Volume 2 NO VON
Number 1
1992
A New Species of Dicranodontium (Musci: Dicranaceae) from Panama
Bruce Alien
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT. The presence of an autoicous species
in Dicranodontium (Dicranodontium interme¬
dium , species nova) further supports Williams’s tax¬
onomic decision to unite Atractylocarpus and Di¬
cranodontium.
Dicranodontium is a group of medium-sized
mosses typically having narrow to setaceous leaves,
broad costae, linear leaf cells, nonsheathing peri-
chaetial leaves, and cygneous setae. The genus is
gametophytically identical to Atractylocarpus: the
genera differ only in sexuality (dioicous in Dicran¬
odontium, autoicous in Atractylocarpus) and the
stance of their moist setae (cygneous in Dicrano¬
dontium, erect to flexuous in Atractylocarpus).
Based on his observations on the variability of setae
curvature within identical species, Williams (1913)
synonymized the two genera. The following new
species of Dicranodontium, which has a cygneous
seta but is autoicous, adds further support to Wil¬
liam’s taxonomic view of these two genera.
Dicranodontium intermedium Allen, sp. nov.
TYPE: Panama. Chiriqui: Volcan Baru, 8°45'N,
82°30'W, Allen 9123 (holotype, MO; isotypes,
NY, PMA). Figures 1-7.
Species nova D. meridionali affinis, a qua differt facie
grossiore, sexuali statu autoico et cellulis interioribus bas-
alibus foliorum incrassatis, porosis.
Plants medium sized, in loose, erect, greenish
yellow tufts, epiphytic or occasionally terrestrial.
Stems moderately to densely white or red-tomentose,
up to 5 cm long, branching irregularly; epidermal
cells thick-walled, red-brown. Leaves evenly and well
spaced, erect-spreading to falcate; concave below,
tubulose above, narrowly lanceolate from a shortly
ovate base, 8-11 mm long, apex long setaceous,
sharply and densely denticulate above. Upper leaf
cells linear, firm-walled; median leaf cells long-rect¬
angular, incrassate and porose, occasionally smooth-
walled; basal cells near the costa rectangular, in¬
crassate and strongly porose, the outer basal cells
longer and narrower forming a short, hyaline border;
alar cells well developed, red-brown, occasionally
fugacious. Autoicous. Perigonia terminal on short
branches (1.5-5.0 mm) below the perichaetia. Setae
yellow becoming red, smooth, 10-13 mm long, red,
twisted and erect when dry, cygneous when wet.
Capsules erect and symmetrical, smooth or weakly
furrowed when dry, oblong, 2.0-2.5 mm long, light¬
ly roughened at base; peristome yellowish becoming
dark red; teeth 0.5 mm long, deeply divided (at
times almost to the base), dorsal surface vertically
striate. Opercula 1.5 mm long. Calyptrae 3.0 mm
long, entire at base. Spores lightly papillose, yellow,
20 /urn.
Paratypes. Panama. CHIRIQUI: Volcan Baru, Allen
9127, 9132, Davidse & D’Arcy 102901 (all MO).
Epiphytic on branches in subcanopy, on tree
trunks, fallen logs and occasionally terrestrial be¬
tween 2,300 and 3,505 m.
Dicranodontium intermedium has broad leaves
that are atypical for the genus. In gross aspect it
looks very much like a species of Campylopus.
Furthermore, its well-developed alar cells and in¬
crassate, porose basal leaf cells are common features
of that genus. However, its linear upper leaf cells
and autoicous sexual condition cannot be accom¬
modated in Campylopus. Three species of Dicran¬
odontium (sensu Williams, 1913) are now known
from Central America. They are separated in the
following key.
Novon 2: 1-3. 1992.
2
Novon
Volume 2, Number 1
1992
Allen
Dicranodontium intermedium
3
Key to the Species of Dicranodontium in Central
America
la. Inner basal leaf cells enlarged, thin-walled, non-
porose and strongly contrasting with narrow,
incrassate and porose outer basal leaf cells; di-
oicous . D. meridionale Bartr.
lb. Inner basal cells not differentiated from the
outer basal cells, either firm-walled or incrassate
and porose throughout; autoicous.
2a. Leaves broadly ovate at base, basal and
median leaf cells incrassate and porose,
alar cells well developed; setae cygneous
when wet . D. intermedium Allen
2b. Leaves linear to shortly ovate at base, basal
and median cells firm-walled straight or
weakly porose, alar cells poorly developed,
fugacious; setae erect to flexuous when wet
. D. longiseta (Hook) Williams
Acknowledgment. I thank Jan-Peter Frahm for his
opinions and comments on this species.
Literature Cited
Williams, R. S. 1913. Dicranaceae. N. Amer. FI. 15:
77-158.
Figures 1-7. Dicranodontium intermedium Allen. —1. Sporophyte. —2. Leaf apex. —3. Leaf. —4. Lower leaf
cells. —5. Peristome teeth, outer (dorsal) surface. —6. Upper leaf cells. —7. Alar cells. Scale bars in mm. Drawn
from the holotype.
Draba barclayana (Brassicaceae), a New Species from Colombia
Ihsan A. Al-Shehbaz
Missouri Botanical Garden, P. 0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT. Draba barclayana , a new Colombian
species endemic to the Cordillera Oriental, is de¬
scribed, and the characters distinguishing it from
the nearest relative, the Ecuadorian endemic D.
spruceana, are given.
Of the approximately 65-70 species of Draba
indigenous to South America (Al-Shehbaz, 1991),
at least 21(16 endemic) grow in Colombia (author’s
compilation). Several of these have been described
only recently (Al-Shehbaz, 1989; Rangel & San¬
tana, 1989), whereas others belong to species com¬
plexes in need of critical study. The following spe¬
cies, named after Harriet G. Barclay, one of its
collectors, belongs to a difficult group, which Schulz
(1927) recognized as section Adenodraba O. E.
Schulz. As indicated by Fernald (1934) and Al-
Shehbaz (1987), however, Schulz’s (1927, 1936)
sectional classification of Draba is evidently artifi¬
cial, and a thorough study of the entire genus is
needed before meaningful infrageneric divisions are
recognized.
Draba barclayana Al-Shehbaz, sp. nov. TYPE:
Colombia. Boyaca: Cordillera Oriental, Sierra
Nevada del Cocuy, Alto Ritacuva, wet areas
with cushion plants ( Distichia) surrounded by
Kspeletia slopes, SW facing, alt. 4,400 m, 16
Apr. 1959, Barclay & Juajibioy 7398 (ho-
lotype, GH).
Herba perennis prostrata; folia basalia rosulata, oblonga
vel oblanceolato-spathulata, 8-15 mm longa, 3-4.5 mm
lata, glabra, ciliata, pilis simplicibus et furcatis; infruc-
tescentia subumbellata, ebracteata, usque ad 8 mm longa;
pedicelli fructiferi divaricati, 4-6 mm longi; petala alba,
obovata, 3.5-4.5 mm longa; antherae ovatae, subapi-
culatae, 0.6 0.7 mm longae; siliquae ovato-lanceolatae,
5-7 mm longae, 2-2.2 mm latae; semina ignota.
Prostrate perennials. Stems branched, to 7 cm
long, the branches slender, terminating in clusters
of subrosulate leaves, sparsely pubescent when young,
soon glabrescent. Basal leaves oblong to oblanceo-
late-spatulate, 8 15 mm long, 3 4.5 mm wide, sub-
sessile, base cuneate-attenuate, margin entire or ob¬
scurely 1-toothed on each side, ciliate with simple
or long-stalked furcate trichomes to 1 mm long,
apex obtuse to subacute; cauline leaves similar to
basal ones, gradually reduced in size upward, when
young minutely pubescent on both surfaces with
simple or furcate trichomes to 0.1 mm long, soon
glabrescent. Inflorescences ebracteate, corymbose
racemes, hardly elongated in fruit; infructescences
subumbellate, 3 8 mm long; fruiting pedicels di¬
varicate, straight, minutely pubescent, 4-6 mm long.
Sepals ovate, 2-2.5 mm long, ca. 1.5 mm wide,
erect, nonsaccate, sparsely pubescent with simple
and furcate trichomes to 0.1 mm long. Petals white,
ohovate, 3.5-4.5 mm long, ca. 2 mm wide, sub¬
truncate, attenuate to a clawlike base. Filaments
subequal, conspicuously dilated at base, ca. 2.5 mm
long; anthers ovate, 0.6-0.7 mm long, subapiculate.
Nectar glands ringlike, subtending the bases of all
filaments. Fruits ovate-lanceolate, 5-7 mm long, 2-
2.2 mm wide; style less than 1 mm; valves and seeds
not seen.
Draba barclayana , which is so far known only
from the type collection, is most closely related to
the Ecuadorian endemic /). spruceana Weddell. It
is readily distinguished by its ebracteate, subum¬
bellate infructescences to 8 mm long, oblong to
oblanceolate-spatulate leaves with glabrous surfaces
and ciliate margins with simple and long-stalked
furcate trichomes, obovate, subtruncate petals 3.5
4.5 mm long and ca. 2 mm wide, and anthers 0.6-
0.7 mm long. In contrast, D. spruceana has brac-
teate (at least along the lower third), racemose in¬
fructescences to 4 cm long, ovate to ovate-oblong
leaves with sparsely pubescent surfaces and margins
with long-stalked cruciform trichomes, spatulate,
shallowly emarginate petals 2.2-2.8 mm long and
0.8-1.2(-1.5) mm wide, and anthers 0.3-0.4 mm
long.
Literature Cited
Al-Shehbaz, I. A. 1987. The genera of Alysseae (Cru-
ciferae; Brassicaceae) in the southeastern United
States. J. Arnold Arbor. 68: 185-240.
-. 1989. New or noteworthy Draba (Brassica¬
ceae) from South America. J. Arnold Arbor. 70:
427-437.
-. 1991. Novelties in Draba (Brassicaceae) from
Venezuela, Ecuador, and Peru. Novon 1: 67-70.
Fernald, M. L. 1934. Draba in temperate northeastern
America. Rhodora 36: 241 261, 285-305, 314—
344, 353-371, 392-404.
Novon 2: 4-5. 1992.
Volume 2, Number 1
1992
Al-Shehbaz
Draba barclayana
5
Rangel, J. O. & E. Santana. 1989. Estudios en Draba
(Cruciferae) de Colombia. 1. Cuatro especies nuevas
de la Cordillera Oriental. Revista Acad. Colomb. Ci.
Exact. 17: 347-355.
Schulz, O. E. 1927. Cruciferae — Draba et Erophila.
In: A. Engler (editor), Pflanzenreich IV. 105(Heft
89): 1-396.
-. 1936. Cruciferae. In: A. Engler & K. Prantl,
Natiirlichen Pflanzenfamilien, ed. 2. 17B: 227 658.
Three New Species of Masdevallia (Orchidaceae) from South America
Carlyle A. Luer
Missouri Botanical Garden, l\0. Box 200, St. Louis, Missouri 63166, U.S.A.
ABSTRACT. The following three species described
as new in Masdevallia represent three sections of
subgenus Masdevallia. Masdevallia goliath is a
successively flowered, round-stemmed species of
section Coriaceae subseclion Durae. Masdevallia
mascarata is a member of section Polyanthes sub¬
section Alaticaules, distinguished by triquetrous pe¬
duncles. Masdevallia ricii is a member of subsection
Saltatrices, noted for the long sepaline tubes, of
section Masdevallia.
Masdevallia goliath Luer & Andreetta, sp. nov.
TYPE: Ecuador. Prov. of Morona-Santiago:
epiphytic, near Bomboiza, alt. 1,700 m, col¬
lected by A. Andreetta, flowered in cultivation
at Paute, 24 May 1088, C. Luer 13629 (ho-
lotype, MO). Figure 1.
Species haec Masdevalliae ayabacanae Luer affinis,
sed sepalorum caudis longioribus illis lateralium approxi-
matis, petalis minute asperatis et labelli marginibis api-
calibus valde revolutis differt.
Plant medium to large, epiphytic, caespitose; roots
coarse. Ramicauls stout, erect, 2-3 cm long, en¬
closed by 2-3 tubular sheaths. Leaf erect, thickly
coriaceous, narrowly obovate, obtuse, 10 23 cm
long, 2.5-3.5 cm wide, gradually narrowed below
into an indistinctly petiolate base. Inflorescence a
congested, successively few-flowered raceme borne
by a terete, purplish, ± horizontal to ascending or
descending peduncle 20 35 cm long, with 2-3 close
bracts below the middle, from low on the ramieaul;
floral bract tubular, 15 mm long; pedicel 10-15
mm long; ovary 10 mm long; sepals fleshy, dark
red-purple, glabrous externally, minutely glandular-
pubescent within, the dorsal sepal narrowly trian¬
gular-ovate, 20-30 mm long, 12-15 mm wide, the
acute apex attenuated into a slender, erect, orange
to green tail 6-9 cm long, connate to the lateral
sepals for 6-9 mm to form a short, conical tube,
the lateral sepals ovate, oblique, ca. 35 mm long,
± prominently veined within, connate 17-23 mm
into a concave lamina, 26 34 mm wide, the acute
apices approximate and attenuated into tails 5-9
cm long; petals cartilaginous, yellow, dotted with
purple, minutely asperate-verrucose, broadly ob¬
long, shortly unguiculate, 5.5 mm long, 3.25 mm
wide, the truncate apex bilobed with a thick, obtuse
apiculum in the sinus, both halves callous, shallowly
sulcate centrally; lip yellow-orange, dotted with red-
purple, thick, oblong-ligulate, 6 mm long, 2.75 mm
wide, the apical third thickened, verrucose, with
revolute margins, the apex rounded with a thick,
verrucose callosity beneath, the disc shallowly sul¬
cate, the base subcordate, hinged beneath; column
cream-colored, stout, semiterete, 4 mm long, the
foot thick with a very short, incurved extension.
Other specimens examined. Peru. AMAZONAS: near
Tarapoto, collected by M. Arias, flowered in cultivation
by W. Rhodehamel at Hoosier Orchid Co., Indianapolis,
Indiana, W. Rhodehamel s.n. (MO).
Htymology. Named for the Philistine giant Go¬
liath.
This large, robust species occurs locally in cloud
forests of southeastern Ecuador and northeastern
Peru. It is closely allied to the Peruvian Masdevallia
ayabacana from the mountains of northwestern
Peru, but M. goliath is distinguished by the flowers
with longer sepaline tails. The apices of the lateral
sepals are approximate so that the bases of the tails
touch in the natural position. The angle between the
apices of the lateral sepals of M. ayabacana is
obtuse.
The cartilaginous petals are minutely exasperate
on the margins and nearby. The petals of M. ay¬
abacana are smooth. The margins of the verrucose
apical third of the lip are acutely revolute resting
against a prominent tuberosity beneath the apex.
Masdevallia mascarata Luer & Vasquez, sp. nov.
TYPE: Bolivia. Santa Cruz: epiphytic in moist
forest south of Yapacani, alt. 650 m, 7 Sep.
1991, C. Luer. J. Luer, L. Moreno it- D. Ric
15402 (holotype, MO; isotypes, AMES, LPB).
Figure 2.
Species haec Masdevalliae auropurpureae Reichb. f.
affinis, sed pedunculo ascendenti vel horizontali, synsepalo
ovato et labello lateriore differt, etiarn M. zahlbruckneri
Krzl. persimilis, sed pedunculo triquetro, petalis oblongis
et labello plicato differt.
Plant small to medium in size, epiphytic, caes¬
pitose; roots slender. Ramicauls slender, erect, 0.5
1.5 mm long, enclosed by 2-3 thin, tubular sheaths.
Novon 2: 6-11. 1992.
Volume 2, Number 1
1992
Luer
South American Masdevallia
7
8
Novon
Figure 2.
Masdevallia mascarata Luer & Vasquez.
Volume 2, Number 1
1992
Luer
South American Masdevallia
9
Leaf erect, coriaceous, subpetiolate, 3-7.5 cm long
including the petiole 0.5-1.5 cm long, the blade
narrowly elliptical, subacute, 0.8-1.5 cm wide, the
base narrowly cuneate into the petiole. Inflorescence
a solitary flower, produced successively in a loose,
few-flowered raceme, borne by a suberect, ascending
to horizontal, triquetrous peduncle, up to 5 cm long
including the peduncle 1.5-3 cm long, with a bract
at the base, from the base of a ramicaul; floral bract
5-7 mm long; pedicel 7-13 mm long; ovary 2-3.5
mm long; sepals yellow, microscopically sparsely
pubescent within, the dorsal sepal obovate, 7-8 mm
long, 4-6 mm wide, connate to the lateral sepals
for 4.5-6 mm to form a short, sepaline tube, the
apex broadly obtuse, contracted into an erect, stout,
yellow tail 12-23 mm long, the lateral sepals dif¬
fusely dotted with red-purple, coalescing toward the
base, suffused with red-purple externally, 12-20
mm long, connate 10-17 mm into an ovate, bifid
lamina 10-16 mm wide, the apices acute, contract¬
ed into subapproximate, slender tails 6-9 mm long;
petals yellow, oblong, 5-6 mm long, 1.5 mm wide,
the apex subtruncate, obscurely bilobulate, the la-
bellar half longitudinally thickened ending in a low,
rounded callus above the base; lip yellow, dotted
with purple, oblong-subpandurate, 4.5 5 mm long,
1.5-2.25 mm wide, with oblique marginal folds above
the middle, the apical portion broadly ovate, obtuse,
minutely subverrucose and erose, the lower portion
oblong, the disc shallowly channeled, the base trun¬
cate, thick and deep, presenting a flat, subquadrate
surface on the end, hinged beneath; column yellow,
stout, semiterete, 4-5 mm long, with a foot 2-3
mm long including a short extension.
Other specimens examined. Bolivia, near Santa cruz
WITHOUT specific locality: collected by L. Moreno, cul¬
tivated by Springdale Orchids, Cincinnati, Ohio, flowered
in cultivation by Hoosier Orchids, Indianapolis, Indiana,
July 1991, C. Luer 15506 (MO).
Etymology. From the Greek maskaratos,
“masked,” in allusion to the confusing identity.
This species occurs in the warm, moist lowlands
of central Bolivia. Although it appears deceptively
similar to Masdevallia zahlbruckneri, known to
occur farther west in central Bolivia, it is more
closely related to M. auropurpurea, previously
known only from the lower altitudes in the Eastern
Cordillera of Colombia. However, M. auropurpurea
has recently been discovered in the Alto Beni prov¬
ince of La Paz, Bolivia. The peduncle of M. mas-
carata is triquetrous (triangular in cross section),
placing it in section Polyanthes. The oblong petals
with a low, basal callus, and the plicated lip confirm
placement in this section.
Masdevallia mascarata is distinguished by a loose,
successively few-flowered raceme, similar to that of
M. zahlbruckneri, but more erect with the peduncle
triquetrous as in M. auropurpurea. The peduncle
of M. zahlbruckneri is round in cross section. Su¬
perficially, the yellow, purple-dotted flower of M.
mascarata looks exactly like that of M. zahlbruck¬
neri, but when bleached colorless, it is similar to the
flowers of M. auropurpurea or M. bicolor and their
relatives.
The lip of M. zahlbruckneri is oblong with a pair
of low, longitudinal carinae, and a subcordate base.
The lip of M. mascarata is plicate above the middle,
and the base is thick and more or less flat on the
end, a curious feature also found in other Bolivian
species (e.g., M. ricii), but not necessarily in the
same section.
Masdevallia ricii Luer & Vasquez, sp. nov. TYPE:
Bolivia. Santa Cruz: epiphytic in moist forest
above Mairana, La Yunga, alt. 2,090 m, 8
Sep. 1991, C. Luer, J. Luer, H. Vasquez cS:
D. Ric 15416 (holotype, MO; isotype, LPB).
Figure 3.
Inter species subsectionis Saltatrices species haec tubo
sepalorum flavo aureosuffuso arcuato non ventricoso, pe-
talorum dente longo incurvo, labelli arcuati base profunda
differt.
Plant medium in size, epiphytic, caespitose; roots
coarse. Ramicauls blackish, slender, erect, 1.5-3
mm long, enclosed by 2-3 thin, tubular sheaths.
Leaf erect, coriaceous, petiolate, 5-12 cm long in¬
cluding the petiole 1.5-3 cm long, the blade nar¬
rowly elliptical, subacute, 1.2-2.2 cm wide, the base
narrowly cuneate into the blackish petiole. Inflores¬
cence a solitary flower, borne by a slender, erect
peduncle 1.5-4 cm long, with a bract above the
base, from low on the ramicaul; floral bract 7-8
mm long; pedicels 5-10 mm long; ovary 5 mm long;
sepals pale yellow, suffused with orange and with a
few small, red spots toward the base, glabrous ex¬
ternally, pubescent within above the middle, the
trichomes simple or caespitose, the dorsal sepal nar¬
rowly obovate, concave, 14-15 mm long, 5 mm
wide expanded, connate to the lateral sepals for 13-
14 mm to form an arcuate, sepaline tube, the apex
broadly obtuse, abruptly contracted into a forwardly
directed, slender, olive-colored tail 25-27 mm long,
the lateral sepals 15 mm long, connate 10 mm into
a bifid, ± obovate lamina, the free portions 6 mm
wide, obtuse, contracted into tails similar to that of
the dorsal sepal; petals light yellow-orange, oblong-
obovate, 4.5 mm long, 1.5 mm wide, the apex
10
Novon
Figure 3. Masdevallia riccii Luer & Vasquez.
Volume 2, Number 1
1992
Luer
South American Masdevallia
11
rounded, minutely erose, lightly notched, with a
longitudinal Carina along the labellar margin ter¬
minating at the base in a broad, thick, descending
process with the tip incurved; lip yellow-orange,
oblong, arcuate, 4 mm long, 1.75 mm wide, the
sides undulate and decurved, the apex obtuse and
decurved, the disc shallowly channeled, the base
thick and deep, presenting a flat, subquadrate sur¬
face on the end, hinged beneath; column yellow-
orange, stout, semiterete, 3 mm long, with a foot
equally long including a slender extension 1.5 mm
long.
Etymology. Named in honor of Darwin Ric of
Montero, Bolivia, co-discoverer of this species.
This species occurs in cool moist forests of central
Bolivia where it grows with M. heideri Koniger, but
much less abundantly. Unfortunately, these forests
are currently in the process of being cut and burned.
Among the species of subsection Saltatrices, M.
ricii is distinguished by the light yellow, long, ar¬
cuate, nonventricose, sepaline tube, suffused with
orange and lightly dotted with red toward the base.
The free portions of the sepals are rounded and
contracted into slender, forwardly pointing tails. The
petals are distinguished by a long, thick, basal pro¬
cess. The lip is arcuate and thick at the base with
a broad, flat surface on the end.
Dos Nuevas Especies de Nototriche (Malvaceae) del Peru
Magda Charico
Herbario San Marcos, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos,
Apartado 14-0434 Lima, Peru
Resumen. Se describen y se ilustran dos nuevas
especies del genero Nototriche: N. antoniana y N.
peruviana. Caracteres morlologicos, incluyendo una
estuctura a manera de “corona,” que las diferencian
de las demas especies del genero se presentan en
una tabla.
ABSTRACT. Two new species of the genus Noto¬
triche are described and illustrated: N. antoniana
and N. peruviana. Morphological characters, in¬
cluding a “corona”-like structure, which differen¬
tiate them from the other species in the genus, are
presented in a table.
Durante la revision del material peruano del ge¬
nero Nototriche Turcz. depositado en el Herbario
San Marcos (l)SM) observe un ejemplar proveniente
del distrito La Union, departamento de Huanuco,
cuyas llores presentaban en la base del tubo estam-
inal una estructura a manera de “corona.” Este
caracter habia sido observado hasta entonces en una
sola especie peruana proveniente del departamento
de Puno, N. cupuliforme Krapovickas (1953).
En un viaje que realice a la localidad de Huayllay,
departamento de Pasco, colecte otra especie de No¬
totriche con estructura similar en la base del tubo
estaminal. A1 ser cuidadosamente estudiados los
ejemplares indicados, pude apreciar que tenian ca-
racteristicas unicas, que los diferenciaban de las de-
mas especies conocidas del genero.
Por lo tanto, las dos nuevas especies se describen
aqui a continuacion.
Nototriche antoniana sp. nov. TIPO: Peru.
Huanuco: Provincia Dos de Mayo, La Union,
piso subnival, sep. 1956, A. Cardich s.n. (hol-
otipo, USM 42520). Figuras 1-3.
Herba perennis, acaulis, basi rosulata, caudex subter-
raneus crassus, lignosus, ramificatusque, folia plurima arete
aggregata. Stipulae petiolo adnatae et cum eo vaginam
5 6 mm longam et 4 mm latam formantes, pars libera
stipularum linearis, apice subacuto, 3-6 mm longo, 1 mm
lato. Lamina flabellata, 5-7 mm longa, 7-11 mm lata,
5-7-lobata, lobis primariis trilobis, supra subtusque in-
cano-tomentosa, interdum subtus glabra (pilis caducis ?).
Petiolus pilis albis stellatis obsitus, stipulae et vagina prae-
ter marginem glabrae. Flos solitarius subsessilis petiolo
infra medium inserto. Caliculus nullus. Calyx campanu-
latus, 10-12 mm longus, dentibus 5 mm longis, 2.5 mm
latis in apicibus subacutis, extus tanquam lamina pilis
obsitus, intus glaber, praeter apices. Nectaria interiora 5,
libera 1 mm longa; 1.5 mm lata. Corolla glabra. Petala
10 12 mm longa et 6 8 mm lata, apice retusa, basi in
tubum coalita, tubus supra patelliformis. Antherae in cap-
ite globoso congestae. Stigmata 10, capitata. Carpella 10,
monosperma, pilis longis stellatis valde obtecta. Fructus
non vidi.
Hierba perenne, acaule, arrosetada, de 11-25
cm de longitud; eje subterraneo lenoso, ramificado,
en cuyos apices de las ramas se agrupan numerosas
hojas formando rosetas de 3 cm de diametro. Es-
tipulas y peciolo forman una vagina de 5-6 mm de
longitud y 4 mm de ancho en la base, disminuyendo
ligeramente hacia el apice. Parte libre de las esti-
pulas linear, de apice subagudo, 3-6 mm de lon¬
gitud, 1 mm de ancho. Lamina flabeliforme, 5-7-
lobada, segmentos trilobados; 5-7 mm de longitud,
7-11 mm de ancho. Epifilo de la lamina incano-
tomentoso, hipofilo igualmente piloso, en algunas
hojas es glabro (pelos caducos ?). Superficies del
peciolo pilosas, pelos estrellados blanquecinos; su¬
perficies de las estipulas y vagina glabras menos los
hordes. Flor solitaria subsesil, ubicada cerca al punto
de separacion de las partes fibres de las estipulas.
Caliculo nulo. Caliz campanulado, 10-12 mm de
longitud; dientes 5 mm de longitud, 2.5 mm de ancho
en la base, apice subagudo; exteriormente todo pi¬
loso, interior glabro menos los apices de los dientes.
Nectarios internos basales 5, aislados, mas anchos
que largos (escutiformes), cerca de 1.5 mm de an¬
cho, 1 mm de longitud. Corola azulada (blanquecina
en el material herborizado). Petalo, 10 12 mm de
longitud, 6-8 mm de ancho, con una pequena es-
cotadura apical, glabro; cada petalo se continua
sobre el tubo de la corola formando dos pequenas
“alas”; tubo de la corola de 3 mm de longitud. En
el punto de insercion de los petalos sobre el tubo de
la corola se desarrolla una estructura pateliforme
de contorno onduiado, carente de venacion, a ma¬
nera de “corona,” que contiene al tubo estaminal,
de 1.5 mm de alto y 4 mm de diametro. Anteras
dispuestas en masa globosa en el apice del tubo
estaminal; este glabro, mas corto que los petalos, de
5 mm de longitud. Estigmas capitados 10; carpelos
Novon 2: 12-15. 1992.
