name of C. spruceana.” No doubt he bases this on the fact that the plate
in Hayne’s Arzneigew (11: pl. 17. 1839), labeled Trachylobium martianum is a
Cynometra.

[Vol. 45
334 ANNALS OF THE MISSOURI BOTANICAL GARDEN

However, the binomial C. martiana (Hayne) Macbr. is illegitimate, being a
homonym of C. martiana (Hayne) Baillon which is applied to a valid species of
Hymenaea. That this name has not been applied in the past to a Cynometra but
to a species of Hymenaea makes no difference, the binomial C. martiana being
already published.

After examination of type collections of Bentham's two varieties, 9. procera
and y macropbylla, I am unable to find any strong characters to substantiate
giving them varietal status. Examination of the type material of var. procera
deposited in Vienna shows leaflets of obviously different ages; the more mature
leaflets are thicker with the secondary veins immersed above, and the venules
inconspicuous beneath, while the more juvenile leaflets are obviously more reticulate
beneath with the secondary veins prominulous above. Since Bentham distinguishes
var. procera by "foliolis quam in forma typica minus coriaceis," it seems unwise
to recognize this variety without additional supporting collections. As for var.
macrophylla, the type collection deposited in Vienna has leaflets which are less
elongate than those described by Bentham (‘3-4 pollicaribus") ; another character
used by Bentham, the length of the racemes and the pedicels, is so variable in the
typical material that it cannot be regarded as a strong one.

While the pedicels and rachises of the racemes are constantly filiform, there is
considerable variation in their length; T'essmann 3636, for example, has elongate
rachises but relatively short-pedicelled flowers; Ule $864 has both elongate rachises
and elongate pedicels; Pittier 216 has both short rachises and short pedicels.

Two excellent plates of C. spruceana are available, one in Hayne's Arzneigew
11: pl. 17. 1830, and the other in Martius Flora Brasiliensis 152:247. pl. 65. 1876.

14b. CYNOMETRA SPRUCEANA Var. o. (Hayne) Dwyer, comb. nov.

Vouapa phaselocarpa Hayne, in Flora 10(11) :745.
Cynometra racemosa Benth. in Hook gei Jour. Bot 108: 1840. (T.: Spruce s. n.!)
Cynometra obtusa Benth. in Pl. Spruce exs. Nom
ie wów phaselocarpa (Hayne) Mese. in Conti Gray Herb. 59:19. 1919.
AMAZONAS: Barra, Spruce 1428 (K); Barra, 2 Negro, Spruce s. , NY,
W); Santa nn Dacie 509 (A, NY, MO, US) 25 5 (G,P, U, US); fe Maat Rio
e pey Ducke 676 (F, MO, US), Darke Ioro (MO, U ŚW
The obtuseness of the apex of the leaflets does not seem to be sufficient grounds
to warrant the retention of C. pbaselocarba as a distinct species. In all other re-
spects it bears a striking resemblance to C. spruceana var. spruceana, especially in
the character of the lax racemes with elongate rachises and filiform pedicels.
Ducke's collection 676 is important inasmuch as it bears legumes. The common
name of the variety is "jutahy-rana" (fide Ducke).

15. CYNOMETRA LONGICUSPIS Ducke, in Bol Técn. do Instit. do Norte. 2:11.
1944. (T.: Ducke 1593!).

Medium-sized trees. Branchlets terete, ultimately angular and arcuate, smooth,
glabrous, drying purple to ashen-white. Petioles up to 8 mm. long, about 1.5 mm.

1958]
DWYER—NEW WORLD CYNOMETRA $55

wide, slender, glabrous to scattered-puberulent; petiolules 0.5-1 mm. long. Leaf-
lets oblong, up to 8 cm. long, up to 3 cm. wide, the acumen up to 1 cm. long,
about 2.5 mm. wide in middle, ultimately emarginate, the sinus about 0.5 mm. deep,
obtuse at base on wider side, the costa prominulous above and below, the secondary
veins + 10, immersed or plane, not markedly arcuate-ascending, the ultimate
reticulations evanescent, the lamina coriaceous, lustrous above and below, rugulose
above and below (sub mag.), inequilateral, glabrous (except possibly on costa);
inflorescences, bracts, bracteoles and complete flowers not seen. Sepals (one seen)
suboblong, 4 mm. long, about 1.3 mm. wide, glabrous; petals not seen; filaments
(two seen) linear, about 7 mm. long, the anthers not seen; rachis of fruit woody,
2-5 mm. long, about 1.8 mm. wide, ligneous, glabrous. Pedicels + 5 mm. long,
somewhat smooth, apparently glabrous; legume obliquely oblong to oblong-rotund,
up to 8 mm. long, up to 6 mm. wide, plump puberulent, rough and crustulose, the
stipe up to 1.5 mm. long.

BRAZIL: AMAZONAs: Tocantins beyond Villa Velha, Ducke 1593 (F, NY).

Ducke in his brief diagnosis of this species related it to C. baubiniaefolia and
to C. spruceana. He mentions the "poor fragments of old flowers [which] do not
show differences from those of the widely distributed C. baubiniaefolia.” A study
of the type material suggests that C. longicuspis is probably closely related to
C. spruceana. 'The fruits of the latter species are more rugose.

Ducke states (loc. cit.) that it is a rather large tree found in very humid
places at Tocantins and Sio Paulo de Olivenca, at Sio Gabriel and its tributary,
Rio Vaupes.

Two collections made by Froes (22147 and 20723), labeled C. longicuspis, are
obviously misidentified. The former collection (in fruit) appears to be a
Peltogyne.

16. CYNOMETRA LONGIFOLIA Huber, in Bol Mus. Goeldi 5:384. 1908. (T.:

Ducke 9083!) .

Tree? Branchlets zig-zag, subterete, puberulent; the lenticels conspicuous.
Petioles up to 0.7 cm. long, puberulent. Leaflets narrow-oblong, 7-12 cm. long,
2.5—2.8 cm. wide, tapering into an obvious acumen (up to 1 cm. long), subauricu-
late at base on wider side, the costa prominulous above, glabrous (except hairs
occasionally scattered on under surface), lowermost pair of secondary veins usually
arising sharply from one point at base of costa, the others well-spaced along length,
prominulous to evanescent above, prominulous below, the lamina rugulose (under
mag.), coriaceous, inequilateral, glabrous (except costa beneath). Bracts ovate,
up to 5 mm. long, occasionally wider than long, often persistent, pubescent dorsally,
the marginal hairs often tufted; bracteoles oblong, up to 6 mm. long, about 1 mm.
wide, located near middle of pedicel, the hairs ferrugineous, more dense along
median axis, tufted apically, the pedicels slender, about 1 cm. long, about 0.5 mm.
wide, Racemes apparently solitary in the axils, the rachises up to 6 cm. long, up
to 2 mm. wide, ferrugineo-villose, 30- to 50-flowered; sepals 4, widely to narrowly

[Vol. 45
336 ANNALS OF THE MISSOURI BOTANICAL GARDEN

oblong, 3-4 mm. long, 2-3 mm. wide, puberulent on outside especially below
middle; petals lanceolate, 4—4.8 mm. long, 1.3—1.4 mm. wide, acuminate; filaments
up to 8 mm. long, the anthers about 1 mm. long; ovaries (here in young fruit)
about 4 mm. long, about 2.7 mm. wide, pilose. Fruit oblong, up to 2 cm. long,
up to 1.4 cm. wide, obviously rugulose.

BRAZIL: PARA: Santarem, Spruce s.n. (W); Rio Mapuera, Maloquinha, Ducke 9083
(G).
C. longifolia is obviously related to C. spruceana as evidenced by its narrowly
oblong leaflets rugulose above, elongate rachises of the racemes, filiform pedicels,
and bracteoles attached near the middle of the pedicels. It is readily distinguished
by its large leaflets, persistent bracts, unusually elongate rachises, and multi-
flowered racemes.

Huber originally described the ovary as sparsely villose; my dissections indicate
that it is densely villose.

17. CYNOMETRA duckei Dwyer, sp. nov. (T.: Ducke 7165b (G)!).

Arbores. Ramuli ultime puberuli cortice in siccitate cinereo. Petioli ad 0.2
cm. longi, glabri vel puberuli. Foliola sessilia oblique oblongo-rotunda vel semi-
ovata, 0.7—2.5 cm. longa, 0.5—1.5 cm. lata, apice obtusa emarginata, basi latiore
latere obtuso, costa supra prominula gradatim ad apicem evanescente infra plana,
venis secundariis == 10 supra inconspicuis lamina nullo modo reticulata vel
papillosa (sub mag.) inaequilaterale latiore latere duplo triploque plure, margine
evidenter calloso; bracteae, bracteolae, rhachides, flores non visa; pedicelli (hic
fructuum) ad 1.5 cm. longi, pubescentes graciles. Legumina, subrotunda, 1.2 cm.
longa, 1 cm. lata, turgida evidenter tuberculata vel rugosa crustulis parvis deciduis-
que ornata.

BRAZIL: AMAZONAs: Rio Negro, Ducke 7165b (G, US).

After considerable deliberation I have decided to describe this as a new species.
Ducke labeled the material C. parvifolia. Its foliage bears a closer resemblance to
that of C. spruceana, among the Amazonian species and, oddly enough, to C.
cubensis. The leaflets are obviously much smaller and completely different in shape
(due in part to the marked inequilateral condition) than those of C. spruceana.
The most striking feature of the new species is the smoothness of the leaflets on the
upper surface, with no evidence of these being reticulate or papillate; the secondary
veins are scarcely visible and the costa tapers above the middle, disappearing toward
the apex of the lamina. Only one legume was seen.

18. CYNOMETRA BAUHINIAEFOLIA Benth. in Hook. Lond. Jour. Bot. 2:99. 1840.

Shrub or tree. Branchlets smooth, glabrous to minutely puberulent, the lenti-
cels conspicuous. Petioles 0.2-0.6 cm. long, glabrous to densely pilose. Leaflets
obliquely ovate-oblong to ovate-lanceolate, 1-4 cm. long, 0.5-2 cm. wide, the
costa prominulous above, the secondary veins 7-8, of these 2-3 obviously spreading

1958]
DWYER—NEW WORLD CYNOMETRA 337

flabellately from base of costa on wide side and scarcely distinguishable from costa
basally, the lamina reticulate, vaguely papillate to scarcely papillate below, thin-
coriaceous, obviously inequilateral, glabrous to minutely puberulent. Bracts oblong
to ovate, 1-2.3 mm. long; bracteoles oblong, up to 2 mm. long, carinate, densely
pilose. Racemes compressed in axils, the rachises apparently usually several per
axil, 1-4 mm. long, pubescent, usually 4- to 9-flowered; pedicels up to 7 mm. long,
puberulent; sepals 3-4, widely oblong, ovate-oblong or obovate-oblong, 2.5-3 mm.
long, 1-3.5 mm. wide; petals ovate to obovate-oblong, 2-3.6 mm. long, obtuse to
acuminate at apex, obtuse to cuneate at base; filaments of stamens 4-9 mm. long,
the anthers 0.5-0.7 mm. long; ovaries oblong to oblong-rotund, 2-3.5 mm. long,
1.5-2 mm. wide, completely pubescent, the style up to 5 mm. long. Legume
oblong to subrotund, up to 2.5 cm. long, up to 2 cm. wide, rugulose or tuberculate,
minutely and densely tomentose, the wall of pericarp up to 2 mm. thick.

KEY TO THE VARIETIES

ded or trees; azi a iu reticulate, the margin not curled; flowers wh
> ets <w m. long, 0.5—2.0 cm. wide, glabrous to minutely bii ung on costa
up Ww 2.8 Si ge 18a. C. baubinia n me beable
tides 4-6 cm. dee vp to 3 cm. wide, completely perle petals up mm.
C. baubiniaefolia var. E PR
ap trees; leaflets lustrous, not conspicuously reticulate, the magi cesta rled; flowers
yellow Gi baubiniaef olia var. meridiana.

18a. CYNOMETRA BAUHINIAEFOLIA Var. BAUHINIAEFOLIA.

GUATEMALA: ALTA VERAPAZ: Rio Icvolay between Rio Apia and Río Soctela, Steyer-

mark 45034 | (F).
URINAM: without specific Rim Schweintz s. n. (NY); Commewijne, B. W. 1137

(U); Sania River, Mell 17 (U

BRITISH GUIANA: without spale locality, Richard wiec s.n. (G, > Tian,
Schomburgk 241 (W); Berbice, Schomburgk 230 or 231 (F, , W); Rio Esse
quibo, Herb. N. Y. Bot Gard. Field no. G 126, Record 621 ^ (EM la nor.
British ater Forest Service 6217 (G, RE Ux 6126 (G); banks of Cerenlyne Epiro, Im
Thurn s.n. (K); Ta pe ag . Lethem, —— District, Irwin 814 (U

VENEZUE
Pargua, Killip 37611 > ‘NY, U US, V). AMAZONAS: El Bagre, Pto. lisi dug due
Williams 1 307 (U j:

CoLOMBIA: Rio Guaviare, San José del stopie: Cuatrecasas 7754 (U

S).

RAZIL: who specific locality, Newman (G); Burchell s (K); Burcbell
995I (NY, P); AMAZONAS: Jaru, Rio Branco, Kobiece 2919 ( io Branco,
S. Bento, Ule 7776 (G, K, US); Rio Branco, Da, = Bento, Black. b -I 13987 (US); Rio
Sinho, Juruema, Rio Jutuhy, Froes 21043 (F, K US); Bóa Vista, Rio Mae , Ducke
1355 (K, NY, (only part of sheet), US); Tefé, WE 1327 (NY). PARA: Tap , Boa
Vista, Capucho 529 (F); Faz. Tuiuiu, Rio Ara Black, Ledaux & Stegemann Sin
=. RIO DE JANEIRO: Quinta de Sao Cini Si ać P).

U: SAN ci : Usa Tessmann 3444 ( LORETO: tee. Haya, Llewelyn
Willions 40 (F); Amazon River, Llewelyn stă 1984 AF , US); Rio Mazan
near taos Level n Williams 8142 (F).

DA res: Mercedes, Rodrigo 782 (NY).

Cisti sila paki locality, McJobn Style s. n. (G).
C. baubiniaefolia with the most extensive geographical range among the New
World cynometras is readily recognized by the arrangement of the secondary veins

[Vol. 45
338 ANNALS OF THE MISSOURI BOTANICAL GARDEN

of the leaflets inasmuch as the lowermost 2 or 3 (on the wider side) arise flabel-
lately from the proximal part of the costa. These are as prominent near their
point of origin as the adjacent costa. A striking feature is the thick-walled fruit
which is densely tomentose.