Volume 2, Number 1
1992
Chanco
Nototriche del Peru
13
Figuras 1-3. Nototriche antoniana Chanco. —1. Hoja. —2. Cara interior del caliz, mostrando un nectario. —3.
Corola con 4 petalos seccionados, mostrando la “corona” en la base del tubo estaminal. Dibujado del holotipo.
10 uniovulados, pilosos desde su base, con pelos
estrellados largos. Fruto maduro desconocido.
Material adicional estudiado. PERU. HUANUCO: Prov-
incia Dos de Mayo, cumbre de Pachacoto-Wansala, 24
feb. 1980, M. Weibel 6 (USM); Pachacoto-La Union,
19 Mar. 1983, O. Tovar & S. Rivas 9779, 9786, 9788
(USM); Valle de Huallanca, O. Tovar & S. Rivas 9913
(USM).
Distribucion geogrdfica. Esta especie ha sido
coleccionada hasta la fecha en el departamento de
Huanuco, entre los 4,200 y 4,780 m.
La autora ha dedicado esta especie al eminente
cientifico Antonio Krapovickas, en reconocimiento
a su constante estudio de las Malvaceas, aportando
para el Peru numerosas publicaciones de generos y
especies nuevos.
Tabla 1. Caracteres que distinguen las tres especies de Nototriche.
N. cupuliforme
N. antoniana
N. peruviana
Lamina
Pinatifida
Flabeliforme
Flabeliforme
Indumento de las hojas
Glabro
Piloso
Piloso
Caliz (longitud)
4-5 mm, glabro
10-12 mm, piloso
7-10 mm, piloso
Nectarios
Semicirculares, 0.5 mm
ancho
Escutiformes, 1.5 mm
ancho
Triangulares, 1 mm ancho
Corola
Rojo-vinosa
Azulada
Rojo-vinosa
“Corona” (diametro)
Cupuliforme, 2.5 mm
Pateliforme, 4 mm
Pateliforme, 3 mm
Tubo de la corola (longi-
1.5 mm
3 mm
1-1.5 mm
tud)
Tubo estaminal (longi-
2-2.5 mm
5 mm
1.5 mm
tud)
Estigmas
6 capitados
10 capitados
8 capitados
Distribucion
Puno: Carabaya
Huanuco: La Union
Pasco: Huayllay
14
Novon
Figuras 4-6. Nototriche peruviana Chanco. —4. Hoja. —5. Cara interior del caliz, mostrando tres nectarios.
6. Corola con 4 petalos seccionados, mostrando la “corona” en la base del tubo estaminal. Dibujado del holotipo.
Las especies mas proximas a TV. antoniana son
N. cupuliforme Krapov. y N. peruviana. (Ver Tabla
1 .)
Nototriche peruviana sp. nov. FIFO: Peru. Pas¬
co: Provincia de Pasco, Huayllay piso subnival,
10 mayo 1976, M. Chanco 368 (holotipo, USM;
isotipo, CTES ). Figuras 4-6.
Herba perennis, rosulata, caudex subterraneus, ligno-
sus. Stipulae petiolo adnatae et cum eo vaginam 5-7 mm
longam et 2 mm latam formantes, pars libera stipularum
linearis, apice obtuso, 3.5-5 mm longo, 0.5 mm lato.
Lamina flabelliforme 7-lobata. Flos solitarius subsessilis
petiolo infra medium insertus. Caliculus nullus. Calyx cam-
panulatus, 7-10 mm longus, dentibus 4-5 mm longis, 2-
3 mm latis in apicibus obtusis, extus tamquam lamina pilis
obsitus, intus praeter apices glaber. Nectaria interiora 5,
libera 2 mm longa, 1 mm lata. Corolla glabra, vinosa.
Petala obovata, 8 mm longa et 6 mm lata, apice retusa,
basi in tubum coalita, tubus supra patelliforme. Antherae
in capito globoso aggregatae. Stigmata 8, capitata. Fruc-
tus maturus non visus.
llierba perenne, arrosetada; eje subterraneo len-
oso, subcilindrico, 15-30 cm de longitud y cerca
de 1 cm de ancho en la parte superior, decrece
notablemente en diametro hacia el extremo inferior,
ramificado apicalmente, con numerosas hojas dis-
puestas a manera de rosetas de 1.5-3 cm de dia¬
metro. Estipulas y peciolo soldados formando una
vagina de 5-7 mm de longitud, 2 mm de ancho,
parte libre de las estipulas linear, de apice obtuso,
con nervio medio conspicuo, 3.5-5 mm de longitud,
O. 5 mm de ancho. Lamina flabeliforme 7-lohada,
segmentos 2-3-lobados, apice de los lobulos semi-
circulares; 4-5 mm de longitud, 7-8 mm de ancho,
superficie inferior de toda la hoja densamente pilosa.
Superficie superior del peciolo y parte basal de la
vagina con pelos estrellados, el resto glabro; super¬
ficie superior de las estipulas glabra menos el apice
que es piloso. Flor solitaria subsesil, ubicada por
debajo del punto donde se separan las partes fibres
de las estipulas. Caliculo nulo. Caliz campanulado,
acrescente, 7-10 mm de longitud, cubierto exter-
iormente por pelos estrellados similares a los de la
hoja, 5 dentado, los dientes 4-5 mm de longitud,
2-3 mm de ancho, obtusos, el interior glabro menos
los apices y hordes de los dientes. Nectarios internos
aislados, triangulares, de apice obtuso, 2 mm de
longitud, 1 mm de ancho. Corola rojo-vinosa; tubo
de la corola de 1.5 mm de longitud; petalos obovados
con apice ligeramente escotado, glabros, 8 mm de
longitud, 6 mm de ancho, cada petalo se continua
sobre el tubo de la corola formando dos pequenas
“alas”; sobre el punto donde se unen los petalos al
tubo de la corola, se desarrolla una estructura car-
nosa 5 lobada a manera de “corona” (pateliforme),
que contiene al tubo estaminal, de 3 mm diametro
y 1 mm de alto. Tubo estaminal de 1.5 mm de
longitud. Anteras reunidas en una masa globosa.
Estigmas capitados en nurnero de 8. Ovario conico,
piloso, pelos largos desde la base. Fruto maduro no
visto.
Material adicional estudiado. Pf.rCi. PASCO: Huayllay,
pajonal del Bosque de Piedra, N. Urquizo s.n. (USM).
Distribucion geogrdfica. Solo se conoce esta es-
pecie de la localidad tipo.
Volume 2, Number 1
1992
Chanco
Nototriche del Peru
15
Por la presencia de una estructura a manera de
“corona” en la base de insercion de los petalos al
tubo estaminal, esta especie tiene afinidad con N.
cupuliforme y /V. antoniana. Las caracterlsticas de
diferenciacion de las 3 especies se resumen en la
Tabla 1.
Agradecimiento. Parte de este estudio se realizo
durante mi estadia en el Departamento de Botanica
y Ecologia de la Universidad Nacional del Nordeste,
Corrientes, Argentina. Expreso mi sincera gratitud
a Antonio Krapovickas, Director del Instituto de
Botanica, por el asesoramiento y las facilidades brin-
dadas en el estudio de la familia Malvaceae duranto
mi estadia en Corrientes. A Jaroslav Soukup, mi
agradecimiento postumo por la diagnosis en latin de
las nuevas especies. Al personal directivo de la Com-
paiiia Minera de Huaron, por facilitarme la estadia
durante mi trabajo de campo en Huayllay.
Literature Cited
Krapovickas, A. 1953. Notas citotaxonomicas sobre
Nototriche (Malvaceae) II. Bol. Soc. Argent. Bot.
5(1-2): 51-74.
Two New Species of Jungia (Compositae) from Peru
Gunnar Ha rling
Department of Systematic Botany, University of Goteborg, Carl Skottsbergs Gata 22,
S-413 19 Goteborg, Sweden
ABSTRACT. Two new species of Jungia, J. schuer-
ae and ,/. gracilis, from the Peruvian Andes are
described and illustrated, and their affinities are brief¬
ly discussed.
The genus Jungia L. f. (Compositae: Mutisieae:
Nassauviinae) comprises 25 30 species, ranging from
southern Mexico through Central America and fol¬
lowing the South American Andes to northern Ar¬
gentina; a couple of species are known from south¬
eastern Brazil and Paraguay. In connection with my
work on a monograph of the genus, I have found
the following two species to be new to science.
Jungia schuerae Barling, sp. nov. TYPE: Peru.
Lima: Prov. Huarochiri, Matucana, dry slope,
2,400 m, 26 May 1040, Asplund 11101 (ho-
lotype, S). Figure 1A.
Frutex a basi ramosus, erectus vel reclinatus, 1-1. 5(-
2.5) m altus, ramis sparse vel dense pubescentibus. Folia
exstipulata, petiolata, laminis chartaceis, ambitu cordato-
suborbicularibus, 3.5-5 cm longis, 3.5-6 cm latis in foliis
superioribus, usque ad 10 cm longis, 11 cm latis in foliis
inferioribus, 5-7-lobatis, lobis acutis vel obtusis vel ro-
tundatis, indistincte serratis vel dentatis, supra sparse vel
satis dense strigosis, subtus dense villosis; petiolus l-3(-
4.2) cm longus, dense pubescens. Synflorescentia dense
paniculata, polycephala. Involucrum campanulatum, 6
7 mm longum, 7-8 mm latum, phyllariis circiter 12,
oblongis vel lanceolatis, acuminatis, 5-6 mm longis, 1-
1.2 mm latis, basi plusminusve callosis, dorsaliter tomen-
tosis. Flores circiter 17-25, corolla sulphurea, tubo 4-5
mm longo, labio exteriore tridentato, in floribus margin-
alibus liguliforini, 3-3.5 mm longo, in floribus centralibus
revoluto, 2.5-2.8 mm longo, labio interiore profunde bi¬
partite, 1.6-2.3 mm longo, segmentis convolutis. Ach-
enium subcylindricum vel fusiforme, erostratum, circiter
4 mm longum, sparse strigosum, pappi setis niveis, 30
35, circiter 6 mm longis, ciliatis vel breviter plumosis.
Erect or slightly reclinate shrub 1 — 1 -5(—2.5) rn
high, branched from the base. Branches terete to
slightly angular, light brown, sparsely to densely
white-pubescent, in older stages glabrescent, often
with dark brown to black lenticels, eglandular like
the rest of the plant. Leaves exstipulate, petiolate;
lamina chartaceous, suborbicular-cordate in outline,
3.5-5 cm long and 3.5-6 cm wide in upper leaves,
to 10 cm long and 1 1 cm wide in lower leaves, 5-
7-lobed, lobes acute to obtuse or rounded, indis¬
tinctly serrate or dentate with mucronate, dark teeth,
rarely slightly lobulate, sparsely to rather densely
strigose above, moderately to very densely white-
villous beneath, rarely subglabrous on both sides,
veinlet reticulum beneath dark-colored, usually with
reddish brown thickenings; petiole 1 —3(—4.2) cm
long, slightly sheathing at base, densely white-pu¬
bescent. Synflorescence usually a very dense, oc¬
casionally ± open panicle; bracts similar to ordinary
leaves but gradually smaller, seldom rounded and
unlobed; peduncles 0.2—1,3(—3) cm long, white-pu¬
bescent, with 0-2 small, linear bracteoles. Involucre
campanulate, 6-7 mm long, 7-8 mm wide; acces¬
sory bracteoles usually 3-6, 2-3 mm long, ca. 0.5
mm wide, linear to narrowly oblanceolate, white-
pubescent; phyllaries (10—) 12(— 13), 5-6 mm long,
1-1.2 mm wide, oblong to lanceolate, acuminate,
± callose at base, moderately to densely white-
tomentose dorsally, green, occasionally with purple
tips; paleae similar to phyllaries but thinner and
pubescent only apically and along midnerve. Florets
17-25, corolla pale yellow, often fading to white,
tube 4-5 mm long, gradually widening toward mouth,
outer lip tridentate, apically slightly pilose, in mar¬
ginal florets liguliform, 3-3.5 mm long, straight to
slightly revolute, in central florets 2.5-2.8 mm long,
revolute, inner lip deeply bifid, 1.6-2.3 mm long,
the segments curled. Achenes subcylindric to fusi¬
form, without beak, ca. 4 mm long, 5-ribbed, dark
between the ribs, with short, hyaline hairs; pappus
white, bristles ca. 30-35, ca. 6 mm long, ciliate to
short-plumose.
Jungia schuerae is a rather uniform and easily
recognized species, distantly related to J. paniculata
(DC.) Gray. It seems to be locally common in Ca-
jamarca. La Libertad, Ancash, lluanuco, and Lima.
The habitat is, according to the collectors, dry, grav¬
elly, or stony open slopes or dry thickets at 2,200-
3,600(-4,000) m elevation. Currently about 50
collections, most of them from Ancash and Lima,
are known to me. A complete list will be published
in my forthcoming monograph of the genus.
Cerrate (1951), who treated the Peruvian species
ol Jungia, confused this species with J. paniculata.
Novon 2: 16-18. 1992.
Volume 2, Number 1
1992
Harling
Jungia from Peru
17
Holotypes of Jungia schuerae Harling (A) and Jungia gracilis Harling (B).
18
Novon
The latter species is, however, an entirely different
plant with stipulate leaves, open synflorescences,
usually glandular peduncles and phyllaries, white
florets, and a distinctly plumose pappus. The true
J. paniculata is common in Ecuador and Colombia,
but seems to be rare in Peru. Apart from the type,
I have so far seen only three collections, all from
Cajamarca.
It is with great pleasure that I name this species
in honor of Mrs. Gudrun Nilsson, nee Schuer, who
during her careful, but unfortunately unpublished,
studies of the genus pointed it out as new (Schuer,
in sched.).
Jungia gracilis Harling, sp. nov. TYPE: Peru.
Amazonas: Prov. Chaehapoyas, Hio Utcubam-
ba Valley, 31 km along road S of Tingo, ca.
1,700 m, 21 Jan. 1983, King & Bishop 9278
(holotype, US; isotypes, G, K. MO). Figure IB.
Suffrutex sparse ramosus, erectus vel reclinatus, raro
subscandens, 1-1.5 in altus, rainis lends, glabris. Folia
exstipulata, petiolata, laminis membranaceis, ambitu cor-
dato-suborbicularibus, 4-9.5 cm longis, 4.5-10 cm latis,
5 7-lobatis, lobis acutis, indistincte serratis vel dentatis,
supra glabris, subtus sparse vel mediocriter villosis; pe-
tiolus 1.5 5(—7) cm longus, glaber vel sparse puberulus.
Synflorescentia laxissiine paniculata, polycephala, ambitu
ad 25 x 30 cm. Involucrum turbinaturn, 4-5 mm lon-
gum, 4-5.5 mm latum, phyllariis circiter 12, oblongis vel
oblanceolatis, acuminatis, 4.5-5 mm longis, 1-1.2 mm
latis, basi distincte callosis, dorsaliter glabris vel sparse
puberulis apice penicillatis. Flores 16-20(-23), homo-
morphi, corolla nivea, tubo 2.6-3 mm longo, labio ex-
teriore tridentato, 1-1.3 mm longo, revoluto, labio inter-
iore profunde bipartito, 0.5-0.8 mm longo, segmentis
convolutis. Achenium subcylindricum vel ampullaceum,
erostratum, 3.5-4 mm longum, sparse strigosum, pappi
setis niveis, circiter 30, 4.5-5 mm longis, ciliatis.
Erect or reclinate, seldom clambering subshrub
1 1.5 m high. Stem and branches terete, slightly
fluted, rather weak, reddish brown, glabrous, eglan-
dular like the rest of the plant. Leaves exstipulate,
petiolate; lamina membranaceous, suborbicular-eor-
date in outline, 4-9.5 cm long, 4.5-10 cm wide,
5-7-lobed, lobes acute, indistinctly serrate or den¬
tate with mucronate, dark teeth, glabrous to sub-
glabrous above, sparsely to moderately villous be¬
neath, veinlet reticulum beneath dark-colored with
partial thickenings; petiole 1.5—5( 7) cm long,
slightly sheathing at base, glabrous to minutely pu-
berulous. Synflorescence an open, many-headed
(heads up to 200 or more) panicle, up to 25 x 30
cm in extent; branches numerous, thin, slightly pu¬
bescent; lower bracts similar to ordinary leaves but
gradually smaller, upper bracts ovate to elliptic, 1-
3 cm long, 0.6-1.4 cm wide, pubescent, margin
entire or slightly serrate; peduncles (l-)2-3.6 cm
long, slightly pubescent. Involucre turbinate, 4-5
mm long, 4-5.5 mm wide, accessory bracteoles 2-
3(-4), narrrowly linear, 2-3 mm long, ciliate at
margin; phyllaries ca. 12, 4.5-5 mm long, 1-1.2
mm wide, oblong to oblanceolate, acuminate, dis¬
tinctly callose at base, glabrous to sparsely pubescent
dorsally, pilose apically, green with reddish nerves;
paleae similar to phyllaries but thinner, glabrous
except pilose apex. Florets 16 20( 23), homomor-
phous, corolla white, tube 2.6-3 mm long, gradually
widening toward mouth, outer lip tridentate, 1-1.3
mm long, revolute, inner lip deeply bifid, 0.5-0.8
mm long, the segments curled, both lips slightly
pilose apically. Achenes subcylindric to slightly am-
pullaceous but without distinct beak, 3.5-4 mm
long, 5-ribbed, light brown, with short hyaline hairs;
pappus white, bristles ca. 30, 4.5-5 mm long, ciliate.
Jungia gracilis is endemic to the Ghachapoyas
area in Amazonas. It has been collected in rather
dry and open vegetation at 1,600-2,000 m ele¬
vation. In all, six collections are known at present.
The closest relative of J. gracilis is probably J.
spectabilis I). Don, known from Ecuador and north¬
ern Peru. Both species are exstipulate, eglandular
subshrubs; they also have in common the leaf shape,
the scarce indumentum, and the conspicuously cal-
lose-based phyllaries. But J. gracilis is smaller and
more slender in all parts, has an extremely widely
spaced synflorescence (vs. dense and partially clus¬
tered), and the veinlet reticulum beneath is entirely
dark-colored (vs. light-colored with black dots).
Acknowledgments. I thank Uno Eliasson for his
critical reading of the manuscript and for photo¬
graphic assistance.
Literature Cited
Cerrate, E. 1951. Revision de las especies peruanas
del genero Jungia. Publ. Mus. Hist. Nat. “Javier
Prado,” Ser. B. Bot., No. 4: 1-24.
Compositae of the Guayana Highlands—VI.
Huberopappus maigualidae (Vernonieae), a New
Genus and Species from Venezuela
John F. Pruski
Institute of Systematic Botany, New York Botanical Garden,
Bronx, New York 10458-5126, U.S.A.
Abstract. Huberopappus maigualidae, a new
genus and species of Compositae tribe Vernonieae
from Sierra de Maigualida, Venezuela, is described
and illustrated. Huberopappus is the only genus of
the tribe with a coroniform pappus and dimorphic
achenes. By its pappus Huberopappus is similar to
Struchium and especially to Ekmania and Gor-
ceixia, the latter two being taken as its closest rel¬
atives. Huberopappus, Ekmania, and Gorceixia
seem to form a generic group and are placed near
Pollalesta in the Vernonieae subtribe Piptocarphin-
ae.
Among collections of Compositae submitted to me
for determination by Otto Huber and Paul Berry
from their recent expeditions to the previously unex¬
plored, granitic Sierra de Maigualida in Guayanan
Venezuela were specimens that could not be assigned
to any genus of the family. These plants are referable
to the cichorioid tribe Vernonieae, in which only
three described neotropical genera have a strongly
coroniform pappus. The plants from Sierra de Maig¬
ualida, however, are the only Vernonieae with both
a crownlike pappus and dimorphic achenes. They
are thus described as a new genus and species in
conjunction with my treatment of the family for the
Flora of the Venezuelan Guayana by Julian Stey-
ermark and collaborators.
Huberopappus maigualidae Pruski, gen. et sp.
nov. TYPE: Venezuela. Bolivar: Depto. Ced-
eho. Sierra de Maigualida, sector nor-oriental,
altiplanicie tepuyana disectada sobre granito en
las cabeceras del Rio Chajura, afluente occi¬
dental del Rio Erebato, aprox. 100 km (en linea
recta) al SW del Campamento Entrerios,
05°33'N, 65°13'W, 2,100 m, 18 Nov. 1988,
Otto Huber & Liz Izquierdo 12791 (holotype,
VEN; isotypes, MY, NY, US). Figures 1, 2.
Frutex 1-2 m altus pauciramosus; caules subteretes
vel angulati pubescentes vel dense pubescentes nigri; folia
simplicia alterna petiolata, lamina subcoriacea vel coriacea
elliptica vel obovata (2-)3-6 x (1.3—)1.7—3.6 cm cu-
neata apice acuta vel rotundata integra, pinnatim venosa
supra glandulosa punctata subtus trichomatibus subap-
pressis tomentella, petiolo 4-7 mm longo; capitulescentia
terminalis cymosa; pedunculi 0-7 mm longi; capitula
homogaina epaleacea, flosculis 14-22; involucrum cam-
panulatum 5-7 x 3-4 mm 3-4-seriatum; phyllaria di-
morpha imbricata, quinque externa plerumque coriacea
nigra deltoidea vel elliptica dense pubescentia; corollae
discoideae quinquelobatae 4.5-5 mm longae glandulosae,
tubo 2.6-3.1 mm longo, lobis adscendentibus linearibus
ca. 1.8 mm longis; aehaenia turbinato-prismatica 2.6-
3.2 mm longa glabra, pappo in corpus coroniforme ser-
rulatum 1.7-2.1 mm longum confluenti vel arista singula
4 mm longa usque praedito.
Subshrub 1-2 m tall; stems erect, few-branched,
subterete below, irregularly angled distally, pubes¬
cent to densely so, this pubescence imparting a
blackish color to the stems; internodes 0.5 2.5 cm
long. Leaves simple, alternate, petiolate; blade sub-
coriaceous to coriaceous, elliptic to obovate, (2-)3-
6 cm long, (1.3—) 1.7-3.6 cm wide, basally cuneate,
apically acute to rounded, entire, venation pinnate,
reticulate, upper surface dark green, glandular-
punctate, otherwise glabrous, secondary veins slight¬
ly raised, lower surface sometimes glandular, to-
mentose with a dense mat of appressed trichomes,
veins darker than areoles, the lower surface thereby
reticulate; petiole 4-7 mm long, densely pubescent.
Capitulescence cymose, sometimes partially ob¬
scured by juvenile leaves, of ca. 3-11 closely spaced
capitula, these terminal on axillary branches from
the uppermost nodes or from stem apex; peduncles
densely pubescent, the lateral ones 2-7 mm long,
the central ones often lacking; capitula homoga-
mous, discoid, 14-22-flowered, often subtended by
reduced, densely pubescent, black foliar bracts; in¬
volucre campanulate, 5-7 mm high, 3-4 mm broad,
3- or 4-seriate; phyllaries 14-17, imbricate; outer
series deltoid to broadly elliptic, to ca. 7 x 3 mm,
occasionally slightly longer than the next series,
coriaceous, densely pubescent, blackish, apically
acute to rounded, margins entire; innermost series
elliptic to lanceolate, 6-7 mm long, 1.5-2 mm wide,
proximally glabrous, stramineous, stiffly charta-
Novon 2; 19-25. 1992.
20
Novon
Figure 1. Huberopappus maigualidae Pruski. —a. Habit. —b. Achenes with serrulate coroniform pappus (left)
and entire coroniform pappus (right); 5-lobed glandular corolla (right). —c. One-awned achenes. —d. Style. The
slightly enlarged basal node of style not shown. —e. Anther showing spurred base. (Drawn from the holotype, Huber
& Izquierdo 12791, VEN.)
Volume 2, Number 1
1992
Pruski
Huberopappus maigualidae
21
Figure 2. Huberopappus maigualidae Pruski showing the closely spaced, ovate, subcoriaceous to coriaceous leaves
(Huber & lzquierdo 12791, the type collection). Photo taken by Otto Huber.
ceous, distally black, densely pubescent, apex round¬
ed, margins entire; receptacle flat, ca. 2 mm diam.,
naked, foveolate, inner walls occasionally elongate
or pointed. Corollas actinomorphic, homogamous,
5-lobed, 4.5-5 mm long, abaxially glandular, es¬
pecially toward apex of lobes, violet; tube (not dif¬
ferentiated from throat) 2.6-3.1 mm long; lobes
ascending, ca. 1.8 mm long, linear; anthers not
exserted, cream-colored, ca. 2.1 mm long, glabrous,
apical appendage lanceolate, rounded apically, base
calcarate, spurs fertile, 0.4-0.5 mm long; style to
5.5 mm long, hispidulous in the upper half, the 2
branches ascending to somewhat recurved, 1-1.5
mm long, apex rounded to attenuate, basal node
only slightly expanded. Achenes dimorphic, turbi¬
nate, prismatic, 3-5-angled, 2.6-3.2 mm long, light
brown, glabrous, ca. 10-striate; carpopodium slight¬
ly stipitate, neither sculptured nor well differenti¬
ated; pappus thinly coroniform, entire or serrulate
apically, stramineous, ca. 1.7-2.1 mm long, but in
some achenes often weakly lacerate or with an elon¬
gate outer awn to 4 mm long, this protruding from
an abaxial strengthened angle of achene. Pollen 36-
40 gm diam. (excluding spines), tricolporate, mi-
croperforate, subechinolophate (pollen type A sensu
Keeley & Jones, 1979).
Table 1. Neotropical Vernonieae with a coroniform or annular pappus.
22
Novon
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texture coriaceous coriaceous scarious scarious scarious marginally
membra-
Table 1. Continued.
Volume 2, Number 1
1992
Pruski
Huberopappus maigualidae
23
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24
Novon
Paratypes. VENEZUELA. TERRITORIO FEDERAL AMAZONAS:
Depto. Atures, Sierra de Maigualida, NW sector, small
valley along an upper tributary of Cano Iguana, 05°30'N,
65°15'W, 2,000 m, 28 Feb.-3 Mar. 1991, Paul Berry,
Otto Huber & Judith Rosales 4812 (MO, MYF not seen,
NY, VEN not seen). BOLlVAR: Depto. Cedeno, Sierra de
Maigualida, sector nor-oriental, altiplanicie tepuyana ubi-
cada en las cabeceras del Rio Chajura, afluente occidental
del Rio Erebato, aprox. 100 km (en linea recta) al SW
del Campainento Entrerios, 05°33'N, 65°13'W, 2,100
m, 28 Mar. 1988, Otto Huber 12745 (K, FIS, VEN).
Distribution. In savannas from 2,000 to 2,100
m in the northern sector (05°30-33'N, 65°13-
15'W) of Sierra de Maigualida, a granitic mountain
range bordering Territorio Federal Amazonas and
Estado Bolivar, Venezuela. Known to flower in No¬
vember and from February to March.
I take great pleasure in naming this new genus
after my friend and colleague Otto Huber, who first
collected it and recognized it as a new species. The
epithet refers to the Sierra de Maigualida, where
this species appears to be endemic.