18b. CYNOMETRA BAUHINIAEFOLIA var. grandiflora Dwyer, var. nov. (T.:
Ducke 16696 (U)!)
Foliola ad 6 cm. longa, omnino puberula; petala ad 5.5 mm. longa.
BRAZIL: PARA: Botanical Garden, Belem do Pará, Ducke 16896 (P, U, US).
The new variety is readily distinguished by its unusually large leaflets which
are entirely puberulent. The flowers are strikingly large.

18c. CYNOMETRA BAUHINIAEFOLIA var. meridiana Dwyer, Var. nov. (T.:
Jorgensen 2124 (MO)!)
Foliola laevia nitida venis secundariis evanescentibus marginibus conspicue
pue flores lutei.

TINA: Formosa, Jorgensen 2124 (MO, U); Formosa, Río Coltapick, Colonia
wae Boca del Bermejo, Pedersen 1295 (US).

19. CYNOMETRA MICROFLORA Cowan, in Mem. N. Y. Bot. Garden 8(2):114-

115. 1953. (T.: Maguire & Maguire 29010!)

Small trees; branchlets glabrous to puberulent; petioles up to 3 mm. long; leaf-
lets ovate-rotund or oval-lanceolate, up to 2 cm. long, up to 1.5 cm. wide,
conspicuously obtuse above, more rounded (on wider side) basally, the secondary
veins 3-4, prominulous, somewhat evanescent above, the lowermost pair arising
flabellately from base of costa and resembling the latter, thin-coriaceous, glabrous,
smooth, obviously inequilateral; racemes compressed, the rachises scarcely measur-
able, few-flowered; pedicels filiform, glabrous, up to 5 mm. long; sepals oblong,
up to 2 mm. long, I mm. wide; petals lanceolate, 2—2.5 mm. long, up to 0.8 mm.
wide; filaments up to 2 mm. long, the anthers up to 0.5 mm. long; ovary oval, up
to 1.5 mm. long, densely puberulent, the style up to 2 mm. long; legumes oblong-
lanceolate up to 1 cm. long, up to 0.6 cm. wide, somewhat rugulose, densely
puberulent.

VENEZUELA: La Urbana, Orinoco River, Maguire & Maguire 29010 (NY, V).

This species is very closely related to C. baubiniaefolia and may well be inter-
preted in the future as a variety of the same. Its small leaflets and flowers, as well
as its glabrous pedicels distinguish it from its ally.

20. CYNOMETRA PARVIFOLIA Tul. in Arch. Mus. Paris. 4:181. 1844. (T+
Hostmann 204!).
Small or medium-sized tree. Branchlets smooth, densely puberulent. Petioles
about 0.1 cm. long, densely pilose. Leaflets narrow-lanceolate, 1-1.8 cm. long,
0.2-1 cm. wide, the costa prominulous above, the secondary veins — 7, the lower-

1958]
DWYER—NEW WORLD CYNOMETRA 339

most spreading flabellately from base of costa, prominulous but inconspicuous
above, the lamina reticulate, rugulose (under mag.), thin-coriaceous or coriaceous,
inequilateral, głabrous. Bracts compressed-rotund to ovate-oblong, up to 2 mm.
long; bracteoles located at base of pedicels, oblong, up to 1 mm. long, carinate, the
hairs few, deciduous; racemes compressed, axillary, the rachises up to 5 mm. long,
densely pilose, the flowers 4—6 per raceme; pedicels 0.4-1 cm. long, ferrugineo-
villose. Sepals 4-5, ovate-oblong, oblong-rotund or subrotund, 3—4.6 mm. long,
1-2.6 mm. wide, glabrous or puberulent dorsally and marginally; petals oblong to
obovate-oblong, 3.5—4.6 mm. long, 0.8—1.8 mm. wide, glabrous, or often with
several weak hairs at apex, obtuse or clawed at base; filaments 4-10 mm. long, the
anthers oblong, 0.4—1 mm. long; ovaries oblique-oblong, 1.3-2.5 mm. long, 1-2.5
mm. wide, densely pilose, the style + 4 mm. long. Legume rotund, coccoid, up to
0.9 cm. long, 0.9 cm. wide, plump, the surface minutely crustulose, the wall of
pericarp about 1 mm. thick.

SURINAM: without locality, Hostmann 204 (NY); Marowyne and Gonine Rivers,
Gonggryb 3751 (U); Surinam River, Kappler 1905 (U); Marowyne River, Lanjouw 6
Lindemann 3471 (

BRAZIL: AMAZONAS: Caraca rahi, Rio Branco, Ducke 23788 (G, P, U, US); Béa Vista,
Rio Branco, Ducke 1355 (NY, in part only).

C. parvifolia is readily distinguished by its very small leaflets (not exceeding
2 cm. in length) which are narrow-lanceolate. It appears to be closely related
to C. marginata and to C. baubiniaefolia.

21. CYNOMETRA HOSTMANNIANA Tul. in Arch. Mus. Paris 4:180. 1844. (T.:

Hostmann 169!).

Medium-sized tree. Branchlets wiry, smooth, glabrous, the lenticels often in-
conspicuous. Petioles about 0.5 cm. long, glabrous to pubescent. Leaflets equi-
lateral to subequilateral, ovate-lanceolate, oblong-lanceolate, 6-12.5 cm. long,
2.5—8 cm. wide, tapering sharply toward apex, usually short-acuminate, the wide
side cuneate or sub-rotund at base, the costa prominulous above, the secondary
veins + 20, strict or scarcely arcuate, the lamina obviously reticulate, non-
rugulose (under mag.), thick-chartaceous to coriaceous, subequilateral, glabrous.
Bracts ovate-rotund to rotund, 1-1.3 mm. long, 1-1.4 mm. wide, puberulent
dorsally and marginally; bracteoles absent. Racemes often umbelloid in shape, 1-4
per axil, the rachises up to 7 mm. long, puberulent, the pedicels slender, up to 1.5
cm. long, glabrous or sparingly pubescent. Sepals ovate-oblong to oblong, 2.8—4
mm. long, 1.1-1.9 mm. wide, glabrous or puberulent dorsally, puberulent at apex.
Petals narrow-oblong, up to 4.5 mm. long, 0.4—1.5 mm. wide, acuminate at
apex, obtuse at base; filaments up to 4 mm. long, the anthers oblong to oblong-
rotund, 0.5-0.7 mm. long; ovaries oblong to oblong-rotund, about 2 mm. long,
densely pubescent, the style up to 3.8 (5?) mm. long. Legumes oblong, up to 4.5
cm. long, up to 3.5 cm. wide, the surface obviously rugulose, the rugae minutely
granulose, the wall of the pericarp up to 0.5 cm. thick, the sterile pods deeply
wrinkled and without any locule.

FRENCH GUIANA: without specific locality, Melinon s. n. (A, F, NY, P, US); Vaillant
IOI (P).

[Vol. 45
340 ANNALS OF THE MISSOURI BOTANICAL GARDEN

SURINAM: without specific locality, — sw. (U); — 1438 (B), 1656
(B); B. W. 198 (U); B. W. 1984 (U), B. W. 476 (U), B. W. 3878 (U); Lanjouw
Lanjouw 1891 (U); Gonggryp 457 b lento 308 (U); Watamiri River, B. W.
1508 (US); Watamiri, B. W. 1958 (U), 3861 (U); Maroni, Sagot s. n. (W); Sagot 1093
(U); Saramacca, Stabel 198a (NY, U, Y); Marowyne near Poeloegoedoe, Versteeg 552
(U), 576 (U); Marowyne River, Gonggryp 5357 (U); Coppename River, B. W. 155 (U);
Coppename, Gonggryp 2568 (MO, U); banks of Surinam River, Mell s. ». (NY, US);
pe porii Hostmann e RA W).