The Vernonieae, a tribe of about 1,500 species
and 80 genera, have long been thought to be a fairly
homogeneous group because of the uniformity of
the style, anther, and corolla (see Gleason, 1906,
1922; Jones, 1977; Robinson et al., 1980; Robinson
& Kahn, 1986; Robinson & Funk, 1987; Keeley
& Turner, 1990). However, many vernonian genera
differ from the prototype of the tribe by rare or
unique features. Huberopappus Pruski is yet an¬
other morphologically distinctive genus, the only one
in the Vernonieae with both a coroniform pappus
and dimorphic achenes.
Seven neotropical genera previously were known
to have a connate (at least outer) pappus, which
ranges from deeply lacerate to entire and from
strongly coroniform to annular. These genera differ
greatly from typical Vernonieae, which have a single
or double pappus of free awns, scales, or bristles.
Achene dimorphy in the Vernonieae is rare, pre¬
viously noted (H. Robinson, 1979, pers. comm.)
only in the distantly related and noncoroniform Het-
erocypsela H. Robinson, Lychnophoriopsis Schultz-
Bip- sens, str., Pithecoseris Martius of Brazil, and
Dipterocypsela S. F. Blake from Colombia. All neo¬
tropical genera of Vernonieae with a coroniform
pappus, as well as those with dimorphic achenes
(both ol these conditions are taken as derived), are
unispecific.
Although the presence of a coroniform pappus is
useful taxonomically, it is not necessarily an indi¬
cator ol phylogenetic relationships. Nevertheless,
Ekmania Gleason, native to limestone outcrops that
arise from a serpentine bed in the mountainous
Cuban Oriente (Borhidi & Muniz, 1986; Borhidi,
pers. comm.) and Gorceixia Baker, of the central
Brazilian highlands, seem to be the closest relatives
of Huberopappus. Ekmania and Gorceixia are re¬
lated to Huberopappus by a terminal capitulesc-
ence, a strongly coroniform pappus, entire subcor-
iaceous leaves, primitive type A pollen
(Lychnophora- type), and angled young stems. Hub¬
eropappus and Ekmania seem more closely related
to each other on the basis of leaf shape, leaf pu¬
bescence, and pedicellate heads. Key traits that dis¬
tinguish Huberopappus from Ekmania are dimor¬
phic (not isomorphic) achenes, black (not gray) young
stems, commonly rounded (not narrowly acute to
acuminate) outer phyllary tips, and a single (not
double) pappus without (vs. with) an inner row of a
few deciduous pappus bristles. By its strongly co¬
roniform pappus Struchium P. Br. is similar to the
former three genera, but differs in many other fea¬
tures (see Table 1) and its affinities clearly are else¬
where.
Huberopappus , Ekmania , and Gorceixia form
a tightly knit generic group that is somewhat re¬
moved from both New World and Old World co¬
roniform genera, as well as the remaining genera of
the Vernonieae. These three genera appear to be
closest to Pollalesta HBK of the Piptocarphinae (as
suggested by H. Robinson, pers. comm.), based on
terminal capitula, type A pollen, angled young stems,
leaf pubescence, and calcarate (not caudate) anther
bases. However, Pollalesta (see Stutts, 1981) clear¬
ly differs from the former three genera by a generally
double pappus with many free, rarely paleaceous or
pseudocoroniform outer bristles. Nevertheless, it is
perhaps best to stretch the boundaries of the Pip¬
tocarphinae (Vernoniinae sensu Jones, 1982) to in¬
clude the former three genera.
The salient features that distinguish Huberopap¬
pus from other neotropical Vernonieae genera with
a basally connate or annular pappus are summarized
in Table 1, which includes the initial report of pollen
types (see Keeley & Jones, 1979, for terminology)
in the first five genera. The pollen types of Pa-
courina Aubl., Rolandra Rottb., and Struchium
given in Stix (1960) and Robinson et al. (1980) are
herein confirmed. Some of the information in I able
1 is from Aristeguieta (1964), Badillo (1989), and
Baker (1873).
The last five genera in Table 1 tend to exhibit
the combination of herbaceous to subshrubby habit,
subterete stems, chartaceous often serrulate leaves,
axillary sessile heads, aristate phyllaries, and lophate
pollen and have been variously placed within the
Vernonieae. Bentham (1873) and Baker (1873) re¬
ferred Heterocoma DC., Pacourina, and Struchium
Volume 2, Number 1
1992
Pruski
Huberopappus maigualidae
25
to the Sparganophoreae, whereas Robinson et al.
(1980) placed them in the Vernoniinae. Harleya
Blake, which was not positioned by Robinson et al.
(1980), also seems to be best placed in the Ver¬
noniinae. Rolandra has been placed in both the
Lychnophoreae (Bentham, 1873; Baker, 1873) and
the Rolandrinae (Robinson et al., 1980). Therefore,
by the occurrence of coroniform taxa in each of
these subtribes, it is apparent that pappus fusion, a
derived vernonian condition (Keeley & Furrier,
1990), has arisen independently in several different
lineages.
Acknowledgments. I thank Otto Huber for set¬
ting aside his initial collections for my study during
a visit in 1989 to Caracas and for use of his pho¬
tograph in Figure 2; Bruno Manara for drawing
Figure 1 and for correcting the Latin diagnosis;
Harold Robinson for his determination of the pollen
type of Heterocoma; and Victor Badillo, Rupert
Barneby, Paul Berry, Charles Jeffrey, Sterling Kee¬
ley, and Harold Robinson for many helpful com¬
ments. My herbarium studies in Venezuela have
been financially supported by the Missouri Botanical
Garden, Flora of the Venezuelan Guayana project.
Literature Cited
Aristeguieta, L. 1964. Compositae. Flora de Venezuela
10(1): 1-487.
Badillo, V. M. 1989. Enumeracion de las Vernonieae
de Venezuela. Ernstia 54: 1-54.
Baker, J. G. 1873. Compositae I. Vernoniaceae. Pp.
1-179 in C. F. P. Martius (editor), Flora Brasiliensis
6(2). Munich.
Bentham, G. 1873. Compositae. Pp. 163-533 in G.
Bentham & J. D. Hooker (editors), Genera Plantar-
urn, vol. 2, part 1. L. Reeve, London.
Borhidi, A. & O. Muniz. 1986. The phytogeographic
survey of Cuba II. Floristic relationships and phy¬
togeographic subdivisions. Acta Bot. Hungarica 32:
3-48.
Gleason, H. A. 1906. A revision of North American
Vernonieae. Bull. New York Bot. Gard. 4: 144-243.
-. 1922. Carduaceae Tribe 1. Vernonieae. N.
Amer. Flora 33(1): 47-110.
Jones, S. B., Jr. 1977. Vernonieae — Systematic Re¬
view. Pp. 503-521 in V. H. Heywood, J. B. Har-
borne & B. L. Turner (editors), The Biology and
Chemistry of the Compositae. Academic Press, New
York.
-. 1982. The genera of Vernonieae (Compositae)
in the southeastern United States. J. Arnold Arbor.
63: 489-507.
Keeley, S. C. & S. B. Jones, Jr. 1979. Distribution of
pollen types in Vernonia (Vernonieae: Compositae).
Syst. Bot. 4: 195-202.
-& B. L. Turner. 1990. A preliminary cladistic
analysis of the genus Vernonia (Vernonieae: Aster-
aceae). PI. Syst. Evol. [Suppl. 4]: 45-66.
Robinson, H. 1979. Two new genera of Vernonieae
(Asteraceae) from Brasil, Heterocypsela and Pseu-
dostifftia. Phytologia 44: 442-450.
-. 1988. Studies in the Lepidaploa complex
(Vernonieae: Asteraceae) VI. A new genus, Aynia.
Proc. Biol. Soc. Wash. 101: 959-965.
-& V. A. Funk. 1987. A phylogenetic analysis
of Leiboldia, Lepidonia, and a new genus Stra-
mentopappus (Vernonieae: Asteraceae). Bot. Jahrb.
Syst. 108: 213-228.
- & B. Kahn. 1986. Trinervate leaves, yellow
flowers, tailed anthers, and pollen variation in Dis¬
tep hanus Cassini (Vernonieae: Asteraceae). Proc. Biol.
Soc. Wash. 99: 493-501.
-, F. Bohlmann & R. M. King. 1980. Chemosys-
tematic notes on the Asteraceae. III. Natural sub¬
divisions of the Vernonieae. Phytologia 46: 421-
436.
Stix, E. 1960. Pollenmorphologische Untersuchungen
an Compositen. Grana Palynol. 2(2): 39-114, pis.
10 - 21 .
Stutts, J. G. 1981. Taxonomic revision of Pollalesta
H.B.K. (Compositae: Vernonieae). Rhodora 83: 385-
419.
Mosiera (Myrtaceae) in Mexico and Mesoamerica
Leslie R. Landrum
Department of Botany, Arizona State University, Tempe, Arizona 85283-1601, U.S.A.
ABSTRACT. The history and relationships of the
mainly Caribbean genus Mosiera are discussed, and
it is comprehensively described. Two Mexican and
Mesoamerican species, Psidium contrerasii and P.
ehrenbergii, are transferred to Mosiera.
Ceneric limits among the American Myrtinae
(Myrtaceae) have long been obscure. Historically
the greatest problem has been definition of the limits
of Psidium L., a large, diverse, and poorly defined
group to which many species of Myrtinae (e.g., P.
contrerasii Lundell and P. ehrenbergii (Berg) Bur-
ret) have been assigned for want of a better place.
Recently, Landrum & Sharp (1989) have described
and illustrated seed coat characters that can he
correlated with floral characters to separate Psidium
from related genera, but the above two species are
among a lew that must he relocated. Psidium con¬
trerasii and P. ehrenbergii are here assigned to
Mosiera, a genus that mainly has been ignored since
its description by Small (1933).
History of Mosiera
Small (1933) created Mosiera to contain two
species from southern Florida that he had earlier
(1913) recognized as belonging to Anamomis Gri-
seb. (= Myrcianthes Berg). I agree with McVaugh
(1973) that Small’s original two species are best
considered as one. Small assigned both Anamomis
and Mosiera to the tribe Eugeniinae, where only
the first genus belongs. Small recognized the species
of Mosiera he had before him as being distinct from
Anamomis because of characteristics of the fruit
and inflorescence. Apparently unknown to him was
the striking difference in embryo structure, the char¬
acter of primary importance in differentiating sub¬
tribes of Myrteae.
Burret (1941), in his worldwide revision of Myr-
tus L. and segregate genera, considered Mosiera to
be a synonym of Myrtus. Burret’s concept of Myrtus
included the Mediterranean type, M. communis L.,
a north African species, and a complex of Caribbean
species, including the two that Small had originally
placed in Mosiera.
McVaugh (1968) considered the Caribbean com¬
plex that Burret had placed in Myrtus to be more
closely related to American genera (e.g., Calycolpus
Novon 2: 26-29. 1992.
Berg) than Myrtus, but he did not then transfer
them to Mosiera or any other American genus,
considering that a more thorough knowledge was
required before a final position could he assigned.
Later (1973), he transferred the type species of
Mosiera (M. longipes (Berg) Small) to Psidium.
In a posthumous publication, Bisse (1985) trans¬
ferred several Cuban species of Myrtus, in the sense
of Burret, and some species of Psidium to Mosiera.
His inclusion of Psidium guineense Sw. in Mosiera
leads me to believe that his concept of the genus
was quite different from mine. I consider Psidium
guineense to he a typical species of Psidium and,
in fact, a close relative of P. guajava L., the type.
A revision of Mosiera is underway, but 1 believe
that it is important that Psidium contrerasii and I*,
ehrenbergii he transferred now so that the correct
names can he used in other publications and to bring
attention to a genus until now unknown in Mexico
and Central America. Mosiera is here for the first
time critically compared to other genera and com¬
prehensively described.
Mosiera Small, Man. S. E. FI., 936. 1933. LEC-
TOTYPE: Mosiera longipes (Berg) Small (=
Eugenia longipes Berg) selected by McVaugh
(1956).
Shrubs or trees to 15 m high. Hairs whitish,
yellowish, or reddish brown, unicellular, simple, to
ca. 0.5 mm long. Leaves persistent, coriaceous, the
venation brochidodromous, with few to ca. 10 pairs
of lateral veins that are united by a marginal vein
that arcs between them near the leaf margin. Inflo¬
rescence a solitary axillary flower (infrequently a 3-
flowered dichasium) or an axillary bracteate shoot
with 1-3 decussate pairs of flowers. Flowers tetram-
erous; calyx lobes oblong to hemiorbicular, the calyx
usually not fused or only slightly fused beyond the
ovary’s summit, tearing slightly or not at all between
the lobes at anthesis; petals submembranous, white;
bracteoles usually small, ovate-triangular, foliose in
one species, caducous at about or before anthesis;
stamens 60-200, folded toward the center in the
bud; anthers globose, 0.3-0.7 mm long, usually with
one terminal gland in the connective; ovary 2-loc-
ular; ovules 3-40 per locule, the placenta not prom¬
inent, the locules sometimes connected by an open-
Volume 2, Number 1
1992
Landrum
Mosiera
27
ing. Fruit subglobose; seeds few, subreniform, the
seed coat hard, lustrous, or in one species leathery
and glandular, the external wall one to a few cells
thick, the surface cells rounded to elongate, the
central portion of the seed sometimes soft; embryo
oily, whitish, C-shaped, the cotyledons reflexed or
straight, less than !4 the length of the embryo.
Relationship to Eugenia (Eugeniinae) and Other
Genera of Myrtinae
Mosiera has been submerged in Myrtus and Psid-
ium and is also probably closely related to Caly-
colpus. It is now clear that Mosiera is tetramerous
while the others are all typically pentamerous, so
there should be no confusion. On the other hand,
Mosiera is superficially similar to Chamguava Lan¬
drum (Myrtinae) and Eugenia L. (Eugeniinae) and
can easily be confused with either. Examination of
embryos is the only certain way to distinguish these
three genera. The synoptic key below summarizes
the differences among all the above-mentioned gen¬
era.
la. Embryo a solid mass, made up of the fused cotyledons, the hypocotyl usually not distinguishable; flowers
tetramerous; inflorescence a solitary flower, or a bracteate shoot of 2 to a few decussate pairs of flowers
. Eugenia
lb. Embryo at least half hypocotyl, the cotyledons distinguishable as erect or reflexed flaps at the apex; flowers
tetramerous or pentamerous; inflorescence various.
2a. Seed coats membranous, thin, never verrucose; seeds to ca. 6-10 mrn long; embryo starchy, the
hypocotyl swollen, much wider than the cotyledons, a central core of vascular tissue clearly distin¬
guishable from the surrounding cortex (in section); placenta often a protruding peltate structure; flowers
tetramerous . Chamguava
2b. Seed coats hard, bony, or in Mosiera ekrenbergii leathery, verrucose; seed often less than 6 mm long;
embryo oily, the hypocotyl not greatly swollen, about the same width as the cotyledons, a central core
of vascular tissue not clearly distinguishable from the surrounding cortex; placenta protruding or not;
flowers tetramerous or pentamerous.
3a. Flowers tetramerous; seed coat smooth, lustrous, or leathery-verrucose; cotyledons much shorter
than the hypocotyl; anthers usually with a single terminal gland in the connective . Mosiera
3b. Flowers pentamerous; seed dull to rough (Psidium) or smooth, lustrous (Calycolpus, Myrtus );
cotyledons about the same length as the hypocotyl (Myrtus), or much shorter (Calycolpus,
Psidium ); anthers with a few to several glands in the connective (occasionally with a single terminal
gland in Myrtus).
4a. Seed coat dull or rough, not lustrous, covered with a thin layer of pulpy tissue when wet (or
a glaze or crusty layer when dry), the hard portion of the seed coat (5-)8-30 cells thick at
narrowest point; calyx tearing irregularly or between the lobes at anthesis; peduncles rarely
in pairs on very short axillary shoots . Psidium
4b. Seed coat lustrous, smooth, not covered with a thin layer of pulpy or crusty tissue or a glaze,
the hard portion of the seed coat 1-5 cells thick at narrowest point; calyx tearing between
the lobes or not at all at anthesis; peduncles often in pairs on very short axillary bracteate
shoots.
5a. Hypocotyl about equaling the cotyledons in length; calyx lobes triangular, unfused or
scarcely fused beyond the summit of the ovary, without an apical appendage; anthers
oblong, to ca. 1 mm long; ovary 2-3-locular; Mediterranean . Myrtus
5b. Hypocotyl much longer than the cotyledons; calyx lobes variously shaped, often fused
well beyond the summit of the ovary, often with an elliptic or linear apical appendage;
anthers oblong to linear, to ca. 2.5 mm long; ovary 2-6-locular; Central America and
northern South America . Calycolpus
Mosiera contrerasii (Lundell) Landrum, comb,
nov. Basionym: Psidium contrerasii Lundell,
Wrightia 5(4): 84. 1975. TYPE: Guatemala.
Peten: La Cumbre, in zapotal on top of hill, W
of km 141-142 on the Peten-Izabal road, 12
Mar. 1975 (fl), Lundell Contreras 19085
(holotype, LL).
Tree 7-15 m high, glabrous or essentially so
except for minute, obscure pubescence on youngest
twigs and buds and on calyx lobes; hairs less than
0.3 mm long, whitish or reddish brown; young twigs
light reddish brown or greenish, smooth, within about
1 year becoming gray to gray-tan, remaining smooth
or becoming slightly flaky. Leaves ovate to lanceo¬
late, 3-7.3 cm long, 1.2-2.5 cm wide, 2-3 times
as long as wide, the margin slightly revolute when
dry; apex bluntly acuminate; base rounded, cuneate,
or acuminate, sometimes oblique; petiole channeled,
3-7 mm long, 0.5-1 mm thick; midvein impressed
slightly above, prominent below; lateral veins about
7-10 straight pairs that extend from the mid vein
28
Novon
to the marginal vein, alternating with weaker,
branching veins that appear to arise in the marginal
vein and extend back toward the midvein, sometimes
impressed slightly above; blades subeoriaceous, dry¬
ing gray-green to dark yellow-green, usually some¬
what lustrous above, densely glandular. Peduncles
uniflorous, 0.4-2 cm long, 0.5-1 mm wide, usually
at leafless nodes, solitary or less often in pairs on
short shoots; bracteoles caducous before anthesis,
foliose, elliptic-linear, 4-6 mm long; calyx lobes
ovate-oblong, ca. 2 mm long and wide, concave,
pubescent within, glabrous proximally to pubescent
distally without; petals suborbicular, ca. 4 mm long;
hypanthium campanulate, 1.5-2 mm long; disk 1.5-
2 mm across; stamens 60-70, to ca. 4 mm long;
anthers ca. 0.3 mm long; style ca. 4 mm long; ovary
2-locular; ovules 7-11 per locule, biseriate, the
placenta central; Iruit globose, ca. 8 mm wide; seeds
4-10 per fruit, subreniform, ca. 4 mm long, lustrous
light yellow.
Phenology. Flowering from March to June; fruit
maturing May to September.
Habitat. As far as is known, growing in seasonally
dry, partially deciduous tropical forest on limestone.
Additional specimens examined. Guatemala. PETEN:
La Cumbre, zapotal, W of km 141-142 of Peten-Izabal
road, on top of hill, 12 Mar. 1975 (young fr), Lundell
& Contreras 19086, 19089, 10 May 1975 (fr) 19278,
11 Sep. 1975 (fr), 19847 (all LL). Mexico, quintana
ROO: km 33 del camino Carrillo Puerto-Vigia Chico, 29
July 1983 (fr), Duran & Olmsted 881 (MO); 1 km al S
del camino que nace 8 km al W de Vigia Chico, 5 Aug.
1983 (fr), Duran et al. 400 (MO); Carrillo Puerto, Si-
anka’an, 14 km al SE del crucero de Chumpon, 7 June
1984 (fl), Duran & Sanchez 990 (MEXLI).
Mosiera ehrenbergii (Berg) Landrum, comb. nov.
Basionym: Myrtus ehrenbergii Berg, Linnaea
27: 404. 1856. Psidium ehrenbergii (Berg)
Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15:
483. 1941. TYPE: Mexico. Ehrenberg 1039
(holotype, B destroyed).
Shrub or small tree 0.5-4 m high, smooth or
scaly barked; hairs whitish, erect to antrorse, ca.
0.1 mm long, essentially confined to young twigs
and margins of petals and calyx lobes; young twigs
reddish brown to gray-brown, densely to sparsely
covered with hairs, these persisting until the first
bark falls, the older twigs smooth or the bark be¬
coming rough and flaky. Leaves elliptic, oblong, or
elliptic-lanceolate, 0.9-3 cm long, 0.4-1 cm wide,
1.8-4 times as long as wide, glabrous, the margin
often slightly revolute; apex obtuse or emarginate;
base acute; petiole 1-2 mm long, ca. 0.5 mm thick;
midvein flat or slightly raised above, prominent be¬
low; lateral veins 2-7 pairs, weak, a marginal vein
usually not detectable; blades subeoriaceous, usually
drying gray-green, nearly concolorous, densely glan¬
dular. Peduncles uniflorous, 315 mm long, ca. 0.7
mm wide, flattened, solitary or more often in pairs,
one opposite the other on short, leafless axillary
shoots; bracteoles oblong-linear, 1-2.5 mm long,
usually caducous before anthesis; calyx lobes ovate
to hemiorbicular, 1.5-2 mm long and wide, glabrous
or with a ciliate margin, glandular, often tinged with
purple; petals suborbicular, ca. 3.5 mm long, often
tinged with purple; hypanthium campanulate, 2-3
mm long, tearing slightly between the calyx lobes;
disk 2-5 mm across, the staminal ring sparsely
pubescent; stamens 90 130, 3-4 mm long; anthers
0.5-0.7 mm long; style 3 6 mm long; ovary
2-locular; ovules 3 1 1 per locule, biseriate or mul-
liseriate; Iruit subglobose, turning dark purple-black
at maturity, 6-10 mm wide; seeds 1-5 per fruit,
ca. 4 mm long, the seed coat not operculate, leath¬
ery, 0.1-0.2 mm thick, strongly glandular.
Phenology. Flowering mainly from May to Sep¬
tember; fruiting mainly from August to October.
Habitat. Found growing in subtropical, seasonally
deciduous scrub forest and in desert.
No type specimen has been found for Myrtus
ehrenbergii, but the protologue leaves little doubt
about the identity of the species. Isotypes may exist
and should be searched for in European herbaria.
Representative specimens examined. Mexico. HI¬
DALGO: Cardonal, Cerro Boludo de Tolantango, 42 km al
NE de Ixmiquilpan, 31 Aug. 1976 (fr), Gonzalez Med¬
rano 9623 (MEXU). san I.uis poTOsfi 2 km E of El Abra
on road from Ciudad Valles to Tampico, 130 in, 27 Oct.
1985 (st), Bartholomew, Landrum et al. 3502 (ASU,
CAS); 4 km E of Charco Blanco on road to Guadalcazar,
1,600 m, 28 Oct. 1985 (fr), Bartholomew, Landrum et
al. 3568 (ASU, CAS); San Jose Pass, 10 Oct. 1890 (fr),
Pringle 3524 (A, CH); Minas de San Rafael, May 1911
(fl), Purpus 5211 (F, CH, MEXU, MICH, MO, NY, UC);
4 km al W de Lagunillas, 900 m, 29 Apr. 1965 (fl),
Rzedowski 7608 (MICH), tamaulipas: 18 km al SO de
Miquihuana, 1,700 m, 15 Aug. 1972 (fr), Gonzalez
Medrano 4723 (MEXU); 6 km al E del Sauz, al NE de
Tula, despues de Francisco Medrano, 1,600 m, 7 Dec.
1976 (fr), Gonzalez Medrano 10117 (MEXU); Tula, 3
km al W del Capulin. 2,000 m, Oct. 1976 (fr), Gonzalez
Medrano 10106 (MEXU); 2 km al N de la Verdolaga,
cerca de Las Antonias, 12 Aug. 1972 (fr), Gonzalez
Medrano et al. 4558 (MEXU, MO); 2 km al SE de
Magdaleno Aguilar, 1,600-1,800 m, 20 Sep. 1976 (fr),
Gonzalez Medrano et al. 9888 (MICH); Tula, 5 km al
W del Ejido El Saltillo, 1,500 m, 3 July 1985 (young
fr), Hiriart 758 (MEXLI); 18 km SE of Bustamante
toward Presita and Tula (99°40'W, 23°21'N), 1,700 m,
20 May 1973 (fl), Johnston et al. 11161 (ASU, CAS,
MEXU, NY); 6 mi. SW of Palmillas (23°18'N, 99°39'W),
5,300 ft., 1 Aug. 1976(A), W ebster & Armbruster 20508
(MEXU).
Volume 2, Number 1
1992
Landrum
Mosiera
29
Placement of Myrtus ehrenbergii in any genus
is somewhat difficult because of its unusual seed
coat, which is leathery, not operculate, and strongly
glandular. The only other species of Myrtinae known
to me with similar seed coats belong to the Aus¬
tralasian genus Pilidiostigma, which has a very
different embryo similar to that of Chamguava.
McVaugh (1968) first suggested that Myrtus eh¬
renbergii was related to the complex of Caribbean
species Burret referred to Myrtus, and that I con¬
sider to be Mosiera. It shares with them 4-merous
flowers, oil-storing embryos with the cotyledons much
shorter than the hypocotyl, ovaries with nonpro¬
truding placentae, and inflorescences of solitary
flowers or abbreviated bracteate shoots. The locules
are often incompletely closed as they usually are in
the ca. 10 additional species of Mosiera.
Another reason for believing Myrtus ehrenbergii
belongs to Mosiera is its geographic position. The
only other genera of Myrtinae that are geographi¬
cally close are Pimenta and Psidium, both of which
are dissimilar in many ways.
Mosiera ehrenbergii differs from the other spe¬
cies of Mosiera most notably in the structure of the
seed coat. Similar situations exist in Acca and Pi¬
menta, both of which have single species with un¬
usual seed coats (A. sellowiana (Berg) Burret; P.
pseudocaryophyllus (Gomes) Landrum) (Landrum,
1986).
Acknowledgments. I thank two anonymous re¬
viewers for helpful suggestions. Specimens from the
following herbaria were kindly made available for
this study: A, ASU, CAS, F, GH, LL, MEXU, MICH,
MO, NY, and UC. Field studies in Mexico were
made possible through NSF grant INT-8407889 to
Bruce Bartholomew and a National Geographic So¬
ciety grant to the author.
Literature Cited
Bisse, J. 1985. El genero Mosiera Small (Myrtaceae-
Myrtoideae) en Cuba 1. Rev. Jard. Bot. Nac. (Cuba)
6(3): 3-6.
Burret, M. 1941. Myrtaceen-Studien. Notizbl. Bot.
Cart. Berlin-Dahlem 15: 479-500.
Landrum, L. R. 1986. Campomanesia, Pimenta,
Blepharocalyx, Legrandia, Acca, Myrrhinium, and
Luma. Flora Neotropica 45: 1-179.
- & W. P. Sharp. 1989. Seed coat characters
of some American Myrtinae (Myrtaceae): Psidium
and related genera. Syst. Bot. 14: 370-376.
McVaugh, R. 1956. Nomenclatural notes on Myrtaceae
and related families. Taxon 5: 133-147, 162-167.
-. 1968. The genera of American Myrtaceae-
An interim report. Taxon 17: 354-418.
-. 1973. Notes on West Indian Myrtaceae. J.
Arnold Arbor. 54: 309-314.