BRITIS wegt, Rerbice River, British Guiana Forest Dept. 6365 (U);
without cage locality di 1659 UJ).

BRAZIL ZONAS: Amapa, Rio Oiapoque, Froes 25771 (US); PARA: Trombetas
(Cachoeira Paten e Rio Cuminá-mirim), Ducke 10911 (R).

A number of substantial characters serve to distinguish this species: the con-
sistently equilateral leaflets, the minute bracts, the absence of bracteoles, and the
often glabrous pedicels.

It is difficult to relate this species to the other New World cynometras. Its
usually umbelliform racemes are suggestive of those of C. źrinitensis and C. retusa.

A fragment of a leaflet from Lanjouw & Lanjouw 1891 (U) was found to
measure 8 cm. in width, indicating that the intact leaflets reach up to 20 cm. or
more in length, thus presumably possessing the largest leaflets among the New
World cynometras.

Despite the numerous collections of C. bostmanniana very few were found in
fruit; the legumes when semeniferous, are deeply wrinkled on the surface and are
thin-walled (the pericarp up to 0.5 mm. thick), a noteworthy fact considering
that the valves are probably the largest (up to 4.5 cm.) among the New World
species. The sterile fruits are very thick walled (cf. remarks in introduction,
p. 317). Amshoff (Fl. Surinam) makes the interesting comment that the ‘infertile
Boa, REL DS splitting into 2, showing no cavity.”

names assigned to the species include "Sakabali" (British Guiana),
“ate (Surinam), “Malako” (Surinam), and “Makraka” (Surinam).

22. CYNOMETRA FISSICUSPIS (Pittier) Pittier, Arbol y Arbust. Venez. Legum.
1:132. 1928,

prs quit: Pittier, Arbol y Arbust. Venez. Dec. 2-3:25. 1923. (Ti: Pittier

10933!
— fissicuspis (Pittier) Léonard, in Bull. Jard. Bot. de l'Etat Bruxelles 19 4:402.

Trees 25-30 m. high. Trunk erect with elongated crown. Twigs smooth and
glabrous below, conspicuously lenticellate, the tips angular, puberulent. Petioles
0.5-2 mm. long, 0.1 cm. wide. Leaflets sessile, ovate to ovate elliptic, 2-4.5 cm.
long, 0.8—2 cm. wide, the narrow side of lamina 0.4—0.8 cm. wide, the wide side
0.5-1.1 cm. wide, obtuse and emarginate, the sinus about 0.5 mm. deep, otten
apiculate at base of sinus, the narrow side of lamina attenuate toward base, the
wide side of lamina auriculate at base, the costa prominulous above and below; the
principal secondary veins 10—20, slender, prominulous, the lamina drying light
green to light brown, subcoriaceous, glabrous, subequilateral to obviously inequi-

1958]
DWYER—NEW WORLD CYNOMETRA 341

Fig. 1. Cynometra fissicuspis
Flower X7; flowering twig X2; fruit X2.

lateral, reticulate. Racemes 1-2 per axil, the rachises up to 2 mm. long, the
flowers + 6 per rachis. Pedicels up to 6 mm. long, about 0.3 mm. wide, puberu-
lent. Bracts up to 2 mm. long, concave, glabrous. Bracteoles narrowly oblong,
up to 2 mm. long, about 0.5 mm. wide, vaguely carinate, sparsely hairy above middle
to glabrous. Sepals 4—5, oblong, up to 3 mm. long, up to 2 mm. wide, glabrous,

in. Petals lanceolate, up to 3 mm. long, 0.5-1.1 mm. wide, acute or subobtuse
at apex, glabrous; filaments up to 6 mm. long, about 0.15 mm. wide, the anthers
up to 0.7 mm. long; ovary rotund, up to 2 mm. long, glabrous, the style up to
3 mm. long, the stigma capitate, the ovules solitary. Legumes obliquely orbicular,
1-2 cm. in diameter, falcately apiculate, indehiscent, the valves reticulate, and
papillate or rugulose (sub mag.), a slender keel (up to 0.5 mm. wide) on dorsal
side, the wall of pericarp about 0.5 mm. thick, the seeds solitary, filling the locule.

0933 (GH, NY, US)

VENEZUELA: ZULIA: Río Lora, Santa Ana, Pitt O
LENA: Cucharo, Castañeda 641

ier I
COLOMBIA: Rio Zulia, Nicéforo s.n. (US); MADGA
(MO).

[Vol. 45
342 ANNALS OF THE MISSOURI BOTANICAL GARDEN

This is the only species among the New World cynometras which has had a
complicated taxonomic history. This is due to the fact that the type collections
distributed to herbaria are all sterile and secondly due to Pittier's description of
the leaflets as pellucid-punctate. The leaflets are not so characterized although
some show (under magnification) areas which have become thinner, probably due
to drying.

The flowering and fruiting material of Nicéforo s. n. provided a solution to
the difficulty. The flowers are obviously those of a Cynometra. Their glabrous
condition is unique for the New World species. Perhaps more noteworthy are
the coin-like (thus section Nummus) legumes, smooth and reticulate on the
surface, and the leaflets auriculate on the wide side at the base. The common name
of C. fissicuspis is “Aracito” (fide Pittier).

SPECIES TRANSFERRED OR EXCLUDED

I am transferring two species of Cynometra, apparently known only from
Minas Geraés, Brazil, to the genus Peltogyne.

PELTOGYNE glazovii (Taub.) Dwyer, comb. nov.
Cynometra Glazovii Taub. Flora 75:76. 1892.
Type Locality: Minas Geraés, Brazil.
Distribution: Known only from the State of Minas Geraés, Brazil.

BRAZIL: MINAS GERAfS: sans locality, Glaziou 13725; Glaziou 14617 (NY, P, US).

PELTOGYNE riedeliana (Harms) Dwyer, comb. nov.
Cynometra riedeliana Harms, Fedde Report 24. 209. 1928.
Type Locality: Minas Geraćs, Brazil.

Distribution: Known only from Brazil.

BRAZIL: without locality, Dorsett, Shamel & Popenoe 136b (U); MINAS GERAES: Serro
do Corcovado, Riedel 6 Luschnath 1182 (A, US); locality and State not decipherable,
Riedel 601 (?) (F, photo. and frag. of syntype collection of C. riedeliana ?).

'Two important characters make it clear that the two species are referable to
Peltogyne: the paniculate inflorescence and the glandular-punctate nature of the
sepals and leaflets. The flowers of all of the cynometras, including the many species
of the Old World, are in racemes and have non-glandular sepals. Another char-
acter of importance is the pubescence of the sepals on the inner surface; in all o
the New World cynometras the sepals are glabrous within, except rarely at the
very base. Unfortunately the fruits of Peltogyne riedeliana and P. glazovii are
unknown.

Ducke in a recent paper (1949) has contributed valuable data on the Ama-
zonian species of Peltogyne. From a study of his paper, as well as from a study
of specimens it appears that P. glazovii and P. riedeliana belong to the P. pauciflor 4;
P. floribunda, and P. discolor complex, segregated by Ducke in his key (loc. cit.).