Small, J. K. 1913. Flora of the Florida Keys. Published
by the author, New York.
-. 1933. Manual of the Southeastern Flora. Pub¬
lished by the author, New York.
Three New Species and a New Combination in the Asclepiadaceae
from the Venezuelan Guayana
Gilberto Morillo
Fundacion Instituto Botanico de Venezuela, Apartado 2156, Caracas 1010-A, Venezuela
Abstract. In preparation of the Asclepiadaceae
treatment for the Flora of the Venezuela Guayana,
one new species of Matelea, M. stergiosii, and two
new species of Ditassa, I). ottohuberi and I). ver-
ticillata , are herein described. A new combination
is also made in Ditassa, l). bolivarensis.
Matelea stergiosii Morillo, sp. nov. TYPE: Ven¬
ezuela. Estado Bolivar: moist forest on rocky
slopes, near Campamento Auraima, Rio Par-
agua, 6°22'30"N, 63°33'30"W, 330 m, May
June 1987, Stergios 10450 (holotype, VEN;
isotypes, MO, MY, NY, PORT). Figure 1.
Species haec Matelea squiresii (Rusby) Morillo prox-
ima, sed recedit: foliis late ovato-ellipticis vel obovato-
ellipticis, corolla 7.8 10 mm diametro, corollae lobis, 3-
3.4 mm longis, 2.4-3.2 mm latis, intus breviter et dense
puberulentis, gynostegio, 1.6-1.65 mm alto, 2.3-2.6 mm
diametro (0.95-1.2 mm alto et 1.55-1.95 mm diametro
in M. squiresii ); antheris 1.4 mm latis; coronae segmentis
basi 1.2-1.25 mm latis.
Twining vine; stems with short and inconspicuous
pubescence in two rows; leaves opposite; petioles
0.5-1.3 cm long, puberulent, blades broadly ovate-
elliptic to obovate-elliptic, 3-5.5 x 1.2-3.3 cm,
apex usually ± abruptly acuminate, base obtuse to
cuneate, glabrous except for a few hairs along the
midvein above, 5 or 6 pairs of lateral veins, 4
nectaries at the base. Inflorescence subaxillary, 2-
or 3-flowered; peduncle 1-2 mm long, bracts ovate,
0.6-0.7 mm long, ciliate along the margins; pedicels
2-2.5 mm long, glabrous. Calyx lobes ovate-elliptic,
1.8-2 x 1.1 —1.3 mm, with few short, curved hairs
outside and short cilia along the margins, the apex
narrowly obtuse. Corolla rotate, 7.8-10 mm diam.,
the lobes 3-3.4 x 2.4-3.2 mm, apparently green
and brown, densely short-appressed or erect-puber-
ulent inside, with few short, curved hairs along a
medial line outside, the apex obtuse. Gynostegium
shortly stipitate, 1.6- 1.65 mm long (tall) and 2.3-
2.6 mm diam.; anthers horizontal, 1.3-1.4 mm
wide. Pollinia horizontal, irregularly and narrowly
ovate-truncate, 0.45 x 0.3 mm, caudicles 0.2-
0.25 mm long, corpusculum narrowly sagittate, 0.2
x 0.1-0.12 mm. Corona formed by 5 column-
shaped segments, which are basally expanded, the
Novon 2: 30-32. 1992.
basal part 1.2-1.25 mm wide, margin almost entire.
Fruits unknown.
This species belongs to a difficult complex with
Matelea squiresii, which is known from southern
Venezuela and the Guianas (also probably in Brazil)
from lowland moist forests. Matelea stergiosii dif¬
fers clearly in the somewhat wider leaf blades, much
wider corolla, and gynostegium and inner face of
the corolla lobes with denser pubescence.
Ditassa ottohuberi Morillo, sp. nov. TYPE: Ven¬
ezuela. Territorio Federal Amazonas: summit
of Cerro Coro-Coro, NW headwaters of Rio
Manapiare, NW section of Serrania de Yutaje,
5°46'N, 66°12'W, 2,200 m, Nov. 1987, Hu¬
ber 12290 (holotype, VEN; isotypes, MO, MY,
NY, RB, TFAV).
Species nostra Ditassae acerifoliae Lasser et Maguire
affinis, sed differt: corolla subcampanulata, 1.9-2 mm
diametro, lobis 1.3-1.4 x 0.8-0.9 mm; gynostegio brev¬
iter stipitato 0.6-0.65 mm longo; antherae alis 0.26-
0.32 mm longis, corona exteriori segmentis oblongis vel
anguste triangulari-oblongis, 0.65-0.75 x 0.2 mm, co¬
rona interiori segmentis 0.2 mm longis.
Erect suffruticose plant, 20-30 cm high; stems
shortly branched, branches rugose, the lower part
with conspicuous persistent petiole bases, internodes
0.5-1 mm long, minutely erect-puberulent. Leaves
apparently disjunctly verticillate or subspirally ar¬
ranged, erect, up to 8 per whorl; petioles 0.6-1.8
mm long, decurrent, glabrous; blades fleshy, ± lin¬
ear, roundly triquetrous in cross section, 0.8-1 x
0.05 0.07 cm, the apex mucronate, the base grad¬
ually cuneate, the surface glabrous, veins and nec¬
taries not evident. Inflorescences axillary or inter-
petiolar, cymose, 1-3-flowered; peduncles 0.6-1.6
mm long, glabrous, bracts and bracteoles narrowly
ovate, 0.6-1 mm long, glabrous; pedicels 2.5-2.6
mm long, glabrous. Galyx lobes narrowly ovate, 1.1-
1.5 x 0.45-0.55 mm, obtuse. Corolla subcampan-
ulate, 1.9-2 mm diam., the tube 0.5-0.6 mm long,
glabrous, the lobes obliquely ovate, somewhat con¬
cave, suberect in anthesis, 1.3-1.4 x 0.8 0.9 mm,
glabrous outside, puberulent-papillose inside, the apex
narrowly obtuse. Gynostegium shortly stipitate, 0.6-
Volume 2, Number 1
1992
Morillo
Asclepiadaceae from
Venezuelan Guayana
31
Figure 1. Matelea stergiosii. —A. Habit, with detail of glands at base of leaf blade. —B. Detail of flower.
0.65 mm long (tall), 0.7-0.8 mm diam., stipe 0.1-
0.15 mm long; anther wings 0.26-0.32 mm long.
Pollinia pendulous, narrowly ovate, 0.25-0.26 x
0.06 mm, caudicles curved, 0.05 mm long, cor-
pusculum oblong-obovoid, 0.4 x 0.06 mm. Corona
double, outer segments narrowly triangular, 0.7-
0.75 mm long x 0.2 mm wide at base, slightly
longer than gynostegium, united by a short ring of
tissue, inner segments scalelike, irregularly ovate,
0.2 mm long. Fruits erect, narrowly ovate-fusiform,
2.9-3.2 x 0.3-0.4 cm, long-attenuate, glabrous,
with persistent calyx; seeds oblong ovate, 4-4.5 x
1.2-1.5 mm, brown-orange, margins entire, coma
ca. 20 mm long.
The general aspect of this plant is nearly identical
to that of Ditassa acerifolia Lasser & Maguire, a
species endemic to Cerro Yavi, but the new species
differs by the characters included in the diagnosis.
These two related species of Ditassa are distin¬
guished from the other Guayanan and Guianan spe¬
cies of the genus in possessing erect, woody stems
and dense acicular, spirally arranged leaves.
Ditassa verticillata Morillo, sp. nov. TYPE: Ven¬
ezuela. Territorio Federal Amazonas: Serrania
Yutaje, Cerro Yutaje, occasional on rocky
slopes, middle section of main branch, lefthand
fork of Cano Yutaje, 1,300-1,400 m, Feb.
1953, Maguire & Maguire 35243 (holotype,
VEN; isotype, NY).
Species nostra Ditassae taxifoliae Decne. proxima sed
notabilis: foliis verticillatis, coronae interioris lobis lineari-
oblongis, 0.9-0.95 mm longis; corollae lobis intus basi
pilis erectis instructis.
Suffruticose plant, stems mostly erect, sometimes
slightly twining, 30-55 cm long, shortly and densely
puberulent throughout. Leaves 3- or 4-verticillate,
rarely opposite in the upper nodes; petioles 0.2 0.3
cm long, with few hairs at the apex and margins;
blades linear, 1.5-2.1 x 0.1-0.3 cm, coriaceous,
the apex acute to mucronate, the base shortly cu-
neate, midvein prominent below, secondary veins
not evident, surface glabrous throughout, margins
slightly to strongly revolute; nectaries absent. Inflo¬
rescence subaxillary cymes, 1- or 2-flowered; pe¬
duncle 0.7-1.1 mm long, thick and rugose, rachis
bracteate, 0.1-2 mm long, bracts ovate to deltoid,
0.5-0.7 mm long, pedicels 1.7-2 mm long, gla¬
brous. Calyx lobes ovate, 0.9-1.1 x 0.7-0.8 mm,
obtuse, glabrous, eciliate. Corolla campanulate, the
tube 0.8-0.9 mm long, glabrous, the lobes ovate,
1.7-1.8 X 1.2-1.3 mm, glabrous outside, with
short, appressed hairs in the upper Vs and short,
erect hairs in the lower Vs inside, the apex narrowly
32
Novon
obtuse. Gynostegium sessile, 1.1 1.2 mm long (tall),
0.9-1 mm diam., anther wings 0.8 mm long. Pollinia
narrowly ovoid-ellipsoid, 0.3 X 0.09-0.1 mm, cau-
dicles 0.09 mm long, corpusculum narrowly elliptic
or narrowly ovate, 0.23 x 0.09 mm. Corona double,
outer segments rhombic-ovate and long-acuminate,
1.75 mm long, 0.7 mm wide at base, united with
each other in the lower 0.3 mm, inner segments
linear to oblong, 0.9-0.95 mm long, curved above
the anthers. Fruits unknown.
Ditassa verticillata is closely related to I), tax-
ifolia , with which it was originally confused, but
differs in the mostly 3- or 4-verticillate leaves, pe¬
duncles to 1.1 mm long, corolla lobes with erect
trichomes in the lower, inner half, and internal seg¬
ments of corona linear-oblong and longer than the
anther membranes.
Ditassa bolivarensis (R. Holm) Morillo, comb,
nov. Cynanchum bolivarense R. Holm, Field-
iana, Bot. 28: 504. 1953. TYPE: Venezuela.
Bolivar: in woods bordering streams draining
into Rio Kukenan, base of Mount Roraima,
1,185—1,280 m, Steyermark 58555 (holo-
type, F; isotype, VEN).
Transfer of Crinum nerinoides to Ammocharis (Amaryllidaceae)
David J. Lehmiller
Doctors’ Building, Suite 316, 3155 Stagg Drive, Beaumont, Texas 77701, U.S.A.
ABSTRACT. Lack of field observations previously
hampered taxonomic discussions of Crinum neri¬
noides. Several investigators suggested that this plant
might be Ammocharis rather than Crinum; how¬
ever, definitive reclassification was impossible be¬
cause pressed specimens often provide inadequate
material to differentiate between the two genera.
During a Namibian expedition in 1991, this taxon
was observed in flower and in fruit; the morpholog¬
ical features were specific for Ammocharis.
Baker (1903) worked from a single incomplete
specimen when he described Crinum nerinoides Ba¬
ker. The pressed specimen originated in Namibia
(formerly South West Africa) 11 years prior to
Baker’s account and consisted only of a scape bear¬
ing an umbel of seven flowers; no drawings or de¬
scriptive field notes accompanied the specimen.
After Milne-Redhead and Schweickerdt examined
the type specimen, they attached a determinant slip
annotated with “probable Ammocharis coranica .”
However, in revising the genus Ammocharis, Milne-
Redhead and Schweickerdt (1939) made the follow¬
ing statement about C. nerinoides: “This is most
probably an Ammocharis, but the type material is
too incomplete to be placed with certainty.” Soldi
(1969) discussed C. nerinoides under the heading
“dubious species” of Ammocharis, but he also con¬
cluded that the type material was too incomplete to
be classified generically.
Verdoorn (1973) recognized Crinum nerinoides
in her revision of the southern African species and
cited two new herbarium collections of the species,
both also from Namibia. Roessler (1974) followed
Verdoorn's treatment in his review of Namibian
Crinum. The paucity of herbarium materials led
Roessler to remark that C. nerinoides was rare.
Duncan (1982) cultivated and propagated bulbs of
C. nerinoides collected in Namibia; these were il¬
lustrated by several artists, most recently Brodley
(Plessis & Duncan, 1989).
Ammocharis and Crinum are closely related gen¬
era and can be confused in herbarium mounts. The
outstanding morphological feature that separates
these genera is the leaf arrangement: distichous and
biflabellately arranged in Ammocharis; rosulate or
distichous but never biflabellately arranged in Cri¬
num. Leaves are sheathed at the base in Crinum
but not in Ammocharis. Unfortunately, the inves¬
tigators who dealt with C. nerinoides were unaware
of its leaf arrangement, a character best seen in the
field.
During a Namibian expedition in January-!eb-
ruary 1991, 1). Hardy of the National Research
Institute (PRE) and I came upon a huge colony of
C. nerinoides growing in a pasture at Farm Okaseko
in the Gobabis District; these hulbs carpeted an area
approximately 200 x 20 m (estimated number in
excess of 20,000). By chance, about 25 bulbs were
blooming late, since this species ordinarily blooms
in December after the onset of the rainy season. All
of the plants had well-developed foliage because local
rainfall had been unusually abundant. The leaves
were clearly distichous and biflabellately arranged
(Fig. 1A), and sheathing of leaves was absent at the
base; the plant was an Ammocharis, not a Crinum.
Also, the fruiting bodies were partitioned by three
shallow furrows in the pericarp, differing from fruit¬
ing bodies of Crinum, which lacked furrows (Her¬
bert, 1837). We found a second cluster of blooming
bulbs surrounding a limestone pan near the town of
Aminuis (Fig. 1B).
Ammocharis nerinoides (Baker) Lehmiller, comb.
nov. Basionym: Crinum nerinoides Baker, Bull.
Herb. Boissier ser. 2, 3: 666. 1903. TYPE:
Namibia. Gobabis District: Hereroland, 1892,
Dove s.n. (holotype, Z).
Bulb ovoid, covered with a brown papery tunic,
20-51 mm diam., tapering into a neck 50 85 mm
long. Leaves 4-10, distichous and biflabellately ar¬
ranged, arching or sprawling on the ground, linear,
mildly channeled, minutely serrated on the margins,
green, all but new leaves growing out with truncated
ends, 130-380 mm long and 3-10 mm wide. Scape
compressed, reddish green, 40-115 mm long. Spathe
valves papery at anthesis. Umbel 1-7-flowered;
flowers actinomorphic, scented. Pedicels 8-19 mm
long. Ovary a small swelling 3-5 mm long and 3-
4 mm wide. Perigone tube not curved, red, 13-20
mm long. Perigone segments lanceolate, distally re¬
curved, pink with a broad dull red stripe on the
dorsal keel, unequal with the inner narrower, 43-
Novon 2: 33-35. 1992.
34
Novon
Figure 1. Ammocharis nerinoides (Baker) Lehmiller. —A. Distichous leaves, biflabellately arranged. Namibia,
Gobabis District, Farm Okaseko, 31 January 1991. —B. Blooming bulb. Namibia, Gobabis District, Hereroland East
#2, 2 km north of Aminuis, 1 February 1991.
47 mm long and 8-11 nun wide. Filaments pink,
unequal with the inner longer, 38-41 mm long;
anthers white, turning dark tan and becoming curved
at maturity, 3 mm long; pollen gray. Style pink,
37-45 mm long, with a tiny capitate stigma. Fruit
subglobose, marked with three furrows and appear¬
ing trilobulated, often with a short apical heak to 5
mm long, reddish hrown, 1 8-20 mm diam., inde-
hiscent; seeds 3-1 2 per fruit, ovoid or angled by
pressure to resemble parts of a sphere, smooth, light
green, to 1 2 mm diam.
Habitat. Arid ecology with an erratic, summer
rainy season. Open grasslands in low places or near
limestone pans; clayish calcareous soils. Flowering
induced by summer rains.
Additional specimens examined. Namibia. gobabis
district: Farm Nico, 1958, MerxmiiUer & Giess 1176
Volume 2, Number 1
1992
Lehmiller
Ammocharis
35
(PRE); Farm Eskadron, 1969, Mason & Boshoff 2516
(PRE); Farm Combumbi, 1977, Giess 14919 (WIND);
Farm Okaseko, 1991, Hardy 7231 (PRE); Hereroland
East #2, 2 km N of Aminuis, 1991, Hardy 7235 (PRE).
ETOSHAPAN: Adamax, 1974, le Roux 596 (PRE); same
locality, 1975, Muller 196 (WIND).
Ammocharis coranica (Ker-Gawl.) Herb., which
is epidemic in Namibia, is a huge bulb in comparison
to A. nerinoides and is easily distinguished among
herbarium mounts on the basis of bulb diameter,
leaf width, scape dimensions, and umbel size; the
floral parts are remarkably similar. In the field, the
many-flowered bright-red umbels and wide leaves of
A. coranica could never be confused with the di¬
minutive, few-flowered pink umbels and narrow leaves
of A. nerinoides. The specific epithet “nerinoides”
is most appropriate since, when casually observed,
A. nerinoides resembles a species of Nerine.
While in Windhoek I visited W. Giess, who was
curator of the Namibian Herbarium (WIND) for 40
years and is now retired. During the interview Giess
kindly projected 35-mm color slides recording A.
nerinoides in bloom at Farm Nico and Farm Com¬
bumbi (see list of specimens examined). Giess also
permitted me to examine field-collected bulbs, which
he cultivated in his garden; these too exhibited bi-
flabellately arranged, distichous leaves.
Acknowledgments. I thank the following herbaria
for use of their facilities: PRE, WIND, K. The
National Research Institute, Pretoria, South Africa,
sponsored D. Hardy’s participation.
Literature Cited
Baker, J. G. 1903. Amaryllidaceae. In: H. Schinz (ed¬
itor), Beitrage zur kenntnis der Afrikanischen flora.
Bull. Herb. Boissier ser. 2, 3: 665-667.
Duncan, G. 1982. Crinum nerinoides at Kirstenbosch.
Indigenous Bulb Growers Association of South Africa
Bulletin 32: 3-4.
Herbert, W. 1837. Amaryllidaceae. J. Cramer Reprint,
Lehre.
Milne-Redhead, E. & H. G. Schweickerdt. 1939. A
new conception of the genus Ammocharis Herb. Bot.
J. Linn. Soc. 52: 159-197.
Plessis, N. du & G. Duncan. 1989. Bulbous Plants of
Southern Africa. Tafelberg Publishers, Capetown.
Roessler, H. 1974. Ubersicht fiber die heute aus Sfid-
westafrika bekannten Crinum- arten. Mitt. Bot. Mfin-
chen 11: 523-544.
Solch, A. 1969. Amaryllidaceae. In: H. Merxmfiller
(editor), Prodromus einer Flora in Sfidwest-Afrika
150: 2-8.
Verdoorn, I. C. 1973. The genus Crinum in southern
Africa. Bothalia 11: 27-52.
Two New Species of Aristida (Poaceae) from Brazil
Hilda Maria Longhi-Wagner
Departamento de Botanica UFRGS-Av. Paulo Gama, s.n.-90049 Porto Alegre,
Bio Grande do Sul, Brazil
Abstract. Two new species of the “Ripariae”
group of Aristida are described and illustrated: A.
brasiliensis is distinguished by the lower glume,
which is shorter than the upper one, and by the
pubescent collar; A. constricta by the articulation
at the base of the long column of the lemma. A key
to the group is provided.
The genus Aristida L. comprises about 280 spe¬
cies from tropical and subtropical areas. During the
revision of this genus for Brazil, two new species
were found and are described in this paper. Speci¬
mens ot both species had been previously identified
as different species by earlier authors and can be
included in the “Ripariae’’ group discussed by llen-
rard (1928, 1932). According to Henrard, this group
includes A. riparia Trin. & Rupr., A. megapotam-
ica Sprengel, A. oligospira Trim, and A. trinii
Henrard, and is characterized by the spikelike pan¬
icles and the lower glume longer than the upper
one. Actually, A. trinii does not belong to this group
due to its linear and subdense, rather than spikelike,
panicle and shorter spikelets. On the other hand,
the inclusion of A. brasiliensis in this group, based
on its habit and spikelike panicle, enlarges the cir¬
cumscription of the “Ripariae” because A. bras¬
iliensis has the lower glume shorter than the upper
one. These species can be distinguished by the fol¬
lowing characteristics.
Key to the Taxa of Aristida Group Ripariae
la. Callus of the lemma obtuse . 4. oligospira
lb. Callus of the lemma acute or bifid.
2a. Lower glume shorter than the upper one;
collar pubescent . A. brasiliensis
2b. Lower glume longer than the upper one;
collar glabrous.
3a. Callus of the lemma bifid .... A. riparia
3b. Callus of the lemma acute.
4a. Lemma articulated at the base of
the 10-15-mm-long column . . .
. A. constricta
4b. Lemma not articulated; column
29-60 mm long.
5a. Awns of the lemma contort¬
ed at the base .
. A. megapotamica
var. megapotamica
5b. Awns of the lemma straight
at the base . . A. megapotamica
var. brevipes
Aristida brasiliensis Longhi-Wagner, sp. nov.
TYPE: Brazil. Minas Gerais: ca. 12 km from
Barao de Cocais, base of Serra do Caraga, 28
Jan. 1971, H. S. Irwin et al. 29325 (holotype,
SP; isotypes, MO, NY, UB). Figure 1.
Perennis, 65- 120 cm alta. Laminae supra glabrae
saepe pilis sparsis longis ad basin praedictae, 25-50 cm
longae, 0.25-0.35 cm latae, culmorum inferne planae,
recurvatae, superne convolutae; ligula ciliata, ca. 0.2 mm
longa; collum pubescens. Panicula spiciformis, densa, 16-
23 cm longa. Glumae uninervatae, scabrae, inferiores
15.5-20 mm longae (aristae 5-8 mm longae computatae),
superiores 23.5-28 mm longae (aristae 7-10 mm longae
computatae). Lemma 33-39 mm longum (callo et col-
umna computatis), columna 23-29 mm longa, callo acuto,
1 mm longo; aristae rectae, centralis 47 59 mm longa,
laterales 40-53 mm longae.
Perennial, caespitose, 65 120 cm tall. Sheaths
longer than the internodes, glabrous, sparsely long
bearded at the summit especially in the innovations,
with deciduous hairs to 2 mm long, collar pubescent;
ligule ciliate, ca. 0.2 mm long; blade 25-50 cm
long, 0.25-0.35 cm wide, abaxially scabrous, gla¬
brous or sparsely hairy near the base (hairs ca. 2
mm long), flat and recurved at the base, convolute
upward. Panicle dense and spikelike, 16 23 cm
long, 3-6 cm wide (including the awns), erect. Glumes
lanceolate, acuminate, awned, 1-nerved, the lower
15.5-20 mm long (including the 5-8-mm-long awns),
0.5-0.8 mm wide (lateral view), densely scabrous,
the upper ones 23.5-28 mm long (including the 7-
10-mm-long awns), 0.5-0.7 mm wide, minutely sca¬
brous. Lemma cylindrical, smooth, 33-39 mm long
(including the twisted 23-29-mm-long column and
the acute and hairy 1-mm-long callus); awns straight,
the central one 47-59 mm long, the lateral ones
40-53 mm. Palea hyaline, 2 mm long, 0.5 mm
wide, acute, 2-nerved. Lodicules 2, hyaline, 1.5 mm
long, 0.3 mm wide. Stamens 3. Caryopsis 5.5 mm
long, 0.6 mm wide.
Distribution and ecology. Brazil, from Minas
Gerais to Santa Catarina. It is most common in
campo rupestre and cerrado vegetation, up to
1,400 m.
Novon 2: 36-40. 1992.
5cm
Volume 2, Number 1
1992
Longhi-Wagner
Aristida from Brazil
37
Figure 1. Aristida brasiliensis Longhi-Wagner. — A. Habit. —B. Pubescent collar. —C. Anthecium. —D. Glumes,
lateral view. —E. Lemma, awns partially removed. (Irwin et al. 28819.)
2mm
38
Novon
Figure 2. Aristida constricta Longhi-Wagner. —A. Habit. —B. Anthecia. —C. Glumes, lateral view. —D. Lemma,
awns partially removed. —E. Lemma apex with an articulation. (Longhi-Wagner et al. 1562.)
2mm
Volume 2, Number 1
1992
Longhi-Wagner
Aristida from Brazil
39
Paratypes. BRAZIL. MINAS GERAIS: ca. 10 km W of
Barao de Cocais, 22 Jan. 1971, //. S. Irwin et al. 28819
(UB); Diamantina, 8 km E Diamantina, road to Extra 5 ao,
16 Mar. 1970, H. S. Irwin et al. 27687 (MO, NY, UB),
km 996 997.5 antiga ferrovia Diamantina Corinto, 12
Feb. 1982, A. G. Burman T. Sendulsky 767 (SP),
Serra de San Antonio, 27-30 Dec. 1929, Chase 10395
(MO, US); Santa Barbara, Serra do Caraga, 22 Jan. 1972,
A. Macedo 5230 (HB, MBM, MO), sao paulo: campos
de Bocaina, 6 Apr. 1894, A. Loefgren 2364 (SP). SANTA
Catarina: Lages, Morro do Pinheiro Seco, 19 Feb. 1971,
L. B. Smith & Klein 15926 (HBR).
Aristida brasiliensis is closely related to A. rne-
gapotamica var. brevipes Henrard from Brazil,
sharing a similar habit and panicle morphology, but
can be distinguished by the characteristics cited in
the key. In addition, A. megapotamica var. brev¬
ipes has a longer column (29-49 mm long) and
callus (1.2-2 mm long). It is also closely related to
A. spiciformis Ell., from North America. They are
similar in habit, panicle, and relative length of the
glumes. However, A. spiciformis differs in its char¬
acteristic glumes, which are shorter and abruptly
long awned, spreading awns of the lemma, longer
callus (1.5-2 mm long), and the absence of a pu¬
bescent collar. Henrard (1933) cited Chase 10395
as a paratype of A. megapotamica var. brevipes.
It is possible that Henrard did not notice the dif¬
ferences in the length of the glumes and the pu¬
bescent collar. The cited collection agrees perfectly
with the new species described here.
Most of the paratypes now identified as A. bras¬
iliensis were previously referred to A. implexa Trim,
which is a synonym of A. megapotamica.
Aristida constricta Longhi-Wagner, sp. nov.
TYPE: Brazil. Rio Grande do Sul: Santana do
Livramento, Cerro Palomas, 17 Nov. 1987,
Longhi-Wagner et al. 562 (holotype, ICN;
isotype, MO). Figure 2.
Perennis, 24-46 cm alta. Laminae glabrae, 10-33 cm
longae, 0.2-0.35 cm latae, culmorum inferne planae,
recurvatae, superne convolutae; ligula ciliata, ca. 0.2 mm
longa; collum glabrum. Panicula spiciformis, densa, 10-
15 cm longa. Glumae uninervatae, inferiores 16-19.5
mm longae (aristae 4-6 mm longae computatae), scabrae,
superiores 13-15 mm longae (aristae 2-2.5 mm longae
computatae), laeves. Lemma 18-22 mm longum (callo et
columna computatis), in basi columna constrictum, arti-
culatum; columna 10-15 mm longa; callo acuto, 1.2-
1.3 mm longo; aristae rectae, centralis 32-40 mm longa,
laterales 26-32 mm longae.