1958]
DWYER—NEW WORLD CYNOMETRA 343

I have excluded the following species from the main body of the paper for
reasons discussed below.

1. CYNOMETRA ZAMORANA R. E. Schultes, Bot. Mus. Leaflet Harvard 13(10):
301-302. 1949.
C. zamorana is referable to the genus Hymenaea; the type of C. zamorana is
Schultes 5429 (GH) from Comisaria del Vaupes, Rio Macaya, Cachivera del
Diablo, Colombia.

2. CYNOMETRA SPHAEROCARPA Pittier, Enum. de Las Plantas Usuales 184. 1926.

Unfortunately I have not seen the type material (in fruit only) collected at
Caruao, Federal District, Venezuela. Pittier not only fails to indicate the type
collection but also mentions only a few characters which may be of importance:
"las hojas inequilanceadas, largamente atenuadas . . . los frutos son subglobosos,
de 3.5-4 cm. de diametro." Pittier states that the common name is "cobalonga”,
and adds that the wood is excellent for construction and the seeds are "estomá-
quicas y antipalúdicas”.

BIBLIOGRAPHY FOR NEW WORLD SPECIES OF CYNOMETRA
Amshoff, G. J. H. Papilionaceae. In Pulle's Flora ui eu II. ia 2:90-93. 1939.
Bentham, G. os In Mart. Fl. Bras. 15 254.

Britton, N. L. & Rose. lili ju NI In a. American Flora 23 (4):220—221.
1930.

Ducke, È. ie) on the Purple-Heart Trees (Peltogyne) of Brazilian Amazonia. In Tropical Woods

Ducke, A. ometra. In Notas Sobre a Flora Neotropica II. (Legum. Amazonia Brasileira) In
Bol. Te Toit: stata do Norte (Belem) 18:73—75. 1949.

Dwyer, J. T be ral American, West a vd South American Species of Copaifera
Z PARE ds Brit aie 7) 43-172.

Dwyer, J. D. The gto s" Genus Tied m (Caesalpiniaceae). In Ann. Mo. Bot.
Garden 41:223—260.
Dwyer, J. D. The Tropical Aia Genus Sclerolobium Vogel (Caesalpiniaceae). In Lloydia
20(2):67—118. 1957
Léonard, J. Notulae Sys tematicae IV Caesalbiniaceae-Amberstieae Africanae Americanaeque. In
Bull. Jard. Bot. Brux. XIX (4):384—408. 1949; XX (2):270—284. 1950.
Léonard, J. Les Cynometra et Les Genres Voisins en Afrique Tropicale. In Bull. Jard. Bot. Brux.
eru) :373—450. 1951
ard, J. Genera des Cynomecreae et des Amherstieae Africaines. In Mem. Sci. Acad. Royale de
* Baita 30 (fasc. 2):1—314. 1957.
Macbride, J. F. Cynometra in Flora of Peru Vol. XIII, Part 3:1943.
Pittier, H. F. Arboles Arbustos del Orden de las Leguminosas 1:131-134. 1928.
binc R. & Schery, R. et al. Een In Flora of Panama (Leguminosae, second part).
nn. Mo. Bot. Gard. 50) 125,

344

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

INDEX TO EXSICCATAE

zed numbers refer to collectors’ numbers,

s.n. (sine numero) to unnumbered

talic
Ta parenthetical numerals refer to the numbers assigned to the species in this
ograph.

Allen, P. and D. Allen 5264 (5).

Archer, W. A. 7906 (14a), 7043 (14a),
8087 (14a).
Black, G. v -13287 (18

a).
Black, G., P. Ledaux, & Stegeman 52-14-
4 (18 d
British Guiana Forest — Gr26 (18a),
6217 (9a), 6365 (21).
British Honduras Forest Dept. 5 (4).
(€ sa = = Ris r

9a), 1508 (21), 155 (20)
1958 (21), 3861 (21), 3578 (2D, 188.
(9a), 5868 (9a), 6086 (9a).
Capucho 529 (18a).
Cardona, F. SE a», 884 (9b).
Castañeda 641
Ca -— ton Garden (6).

» H. T. s.n. (3), 347 (3), 2048

(3).
Cuatrecasas, J. 7754 (18a).

Delgado 216 (9a) or (14a).
D 509 (14b), 676 (14b), 9083

19
(18a or 20), 7593 (15), 2320 (14a),
3549 (149), zrósb da, v jou
0820 (14a), IOQII
(13), 16896 (18b), I 6308 LG a
(13), 23788 (20), 24247 (142), 24248
(14a), 35415 (14b).
Eggers, H. F. 15752 (11).
Ekman, E. L. H2343 (1), H4563 (1),
pi (3), 12684 (2), 12978 (2),
7375 RICO ).
bali 310 (1).
Ferris, R. 6188 ene? (10).
Froes, R. L. 21043 (18a), 23600 (9a),

=: (21).
Froes, R. L. & Murga 24014 (9c)
nia 1452 (3).
Goeffrey s.n. (11).
Glaziou, A. 13756 | Sa po (18a).
Gleason, H. A. A-2I (3), A-
136 (3)

Gonggryp, J. W. 457 (21), 2568 (21),

3751 (20), 5357 (21).
Gonggryp, J. W. & G. Stahel roro (9b).

Haller s.s. (21).
or: $ 40 [^n

erb. N. Y. Gard. a. 6217 (18a).
tinge. 10893 (
Hostmann, F. W. A “als di (20).
Hottle 23 (4).

Im Thurn, E. F.
Irwin 814 (18a)
Jenmann, G. S. 21).

Jiménez, J. 458 G (69,205 (3).

£183).

n,

Jórgensen, J. hs (18c).

Kappler, A. 1905 (20).
Killip, E. P. ser (18a).
Kne 1448 (14a).
Imann, EE 1067 (13), 2919 (182),
ran
Krukoff, B. i 6439 (12), 6631 (12),
8418 (12).

puis J. 858 (9a), 1219 (92).
Lanjouw, J. & J. Lanjouw 1891 (21).
Lindemann 2964 (3a);

20)
Langlassé, E. s.n. (10), 441 (10).
León, Bro. 13552 (2), 13805 (2).
Leonard, G. M . C. Leonard 12748

Maguire, B 240404 (9a), 32705 (9a).
Maguire, B. & Maguire 4047 (18a), „2304
(9a), 23906 (9a), 24040 (9a), 2
19).

. 21).
Mell, s. z. (21,17 (182).

Nene 7004 (10).
Newman, L. M. s.n. (18a).
Nicéforo, IL tn. (22):

1958]

DWYER—NEW WORLD CYNOMETRA 345
Paul, Bro. 463 (9a). (14a), s.n. (14b), 872
Pedersen, T. 1295 (18c). że ws (14b), 1513 (14a), 3479
Pires, J. M. r327 (182), R vp (14a).
Pires, J. M. & Silva 4210 (14 Stahel, G. 1984 (
Pittier, H. F. 216 (14a), ae Q2). Standley, P. ihe te, 24254 (4), 52915
Purpus, C. A. 7383 (10). (4), 56686 (4).

Standley, P. & Valerio 48799 (5).

Richard, L. C. M. s. s. (13). Steyermark, J. 38671 (4), 44557 (4),
Record, S. 8832 (4). 45034 (18a), 45565 (4).
Rodrigo 782 (18a). Tessmann, G. 3444 (18a), 3636 (14a).