Perennial, caespitose, 24-46 cm tall. Sheaths
longer than the internodes, glabrous, sparsely long
bearded at the summit especially in the innovations,
with deciduous hairs; ligule ciliate, ca. 0.2 mm long;
collar glabrous; blade 10-33 cm long, 0.2-0.35 cm
wide, abaxially and marginally scabrous. Panicle
dense and spikelike, 10-15 cm long, 5-6 mm
wide, erect. Glumes lanceolate, acute, short awned,
1- nerved, the lower 16-19.5 mm long (including
the 4-6-mm-long awns), 0.7 mm wide (lateral view),
densely scabrous, the upper one 13—15 mm (in¬
cluding the 2-2.5-mm-long awns), 0.5 mm wide,
smooth. Lemma cylindrical, constricted and artic¬
ulated at the base of column, 18-22 mm long (in¬
cluding the twisted and 10-15-mm-long column and
the 1.2-1.3 mm, acute callus); callus hairy except
for the upper 0.2-0.3 mm of the apex, which is
glabrous and forms a conspicuous and curved beak;
awns straight, the central one 32-40 mm long, the
lateral ones 26-32 mm long. Palea hyaline, acute,
2- nerved, subequal to the lodicules, 1.4-1.6 mm
long, 0.3-0.4 mm wide. Lodicules 2, hyaline, linear,
1.4 mm long, 0.1 mm wide. Stamens 3.
Distribution and ecology. According to the data
presently available, Aristida constricta has a re¬
stricted distribution in southwestern Rio Grande do
Sul, Brazil. It is common on basalt crevices and on
the top and slopes of hills.
Paratypes. Brazil. RIO grande DO SUL: Santana do
Livramento, Cerro Armour, 17 Nov. 1987, Longhi-
Wagner et al. 1574 (ICN), Cerro Palomas, 25 Nov.
1972, Vails & Barcellos 2527 ( ESAL, ICN), Mar. 1991,
Vails et al. 12773 (CEN, ICN); Sao Francisco de Assis,
23 Nov. 1972, Vails & Barcellos 2470 (ICN).
The most distinctive characteristic of Aristida
constricta is the presence of a constriction at the
base of the lemma column, which represents an
articulation point; this allows the awns and column
to be easily detached from the lemma.
Specimens of A. constricta have been cited as
A. trinii Henrard by Severo (1982) for Rio Grande
do Sul, Brazil. In fact, A. trinii was described from
central Brazil and does not occur in Rto Grande do
Sul. Aristida trinii has a subdense panicle and lacks
an articulation at the lemmas.
Acknowledgments. I express my appreciation to
the Missouri Botanical Garden for the May Fellow¬
ship funded by the May Department Stores Com¬
pany and its Famous-Barr Division, which enabled
me to work at the Garden. I also thank Enrique
Forero and the staff of the Missouri Botanical Gar¬
den. Special thanks to Gerrit Davidse for his critical
review of the manuscript, to Tarciso Filgueiras for
correcting the Latin diagnoses, and to Carlos Reynel
for his help during the processing of the text.
Literature Cited
Henrard, J. Th. 1928. A critical revision of the genus
Aristida. Meded. Rijks Herb. 3(54b): 502-701.
40
Novon
-. 1932. A monograph of the genus Aristida.
Meded. Rijks-Herb. 2(58): 157-325.
-. 1933. A critical revision of the genus Aris¬
tida. Meded. Rijks-Herb. 55c: 703-747 (Suppl.).
Severo, B. A. 1982. 0 genero Aristida L. (Gramineae)
no Rio Grande do Sul. M.S. Thesis, Universidade
Federal do Rio Grande do Sul, Brasil.
Guadua chacoensis (Poaceae: Bambuseae), its
Taxonomic Identity, Morphology, and Affinities
Ximena Londono
Institute* Vallecaucano de Investigaciones Cientificos-INCIVA,
Apartado Aereo 5660, Cali, Colombia
Paul M. Peterson
Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, U.S.A.
ABSTRACT. A new combination, Guadua chacoen¬
sis, is made. Based on morphological evidence, G.
chacoensis appears closely allied to G. trinii. A
detailed illustration of the species is included, and
morphological descriptions and illustrations of the
seedling and fruit are provided.
Guadua was segregated from the genus Bambusa
Schreber and described by Kunth (1822). Important
diagnostic characters that distinguish this genus are
triangular culm leaves in which the margins of the
sheath and blade are usually contiguous, a distinctive
hand of short white hairs above and below the nodal
line, synflorescences with 2- to many-flowered pseu-
dospikelets, a palea of firm texture with prominent
wings emanating from the keels, and the presence
of stomata and papillae on the adaxial surface of
the leaf blades (Soderstrom & Ellis, 1987; Soder-
strom & Londono, 1987). Other characters useful
in distinguishing among the species of the genus are
the shape of culm and foliage leaf, the structure and
shape of pseudospikelets, and the habit of the plant.
Guadua occurs throughout tropical America, from
Mexico to northern Argentina, and contains perhaps
30 species. Along with Guadua, four other genera
are placed in the subtribe Guaduinae: Criciuma
Soderstrom & Londono and Eremocaulon Soder¬
strom & Londono from Brazil, and Olmeca Soder¬
strom and Otatea (McClure & Smith) Calderon &
Soderstrom, from Mexico (Soderstrom & Ellis, 1987).
Guadua chacoensis occurs in northern Argen¬
tina, southeastern Bolivia, and southern Paraguay
and is one of the three southeasternmost species of
the genus. It is frequently misidentified as G. an-
gustifolia Kunth. The species was described in 1918
by the Argentinian botanist Nicolas Rojas Acosta as
Bambusa chacoensis. A type specimen was not
designated but the locality data (General Vedia, Dep-
to. Bermejo, Prov. del Chaco-Argentina, and He de
la Colonia Aquino, Rio Paraguay) and the common
name “tacuara,” distinguish it from two species that
occur sympatrically, G. paraguayana Doell, “pi-
canilla,” and Guadua trinii (Nees) Ruprecht, “ya-
tevo,” “tacuaruzu,” or “tacuara brava” (Nicora &
Rugolo, 1987; Young, 1985).
McClure (1973), in his annotated checklist of
species, included G. chacoensis under the heading
of unresolved specific taxa. Young (1985) treated
this species as one of two subspecies in the variable
Guadua angustifolia Kunth complex and indi¬
cated a neotype. However, Young (1985) pointed
out that “the two subspecies can be separated using
a combination of vegetative and reproductive char¬
acters but since the morphological gap separating
them is smaller than that separating most species
in the subgenus, the taxa are considered here to be
only subspecifically distinct.”
In this study, a comparison of floral and vege¬
tative morphology among G. chacoensis, G. an¬
gustifolia, and G. trinii is presented. Based on com¬
plete material collected by Camilo Quarin, a botanist
from the Universidad Nacional del Nordeste, Cor-
rientes, Argentina, a detailed illustration of the habit
of G. chacoensis and descriptions and illustrations
of the seedling and fruit are provided.
Cuadua chacoensis (Rojas) Londono & Peterson,
comb. nov. Basionym: Bambusa chacoensis
Rojas, Bull. Acad. Int. Geogr. Bot. 26: 157.
1918. TYPE: Argentina. Chaco: Departamen-
to Bermejo, General Vedia, Riberas del Rio de
Oro, 23 July 1974, Quarin et al. 2384 (neo¬
type, designated here, CTES; isoneotypes, F,
G not seen, MO, US). Figure 1.
Woody, thorny bamboo. Rhizomes pachymorph.
Culms 10-20 m tall, 8-15 cm diam., green, erect
below and arching apically; internodes hollow; nodes
solitary; bud solitary, covered by a prophyll, posi¬
tioned 2-4 mm above the nodal line. Culm leaves
Novon 2: 41-47. 1992.
42
Novon
Figure 1. Guadua ehacoensis (Rojas) Londono & Peterson. —A. Flowering branch. —B. Leaf complement. —C.
Leaf ligule. —D. Culm sheath. —E. Detail of culm sheath ligule, adaxial view. —F. Branch complement. Based on:
A, Quarin et al. 2384 , B, C, Schinnini 10956; D. Tressencs & Schinnini 1310 , E, F, Schinnini et al. 6819.
Volume 2, Number 1
1992
Londono & Peterson
Guadua chacoensis
43
coriaceous, light brown to stramineous, deciduous,
the blade l A- l A as large as the sheath; sheath 20-
50 cm long, 8 30 cm wide, abaxially strigose to
glabrescent, the coarse, rigid, brown hairs to 2 mm
long, easily removed, adaxially glabrous, shiny, and
tessellate, auricles and fimbriae absent; blade 4-13
cm long, 3-8 cm wide, triangular, erect, persistent,
strongly mucronate at the apex, abaxially strigose
to glabrous, adaxially conspicuously nerved and pu¬
bescent between the nerves, the margins ciliate with
hyaline, deciduous hairs, up to 2.5 mm long, the
junction with the sheath slightly curved to ± hor¬
izontal; inner ligule 0.8-1 mm long, glabrous, stra¬
mineous, ciliolate along the margin, following the
junction between the sheath and the blade and end¬
ing at the margins; outer ligule absent. Branching
intravaginal; primary branch solitary and armed,
later developing 2 or 3 secondary branches from
the basal proximal nodes. Foliage leaves 7 to 9 per
complement; sheath glabrous, nonauriculate, ciliate
on the margins; fimbriae 2-3 mm long, restricted
to the sheath summit, erect basally, wavy to curly
distally, 3-5 per tuft, white-ivory; blade 10-24 cm
long, 0.5-2.5 cm wide, L:W ratio 11-26:1, ex¬
tremely variable in size, linear, 8-14-nerved, adax¬
ially glabrous to sparsely hispid, the hairs 0.5-1 mm
long, hyaline to brown, antrorse, 3-4 rows of prick¬
le-hairs along one margin, abaxially glabrous, with
sparsely coarse, rigid, hyaline hairs mainly along
the midnerve, the margins strigose; pseudopetiole
2-3 mm long, adaxially pilose to glabrous, abaxially
glabrous; inner ligule 0.2-0.4 mm long, minutely
pubescent, ciliate on the margin with erect, hyaline
hairs, the hairs 0.2 mm long; outer ligule 0.1 -0.2
mm long, minutely ciliolate on the margin. Synflo¬
rescence 30-40(-50) cm long, usually terminating
leafless branches, consisting of 6-9 coflorescences
with 4-7 multiflowered pseudospikelets per coflo¬
rescence; rachis glabrous. Pseudospikelet 2-5( 6)
cm long, 0.4-0.5(-0.6) cm wide, robust and erect,
straight, corpulent, green when young, later brown
to stramineous, consisting of a subtending bract, a
prophyll, 1-2 glumes, 1-2 sterile lemmas, 2-4(-6)
fertile florets, in a terminal rudimentary anthecium;
rachilla 5-7 mm long between florets, hispidulous;
subtending bracts 8-10 mm long, 4-5 mm wide,
ovate to ovate-lanceolate, abaxially glabrous to
sparsely pubescent, adaxially minutely pubescent on
the distal V3, the apex mucronate; prophyll winged,
pubescent between keels and ciliolate on the wings,
occasionally gemmiferous; glumes 5-8 mm long, 3
mm wide, 1-2, usually gemmiferous, broadly ovate
to ovate-lanceolate, 7-9-nerved, mucronate, posi¬
tioned close to the prophyll, abaxially shiny, with
scattered minute transparent hairs on the dorsal
part, adaxially minutely pubescent, with transverse
venations apically, the margins smooth; sterile lem¬
ma 10-14 mm long, 6-10 mm wide, sometimes
enclosing a rudimentary palea, usually stramineous,
mucronate, abaxially shiny and glabrous, adaxially
minutely pubescent, the margins scarious, smooth;
fertile lemma 12-16 mm long, 7-12 mm wide,
ovate-lanceolate, 12-17-nerved, totally embracing
the palea, mucronate, stramineous or purplish, abax¬
ially covered with hyaline, appressed, sometimes
spreading hairs on the proximal l A, otherwise gla¬
brous and shiny, adaxially shiny, minutely pubes¬
cent, the margins scarious, shiny, bearing a tuft of
hairs distally, below the mucro, the mucro 0.8-1
mm long, brown when old; palea 6-14 mm long,
2-4 mm wide, usually shorter than the lemma,
stramineous, abaxially pubescent, adaxially gla¬
brous, the sulcus 2-2.5 mm wide, 3-nerved, apic-
ulate at the apex with a tuft of hairs, the keels
winged, the wings 1-1.5 mm long, 2-3-nerved,
stramineous, prolonged at the apex, ciliate on the
margin, abaxially puberulous close to the keels, oth¬
erwise glabrous and shiny, adaxially glabrous, the
enfolded margins 3-nerved, glabrous. Lodicules 3,
transparent, many-nerved, glabrous, ciliolate on the
upper part of the margin; the anterior pair 3-5 mm
long, 2-3 mm wide, the posterior one 2-3 mm long,
1.5-2 mm wide. Stamens 6; anthers 5-8 mm long,
0.5-1 mm wide, sagittate at the base, apiculate at
the apex of each theca, yellowish brown. Ovary 2-
4 mm long, 1-2 mm wide, fusiform, glabrous and
shiny; style 2-3 mm long, hispidulous, darker than
the ovary with 3 plumose stigmas, the stigmas 3
mm long and purple when young. Fruit 9-12 mm
long, 2.5-5 mm wide, an asymmetric fusiform car-
yopsis, brown, glabrous; embryo 2-3 mm long, 2-
3 mm wide, basal, circular.
Distribution. Known from northern Argentina,
southeastern Bolivia, and southern Paraguay where
it occurs in gallery forests usually along stream and
river edges at elevations of 75-450 m.
Additional specimens examined. Argentina, cor-
RIENTES: Depto. Capital, Arroyo Riachuelo y Ruta 12, 16
Jan. 1974 (fl), Quarin et al. 1859 (US), 15 Aug. 1974
(fl), Quarin 2402 ( MO, US); 21 Dec. 1974 (fl), Maruhak
635 (US); 27 Nov. 1975 (fl), Quarin 3125 (MO, US);
Puente Pesoa, Arroyo Riachuelo, 3 Aug. 1973, Schinnini
et al. 6819 (US); Depto. Concepcion, Rincon de San
Pedro, 17 Dec. 1974 (fl), Burkart et al. 30687 (MO,
US), Quarin 2870 (MO, US); Depto. Empedrado, Ruta
12 y Rio Empedrado, 24 Oct. 1975 (fl), Quarin 3210
(MO, US); Depto. Ituzaingo, Isla Apipe Grande, Pto. San
Antonio, 8 Dec. 1974 (fl), Krapovickas et al. 23851
(MO, US); Rincon Ombu Chico, 4 July 1974 (fl), Kra¬
povickas et al. 25470 (MO, US); Depto. Mercedes, 75
km N de Mercedes, Laguna Trin, 17-24 Oct. 1975 (fl),
44
Novon
Quarin et al. 3182 (US), mlsiones: vicinity of Pto. Aguirre,
100 m, 8-10 July 1914, Curran 676 (F, NY, US); Pto.
Leon, 75 100 m, 21 July 1914 (fl), Curran 677 (NY,
US); Iguazu, 7 Oct. 1910, Rodriguez 486 (MO, NY,
US); San Ignacio, Penon Reina Victoria, 14 Jan. 1976
(fl), Krapovickas <S: Cristobal 28759 (MO, US). CHACO:
Depto. Bermejo, Isla Guascara, 2 July 1981, Tressens
& Schinnini 1310 (US); Casa lata, 28 July 1944 (fl),
Rojas 11719 (NY). TUCUMAN: Cultivado, Jardines Insti¬
tute Miguel Lillo, procedente de Misiones, Iguazu, 26
Aug. 1976 (fl), Cuezzo s.n. (US). Bolivia, santa cruz:
Prov. Sara, Rio Surutu, 1 Oct. 1925 (fl), Steinbach
7233a (F, MO, US); Prov. Ichilo, 15 km SE of Buena
Vista, 375 in, 28 July 1987, Nee et al. 35397 (MO,
NY); Parque Nacional Arnboro, along Rio Cheyo, 400
m, 24 ago. 1985 (fl), Solomon 14021 (MO, US,); 23 km
S of Buena Vista along Rio Chonta, 420 m, 18 Nov.
1988 (fl). Nee 36861 (NY); along Rio Saguayo, 1.5-3
km NE of entrance into first Andean foothills, 17°38'-
39'S, 63°43'W, 375 in, 21 Dec. 1988 (fl), Nee 37301
(MO). Paraguay, central: Ruta 2 y Arroyo Mboiy, 2
Mar. 1975 (fl), Schinnini 10956 (US). PARACUARl: sur
le bord des ruisseaux, Dec. 1883 (fl), Balansa 4346 (P,
US), san PEDRO: Colonia Nueva Germania, 23 Oct. 1916
(fl), Rojas 2311 (US); alto Paraguay, Primavera, 16 June
1957, I Voolston G-112 (NY). CAAGUAZO: Orillas arroyo
Tebicuary, entre Coronel Oviedo y Colonia Independen-
cia, 15 Oct. 1951 (fl), Burkart 18741 (SI, US). flEEMBUcO:
Barranca selvatica del rio Tebicuary-guazu, Jan. 1944
(fl), Pavetti <£■ Rojas 10959 (UTS).
Flowering. The incidence of flowering and fruit¬
ing in bamboos varies from one species to another
(McClure, 1966), and according to records of spec¬
imens available at F, ISC, MO, NY, and US, G.
chacoensis flowered in the following years: 1883,
1914, 1925, 1944, 1951, 1957, 1975-1976,
1981, and 1985-1988. We cannot establish a reg¬
ular flowering cycle for this species; however, it is
evident that after a long period without flowering,
G. chacoensis can flower uninterruptedly for at least
3 years. Observations made by Quarin during 1974-
1975 indicated that the flowering culms in G. cha¬
coensis are usually leafless, and fruits and seedlings
are produced when most of the plant is dry.
The flowering cycle of G. trinii appears to be
unique and well known in the genus (McClure, 1966).
It was reported by Parodi (1955), who personally
observed the flowering cycle from the same hered¬
itary line for a period of 30 years (1923-1953).
After flowering and fruiting uninterruptedly for one
year, the clump died completely (i.e., truly mono-
carpic). The fruits then fell to the ground and ger¬
minated.
The flowering cycle of G. angustifolia has not
been established yet; however, observations of the
flowering cycle of natural stands in Valle del Cauca
and Quindio, Colombia, have been made by tlie first
author every year from 1979 to the present. Not
all the culms in a single clump flowered, and the
flowering culms remain green or yellowish and con¬
tinue to produce new foliage leaves. According to
McClure (1966), a plant of G. angustifolia from
Milagro, Ecuador, established under his direction at
Chocola, Guatemala, and Tingo Maria, Peru, had
flowered annually during the dry season.
Fruit. The fruits of G. chacoensis fall on the
ground, germinate, and grow beneath the parent
plants. Phis suggests that the rate of survival of the
seeds of G. chacoensis is higher than in other species
of the genus, such as G. angustifolia , where it is
comparably low (estimated at less than 1%). In G.
angustifolia , the young ovary is often destroyed by
insects.
Mature fruits of G. chacoensis are usually found
m the middle section of the pseudospikelet and fall
with the rachilla, lemma, and palea attached to the
base. The caryopsis is asymmetrical and flattened
on one side with a short prolongated, hirsutulous
style that persists at the apex (Fig. 2S-2U). The
dorsal surface of the fruit has a conspicuous hilum
about as long as the fruit (Fig. 2U), and the ventral
surface has a prominent embryotegium or covering
at the base (Fig. 2T). A cross section of the fruit of
G. chacoensis reveals the presence of a pericarp,
endosperm, and embryo. The pericarp is thin, brown,
and approximately 0.1 mm thick, whereas the en¬
dosperm is starchy, well developed, creamy-white,
and occupies almost the whole diameter of the fruit
(Fig. 2T). The embryo is V\- X k> the length of the
whole fruit, cream-yellowish, and is located at the
base of the fruit in a lateral position (Fig. 21).
Upon germination of the embryo, the primary
shoot is first to emerge laterally just above and
adjacent to the root, and there is no internode be¬
tween the caryopsis and the coleoptile. The primary
shoot is a segmented axis of clearly positive pho-
totropism and bears a foliar appendage at each node.
The root grows sinuously downward, 12-15 cm,
bearing many secondary adventitious roots through¬
out (Fig. 2W).
The seedling of G. chacoensis consists of, from
the base upward, a coleoptile, the first basal culm
leaf sheath with an elongated internode, the second
basal culm leaf sheath with another elongated in¬
ternode, followed by one culm leaf sheath with a
broad, ovate-lanceolate blade positioned horizontal¬
ly, one or several blades that develop into the first
seedling leaf, and the terminal shoot (Fig. 2V). This
morphological pattern seems to represent the de¬
velopmental sequence of a typical bambusoid seed¬
ling (Calderon & Soderstrom, 1973).
Affinities. Guadua chacoensis appears to be most
closely related to G. trinii, rather than G. angus¬
tifolia. However, in all three species the general
Volume 2, Number 1
1992
Londono & Peterson
Guadua chacoensis
45
Table 1. A selective
morphological comparison
of Guadua chacoensis, G. trinii.
and G. angustifolia.
Character
G. chacoensis
G. trinii
G. angustifolia
Culm leaf
inner ligule
slightly curved and end¬
ing at the margins
more or less horizontal
and ending at the mar-
gins
convex and ending before
the margins
Foliage leaf
blade size (cm)
10-24 x 0.5-2.5
4.5-11 x 0.4-1.3
11-21 x 1-2
l:w ratio
11-26:1
7-12:1
5-18:1
form
linear
linear
linear to lanceolate-ovate
Pseudopetiole
length (mm)
2-3
1-2
2-5
adaxial surface
pilose to glabrous
pilose
short pubescent
abaxial surface
glabrous
pilose
glabrous
Pseudospikelet
size 1 x w (mm)
2-6 x 0.4 0.6
1.8-4 x 0.4-0.6
4-10 x 0.3-0.4
form
robust and erect
robust and erect
slender and acute
Fertile lemma
size 1 x w (mm)
12-16 x 7-12
9-12 x 5-7
9-10 x 5-6
margins
scarious, hairs at the dis
tal end
papery, glabrous distally
papery, glabrous distally
mucro length (mm)
0.8-1
0.5-1
0.5
Palea
size 1 x w (mm)
6-14 x 24
9-11 x 3
7 x 1-1.5
wing width (mm)
1 1.5
0.5
0.5
apex
prolongated
prolongated
not prolongated
Stamens
length (mm)
5-8
5-7
4-5
apex
apiculate
apiculate
obtuse
form of the culm leaves, foliage leaves, and habit
are similar.
Guadua chacoensis and G. trinii share the fol¬
lowing characters: (1) a more or less horizontal junc¬
tion between the culm blade and sheath; (2) a culm
leaf inner ligule ending at the margins (Fig. ID); (3)
erect and robust pseudospikelets (Figs. 1A, 2A); (4)
prolongation of the keels-winged at the apex; and
(5) a monocarpic habit, commonly with a leafless
flowering branch. Guadua chacoensis differs from
G. trinii by having abaxially glabrous pseudope¬
tioles, longer (10-24 cm) leaf blades that are gen¬
erally wider (0.5-2.5 cm), and a lemma that is
longer than the palea (Table 1).
Guadua chacoensis can be distinguished from G.
angustifolia by having: (1) a slightly curved and
continuous inner ligule ending at the margins; (2) a
glabrous leaf sheath (Fig. 1C); (3) a conspicuously
ciliate margin of the inner ligule of the foliage leaf
(Fig. 1C); (4) an adaxially pilose to glabrous pseu¬
dopetiole; (5) erect and robust pseudospikelets; (6)
a scarious margined lemma, with a tuft of hairs at
the apex (Fig. 21); (7) a prolongation of the keeled
wings at the apex of the palea; (8) apiculate apex
of each anther theca (Fig. 2P); (9) an asymmetrical,
fusiform caryopsis (Fig. 2S); and (10) monocarpic,
leafless flowering culms (Table 1). The geographic
distribution of G. chacoensis is completely allopatric
with that of G. angustifolia , which occurs in north¬
western South America and Panama.
Acknowledgments. We are grateful to Jose Cua-
trecasas. Emmet Judziewicz, Lynn Clark, and Lars
Peter Kvist for advice and support during this study.
Special thanks are given to Alice R. Tangerini for
her skillful illustrations. Support for the first author
was provided by a Smithsonian Institution Short-
Term Visitor Grant.
46
Novon
Figure 2. Guadua chacoensis. —A. Pseudospikelet showing prophyll, basal bracts, and florets. —B. Rachilla
segment. —C. Prophyll, dorsal view. —D. Prophyll, ventral view. —E. Basal bract, ventral view. —F. Basal bract,
Volume 2, Number 1
1992
Londono & Peterson
Guadua chacoensis
47
Literature Cited
Calderon, C. E & T. R. Soderstrom. 1973. Morpho¬
logical and anatomical considerations of the grass
subfamily Bambusoideae based on the new genus
Maclurolyra. Smithsonian Contr. Bot. 11: 1-55.
Kunth, C. S. 1822. Voyage aux regions equinoctiales
du Nouveau Continent, fait en 1799 1804, partie
6, Botanique Sect. 5. Synopsis plantarum. [Descrip¬
tion of Guadua angustifolia, 1: 252-253.]
McClure, F. A. 1966. The Bamboos — A Fresh Per¬
spective. Harvard Univ. Press, Cambridge.
-. 1973. Genera of bamboos native to the New
World (Gramineae: Bambusoideae). Smithsonian
Contr. Bot. 9: 1-148.
Nicora, E. G. & Z. E. Rugolo De Agrasar. 1987. Los
generos de gramineas de America Austral. Editorial
Hemisferio Sur S. A., Argentina.
Parodi, L. R. 1955. La floracion de la Tacuara brava
(Guadua trinii). Revista Argent. Agron. 22: 134-
136.
Soderstrom, T. R. & R. P. Ellis. 1987. The position
of bamboo genera and allies in a system of grass
classification. Pp. 225-238 in T. R. Soderstrom, K.
W. Hilu, C. S. Campbell & M. E. Barkworth (editors),
Grass Systematics and Evolution. Smithsonian Insti¬
tution Press, Washington, D.C.
- & X. Londono. 1987. Two new genera of
Brazilian bamboos related to Guadua (Poaceae: Bam¬
busoideae: Bambuseae). Amer. J. Bot. 74: 27-39.
Young, S. M. 1985. The Taxonomy and Natural His¬
tory of the Bambusa guadua Complex (Poaceae:
Bambusoideae). M.Sc. Thesis, University of Florida,
Gainesville.
dorsal view. — G. Lemma with a rudimentary palea below, ventral view. —H. Lemma, dorsal view. —I. Apex of
lemma showing mucro and ciliolate margin. —J. Palea, ventral view. —K. Palea showing the ciliolate winged-keels,
dorsal view. — L. Young palea enclosing androecium and gynoecium. —M. Young androecium and gynoecium
protected by lodicules. —N. Lodicule complement. —O. Androecium with 6 stamens. —P. Anther apex. —Q.
Gynoecium with 3 plumose stigmas. —R. Mature floret enclosing fruit with rachilla attached to the basal point.
S. Fruit with a short stylar column, lateral view. —T. Fruit showing the prominent embryotegium, ventral view. —
U. Fruit showing the hilum, dorsal view. —V. Seedling showing the germinated caryopsis. —W. Detail of the basal
portion of seedling showing the lemma attached to the fruit (1), the coleoptile (cp), the first basal culm-leaf sheath of
the primary shoot (s) and the root (r). Based on: A-Q, Quarin 3210; R-U, Quarin et al. 1859; V, W, Krapovicas
et al. 23851.
Una Nueva Ayenia (Sterculiaceae) con Nectarios Foliares de Costa Rica
Carmen L. Cristobal
Instituto de Botanica del Nordeste, C.C. 209, 3400 Corrientes, Argentina
Nelson Zamora V.
Universidad Nacional de Costa Rica, Escuela de Ciencias Ambientales,
Apdo. 86-3000, Heredia, Costa Rica
Marfa Mercedes Arbo
Instituto de Botanica del Nordeste, C.C. 209, 3400 Corrientes, Argentina
ResuMEN. Se describe y se ilustra Ayenia mas-
tatalensis de Costa Rica. La anatomia de los nec¬
tarios se estudia y se compara con la de las otras
especies del genero con estas glandulas. Se discuten
las relaciones taxonomicas entre las especies de Aye¬
nia con nectarios.
Abstract. Ayenia mastatalensis from Costa Rica
is described and illustrated. The anatomy of its foliar
nectaries is studied and compared with that of the
other species in the genus with these glands. The
taxonomic relationships between the species of Aye¬
nia with nectaries are discussed.
Ayenia mastatalensis Cristobal & Zamora, sp.
nov. TIPO: Costa Rica. San Jose: Zona Protegi-
da La Cangreja, Mastatal de Puriscal, 9°42'N,
84°20'W, 400 m, 4 Dec. 1980, Zamora <&
Jimenez 1312 (holotipo, CR; isotipos, CTES,
F, C, CH, K, MICH, MO, UC). Figuras 1, 2.
Frutex 2-3 m altus, ramis pilis simplicibus vel bifurcatis
parvioribus et pilis setiformibus sparsis. Stipulae persis-
tentes, deltatae, striatae, 2-8 mm longae x 0.7-1.5 mm
latae. Petiolus 4-6 mm longus. Lamina ovalis-lanceolata
ad apicem longe attenuata, 5.7-22 cm longa x 1.5-7.4
cm iata, plerumque integra, subglabra, nectario ovale,
multiaperturato, 2-2.5 mm longo, in vena media hipo-
phylli prope bassim invenitur. Ramuli floriferi 4-11 cm
longi, terminales et axillares ut videtur nudi, 3-4 cymis
umbelliformibus 5-floribus. Petali longe unguiculati, cu-
cullo subtrapeziforme, apice integro; lamina petali linearis
ca. 1.3 mm longa. Androgynophorus 3-4 mm longus.
Stamina 5, antheribus trilocularibus. Staminodia cuccul-
lata margine exteriore acuto. Fructus aculeatus, ca. 11-
12 mm longus x 10 mm latus, coccis dehiscentibus,
monospermis. Semina anguste ovoidea.
Arbusto 2-3 m de alto, ramas delgadas y flexibles,
las terminales con entrenudos de 1-3.5 cm de largo
y 2 3 mm diam., con dirninutos pelos erectos, sim¬
ples o bifurcados y pelos setiformes simples, raro de
mas ramas, dispersos. Estipulas persistentes, coria-
ceas, membranaceas hacia el margen, 2-8 mm de
largo, 0.7-1.5 mm de ancho, deltoideas, a veces
prolongadas en un acumen acicular, cara interna
lisa, cara externa estriada con las venas paralelas y
sobresalientes, base pubescente como el tallo, pelos
glandulares rojizos, muy pequehos y caedizos en el
margen. Peciolo 4-6 mm de largo, 1-1.3 mm diam.,
semiterete, algo abultado y oscuro en la mitad in¬
ferior, pelos pequehos dispersos en la cara interna
sobre la linea media. Lamina membranacea, oval-
lanceolada, subsimetrica, base redondeada, larga-
mente atenuada hacia el apice, 5.7-22 cm de largo,
1.5-7.4 cm de ancho, entera, excepcionalmente 1-
5 dientes en el tercio apical, subglabra, epifilo con
dirninutos pelos simples sobre y a los lados de la
vena media, hipofilo con escasos pelos setiformes y
de dos ramas sobre las venas principales; venacion
sobresaliente en el hipofilo, 8-10 pares de venas
laterales unidas formando arcos cerca del margen;
nectario 2-2.5 mm de largo, 0.7-1 mm de ancho,
ubicado sobre la vena media por encima del primer
par de venas laterales, eliptico, levemente abultado,
glabro, poros poco conspicuos. Ramitas floriferas 4-
Figura 1. Ayenia mastatalensis Cristobal & Zamora (Zamora & Jimenez 1312, CTES). —A. Rama. —B. Flor
completa. —C. Caliz, cara externa. —D. Petalo completo, cara externa. —E. Porcion superior de un petalo, cara
interna. —F. Tubo estaminal. —G. Porcion del tubo estaminal, cara interna. —H. Gineceo. —I. Fragmento de un
coco. —J. Semilla, cara ventral.
Novon 2: 48-52. 1992.
Volume 2, Number 1
1992
Cristobal et al.
Ayenia mastatalensis
49
50
Novon
Figura 2. Ayenia mastatalensis (Zamora Jimenez 1312, CTES). —A. Base de la hoja mostrando el nectario
sobre la vena media. —B. Corte transversal de la vena media a la altura del nectario. —C. Detalle de la epidermis
pluriestratificada y pelos glandulares.
1 1 cm de largo, terminales y axilares, dispuestas
hasta en los 15 nudos apicales, las basales en la
axila de hojas de tamano normal, hacia el apice las
hojas se pueden reducir notablemente hasta 4-6
mm de largo, 1-1.5 mm de ancho; ramitas floriferas
aparentemente desnudas, en cada nudo una hoja
estipulada muy reducida y 3 4 cimas umbeliformes
5-floras, pedunculos 5-10 mm de largo, pedicelos
ca. 4 mm de largo, articulados en la parte media,
glabros o puberulos, bracteas subuladas, verticila-
das, 0.5-1 mm de largo, escasos pelos en la cara
externa y margen; ocasionalmente las ramitas flori-
feras originan en la base ramas de segundo grado
de hasta 4 cm de largo. Alabastros globosos, 2-2.5
mm diam., apice de los sepalos sobresaliente y co-
loreado, cara externa con pelos simples y de dos
ramas, dispersos. Sepalos 5, membranaceos, ver-
dosos, reflexos en la antesis, ca. 3 mm de largo, 1
mm de ancho, oval-lanceolados, agudos, apice rojizo
y algo engrosado, dos pares opuestos unidos hasta
la mitad y el resto de los lados solo en la base, cara
externa con pelos simples y bifurcados largos dis¬
persos, cara interna y acumen con pelos glandulares
breves. Petalos 5, rosado-lilas, largamente ungui-
culados, porcion inferior de la una filiforme, ca. 6
mm de largo, glabra, describiendo un semiaro; por¬
cion superior subtrapezoidal, ca. 2 mm de largo, 2.5
mm de ancho, adosada al horde del tubo estaminal,
membranacea, margen inferior con los lobulos re-
dondeados y minusculos pelos glandulares en el mar¬
gen, apice entero, redondeado, con dos dientes car-
nosos en la cara interna que abrazan la porcion libre
del fllamento estaminal, el resto de la cara liso;
lamina de los petalos reducida a un apendice algo
Volume 2, Number 1
1992
Cristobal et al.
Ayenia mastatalensis
51
adelgazado en el apice, ca. 1.3 mm de largo, inserto
en la mitad superior de la cara externa, glabro.
Androginoforo 3-4 mm de largo. Tubo estaminal
campanulado, 1 mm de largo; estambres 5, oposi-
tipetalos, anteras tritecas en el extremo de una breve
porcion libre de filamento tan larga como la mitad
del tubo estaminal; estaminodios 5, lobulos algo car-
nosos e inflados, cuculados, perpendiculares al eje
de la flor, apice rodeando al estilo, margen externo
agudo. Ovario globoso, cubierto de mamilas; estilo
simple, breve; estigma capitado con 5 lobulos in-
conspicuos. Capsula 5-coca, sublenosa, ca. 11-12
mm de largo, 10 mm de ancho, pedunculo y pedicelo
ca. 9 mm de largo, cocos monospermos, dehiscentes
a lo largo de la sutura ventral y Vs de la vena media,
mitades incurvas y separadas entre si, cara interna
lustrosa con algunos pelos glandulares dispersos, en
la cara externa aculeolos de 0.5-1.5 mm de largo
± conicos, agudos, finamente pubescentes, caedizos
con una porcion de la capa superficial del pericarpo.
Semillas castaiias, angostamente ovoides, 8 mm de
largo, 3.5 mm de ancho, caras laterales planas, algo
reticuladas, cara dorsal convexa, lisa.
Anatomia de los nectarios foliares. En cortes
transversales de hoja se observa que la vena media
esta ligeramente ensanchada a la altura del nectario
(Fig. 2B).
El nectario es multiaperturado (Arbo, 1972) y
consiste en un cojin de epidermis pluriestratificada
de 5-6 capas de espesor, en el que se encuentran
hundidos los pelos glandulares. Las celulas epider-
micas presentan paredes engrosadas, y algunas de
ellas contienen cristales prismaticos (Fig. 2C); ex-
teriormente estan recubiertas por una cuticula gruesa,
que tapiza ademas las cavidades en las que se en¬
cuentran los pelos glandulares. A veces se encuen¬
tran dos pelos juntos en una misma cavidad. Los
pelos glandulares estan formados por una celula
basal voluminosa, una celula del cuello con engro-
samientos perifericos de cutina en las caras tangen-
ciales, y una cabeza aproximadamente conica for-
mada por 3-5 capas de celulas pequenas, con
contenido denso, granuloso. El parenquima subepi-
dermico presenta 7-8 capas de celulas con paredes
delgadas, muchas de ellas con cristales prismaticos.
Se encuentran algunas bolsas de mucilago entre el
haz vascular y el nectario, y numerosas a los cos-
tados.
Los nectarios de Ayenia mastatalensis son muy
similares a los nectarios de A. klugii Cristobal &
Arbo, aunque con menor cantidad de bolsas de mu¬
cilago en el parenquima subglandular. En las otras
especies de Ayenia L. con nectarios con pelos glan¬
dulares, estos son menos diferenciados, presentan
menor numero de capas de epidermis pluriestrati¬
ficada, y las cavidades donde se encuentran los pelos
son cilindricas o infundibuliformes. Tambien la for¬
ma de los pelos secretores es menos diferenciada en
estas especies, pues la cabeza es cilindrica o abultada
en forma de clava, semejante a la de los pelos
glandulares que se encuentran en otros organos
(Cristobal & Arbo, 1971).
Paratipos. Costa Rica, san jos£: Zona Protegida La
Cangreja, Mastatal de Puriscal, 9°42'N, 84°20'W, 400
m, 7 Nov. 1987 (fl), Zamora, Chacon & Solis 1475 (C,
COL, CR, CTES, MICH, MO, NY, SI, VEN), 2 July
1986 (fr), Zamora, Sanchez & Berrocal 1171 (CR,
CTES, MO).
El hallazgo de Ayenia mastatalensis es espe-
cialmente interesante ya que eleva a seis el numero
de especies del singular grupo de Ayenias con nec¬
tarios foliares. Dichas glandulas son caracteristicas
de Byttneria Loefl. (Cristobal, 1976) y Rayleya
Cristobal (Cristobal, 1981; Arbo, 1981), los dos
generos mas relacionados con Ayenia L., de los
cuales se diferencia, entre otros caracteres, por tener
anteras tritecas.
Ayenia seccion Ayenia se divide en cuatro sub-
secciones. Las especies de la subseccion tipo no
presentan nectarios, y las que los poseen estan se¬
paradas en las tres subsecciones restantes, teniendo
en cuenta principalmente los petalos y los estami¬
nodios (Cristobal & Arbo, 1971).
Ayenia mastatalensis comparte la subseccion
Stipulares Cristobal (Cristobal, 1960), con las es¬
pecies colombianas A. cuatrecasae Cristobal (Cris¬
tobal, 1962) y A. stipularis Triana & Planchon,
por tener los estaminodios con el margen externo
agudo y el apice de la una de los petalos entero,
mientras que en el resto de la seccion Ayenia es
escotado.
Las flores de A. mastatalensis son bastante se-
mejantes a las de A. cuatrecasae, pero esta especie
es pauciflora y sus hojas son aserradas desde la base
y pubescentes. Pasa lo contrario si la comparamos
con A. stipularis, ya que ambas tienen un aspecto
general semejante dado por las hojas y la inflores-
cencia terminal. Las diferencias se encuentran en
las flores: en A. stipularis, la porcion superior de
la una de los petalos es redondeada, con los lobulos
convergentes; la cara interna posee, debajo de los
dientes, un area circular, levemente concava y papi-
losa donde se acomoda el estambre; la lamina del
petalo esta ausente; el margen externo de los es¬
taminodios es largamente acuminado, y la porcion
libre del filamento estaminal es casi tan larga como
el tubo.
Con el fin de facilitar la comparacion de Ayenia
mastatalensis con las otras dos especies de la sub-
52
Novon
seccion Stipulares, se resumen las diferencias en
ima clave <]ue complementa la publicada para Aye-
nias con nectarios foliares (Cristobal & Arbo, 1971).
la. Petalos con la lamina reducida a un apendice
inserto en el dorso de la region trapezoidal o
subtrapezoidal de la una.
2a. Lamina foliar aserrada desde la base, pu-
bescente; inflorescencias axilares, pauci-
floras . 4. cuatrecasae
2b. Lamina foliar entera, excepcionalmente con
1-5 dientes en el tercio apical, subglabra;
ramas flonferas terminates y axilares, apa-
rentemente desnudas, multifloras .
. A. mastatalensis
lb. Petalos sin lamina; porcion superior de la una
redondeada; inflorescencia terminal, desnuda,
multiflora; hojas glabras, enteras o con dientes
en el tercio apical . 4. stipularis
A medida que aumenta el conocimiento de las Aye-
nias con nectarios foliares parece aumentar tam-
bien la coherencia entre ellas. Por ejemplo, se pre-
sentan semejanzas de A. mastatalensis con A.
praeclara Sandwith y con A. klugii por los frutos
y los nectarios respectivamente, aun cuando las dos
ultimas especies corresponden por caracteres llora-
les a la subseccion Praeclarae Cristobal.
De las seis especies con nectarios, solo se conocen
los frutos de A. mastatalensis y de A. praeclara
(Peru, Pasco, Oxaparnpa, 24 Sep. 1982, Poster
89 15, CTES). En ambas los frutos poseen aculeolos
aproxiinadamente conicos, a diferencia de los lrutos
de la seccion Ayenia subseccion Ayenia, donde son
lineares. Los aculeolos de las dos especies antes
mencionadas se aproximan por su forma a los de la
seccion Cybiostigma (Turcz.) Griseb., o mas aun a
los del genero Byttneria. Los frutos de A. praeclara,
especialmente, se parecen a los de este genero por
el tamano de los cocos, hasta 2 cm de largo, la
consistencia leiiosa y los aculeolos punzantes. En
cuanto a las semillas, en ambas especies son casi
lisas, muy diferentes a las del resto de la seccion
Ayenia donde casi sin excepcion poseen costillas.
Desde el punto de vista geografico, la nueva es-
pecie aqui presentada parece ser, como sus afines,
una rareza de area restringida. Vive en una mancha
de bosque tropical muy humedo, rica por su diver-
sidad floristica, y cuya exploracion ha dado varias
especies arboreas nuevas para la ciencia. La pre-
sencia de esta Ayenia en Costa Rica marca un
lugar intermedio entre Tamaulipas, Mexico, locali-
dad donde se encontro A. conciliata Cristobal sub¬
seccion Conciliatae Cristobal (Cristobal, 1969) y las
areas de las cuatro especies sudamericanas.
Como conclusion, debe dejarse abierta la posibi-
lidad de unificar taxonomicamente estas Ayenias,
jerarquizando la presencia del nectario foliar y el
gran porte. El grupo podria constituir una seccion
independiente, vinculo entre Ayenia y Byttneria ,
con especies altamente diferenciadas, que responden
a un esquema de variacion muy diferente al de la
subseccion Ayenia.
Los detalles de las Figuras 1 y 2 fueron dibujados
con camara clara; piezas florales y nectario por C.
L. Cristobal, cortes anatomicos por M. M. Arbo,
rama y pasado a tinta por Laura Simon.
Literatura Citada
Arbo, M. M. 1972. Estructura y ontogenia de los nec¬
tarios foliares del genero Byttneria (Sterculiaceae).
Darwiniana 17: 104-158.
-. 1981. Anatomia de tallo y hoja de Hayleya
bahiensis Cristobal (Sterculiaceae). Bonplandia 5(9):
51-62.
Cristobal, C. L. 1960. Revision del genero Ayenia L.
(Sterculiaceae). Opera Lilloana 4: 1-230.
-. 1962. Notas sobre el genero Ayenia. Lilloa
31: 207-212.
-. 1969. Una nueva e interesante especie de
Ayenia (Sterculiaceae) de Mexico. Bonplandia 3(4):
49-52.
-. 1976. Estudio taxonomico del genero Bytt¬
neria Loefl. (Sterculiaceae). Bonplandia 4: 1-428.
-. 1981. Rayleya, nueva Sterculiaceae de Ba¬
hia, Brasil. Bonplandia 5(8): 43-50.
- & M. M. Arbo. 1971. Sobre las especies de
Ayenia (Sterculiaceae) con nectarios foliares. Dar¬
winiana 16(3-4): 603-612.
New Taxa of Lonchocarpus (Fabaceae: Milletieae) from the
Venezuelan Guayana
Hans-Helmut Poppendieck
Institut fur Allgemeine Botanik und Botanischer Garten,
Ohnhorststrasse 18, D-2000 Hamburg 52, Germany
ABSTRACT. Prior to the treatment of Lonchocar¬
pus for the Flora of the Venezuelan Guyana , two
new species endemic to the northern part of Estado
Bolivar, Venezuela, and partly extending into ad¬
jacent Territorio Federal Delta Amacuro, are de¬
scribed: Lonchocarpus crassispermus, related to L.
hedyosmus, and Lonchocarpus imatacensis from
the subgenus IXeuroscapha. In addition, Loncho¬
carpus tubicalyx is validated.
Lonchocarpus crassispermus Poppendieck, sp.
nov. TYPE: Venezuela. Bolivar: 17 km de Upa-
ta sobre la carretera nueva Upata-San Felix,
en las cercanias de los campamentos “El Par-
aiso” y “La Yagua,” 24-28 km NE del caserio
Los Rosos, “mahomo chino,” Nov. 1965 (fr),
C. Blanco 382 (holotype, VEN; isotypes, MO,
NY—2 sheets, US—2 sheets). Figure 1.
Arbor parva. Folia 9-foliolata, foliolis anguste ovatis,
rugosis, marginibus distincte involutis, costa, marginibus
et infra venis lateralibus pilosis. Inflorescentia terminalis,
floribus geminatis. Corolla roseo-purpurea, epunctata.
Vexillum basi distincte callosum. Ovula 4. Pedunculus
fructus articulatus. Fructus 1-2-seminalis, indehiscens,
aliquantum lignosus, lanceolatus ad oblongus, planus sed
conspicue incrassatus circum semina, 7.5-12 cm longus.
Semina reniformia 7-9 mm crassa.
Small evergreen tree. Leaves 9-foliolate; stipules
caducous; petiole with rachis 7-9 cm long, pubes¬
cent. Leaflets firmly membranaceous, epunctate,
narrowly ovate, rugose, margin distinctly involute,
3.5-7 x 1.3-2.3 cm, pilose on margin, central vein
above, and on principal veins beneath. Inflores¬
cences terminal on leafy shoots, strigose, spikes 8.5-
17 cm long, peduncle 2.5-5 cm long. Flowers gem¬
inate; floral peduncle ± persistent after anthesis,
2.8-3 mm long; pedicel 2-3 mm long, with cadu¬
cous subulate bracteoles above the midpoint. Calyx
zygomorphic, 5-dentate with 3 triangular carinal and
2 obtuse vexillar teeth, punctate, strigose outside,
glabrous within, 4-5 mm long. Corolla rose-purple,
wing and standard petal strigose outside, wing and
rarely keel petal sparsely dotted. Standard reflexed,
clawed, lamina ovate to orbicular with prominent
basal callosities, 8x8 mm. Staminal tube straight,
filaments connate for % to ?4, anthers glabrous.
Ovary densely sericeous, ovules 4. Fruit on jointed
peduncle, 1-2-seeded, indehiscent, slightly woody,
flat but abruptly and conspicuously thickened at the
seed region, lanceolate to oblong, base attenuate,
apex rounded and mucronulate, vexillar margin in¬
conspicuously keeled, 7.5-12 cm long and 3.4-4
cm wide, at seed region 0.8-1 cm thick, otherwise
0.5-0.6 mm thick, ochre, slightly tomentose. Seeds
thick, kidney-shaped, 1.1-1.5 cm long and 7-9 mm
diam.
Distribution and phenology. Endemic to the
northeastern part of Estado Bolivar, Venezuela, in
moist to dry forests, 200-300 m; flowering in July
and August, ripe fruits produced in November. Com¬
mon names: Jebe, Mahomo chino.
Additional specimens examined. VENEZUELA. BOL-
IvAR: 6 km S de El Peru, 15 km SE de El Callao, 300
m, 61°48'W, 7°18'N, 3 July 1960 (fl, lvs), Little 17531
(MER); 17 km de Upata sobre la carretera nueva Upata-
San Felix, en las cercanias de los campamentos “El Par-
aiso” y “La Yagua,” 24-28 km NE del caserio Los Rosos
(type locality), Aug. 1964 (fl, lvs), Blanco 226 (NY, US,
VEN); 7 km N de San Pedro de las Bocas, 200 m,
62°54'W, 7°00'N, “mahomo chino,” 27 July 1978 (fl),
Liesner & Gonzalez 5750 (MO); entre Encrucijada y El
Pao, 62°40'W, 8°03'N, 28 Aug. 1954 (fl, fr), Trujillo
247 (MER, S).
The small rugose leaflets, in combination with the
large fruits and the nearly spherical seed, make this
species unique. Flowers and inflorescences, however,
agree well with both Lonchocarpus hedyosmus Miq.,
which extends from the Estado Guarico in Venezuela
to the Guianas, and with the predominantly Colom¬
bian L. macrocarpus Benth., which has its eastern
limit of distribution in Estados Zulia and Merida.
Lonchocarpus imatacensis Poppendieck, sp. nov.
TYPE: Venezuela. Territorio Federal Delta
Amacuro: E de Rio Grande, NE de El Palmar,
cerca de los limites del Estado Bolivar, “Jebe,”
13 Jan. 1965, Marcano-Berti 560 (holotype,
VEN; isotypes, MO, NY, VEN). Figure 2.
Arbor magna sempervirens. Folia 5-foliolata. Inflores¬
centia terminalis, floribus geminatis. Corolla roseo-pur-
Novon 2: 53-57. 1992.
54
Novon
Figure 1. Lonchocarpus crassispermus Poppendieck. —A. Flowering branch. —B. Fruits. —C. Fruit, side view.
— D. Leaf base, from below. —E. Flower, not yet expanded, indumentum left out to show glands on calyx. —F I.
Standard, floral buds, keel, and wing petal. (A from Blanco 22b\ B, C from isotype at NY; D-J from holotype.)
Volume 2, Number 1
1992
Poppendieck
Lonchocarpus from Venezuelan Guayana
55
A,B-5cm
D = 5mm
C,E-H-8mm
Figure 2. Lonchocarpus imatacensis Poppendieck. —A. Flowering branch. —B. Branch with mature fruits. —
C. Details of inflorescence with flower buds. —D. Flower from below, to show indumentum and position of glands on
keel petal; indumentum on calyx left out, as in E and F. —E. Flower, side view. —F. Flower, corolla removed. —
G. Keel and wing petal, natural position. —H. Standard. (A from Blanco 146, after a drawing by B. Manara; B
from isotype at NY, C H from holotype.)
56
Novon
F igure 3. Lonchocarpus tubicalyx Pittier ex Poppendieck. —A. Branch, inflorescences, and new leaf. —B. Mature
leaf. —C. Detail of inflorescence with flower buds. —D. Flower, indumentum on calyx left out to show lactifers. —
E. Flower, standard petal removed. —F. Flower section of lower part. G-I. Standard, wing, and keel petal. —J.
Staminal tube expanded. —K. Calyx expanded. (All from the holotype.)
purea, epunctata praeter 1-2 glandulas apice petalorum
carinaliuin. Vexillum basi distincte callosum. Ovula 8-
10. Pedunculus fructus articulatus. Fructus 1-7-semin-
alis, indehiscens, anguste oblongus ad lanceolatus, planus
sed bullatus circum seinina, (5.5-)l 1-14 cm longus. Sem¬
ina reniformia, castanea, 1.8 mm crassa.
Large evergreen tree up to 26 m tall, trunk
diameter up to 40 cm. Leaves 5-foliolate; stipules
caducous; petiole with rachis 3.5 6.5 cm long. Leaf¬
lets membranaceous, not pellucid-punctate, ovate
to obovate, base cuneate, apex bluntly acuminate,
(2.5-)6-8(-9.5) x 2.5 5 cm, glabrous, veins prom¬
inent above. Inflorescences terminal on leafy shoots,
glabrous, 9-14 cm long, peduncle 2.2-2.8 cm long.
Flowers geminate; floral peduncle ± persistent alter
anthesis, 1.2-1.8 mm long; pedicel 1.3-1.7 mm
long, with ovate, punctate bracteoles at the base of
the calyx. Calyx cyathiform, truncate at base.
epunctate, strigose outside, glabrous within, 3 mm
long, 5 mm wide. Corolla rose-purple, strigose out¬
side, glabrous within, epunctate except for one or
two large glands on the abaxial side of the keel petal.
Standard reflexed, clawed, lamina orbicular to
subquadrate, apically sinuate to of its length, with
prominent basal callosities, 7-8 mm long and 3.2-
3.5 mm wide. Staminal tube ± straight, somewhat
persistent after anthesis, filaments connate for % to
44, anthers glabrous. Ovary densely sericeous, 8-
10- ovulate. Fruit on jointed peduncle, 1-7-seeded,
indehiscent, flat but slightly bullate around the seeds,
narrowly oblong to lanceolate, base stipitate, cu¬
neate, apex rounded, obliquely acuminate, (5.5-)
11- 14 cm long and 1.3-1.8 cm wide, thickness
at seed region 3.3-3.5 mm, dorsally 0.9-1.1 mm
and ventrally 0.4 mm, dark ochre to castaneous,
glabrous. Seed kidney-shaped, castaneous, 9x7
mm, 1.8 mm thick.
Volume 2, Number 1
1992
Poppendieck
Lonchocarpus from Venezuelan Guayana
57
Distribution and phenology. Endemic to north¬
ern Estado Bolivar and the border with Territorio
Federal Delta Amacuro, Venezuela; rainforests, 50-
350 rn; flowering May to July, fruits collected from
November to January.