Rovirosa, J. N. 748 (4). Tresling, J. 300

a).
Trinidad Bot. Gard. (6).
(5).

EP AC Kw (21), 1003 (21),
i de la R. s.n. (2), 307 (2). PDA ER
Schipp, N. A. 1192 (7). Ule, E. 7776 (18a), 8864 (14a).
Schomburgk, Rich. s.n. (9b), s.n. (18a),

Vaillant TOI

(21).
197 (168) (14a), 230 or 231 die cit Versteeg, G. M. 552 (21), 576 (21).

(18a), 777 vo 1533 (9b).
Scho pup t. (Herb Hance 7667) Williams, Llewelyn 40 (18a), 1984 (18a),
ot 182), 14a).
ead c XI 357 (8). E val. T I a), 15593 (14a)
Schultes, R. 8 I. Cabrera 12616 (14a). R. 8 (21), 1656
Schwacke, C. A 358 (14a). villi H. 1438 (21), 165

Sintenis, P. E. E. 5600 (3). Yuncker, Koepper, & H. Wagner $007
Snethlage, E. H. zwał (14a). (4).

GENERAL INDEX TO VOLUME XLV

New sci ientific names of plants and the final members of new combinations are printed

in bold-face ty nonym
n. and all feri matter, in ordinary ty

s and page 2^ having reference to figures and plates, in

a 76; pentagonus, 76, 77
96

Asics: 227; levyana, d 228; purpuras-
cens, 228; rostellata, 22

Acitentheni,. 229; rari 231; quadrata,
229, 230; recurva, dp uniflora, 230

Acrolasia, 32; squalida,

Adamarum, 1 3

na dor e 205; adscendens, 206, 2

Adenaria, 112; floribundi: 113, is var. a,
114, var. australis, 114, var. microphylla,
114, var. parvifolia, 114; grisleoides, 114;
lanceolata, parviflora, 114;

01; pygmea, 100; sagittata,

102; sanguinolenta, 102, subsp. longifolia,
ut subsp. purpurea, 102, subsp. pe
2; stylosa, 102; texana, 102, ramosa,
ite! ramosior, 100, 101, 102; Mec

102
n G. J. H.: Myrtaceae of Panama,

Asini, ^F fragrans, 182
Antbactinia
Asfbérylbee foribindem, 114; grisleoides,

alatum, rd campanu-

dowie 2 T 2z

Asophora, 137

Astepbanantbes, 1

p dire

Augu

Allies jd completens, 304

Aulomyrcia, 170; coumeta,
tosa, 171, 172; zetekiana, 1

173; tomen-
170

Axinaea, 209; costaricensis, 209, 270

Badamia,

Baldwins peltata, 12
Balsamona, 104; pinto, 109.
Barona

ać Li allenii, 62; barbana, 49; brevi-

elliptica, M: nllo 64; > pes, 55»

; fische ibi: 55, 5
osa, 56; A , 46; 47; PAP 54,
55; glabra, 61; allude! 46; guadu-

eydei, 59,
62; csi great de. var deg ; paa er
volucrata, 59, iosa, 60; laur
64; leptopoda, 36; locllat, ga Willa
54; + 5 a, 56; m nticola,

multinervia, 50, 60;
folia, 44; oaxacana, 58, 59, B pilosu la, 58;
opuliflora, 63; ottonis, pi bey slifoli,
54; pittieri, 62; plebeja, 45, 46;
folia, 57; dubite dicella, sp geld 3,
rosea, 56; scandens, 54;
43; PYTA h 66; siini-
ovata, 55, 56; pko 64; serrulato-
ala, 58; spruceana, 56; squarrosa, >
stigmosa, 50; strigillosa, 49, 50; ton nduz.
ov.

ticifolia, 61; uvana, 46; vestita, 51, 52;
illipetiola, 48, 49; s
Begoniaceae of Pan
Begonie
Bellucia, 258; costaricensis, 258, 259
Bergena,
ao

Bert folowie Ma, 239
(347)

348

Besseya, 306
Blakea, 239; BRA, 242; calycosa, 241;
crinita, 241; foliacea, 242; gracilis, 242;
parviflora, 243; Bacio 243; wood:
T

Boykinia, 9i: iudi 100
Buceras, 157; bucida, 157
Buchenavia, 159; capitata, 158, 15
, 157; buceras, 156, 157; SA
9

Bureava, 145

C

Cacoucia, 144; deco, 150

Cactaceae of P

Cactus ble 25 5 pp T 74; elatior,

75; lucidus, 70; nigricans, 75; pendulus,

91; EAE 76; baia 70; phyl-
lantbus, 83; pitajaya, 76; prismaticus, 76;
una B elatior, 75, y nigricans, 75

Calopbylloides, 179

Calopbysa cu 254; tococoidea, 254

Calycampe, 17

Calycolpus, 192; Peritorai, 196; warze-

iczianus, 103,

Calyplectus, 110

Calyptbrantbes, 166

Calyptranthes, 166; SP o 168, 169;
costaricensis, 169; hylobates, 169; pit-
tieri, 167; s ATA dh 167; pi Anali
daliana, 167; tonduzii, 198; tumidonodia,
167; urobbylla, 16

rella, 212; micrantha, 213, 2
Cal M irchedien m, 196; rcrichthalmen,
; sartorianum, 19

Cempylochiton, 1

Carica, 26; chiriquensis, 30, 37; cucurbiti-
folia, 29; dolicbaula, 23; bermapbrodita,
26; mamaja, zi: RY» 26 :27

Caricaceae of P

Carpophyllus, 6

Casbarya, 41; coc bae: 61; columnaris, 61,
p eante 6 1; trachyptera, 61; urticae,

1, 8 hispi
E ies 140; we 142; cubensis, 142;
ign I41, 142, B alba, 142, vy pauci-
, 142; Bees 142; "bodanibe,

Catappa, 155
Catinga, 179

Centradenia, 218 ji inaequilateralis, 219;
maxoniana, 219, 2
Centronia, 212; grandiflora, 212; tomen-
2II,212

Cératosebulum, 1; micrantbum, 15

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Cereus p Mtis ie 76; subgenus
Hylocereus 78; acu tangulus, 76: costa-
ricensis, „30 dus
nitidus, 7 A 76;
83; IAY 76; polyrhizus, 80; pa
76; prismaticus, 76; sirul, 76; trigonus
var. costaricensis, 80; variabilis, 76;
vasmeri, 7

om»yrtus, 180

Chrysostachys, 145; ovatifolia, 151

Cbrysostoma, 32

Chuncboa obovata, 155

Cbuncoa, 153; amazonia, 155; obovata, 155

Cbytraculia, 1

Cbytralia, 166

1; angustifolia, 13; appendiculata,
e

pallida, 13;

pe
pseudo. set 13; suberosa, 12; viridis,

3
era 41
Clidemia, 247; barbinervis, 289; capitellata,
252; SR, 253; densiflora, 255; den

266; taurina, 254; tococoidea, 25
rig mexicanum, 265
Coleophora
Cane w Panama, 143
Combretum, 144; sect. ded S. 145; sect.

tum
Er

jacquinii, 151;
axum, 140, 15
petalum, 147;

*
o
U

xum,

loeflingii 147,

1958]