Additional specimens examined. Venezuela, bol-
fvAR: Altiplanicie de Nuria, on ± level forest on trail
between El Cruzero and slightly beyond pica 101, ESE
of Villa Lola, Jan. 1960 (lvs, fl), Steyermark 86361 (NY,
US, VEN); Sierra Imataca, Rio Toro (Rio Grande), be¬
tween Rio La Reforma and Puerto Rico, N of El Palmar,
14 Dec. 1960 (fr, lvs), Steyermark 88036 (NY, US);
alrededores del campamento “Las Pavas,” arriba del Salto
Para, 6°12'N, 64°15'W, 22 May 1978 (fl, lvs), Stey¬
ermark et al. 117181 (VEN). territorio federal delta
AMACURO: NE de El Palmar, E de Rio Grande, cerca de
los limites del Estado Bolivar (type locality), June 1965
(fl, lvs), Blanco 146 (MO, NY, VEN), 24 May 1964 (fl,
lvs), Marcano-Berti 194 (MO, NY, VEN), 26 May 1964
(fl, lvs), Marcano-Berti 197 (MO, NY, VEN); Sierra
Imataca, Cerro de La Paloma, E side of Rio Cuyubini,
vicinity of large granitic boulders, 18 Nov. 1960 (lvs, fr),
Steyermark 87647 (NY).
This species is peculiar because of the fruit, which
is bullate around the seeds, which distinguishes it
from the southern Brazilian Lonchocarpus nitidus
Bentham, with which it has been misidentified. It
belongs to subgenus Neuroscapha Pittier (= Neu-
roscaphi Bentham), the South American species of
which form a series of closely related allopatric spe¬
cies whose ranges sometimes partly merge.
Lonchocarpus tubicalyx Pittier ex Poppendieck.
TYPE: Venezuela. Apure: San Juan de Payara,
arbusto de 4-5 m, 1 1 Feb. 1941 (fl, lvs). Char-
don 237 (holotype, VEN). Figure 3.
Arbor parva. Folia decidua, 7-11-foliolata, foliolis ova-
tis acuminatis, iuventute discoloribus: supra fuscatis, infra
griseis strigosis glabrescentibus. Florescentia coaetanea,
inflorescentia lateralis, floribus in verticillastris paucifloris
dispositis. Calyx basin leviter inflatus, basin maculis lac-
tiferibus punctatis. Corolla roseo-purpurea. Vexillum basin
attenuatum, ecallosum. Ovula 5. Fructus ignotus.
Small deciduous tree 4-5 m tall. Bark gray. Leaves
7-11-foliolate; stipules caducous; petiole with rachis
10-15 cm long. Leaflets membranaceous, epunc-
tate, ovate, apex bluntly acuminate, 4.5-8 x 2.2-
3.5 cm, glabrous above, strigose to subglabrous be¬
neath, young leaflets discolorous with blackish upper
and grayish lower side. Flowering coaetaneous. In¬
florescences lateral, in axis of old leaves, composed
of 2-3 spikes on short shoots 1.5-2.5 cm long.
Spikes 7.5-11.5 cm long with a sterile basal part
1.5-2.3 cm long. Peduncle reduced, flowers spaced
so as to appear verticillate, pedicel 1.3 mm long,
bracts 0.5-1 mm long, rounded to ovate. Calyx
campanulate, slightly zygomorphic with upper mar¬
gin undulate, lower margin 3-dentate and upper side
slightly inflated, strigose outside, glabrous within,
with prominent latex ducts especially at the base,
4-5 mm long and 3.5-6 mm wide at anthesis.
Corolla purple, only slightly and sporadically pu¬
bescent. Standard orbicular, attenuate at base, with¬
out basal callosities, 5.5-7.5 mm long and 6,5-9
mm wide. Staminal tube 8-11 mm long, straight,
filaments connate for V\ , base broadened, with two
long dorsal slits. Ovary apically slightly bent, gla¬
brous in the apical third of its length, otherwise
strigose, ovules 5. Fruits unknown.
Distribution and phenology. Presently known
from only two collections in Apure and Delta Ama¬
curo, Venezuela, flowering in February.
Additional specimens examined. Venezuela, terri¬
torio federal delta amacuro: Rio Manimo, Vuelta Triste,
20 Feb. 1911 (fl, lvs), Bond et al. 145 (GH, NY, US).
Since Pittier (1944: 102) lacked a Latin diag¬
nosis, it is necessary to validate this name, but be¬
cause of the lack of fruits some uncertainties remain.
Lonchocarpus tubicalyx belongs to an aggregate
of allopatric species with coaetaneous inflorescences,
discolorous juvenile leaves, and purple flowers that
usually can easily be differentiated by their fruits.
The name is here applied to two widely separated
collections, which are unique because of the very
short pedicels by which the flowers appear to be
arranged in verticillasters. This, and the somewhat
smaller flowers, distinguish them from the Brazilian
L. denudatus Bentham. Its Venezuelan and partly
Colombian relatives L. crucisrubierae Pittier and L.
fendleri Bentham (= L. sanctae-marthae Pittier)
have obtuse leaves instead of acuminate ones. The
two cited collections are almost identical except for
slight differences in the size of the flowers, those
from Delta Amacuro being at the upper and those
from Apure being at the lower range of the mea¬
surements given above.
Literature Cited
Pittier, H. 1944. Leguminosas de Venezuela, vol. I.
Bob Tec. Min. Agric. y Cria, Caracas 5: 1-171.
Cryosophila macrocarpa (Palmae), a New Species from
Choco Department, Colombia
Randall J. Evans
University of Michigan Herbarium, Ann Arbor, Michigan 48109-1057, U.S.A.
ABSTRACT. A disjunct population of Cryosophila
recently discovered in the Colombian Choco is mor¬
phologically quite distinct from previously described
Cryosophila species. Unique to the genus are its
large Iruits with large, deeply furrowed seeds. This
new Cryosophila, C. macrocarpa, appears to be
most closely allied to C. guagara of the Golfo Dulce
area of Costa Rica and adjacent Panama.
Cryosophila Blume comprises approximately 11-
1 3 mostly narrowly endemic species of understory,
primarily lowland wet forest palms distributed from
Pacific coastal and southern Mexico to northwestern
Colombia. The most distinctive generic character¬
istic is the usually branched, often numerous and
long spines derived from roots that grow out of the
normally solitary stem. Although the generic limits
of Cryosophila are well defined, its species are poor¬
ly delineated. The difficulty arises because there are
few characters that differentiate species. Most char¬
acters are either invariant (e.g., most floral char¬
acters) or continuously variable (e.g., the pattern
and degree of stem armature and leaf blade splitting)
across the genus. The most important diagnostic
features are inflorescence shape and structure. Few
species can be identified sterile, even in the field.
G. Galeano and R. G. Bernal (both of COL) re¬
cently discovered a disjunct Pacific coastal Cryoso¬
phila population (sterile at the time) near the town
of El Valle in the Department of Choco. While in
Colombia conducting field studies as part of a mono¬
graph of the genus, I collected fertile material from
this population possessing characteristics that clearly
differentiate it from previously described species of
Cryosophila.
Cryosophila macrocarpa R. Evans, sp. nov.
TYPE: Colombia. Choco: ca. 300 m S of Bahia
Solano-El Valle road ca. 2 km from El Valle
(ca. 500 m E of the bridge over Quebrada
Tundo), elev. ca. 20 m, 6°07'30"N, 77°25'W,
6 Mar. 1991, R. J. Evans Ac J. G. Ramirez
213 (holotype, COL; isotypes, JAUM, MICH,
MO). Figures 1, 2.
A congeneribus fructibus majoribus (3.1 —3.6( 3.7 in
statu vivo) cm longis, 2.6-2.9(-3.0 in statu vivo) cm latis)
et seminibus majoribus (2.5-2.7 cm longis, 2.1-2.3 cm
latis) testis profunde sulcatis differt.
Solitary palm. Stem ca. 5 m long, erect or briefly
decumbent, ca. 12 cm DBH, armed with root-spines
distributed uniformly or not along length of stem;
internodes not discernible. Root-spines 0-ca. 100
per 10 cm of stem, most to 18 cm (average 6-8
cm) long, occasional outliers to 55 cm long, almost
always 1-2-times branched, usually descending,
growing into soil at the stem base. Leaves ca. 30
in a spreading crown, ca. 6 distal to youngest inflo¬
rescence, 7-16 dead and pendulous; petiole rounded
abaxially, channeled adaxially, 2.80-3.05 m long,
2.38-2.57 cm wide (fresh) at narrowest point just
proximal to blade, splitting basally with age; basal
sheath cream-colored, densely floccose, disintegrat¬
ing and fraying into elongate fibers over time; hastula
elevated, blunt, very depressed-triangular, 0.1-0.7
cm long, 2.9-3.8 cm wide, dorsally plicate and
carinate; blade induplicately palmate, very broadly
ovate in outline, 0.200 0.225 mm thick (fresh)
adjacent to central margin 10 cm from apex of most
recently fully expanded leaf, adaxial surface gla¬
brous, abaxial grayish pubescent; central segments
126.5-158.0 cm long; lateral segments 85.0-1 10.5
cm long, ca. as long as central segments;
central abaxial split to within (2.5-)4.0-6.0 cm of
base, dividing blade into two ± equal halves of 29-
32 segments each; primary adaxial splits less deep
than central abaxial split with depth of splitting
increasing from inner (ca. to base) to outer
splits, dividing each half into 6-7 sections of 2-10
segments each, with the central sections containing
(5-)8-10 segments each, the middle 3-4 sections
of each half containing (3-)4-5(-6) segments each,
and 2 the lateral-most sections of each half con¬
taining (2-)3-4(-5) segments each; secondary ad¬
axial splits y*- l A(-W) to base; inner ca. 4 sections
of each half fully divided; lateral section undivided;
segments widest (4.1-4.6 cm wide ca. 40 cm from
apex) near central abaxial split, narrowing toward
the lateral segment, sometimes with slight constric-
Novon 2: 58-61. 1992.
Volume 2, Number 1
1992
Evans
Cryosophila macrocarpa
59
Figures 1, 2. Cryosophila macrocarpa R. Evans (Evans & Ramirez 213). — 1. Inflorescence with aborted flowers.
— 2. Infructescence with mature fruit. Scale bar = 20 cm.
tion distally, briefly bifid and often sub-sickle-shaped
and diverging apically; longitudinal veins 10-14 per
half-segment, 0.9-3.3 mm apart at widest portion
(ca. mid-segment) of widest segments; longitudinal
veinlets 3-7 between adjacent veins, 0.2-0.6 mm
apart; irregular transverse commissures connecting
longitudinal veins abundant and conspicuous abax-
ially and adaxially. Inflorescences with 3 orders of
branching, interfoliar, ascending at emergence
through split petiole base, then abruptly deflected;
primary axis 62-78 cm long, cream-colored, dense¬
ly floccose, 1.15-1.29 cm diam. at juncture of
peduncle and rachis; prophyll lanceolate with trun¬
cate, sheathing base and acuminate apex, bicarinate,
23.5-25.5 cm long, persistent; inflorescence bracts
with tubular base and cochleariform, apically acu¬
minate blade, cream-colored, densely floccose, the
2 basal-most bracts ovate when detached and flat¬
tened, ca. 31 cm long, the bracts progressively
smaller, particularly in width, toward apex, apical
bracts very narrowly obovate; peduncle 50.0-57.5
cm long; peduncular bracts 4, persistent; peduncular
bract scars slightly oblique; rachis 12.0 20.5 cm
long; first-order branches ca. 10, broadly flattened
perpendicular to primary axis, irregularly divided,
to 7.5 cm long near base of rachis, shortening toward
apex, each subtended by a deciduous bract; rachis
bracts caducous apically, progressively more per¬
sistent toward base of rachis; rachis bract scars
strongly oblique; rachillae to 5 cm long near base
of rachis, shortening toward apex; rachillae brac-
teoles sheathing basally, irregularly shaped, 4.5-
8.5 mm long, 0.5-1.5 mm wide just distal to basal
sheath. Flowers perfect, 4.9-5.4 mm long, 3.8-4.8
mm diam., borne singly on pedicels 1.0-1.5 mm
long, each subtended by a narrowly triangular to
triangular bracteole 1.0-1.4 mm long, 0.4-0.7 wide;
receptacle 0.6-0.8 mm long; sepals 3, acute, 4.3-
5.3 mm long, connate basally ca. l A~ l A their length;
petals 3, cochleariform, thickened at center, hyaline
at margin, stipitate, 2.6-3.1 mm long, 3.4-4.4 mm
wide and flabelliform when detached and flattened,
imbricate, adnate to calyx basally; stamens 6; fila¬
ments flat, 2.8-3.3 mm long, connate nearly to
their apices in a hyaline, membranous, narrowly
ampulliform tube 1.4-1.8 mm diam.; anthers briefly
bifid at base and apex, 1.6-2.0 mm long, 0.6-0.9
mm wide, cream-colored, dehiscent laterally by lon¬
gitudinal slits; carpels 3; ovary ca. 1 mm long, 0.5-
0.7 mm diam.; stigma + style 1.9-2.8 mm long;
stigma slightly expanded. Fruit oval, 3.1-3.7 cm
long, 2.6-3.0 cm diam., cream-colored at maturity,
60
Novon
Figure 3. Cryosophila fruits (upper row) and seeds. Left to right: C. macrocarpa (Evans & Ramirez 213), note
deeply sulcate testa; C. warscewiczii (B. Jacobs 2351), the second-largest-fruited Cryosophila; C. guagara (Evans
207), note shallow sulci; C. kalbreyeri (Ramirez <£- Echavarria 3580), the only other previously described Colombian
Cryosophila. Scale bar = 2 cin.
epicarp smooth, mesocarp slightly fleshy, endocarp
membranous; seed oval, 2.5-2.7 cm long, 2.1-2.3
cm diain., not adherent to the endocarp, the testa
deeply sulcate.
Common name: Noli, but this name is also used
in the area for Chelyocarpus dianeurus (Burret) H.
E. Moore, a much more common palmate-leaved
palm. Uses: leaves used as thatch.
Additional specimen examined. Same population and
date as holotype, R. J. Evans & J. G. Ramirez 214 (BH,
MICH, NY, US).
Known only from the type locality, an area of
slightly disturbed lowland wet forest in the flood plain
of the Rio Valle near the northern extremities of
the Golfo de Tribuga. Extensive primary forest re¬
mains in the immediate area as well as most of the
entire Golfo de Tribuga coastal region, due to its
remoteness. The area is very poorly known botan-
ically and therefore Cryosophila macrocarpa may
occur throughout the region. However, within its
known bottomland habitat C. macrocarpa is very
rare and localized, apparently occurring only in scat¬
tered small populations. Cryosophila macrocarpa
does not occur on the nearby slopes where Chely¬
ocarpus dianeurus is very common (nor does Chel¬
yocarpus dianeurus occur in the bottomlands with
Cryosophila macrocarpa). One local resident,
knowledgeable enough to distinguish between these
two superficially similar palmate-leaved species, re¬
ported C. macrocarpa to be more common on Cabo
Gorrientes than in the El Valle area. Cabo Corrientes
delimits the southern end of the Golfo de Tribuga
about 70 km to the south.
The large fruits and seeds with deeply sulcate
seed coats of Cryosophila macrocarpa readily dif¬
ferentiate it from all other Cryosophila species (Fig.
3).
Cryosophila macrocarpa is morphologically most
similar to Cryosophila guagara P. Allen of the
Golfo Dulce and adjoining areas of southwestern
Costa Rica and adjacent Panama. Character states
shared by C. macrocarpa and C. guagara are abun¬
dant leaf blade cross-veins that are conspicuous both
Volume 2, Number 1
1992
Evans
Cryosophila macrocarpa
61
adaxially and abaxially; short, broad, and flattened
first-order inflorescence branches; and filaments that
are connate nearly to their apices. Cryosophila ma¬
crocarpa inflorescences differ from those of C. gua-
gara by their fewer peduncular bracts (4 in C.
macrocarpa vs. 7-10 in C. guagara) and their
comparatively shorter rachises containing fewer first-
order branches (rachis Vs—Vi the total inflorescence
length containing ca. 10 first-order branches in C.
macrocarpa vs. rachis V 3 -V 3 the total inflorescence
length containing ca. 30 first-order branches in C.
guagara).
Whether or not the phenetic similarity between
C. macrocarpa and C. guagara represents a close
phyletic relationship should be resolved by a phy¬
logenetic analysis of Cryosophila species (Evans, in
prep.). If C. macrocarpa and C. guagara are dem¬
onstrated to be sister species within the genus, this
would be another example indicating the floristic
affinities between the Pacific lowland wet forests of
the mostly Costa Rican Golfo Dulce area and the
Colombian Choco (Hartshorn, 1983).
Acknowledgments. I am grateful to Alvaro A.
Cogollo (JAUM), Hermes Cuadros (JBGP), Josefina
Espina (CHOCO), and Juan G. Ramirez for their
herbarium and/or field assistance. I thank Rodrigo
G. Bernal and Gloria Galeano (both of COL) for
directing me to the El Valle locality. Fieldwork was
supported by NSF Grant BSR-8914705.
Literature Cited
Hartshorn, G. S. 1983. Plants. Introduction. Pp. 118-
157 in D. H. Janzen (editor), Costa Rican Natural
History. Univ. Chicago Press, Chicago.
New Species and New Combinations of Justicia
(Acanthaceae) from the Venezuelan Guayana
Dieter C. Wasshausen
Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.
ABSTRACT. Recent expeditions in the Venezuelan
Guayana have resulted in additional new taxa in
Justicia (Acanthaceae). The following places these
novelties and new combinations on record for the
convenience of other taxonomists prior to the pub¬
lication of the Flora of the Venezuelan Guayana.
Nine new Venezuelan species of Justicia, J. brete-
leri, J. delascioi, J. huberi, J. monachinoi, J. nu-
riana, J. panarensis, J. parimensis, J. paruana,
and J. uvida, are described, illustrated, and com¬
pared with their closest relatives. In addition, two
new combinations, J. paraguazensis and J. rnor-
itziana, are also necessary. A key to all of the known
species of Justicia from the region (Territorio Fed¬
eral Delta Amacuro, Estado Bolivar, and Territorio
Federal Amazonas) is also provided.
Justicia is the largest and taxonomically the most
complex genus of Acanthaceae, with between 420
(Mabberley, 1987) and 600 or so (Graham, 1988)
species worldwide. The genus, named for eighteenth-
century Scottish horticulturist and botanist James
Justice, is distributed throughout the tropics and
subtropics of both hemispheres, even into the tem¬
perate regions of North America, with one species
extending as far north as Quebec, Canada. The
taxonomic history of the genus is rather complex,
and recently Graham (1988) attempted an overall
schematic delimitation of the taxa in this assemblage
of dissimilar elements. By utilizing characters of
inflorescence, androecium, pollen, and seeds, she
proposed the broadest definition of Justicia yet to
date. Earlier, Nees (1847), in his major monograph
of the family, divided the group into a number of
smaller genera such as Adhatoda, Amphiscopia,
Beloperone, Leptostachya, Rhytiglossa, and Ser-
icographis in the New World. At that time he knew
nothing about the markedly different types of pollen
within the family that Radlkofer (1883) and Lindau
(1895) later emphasized and used extensively in
their taxonomic delimitation. Bremekamp (1948),
after diligently studying the pollen types in the group,
recognized most of the genera proposed by Nees.
In addition, he described a number of new genera
Novon 2: 62-80. 1992.
for the New World such as Calliaspidia, Dys-
pemptemorion, and Psacadocalymma. Currently
the authors of most recent floras reject the delimi¬
tations proposed by Nees and Bremekamp. They
seem to concur with Stearn (1971) that until cor¬
relations between character of inflorescence, floral
characters, and pollen types have been analyzed for
a large number of species throughout their ranges,
and relatively homogeneous natural groups have
been defined, the simplest course is to maintain
Justicia as a large genus while recognizing the pos¬
sibility of its later dismemberment and the revival
of names now huried in its synonymy.
The genus is characterized by the 2 or 3 aper-
turate pollen with a trema area usually traversed
by 2-8 rows of up to 10 insulae. This form of pollen
was called “Knotchenpollen” by Lindau (1893). The
plants are shrubs or herbs with the flowers variously
disposed. In the region here covered, however, two
basic inflorescence types are prominent: one consists
either of a simple spike (sometimes terminal and
axillary forming a cluster of spikes) with 2 flowers
per node (or rarely a simple spike reduced to 1 or
2 sessile axillary flowers); the other is compound,
composed of spicate units, which are usually secund
and arranged along a main axis forming a rather
large panicle. The floral bracts and bracteoles are
of various shapes and sizes, imbricate or distant,
and the calyx is 5-parted with the segments usually
narrow and nearly equal. The corollas are gama-
petalous and zygomorphic, with the tube divided
above into 2 lips, the upper shallowly bilobed and
the lower distinctly trilobed. There are two stamens
and no staminodes; the anthers are variously 2-lobed
with the thecae superposed or 1-lobed. The oblique
connective varies in length and the lower thecae is
usually spurred, apiculate, or muticous, occasionally
much reduced. The fruit in Justicia is a 2-valved
loculicidal capsule with usually 4 but occasionally 2
seeds. The lower portion of the capsule is commonly
sterile and solid (capsule clavate), hut in the Ven¬
ezuelan Guayana we find a species in which the
capsule is completely ovoid, without a solid basal
stipate portion.
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
63
In the Venezuelan Guayana, most taxa of Justicia
are limited to gallery forests and woodlands along
banks of rivulets and swale margins at elevations
between 100 and 1,400 m. Like many Acanthaceae,
they are of local distribution in undisturbed forests
and are rarely collected in secondary growth. This
Key to the Venezuelan Guayana Species of Justicia
la. Capsule ovoid, without a solid basal stipitate portion, abruptly beaked at apex . 3. J. nuriana
lb. Capsule claviforni (unknown for J. huberi, J. delascioi, and J. panarensis, but probably claviform), with
solid basal stipitate portion, acute to obtuse or bifid at apex.
2a. Inflorescence diffuse, with numerous branches; bracts and bracteoles small, less than 2 mm long
and 1 mm wide; low-growing herbs or undershrub; corollas usually less than 15 mm long.
3a. Corolla 4.5-6 mm long; inflorescence axillary and terminal, with branches 3-7 in whorl . . .
. J. comata
3b. Corolla 7-15 mm long; inflorescence mostly terminal, simple or with alternate branches.
4a. Flowers borne in spikes to 5 cm long; capsules ovate-stipate, 17 mm long; seeds subor-
bicular, 5 mm long, 5.5 mm wide, gray or flecked brown . J. laevilinguis
4b. Flowers borne in rather loose panicles 5-17 cm long; capsules clavate, 7.5-8 mm long;
seeds suborbicular, 1.5 mm diam., light or reddish brown.
5a. Leaf blades ovate, decurrent on the petiole from a rounded base; panicles ca. 17
cm long and 15 cm broad; corollas yellow with faint rose on the lower lip, 9-10
mm long .4. J monachinoi
5b. Leaf blades narrowly to broadly lanceolate, acute to obtuse at base, not decurrent
on the petiole; panicles 5-16 cm long and to 6 cm broad; corollas white, lilac, or
purple, to 8 mm long . J. pectoralis
2b. Inflorescence dense, stout (or if somewhat lax, as in J. parimensis, then with corollas ca. 45 mm
long); bracts and bracteoles conspicuous, more than 2 mm long and 1 mm wide; herbs, weak shrubs,
or subshrubs; corollas usually more than 15 mm long (except in J. cataractae, J. polystachya, and
J. moritziana, where corollas can be 10-13 mm long).
6a. Inflorescence a simple spike (sometimes terminal and axillary forming a cluster of spikes) with
2 flowers per node, a 2-3-flowered cyme, or reduced to 1 or 2 sessile axillary flowers.
7a. Inflorescence a congested terminal capitulum, sessile, subtended by 4 ovate leaf blades
. 1. /. paraguazensis
7b. Inflorescence short spikes, cymes or clusters of spikes or thyrse, these pedunculate, not
subtended by 4 leaf blades.
8a. Corollas white to pale purple, about 10 mm long.
9a. Bracts subulate, very narrow, 0.25 mm wide. J. cataractae
9b. Bracts oblanceolate or ovate, 3-7.5 mm wide.
10a. Bracts ovate, imbricate, 6.5-7.5 mm wide; spikes dense, 2-7 cm long,
1.5 cm wide . J. polystachya
10b. Bracts oblanceolate, not imbricate, 3 mm wide; spikes lax, 1-1.5 cm
long, 0.5 cm wide .2. J. moritziana
8b. Corollas red, purple, rose-lavender, glossy orange, never completely white, 15-60
mm long.
11a. Bracts and bracteoles conspicuously deep red or wine-colored.
12a. Flowers borne in dense, terminal spikes 3 cm long, these surrounded
and partially hidden by the upper leaf blades; the blades imbricated,
elliptic-oblong, 2.7-3.2 cm long and 1.4-1.6 cm wide, subcoriaceous;
corollas 40-45 cm long .5. J. huberi
12b. Flowers borne in axillary and terminal spikes 3.5-5 cm long, these
(at least the terminal spikes) exceeding the upper leaf blades; the blades
not imbricated, oblong to lanceolate, 6.5-8 cm long and 1.8-2.3 cm
wide, firmly membranous; corolla 35-40 cm long .7. J. paruana
lib. Bracts and bracteoles not conspicuously colored.
13a. Corollas 15-16 mm long.
14a. Leaf blades oblong, 11.5-16 cm long, narrowed at the base and
decurrent on the petiole, subchartaceous; bracts and bracteoles
6-7 mm long, 0.5 wide; calyx segments 9-10 mm long, 1 mm
wide. J. hylaea
14b. Leaf blades lanceolate to ovate, 5 8 cm long, obtuse or rounded
at base, not decurrent on the petiole, membranous; bracts and
bracteoles 5.5-6 mm long, 0.8-1 mm wide; calyx segments
6.2-8 mm long, 1.4-1.75 mm wide.
pronounced endemism explains why so many of the
new taxa are only known from single collections. In
an attempt to identify the recent collections of Jus¬
ticia from the Venezuelan Guayana, I have found
nine undescribed species of the genus.
64
Novon
15a. Spikes moderately dense, the peduncles 1.8 8.8 cm long,
corolla purple or red, the tube 4 mm wide, the upper lip
cucullate, the lobes of the lower lip 1 mm long .
. J. guianensis
15b. Spikes dense, the peduncles 10 13.5 cm long; corolla
rose-lavender, the tube 3 mm wide, the upper lip flat, the
lobes of the lower lip 2 mm long .8. J. uvida
13b. Corollas 25-60 mm long.
16a. Bracts conspicuous, ovate, oblong or oblanceolate, 2.25 10 mm
wide.
17a. Flowers (2-3) borne in short, axillary and terminal cymes
1.5-2 cm long . 9. J. delascioi
17b. Flowers borne on one to several axillary and terminal
compact spikes 2-7 cm long.
18a. Spikes short, 2-2.5 cm long; bract oblong, 2.25-
3 mm wide; corolla glossy orange, yellow at base,
about 50 mm long .10. J. breteleri
18b. Spikes 2-7 cm long; bracts ovate or oblanceolate,
3-10 mm wide; corolla red, reddish purple, or violet.
19a. Corolla red, 40 mm long, the upper lip oblong
to narrowly ovate, 18 mm long, the tube
narrowly infundibuliform, about 28 mm long,
3 mm wide at base .6. J. panarensis
19b. Corolla reddish purple or violet, the upper lip
triangular ovate, about 10 mm long, the tube
infundibuliform, about 10 mm long, 4 mm
wide at base . J. carthaginensis
16b. Bracts inconspicuous, not readily distinguished from the brac-
teoles and calyx segments, linear lanceolate, 0.5-1.5 mm wide.