INDEX

poro ate 147; marcbii, 151; mi-

150; pulchellum, 151; punctulatum, 152;
sambuense, 149; secundum, 147; spino-
` sum, 140, 152; superbum, 147; terminali-
oides, 151; variabile, 151; viscidum, 151;
warszewiczianum, 14
Conocarpus, e acutifolius, 161; am
160, 161, va 1

y
a, 273; setosa, 270; sororia, 2 s
Th 47 subcrustilita; 274; xala-

Consolea, 75
Copaifera fissicuspis, 340; hemitomophylla,

Cornelia, 10

Couratari, e panamensis, 134, 135

Courou 125; cutteri, 125;
id "|", | 12 i

Creolobus, 5

Cristaria, 1
dorus ay > 270

cn ptotbeca

Cuphea, 104; canti —_ 108; appen
diculata, 110; bals 19% calophylla,

bifolia,

b

Cyatbocnemis, 41

Cynometra, The New World Species of,
313

Cynometra: economic importance, 318;
generic relationships, 313; geographical
similia 318; history of the genus,
313; interspecific relationships, 318; key
to spp., 320; morphological characteris-
tics, 315

349

sect. Comatovaria, 320;
sect. ki pr 520; sect. Nummus, 320;
american debili în: 336, var.
m 337, var. £randiflora, 558,

Cynometra, 319;

» 341;
325; hon-
microflora, 338;

334, var. B procera var. spruceana,
333; Ira a 331; trinitensis, 325;
zamorana, 343

Dactylopetalum, 1

Daphnopsis, 94; serii 94, 95

Davya macro phy Ila,

Decaloba, 1; ce asi E var. major, 7, Var.
mexicana, 7; cyathophora, 11; bolo-
sericea, 11; jacquimi, 14; punctata, 9;
sexflora

Diolena, 236; spicata, 236

4

Diis 1

Distep È
Distephania, 1
Distepbia,

Ditheca, 101

Donaldia, 41; ottonis, 64

Doratometra, 41

Dwyer, John D.: The New World Species
o metra,

Dysosmia, 1; acerifolia, 15; ap 21;
pko 21; foetida, 21; sypii-
folia, bastata 21; bibiscifolia, 21;
la

E
he 125
Embryogonia, 145
Endecaria, 104
Epiphyllum, 82; costaricense, 86; ee
83; gigas, ibe epidocarpum, 8
nius; 83, ea lin

Epleienda, 179

350

Erndelia, 1

130; calyculata, 133; gara-

garae, 133; panamensis, 130; ne 131,
132; reversa, 130; verruculosa,

sp., 187; ini 188,

181; antiquae, 189;

in-smithii, 91; mesi 187;

floribunda, 177; fragrans,
2; 18 3; hiraeifolia, 189; jembos, 176;
177; m

192; ono 186;
aman-
ana, 190; sericiflora, 184; sorki, 182;
tapacumensis, 190; tomentosa, 172; uni-
flora, 182; vallis, 172; vismeaejolia, 179;
warscewiczii, 194; zetekiana, 1

A. W.: Combretaceae of Panama,

F
Falkea, 41
Fatraea, z:
Ficindica
nti 144; laevis, 150

G
Gaerdtia, 41
Gaura Fai icosa
Gimbernatea, Ds ye 5155
Gireoudia, 41; c tl. 43; nelumbii-
folia, 45; stigmosa, 50; strigillosa, 49;
warscewicziana,
Sa H. A.: Melastomataceae of Pana-

Sia rpus, 144; jacquinii, 15

Granadilla, 1; foe tida, 21; PP
18; bero. 12

Greggia, 179

Gr Rhizophoraceae of

Grias, 122; fendleri, 124; pittieri, 123, 124
Grislea, 14

Gronovia, 40; gens: 39, 40

Guajava mollis, 2

Guibourtia ym 340
Gurltia, 41

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Gustavia, 117; brachycarpa, 119; insignis,
20; nana, 120; pleurocarpa, 118; su-
perba, 120, 121, var. salviniae, 120

H
Haag
H uides 144
Hambergia, 14
Hapalocarpum, 101
Hargasseria
Hariota, 89; cassytba
Heeria, 223; cupbeoides,

Henriettea, 256; AS 260; succosa,

257, 25
Henriettella, 260; densiflora, 255; fascicu-
laris, 262; seemanii, 261, 262; tubercu-
osa,
Hesperaster, 32
Heterocentron 223; gwen 221,2
oe 2 ovemnervium, A
um, 277, 2
Heche. 179
Hoffmannella rosea, 56
Horau,
Hadionia, 159; arborea, 159
Wu 36
Huszi
Hadvol pia
Hapus "Ai pros. var. tenuifolia,

ini 78; costaricensis, 80; monacan-
thus, 79, 80; polyrhizus, 80

I

Irm
re send 102
Isopteryx,

J
a 175; domestica, 176; malaccensis,
76; samaragnensis, 177; gu ulgari, 1 176
p Aa 117; superba, 1
Jacaratia, 23; costaricensis, = dolichaula,

aoe
Jossinia, 179
Jussiaea sagittata, 102

K
Klaprothia, 34; mentzelioides, 35
Knesebeckia,
Kniphofia, 153

— : = punicifolia, TII, 112

ap , 162; obovata, 162; racemosa,
“162,
zii e Bipenicllt, 234
Lauchea,

1958]

INDEX

Leandra, 265; consimilis, 268; ai ge
266; melanodesma, 266, 267; mexic
265; strigosa, 268; cubeatiatà; 266; lab
ulata, 268

Lecythidaceae of Panama, 115

Lecythis, 127; ampla, 128; rigori 128;
melliana, Do tuyrana, 128, 129

Lecytbopsis

a 2 "io: gira 142

Lepsia

Ike ei bipinnata, 36; grandis, 38;
rhoeadifolia, 36; rudis, ks. 37; speciosa,

0; brunnescens, 260
Pul i area 10
Lythraceae of Panama, 97
Lythrum catthagenensis, 109

M

Macropbora, 1; sanguinea, 1
Maieta setosa, 254; eser 254

Mangle, 137

Meioperis, 1; angustifolia, 13; bederacea,
13; minima, 13; ball. pe 13; pannosa, 5;
peltata, 13; ‘subero. osa,

Melanium, 104

Melastoma adscendens, 207; albicans, 292;
lata, 252; peras 285;
ir
spin
971

4, 282; nervosum, 29

prasina, 290; purpurascens, 228; rubens,
301; rubiginosa, 293; rubra, 255; strigil-

losa pini ; succosa, 258; tbesezanz; 299;
la apen e, 271
Melastomataceae of Panama, 203
Melfona, 104

Melvilla, 104

Mentzelia, 32; aspera, 33, 34; pediceliata,
34; stipitata, 3

Meriania, 207; macrophylla, 209; panamen-

Amblyarchena, 297;

Jucunda, 282; sect. Miconia, 284; sect.

351

Tamonea, 283; aeruginosa, 286; „albicans,

ciliata, 296; costaricensis, 298;
petiolata, 296; cuspidatissima, 300; dari-
IH. 290; ‘disparilis, 282; gatunensis,

ittieri, 300; prasina, 2%) pteropoda,
0; binsoniana, 2 bens, 301;
rubiginosa, 293; ru f uice 294;

schlimii, 298; septuplinervia, 292; shat-
tuckii, 302; stenostachya, 287; sub-
lado: 274; theaezans, 299; trans-
versa, 296; Ap rg 286

Microjambosa, 17

Mitrantbes Mant. 198

Mitropsidium x. 198

Mitscberlicbia,

Mona dx iras: > angustifolia, ię cori-

10; eracea, 12; minima, 12;

pelata, est suberosa, 12

Monochaetu 214; bracteolatum, 216;
cymosum, ET 5, 216; macrantherum, 216;
rivulare, 216

Monolena, wd ovata, 236, 237

Moschkowitzia, 41

Mouriri, brunneicalyx,

30 "s com-
pak 304; parvifolia, 303,
1

304;
304
Muru
Masa, wł subgenus Aulomyrcia, 170;
rare s costaricensis, nd ki A
coumeta, 173; gatunensis, 175; tede-
a 175; Arrone 175; Diana
82; A 172
cada.
Myrciaria, Ea oriunda 2 178; o'neil-
lii, 177; vismeacfolia,

Myrobalanfere, 1
yrobalanus, 153; ga 157; obovatus,
55

>
Myrtaceae of Panama, 165
Myrtinae, 165
Myrtopsis, 180
gus biflora, 184; fragrans, 182; mari
189; elei. 196; samaragnensis,

352

N

Nephromischus, 41
dis 226; aquatica, 226, 2
Nevling, Lorin I.: seb fos di of Panama,

Nimmoia, 98
Nopals, 72; cochenillifera, 73, 74; dejecta,

N rar mannia, 94

Odostelma, 1

e 179

Opa,

One 79

Opuntia, gee coc degni MER a dejecta, 74;
elatior, M 5; pbyllantbus,

Ortiga, 5

Ossaea, "b: disparilis, 282; diversifolia,
264; micrantha, 263, 264; trichocalyx,
264

Oxydiastrum, 180

Pamea, E

Panel, 1

Papaya, pe sativa, 26; vulgaris,

Parsonsia; los calopkylla, i Miel

07; pinto, 109; utriculos

Passiflora, r zbate În 15; SATEH 17;
sdenopeds; 15; alba, 17; ambigua, 1
angustifolia, 12;

aquatica, 13; capsularis var. geminiflora,
; aponioides, 11; ceratosepala, 15;
ciliata, 21; cinerea, 11; clypeata, 10;

oribunda, 5; foetida, 21; glabra, 12;
glabrata, 7; globo 13; s gonybifoli 21;
Di

ZE ns ih
13; Deere 12; fne 12; longilobis,
9; 1 M » 7, var. costa = 7; macrocebs,

8; miliana, 9; mbranacea, 14;
ricarica, 17; ; microcarpa, 9; min
ma, miraflorensi

;
moritziana, 2
21; nigra, 12; nitida, 20; nympbaeoides,

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

20; obtusifolia, 10; oerstedii, 19; olivi-

formis, 12; orbifo lia, 20; pseudo- d
pulcbella, 14, var

unicea, 15;
pur pusi, 19; pallida, 12; panamensis, 8;
pannosa, 5; parviflora, 12; pediculata, 12;
Sellada 12; populifolia, 19; pseudociliata,

a

torta, 11; translinearis, 9; transversa, 7;

a 13; triflora, 5; MPH
oides, 4; vesicaria, 21; vespertilio, 9 ;

villosa 13; vitifolia, 15; warei, 13; wil-

liamsii, 16

Pássifloraceae of Panama, 1

Peires

Pekea, 1

M > 342; riedeliana, 342

neo 53

peli orcilental 100

Pica. 69; aculeata, 70, 71, B longispina,

sis, 72; beréskio, 70; undulata, 70
Pericodia, 1
ip spade 41
Pbyllocactus, 82; costaricensis, 86; gaillar-
idocarpus, 85; ma acrocarpus
phyllanthus, 83;
nus, 86

Pi

Pesa, n es 122; superba, 120

Platycentrum, 41

Platyclinium, 41

Poivrea, 144; alternifolia, 152; eriopetalum,
152

Poggendorffia, 1
Pogonorbyncbus amplexans, 284

Prosanerpis panamensis, 255

Pseudoeugenia, 180

d 87; bimantoclada, 88

Psidiastrum

Psidium 19 6; araca, 201; sete 196;
friedrichsthalianum, 200; gen 196;
guajava, 200; guineense, 201; walle; 201;

was zywa 196; polycarpon,

1958]

INDEX

ria 200; pyriferum, n salu-
96,

tare; 107; sartorianum, 8, 109;
dw dé

Psilantbus

Pterogastr. Vitia ides,

Pterole pis, 231; udi ji pumila, 232
Putzeysia, 41

Q
ek 98
Quirin
Cai indica, 144
R
Rachia, 41
Rat bea, 1

Reichenbeimia, 41
Resinaria, 153
Rbexia inaequilateralis, 219; limnobios, 231;

longifolia, 234; pumila, 232; recurva,
230; uniflora, 23
Rhipsalis, 89; assytha,

himantoclada, 88; tea, 91; ai
tS,
Edie. 162; alba, 162
Rhizophora, 137; eee 137; brevi-
styla, 140; mangle, 137, var. samoensis,
139; racemosa, 140; wa 138, 139
rę skapnie? of Panama, 136
Rhyacophila, 9
Rh seiner 217; mexicana, 2/7, 218;
18

Rotala, E S unir 100; mexicana,
p. bierniana, 100; subsp. pusilla,

si nc 100, var
100, var. interior, 100; varticilleris,
Rudbeckia, 161

Sagraea epiphytica, 2

Sassea, 41; columnaris, 6 6 1; pet 61; boff-
oe 61; urticae

Saueria, 41

Scbriiwedlchs, 41

Schizocentron, 223

Sch oenobiblus, 95; panamensis, 96

Scbousboea, 144; coccinea, 150; commutata,

162

"eur, pne: G., > e. B. Smith:
Begoniaceae of Pan
liane 224; copies, 224, 225

353

Seguier

pa ów , & Bernice G. Schubert:
sc 2 bunk ama, 41
SR
Spennera rail 228
Sphenocar pus,
Rys met o

1

179; michelii, 18

Stibadotbeca, 41; ee dea 61
179

1
Synthyris ranunculina
Synthyri is, 307; pri Ade and variation
s: 309; rinünculing. number of flower
311

>
Syzygium, 175; jambos, 176; malaccense,
176; samaragnense, 17

T

Tacsonia, 1; buchanani, 15; sanguinea, 15

3; amazonia, ; buceras,
ui nsis, 154,
155; erecta, 16 1; bayesii, 155; bilariana,
159; obovata, 155, 159; odontoptera, 155
Thymelaeaceae of Panama, 95
T 233; Adasia ta, 234; nana,
4; aa 233; lossitolia, 234, 235;
palic

lleida 41

Tococa, 275; RAE 276, 277

, 245; cordata, 244;

ce oif wę 247; mi-
sky ialis, 245; praecox,

246; regeln 246; ale 245

Torrey

ruck. a

Trachelocarpus, 4

Trachylobium edis 333

Trachyphytum, 32

suspi 41

Triolen zag? 8; hirsuta, „230 239
Tripsilina, i; A HE

U
Ugni, 194; warscewiczii, 194, 705

354
V
Vicentia
Vouapa gta 334
W

W ageneria, > deflexa, 54; glabra, 54;
lucida, 54; montana, 54; moritziana, 54
wiem d 81; panamensis, 81, 8

[Vol. 45

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Weibea, 140
Lc 41

ER E Synthyris ranunculina, 305
vu

Wittia, b cinstite 88; himantoclada,
88, 89

88; panamensis,

X

Xerogona, 1

MISSOURI BOTANICAL GARDEN

i

- EDGAR ANDERSON, | —
.. Curator of Useful Plant

e Director