20a. Calyx segments green, 10.5-12.5 mm long, 1.75 mm
wide, the cystoliths conspicuous under a lens; corolla gla¬
brous; bracts and bracteoles puberulent, the trichomes
whitish . J. schomburgkiana
20b. Calyx segments colored, 5 mm long, 0.75 mm wide, the
cystoliths lacking or inconspicuous under a lens; corolla
pubescent; bracts and bracteoles pilose, the trichomes
brown . J. kunhardtii
6b. Inflorescence compound, composed of spicate units that are usually secund and arranged along
one main axis.
21a. Spikes fascicled, forming in aggregate a loose terminal panicle . 11. J. parimensis
21b. Spikes not fascicled, inflorescence composed of spikelike racemes, the latter forming a
rather large terminal panicle.
22a. Upper leaf blades conspicuously large, 24-30 cm long and 8-12 cm wide; panicles
somewhat lax; bracts subulate, 2.5 mm long; corolla rich purple-pink . . J. wurdackii
22b. Upper leaf blades smaller, to 15 cm long and 4.5-6 cm wide; panicles rather
dense; bracts triangular, 1.5 mm long; corolla dull crimson to brick red.
23a. Corolla 50-60 mm long, slightly curved, glabrous; calyx segments linear
and tapering into a long point, 1 mm wide, glabrous or minutely puberulous,
the trichomes, when present, glandular ./. calycina
23b. Corolla about 30 mm long, erect, sparingly puberulous; calyx segments
oblong-lanceolate, 1.25 mm wide, hirtellous with a mixture of glandular and
eglandular trichomes . J. secunda
1. Justicia (sect. Simonisia) paraguazensis Was-
shausen, nom. nov. Replaced name: Chaeto-
chlamys wurdackii Leonard in Mem. New York
Bot. Card. 10(4): 47. 1961, non J. wurdackii
Leonard. 1961. TYPE: Venezuela. Bolivar:
IVurdack & Monachino 40985 (holotype, US).
2. Justicia moritziana Wasshausen, nom. nov.
Replaced name: Amphiscopia lancifolia Taf-
alla ex Nees var. minor Nees in DC. Prodr.
1 1: 357. 1847, non J. minor T. Anders. 1864.
TYPE: Venezuela. Caracas: Moritz s.n. (ho¬
lotype, B destroyed).
3. Justicia nuriana Wasshausen, sp. nov. TYPE:
Venezuela. Bolivar: Altiplanicie de Nuria, E of
Cerro El Picacho, N of Las Nieves and Las
Chicharras, 45 km N of Tumeremo, vicinity of
Deborah, 600-650 m, 5-8 Feb. 1961, Stey-
ermark 89126 (holotype, US; isotypes, NY,
VEN). Figure 1A D.
Herba, caulibus quadrangularibus, glabris vel sursum
puberulis. Foliorum lamina late oblonga vel ovata, acu¬
minata, basi obtusa, membranacea, utrinque glabra vel
subtus in costa et venis lateralibus parce puberula; petioli
glabri vel parce puberuli; spicae graciles, terminales, pe-
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
65
Figure 1. A-D. Justicia nuriana Wasshausen (Steyermark 89126). —A. Habit. —B. Bracteoles and calyx
segments. —C. Calyx segments, cupule, and glands. —D. Bracteoles, calyx segments, and capsule. E-G. Justicia
monachinoi Wasshausen (Wurdack & Monachino 39978). —E. Habit. —F. Calyx, ovary, and pistil. —G. Bracts,
bracteoles, calyx segments, corolla, and stamens.
66
Novon
dunculis subquadrangularibus, bifariam puberulis, rachi-
dibus puberulis; bracteae et bracteolae triangulares, min¬
ute puberulae et ciliatae; calycis segmenta viridia,
lanceolata, minute puberula et ciliata; corolla non visa;
capsulae ovoidae, apice abrupte rostratae, aliquantum
compressae, atropurpureae et nitidae, parce puberulae.
Herb to 1 m tall; stem erect, subquadrangular,
glabrous below, becoming puberulous above, the tri-
chomes retrorsely curved, subappressed, arranged
± in two rows. Leaves on petioles 2.5-6 cm long,
these glabrous or sparingly and inconspicuously pu¬
berulous, the blades broadly oblong to ovate, 16.5-
21.5 cm long, 5.5-10.5 cm wide, acuminate, obtuse
at base, membranous, entire, dark green and gla¬
brous above, dull paler green and glabrous below,
the costa and lateral veins (7-9 pairs) inconspicuous
above, raised and prominent beneath, sparingly pu¬
berulous, the cystoliths conspicuous under a lens
and scattered on the upper surface, less conspicuous
beneath. Flowers borne in slender spikes terminating
the branches, 9-12 cm long, 1.5 cm broad, rather
loose, the internodes 10-20 mm long, the peduncles
10-15 cm long, subcpiadrangular, bifariously pu¬
berulous, the trichomes retrorsely curved, subap¬
pressed, the rachis puberulous, the trichomes re¬
trorsely curved; bracts triangular, 2 mm long, 1
mm wide, minutely puberulous and ciliolate; brac-
teoles triangular, 1.5 mm long, 0.7 mm wide near
the base, puberulous and ciliolate; calyx green, deep¬
ly 5-segmented, the segments lanceolate, 6 mm long,
0.75 mm wide, minutely puberulous and ciliolate,
distinctly 3-nerved; corolla wanting. Capsule ovoid,
13 mm long, 8 mm broad, 2-3 mm thick, abruptly
beaked at apex, somewhat compressed, deep pur¬
plish and nitid when ripe, sparingly puberulous.
Distribution. Steep moist forested slopes over
boulders, cloud forest just below the summit, Estado
Bolivar, Venezuela, between 600 and 650 m. Known
only from the type collection.
Justicia nuriana does not appear to be closely
related to any of the known species of Justicia from
the Cuianas, Venezuela, or Colombia. Unfortunate¬
ly, the new species is known only from fruiting
material. However, since the ovoid capsule of this
taxon is so distinctive, I feel that it is important at
the present time to bring these unusual collections
into the Acanthaceae literature.
4. Justicia monachinoi Wasshausen, sp. nov.
TYPE: Venezuela. Estado Bolivar: Piedra Mar-
imare, E bank of Hio Orinoco opposite head of
Isla El Callo, 200 m, 19 Dec. 1955, Wurdack
cV Monachino 39978 (holotype, US; isotype,
NY). Figure 1E-G.
Suffrutex, caulibus quadrangularibus, glabris vel parce
puberulis. Foliorum lamina ovata, acuminata, basi rotun-
data et in petiolum decurrens, membranacea, utrinque
glabra vel subtus in costa et venis lateralibus parce pub¬
erula; paniculae terminales et axillares, aliquanto laxae,
pedunculis et pedicellis puberulis; bracteae et bracteolae
subulatae, puberulae; calycis segmenta linearia-lanceola-
ta, puberula, ciliolata; corolla flava cum rosea diluta in
labio inferiore, minute pubescens, labio superiore erecto,
triangularo-ovato, minute bilobato, labio inferiore trilo-
bato, lobis ovatis, rotundatis; stamina exserta; lobi an-
therarum superpositi.
Undershrub to 1.5 m high; stems somewhat
branched, subquadrangular, glabrous or sparingly
puberulous, the trichomes retrorsely curved, subap¬
pressed, occasionally gland-tipped. Leaves on peti¬
oles 1-2.5 cm long, these sparingly puberulous with
appressed-ascending trichomes, the blades ovate, 6-
1 1 cm long, 3-5.5 cm wide, acuminate, decurrent
on the petiole from a rounded base, membranous,
entire or undulate, yellowish green, both the upper
and lower surface glabrous or sparingly and incon¬
spicuously puberulous, the costa and lateral veins
(5 or 6 pairs) bearing a few appressed-ascending
trichomes, prominent and conspicuous on the lower
surface, less so above, the cystoliths minute, scat¬
tered. Panicles terminal and axillary, forming a large
complex inflorescence 17 cm long and 15 cm broad,
the ultimate bracts subulate, 1 mm long and 0.4
mm wide, puberulous, the trichomes ± spreading;
bracteoles subulate, 0.8 mm long and 0.25 mm wide,
puberulous, the trichomes ± spreading; rachis and
rachilla puberulous, the trichomes erect, occasion¬
ally gland-tipped; pedicels 0.5-1 mm long, puber¬
ulous; calyx deeply 5-segmented, the segments lin¬
ear-lanceolate, 2.8-3 mm long, 0.5 mm wide near
the base, puberulous and gland-dotted, the trichomes
erect, ciliolate; corolla yellow with faint rose on the
lower lip, 9-10 mm long, minutely pubescent, the
tube about 1 mm broad at base, 4 mm broad at
mouth, the upper lip erect, triangular-ovate, 3.5
mm wide at base, gradually narrowed to tip, this
bilobed, the lobes about 0.2 mm long and wide,
acute, the lower lip ± spreading, cuneate, 5 mm
wide at base of lobes, these ovate, about 1.5 mm
long and 1.5 mm wide, rounded at tip; stamens
exserted about 2 mm beyond the mouth of the
corolla tube, the anthers 2.5 mm long, the lobes
superposed, vertically attached to a connective which
itself is 1 mm long, glabrous, the lower thecae cal-
carate, the spur about 0.4 mm long, blunt; pollen
grains 2-porate, bilateral, 44 X 22 /im, the pores
circular, trema area microreticulate, traversed by
2 rows of 5 peninsulae (Fig. 3A). Capsule clavate,
7.5 mm long, 2.5 mm wide, 1.5 mm thick, puber-
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
67
ulous; seeds suborbicular, about 1.5 mm diam., 0.5
mm thick, light brown, tuberculate.
Distribution. According to label data locally fre¬
quent, Estado Bolivar, Venezuela, at 200 m ele¬
vation. Known only from the type locality.
Justicia monachinoi is perhaps most closely re¬
lated to the widespread lowland species of J. pec-
toralis Jacq., which differs markedly by having a
rather diffuse inflorescence with numerous very slen¬
der branches 5-16 cm long and to 6 cm broad, the
corollas being white, lilac, or purple, to 8 mm long,
with the throat transversely plicate, sometimes spot¬
ted with dark purple and the plants being weak
stemmed herbs. In contrast, J. monachinoi has ax¬
illary and terminal panicles forming a large complex
inflorescence 17 cm long and 15 cm broad, the
corollas yellow with faint rose on the lower lip, 9-
10 mm long and the plants being undershrubs with
branched stems.
5. Justicia huberi Wasshausen, sp. nov. TYPE:
Venezuela. Bolivar: Dist. Cedeno, Sierra de
Maigualida, NE sector, high plateau tepuyana
located on the headwaters of the Rio Chajura,
western affluent of the Rio Erebato, 5°33'N,
65°13'W, 2,100 m, 28 Mar. 1988, Huber
12731 (holotype, NY). Figure 2A-I).
Suffrutex, caulibus subquadrangularibus, glabris vel
puberulis. Foliorum lamina imbricata, elliptico-oblonga,
acuta, basi angustata, subcoriacea, supra glabra vel pub-
erula, minute punctata, subtus in costa et venis lateralibus
parce puberula; petioli breves; spicae terminales, densi-
florae, deminutae foliorum laminae subtentae, pedunculis
subquadrangularibus, dense puberulis; bracteae vinaceae,
ellipticae, glabrae vel parce puberulae et ciliatae; brac-
teolae vinaceae, oblongae; calycis segmenta lanceolata,
glabra vel parce puberula, ciliolata; corolla atrorosea,
glabra, tubo anguste infundibuliformi, labio superiore er-
ecto, oblongo vel anguste ovato, bilobato, labio inferiore
trilobato, lobis oblongis, rotundatis; stamina exserta; lobi
antherarum superpositi.
Shrub 0.5-1.5 m tall; stem erect, subquadran-
gular, glabrous or puberulous, the trichomes re-
trorsely curved, subappressed, sordid. Leaves on
petioles 2-3 mm long, these puberulous with sordid,
appressed-ascending trichomes, the blades imbri¬
cated, elliptic-oblong, 2.7-3.2 cm long, 1.4-1.6 cm
wide, acute, narrowed at base, subcoriaceous, entire,
the margins revolute, the upper surface lustrous,
glabrous or bearing a few appressed trichomes, es¬
pecially along the costa, minutely punctate, the low¬
er surface sparingly puberulous especially along the
costa and lateral veins (5-6 pairs), these obscure
above and prominent and raised beneath, the tri¬
chomes adpressed-ascending, 0.2 mm long. Flowers
borne in dense, terminal spikes 3 cm long and 1-
1.5 cm broad, obtuse or rounded at tip, each flower
cluster subtended by reduced leaf blades, the pe¬
duncles 10-12 mm long, subquadrangular, densely
puberulous with sordid, appressed-ascending tri¬
chomes; bracts wine-colored, elliptic, 15-16 mm
long, 8-9 mm wide, obtuse, narrowed at base, firm,
nitid, glabrous or sparingly puberulous, the margins
revolute, sparingly ciliate; bracteoles wine-colored,
oblong, 10 mm long, 1.5-2 mm wide, glabrous or
sparingly and minutely puberulous, the margins cil-
iolate, the costa and lateral veins prominent; calyx
deeply 5-segmented, the segments lanceolate, 7 mm
long, 1 mm wide, nitid, glabrous or minutely and
inconspicuously puberulous, ciliolate; corolla deep
rose-colored, 40-45 mm long, glabrous, the tube 3
mm broad at base, narrowed to 2.5 mm at 3 mm
above base, thence gradually enlarged to 7.5 mm
at mouth, the lips subequal, 22 mm long, the upper
lip oblong to narrowly ovate, 1 1 mm wide near the
base, 4 mm wide near the tip, bilobed, the lobes 0.5
mm long, rounded, the lower lip ± spreading, 3-lobed,
the lobes oblong, 6 mm long, the lateral lobes 3.5
mm wide, the middle lobe 4 mm wide, all rounded
at tip, delicately veined; stamens almost reaching
the tip of the upper lip, the filaments flattened,
glabrous, the anthers 4 mm long, the thecae 2.5
mm long, superposed, somewhat obliquely attached
to the connective which itself is about 0.5 mm wide,
both thecae terminating at base in a short blunt spur
0.25 mm long; pollen 3-porate, prolate to perpro-
late, 49 x 25 n m, the pores circular, the membrane
reticulate, trema area studded with 2 rows of 5, ±
circular insulae (Fig. 3B, C); ovary glabrous; style
3.5 cm long, glabrous; stigma minute, bilobed. Cap¬
sule wanting.
Distribution. Frequent in grassland. Estado Bo¬
livar, Venezuela, at 2,100 m. Known only from the
type locality.
Justicia huberi is closely affiliated with another
Venezuelan species, J. paruana Wasshausen from
Territorio Federal Amazonas, Cerro Sipapo, and Ser-
rania Paru. In J. paruana the flowers are borne in
axillary and terminal spikes 3.5-5 cm long, they
are not subtended by the reduced leaf blades, the
leaf blades are oblong to lanceolate, 6.5-8 cm long
and 1.8-2.3 cm wide, firm and net-veined and the
corollas are rose-colored, 35-45 mm long, and pu¬
berulous. In contrast, the flowers of J. huberi are
borne in dense, terminal spikes 3 cm long, and each
floral cluster is subtended by reduced leaf blades,
the leaf blades are elliptic-oblong, 2.7-3.2 cm long
and 1.4-1.6 cm wide, subcoriaceous, not net-veined.
68
Novon
Figure 2. A-D. Justicia huberi Wasshausen (Huber 12731). —A. Habit. —B. Bract. —C. Bracteoles and calyx
segments. —D. Corolla and stamens. E H. Justicia panarensis Wasshausen (Boom & Grillo 6315). —E. Habit.
— F. Bract. —G. Bracteoles and calyx segments. —H. Corolla and stamens.
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
69
Figure 3. Scanning electron (SEM) photomicrographs of Justicia pollen. —A. J. monachinoi , equatorial view,
x 2,000 (Wurdack & Monachino 39978). B, C. J. huberi (Huber 12731). —B. Equatorial view, x 1,700. —C.
Equatorial view, x 2,000. —D. J. paruana, equatorial view, x 1,800 (Maguire & Politi 28244).
70
Novon
and the corollas are deep rose-colored, 40-45 mm
long and glabrous.
6. Justicia panarensis Wasshausen, sp. nov.
TYPE: Venezuela. Bolivar: Dist. Cedefio, vi¬
cinity of Panare village of Corozal, 6 km from
Maniapure toward Caicara, 6°55'N, 66°30'W,
90 m, 11 Oct. 1985, Boom & Grillo 6315
(holotype, US). Figure 2E-H.
Suffrutex debiiis, caulibus subquadrangularibus, pub-
erulis. Foliorum lamina oblongo-ovata, acuta vel brevia-
cuminata, basi angustata, firma, supra parce puberula,
subtus puberula, costa sicut venis lateralibus puberula et
pilosa; petioli dense puberuli; spicae solitariae vel plures
axillares et terminates, pedunculis et rachidibus subquad¬
rangularibus, puberulis; bracteae viridae, ovatae, glabrae
vel parce puberulae, parce ciliatae; bracteolae lanceolatae,
glabrae vel parce puberulae, ciliatae; calycis segmenta
lanceolata, ciliolata, parce puberula; corolla scarlatina,
minute pubescens, tubo anguste infundibuliformi, labio
superiore erecto, oblongo vel anguste ovato, bilobato, labio
inferiore trilobato, lobis oblongis, rotundatis; stamina
exserta; lobi antherarum superpositi.
Weak shrub 0.5 m tall; stem erect, subquadran-
gular, puberulous, the trichomes retrorsely curved,
subappressed. Leaves on petioles 8-12 mm long,
these densely puberulous with curved trichomes, the
blades oblong-ovate, 5-7 cm long and 2.5-3 cm
wide, acute to short-acuminate, narrowed at base,
rather firm, entire or undulate, the upper surface
sparingly puberulous, the lower surface moderately
puberulous, occasionally interspersed with a few pi¬
lose trichomes, the costa and lateral veins (5-6 pairs)
puberulous and pilose, the cystoliths often incon¬
spicuous and scattered. Spikes one to several, ax¬
illary and terminal, 5-6 cm long and 2-2.5 cm
broad, obtuse or rounded at tip, the peduncles 0.5
cm long, the rachis puberulous, the trichomes erect,
white, about 0.3 mm long; bracts green, ovate, 19
mm long, 9.5-10 mm wide, acute, obtuse at base,
thin, glabrous or sparingly puberulous, sparingly
ciliate, the veins coarsely reticulate, these and the
costa rather prominent; bracteoles lanceolate, 8 mm
long, 2.5-3 mm wide, glabrous or sparingly and
minutely puberulous, the margins ciliate, the costa
prominent, the lateral veins delicately and coarsely
reticulate; calyx 6 mm long, deeply 5-segmented,
the segments lanceolate, 5.5 mm long, 1.2 mm wide,
ciliolate, minutely and sparingly puberulous, the cos¬
ta prominent; corolla red, 40 mm long, finely pu¬
bescent, the trichomes to 0.2 mm long, the tube 3
mm broad at base, narrowed to 2.5 mm at 3 mm
above base, thence gradually enlarged to 5 mm at
mouth, the lips subequal, 18 mm long, the upper
lip oblong to narrowly ovate, 7.5 mm wide near the
base, 1.5 mm wide near the tip, bilobed, the lobes
0.75 mm long, rounded, the lower lip ± spreading,
3-lobed, the lobes oblong, 14 mm long, 4.5 mm
wide, rounded at tip, delicately veined; stamens ex-
serted 8 mm beyond the mouth of the corolla tube,
the filaments white, flat, glabrous, the anthers 3 mm
long, the thecae 1.5 mm long, superposed, somewhat
obliquely attached to the connective, this about 0.5
mm wide, the upper thecae muticous, the lower
terminating in a short blunt spur 0.25 mm long;
pollen 2-porate, prolate to perprolate, 60 X 30 /am,
the pores circular, the membrane reticulate, trema
area studded with 2 rows of 7-10, ± circular insulae
(Fig. 8A, B); ovary glabrous; style 2.5 cm long,
minutely puberulous at base. Capsule wanting.
Distribution. In semideciduous forest on granitic
slopes and savanna, 90-400 m. Known only from
the type locality.
Justicia panarensis superficially resembles J.
leucerythra Leonard from Colombia. However, in
J. leucerythra the bracts are firm, 8 mm wide, the
corolla is pink or occasionally red, to 33 mm long,
with the subequal lips 11 mm long, and the leaf
blades are larger, 20 cm long and 6 cm wide. In J.
panarensis the bracts are thin, 9.5-10 mm wide,
the corolla is red, 40 mm long, with the subequal
lips 18 mm long, and the leaf blades are much
smaller, 5-7 cm long and 2.5-3 cm wide.
7. Justicia paruana Wasshausen, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas: Ser-
rania Paru, Rio Paru, Cano Asisa, Rio Ven-
tuari, 1,800 m, 1 Feb. 1951, Cowan & If iir-
dack 31116 (holotype, US; isotype, NY). Figure
4A-E.
Suffrutex debiiis, caulibus aliquantum ramosis,
subquadrangularibus, minute puberulis. Foliorum lamina
oblonga vel lanceolata, acuminata, basi angustata, firma
reticulato-venosa, utrinque glabra vel in costa et venis
lateralibus parce puberula; petioli puberuli; spicae ter-
minales et axillares, pedunculis subquadrangularibus, pub¬
erulis; bracteae vinaceae, ellipticae, glanduloso-puberulae
et ciliatae; bracteolae vinaceae, oblanceolatae; calycis seg¬
menta linearia, puberula, ciliolata; corolla rosea, puberula,
tubo anguste infundibuliformi, labio superiore erecto, ova¬
to, bilobato, labio inferiore trilobato, lobis oblongis, rotun¬
datis; stamina exserta; lobi antherarum superpositi.
Weak shrub 1.5-3 m tall, somewhat branched;
stems erect, subquadrangular, minutely puberulous,
the trichomes antrorsely curved, subappressed.
Leaves on petioles 9-15 mm long, these puberulous
with appressed-ascending trichomes, the blades ob¬
long to lanceolate, 6.5-8 cm long and 1.8-2.3 cm
wide, acuminate, narrowed at base, firm, net-veined,
entire or undulate, the margins revolute, the upper
surface subnitid, glabrous or the costa bearing a few
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
71
Figure 4. A-E. Justicia paruana Wasshausen ( Cowan & f Vurdack 31116). —A. Habit. —B. Bract, bracteoles,
and calyx. —C. Bracteole. —D. Calyx segments. —E. Corolla and stamens. F-J. Justicia uvida Wasshausen
(Steyermark & Aristeguieta 35). —F. Habit. —G. Bract, bracteole, and immature corolla. —H. Bracteole. —I.
Calyx. —J. Corolla and stamens.
72
Novon
small trichomes, the cystoliths minute, inconspicu¬
ous, the lower surface glabrous except the costa and
lateral veins (5-6 pairs), these minutely and spar¬
ingly puberulous, on both surfaces the costa and
lateral veins are prominent and conspicuously raised.
Flowers borne both in axillary and terminal spikes
3.5-5 cm long and 1-1.5 cm broad, obtuse or
truncate at tip, the peduncles 10-15 mm long,
subquadrangular, puberulous, the trichomes erect;
bracts wine-colored, elliptic, 14 mm long, 5-5.2
mm wide, acute to short acuminate, narrowed at
base, puberulous with both glandular and eglandular
trichomes, the margins ciliolate; bracteoles wine-
colored, oblanceolate, 12 mm long, 2.5 mm wide,
puberulous, the margins ciliolate, the trichomes oc¬
casionally gland-tipped; calyx deeply 5-segmented,
the segments linear, 4.5 mm long, 0.5 mm wide,
puberulous, ciliolate, the trichomes erect, occasion¬
ally gland-tipped; corolla rose-colored, 35-40 mm
long, sparingly and inconspicuously puberulous, the
tube 2 mm broad at base, narrowed to 1.5 mm at
3 mm above base thence gradually enlarged to 6
mm at mouth, the lips subequal, 17 mm long, the
upper lip ovate, 7.5 mm wide near the base, nar¬
rowed toward tip, bilobed, the lobes 0.5 mm long,
rounded, the lower lip ± spreading 3-lobed, the lobes
oblong. 2.5 mm long, the lateral lobes 2 mm wide,
the middle lobe 1.5 mm wide, all rounded at tip,
delicately veined; stamens exserted 15 mm beyond
the mouth of the corolla tube, the filaments flattened,
glabrous, the anthers 4 mm long, the thecae 2.5
mm long, superposed, somewhat obliquely attached
to the connective which itself is about 0.5 mm wide,
both thecae terminating at the base in a short, white
spur 0.25 mm long; pollen 3-porate, prolate to per-
prolate, 46 x 24 the pores circular, the mem¬
brane reticulate, trema area studded with 2 rows of
4, ± circular insulae (Fig. 3D); ovary glabrous; style
3.2 cm long, glabrous; stigma minute, bilobed. Cap¬
sule clavate, puberulous, 12 mm long, 4 mm broad,
3 mm thick, the solid stipe 5 mm long and 2.5 mm
broad, gland-dotted; seeds not seen.
Distribution. Occasional, in talus forest and
woodland along streams at elevations between 1,600
and 1,800 m, in Territorio Federal Amazonas, Ven¬
ezuela.
Para types. Venezuela, territorio federal amazonas:
Cerro Siapo (Paraque), N branch of Cano Profundo, 1,600
m, Maguire & Politi 28227 (US); upper Cano Negro,
1,600 in, Maguire Politi 28244 (NY, US).
Justicia paruana is closely affiliated with another
Venezuelan species, J. huberi Wasshausen from Es-
tado Bolivar. Its relationship to J. huberi is described
under that taxon.
8. Justicia uvida Wasshausen, sp. nov. TYPE:
Venezuela. Bolivar: vicinity of km 128 S of El
Dorado, NE of Luepa, 800-1,200 m, 6-11
Mar. 1962, Steyermark Aristeguieta 35
(holotype, US; isotype, VEN). Figure 4F J.
Herba, caulibus ramosis, plus minusve patulis, quad-
rangularibus, glabris vel parce puberulis; folii lamina ova-
ta, breviacuminata, basi rotundata, subchartacea, utr-
inque glabra vel subtus secus costam puberula; petioli
breves; spicae terininales vel subterminales, densiflorae,
contractae, pedunculis et rachidibus puberulis; bracteae
et bracteolae lanceolatae, glabrae et ciliatae; calycis seg-
inenta lanceolata, ciliata; corolla roseo-lavandulacea, infra
glabra, sursum pilosa, labio superiore erecto, obovato-
elliptico, apice bilobato, labio inferiore trilobato, erecto
vel reclinato, obovato vel ovato, lobis obovatis, rotundatis;
stamina exserta; lobi antherarum leviter superpositi.
Sprawling, subherbaceous, to 2 m tall; stems elon¬
gated, obscurely quadrangular, shallowly sulcate, at
least above, glabrous or sparingly puberulous, the
trichomes curved and subappressed, about 0.5 mm
long, disposed ± in 2 rows, the internodes of the
stem 3.5-10.5 cm long. Leaves sessile or very short¬
ly petiolate; if petiolate then petioles 12 mm long,
densely puberulous, the trichomes sordid, the blades
ovate, 6.5-8 cm long and 3.5-4.7 cm wide, with
the uppermost pair of leaves conspicuously reduced
(2.5-3 cm long, 1.3-1.4 cm wide) and subtending
the spikes, short-acuminate, obtuse or rounded at
base, firm subchartaceous, entire or undulate, deep
green and sulcate-nerved above, dull green with
raised nerves (about 6 pairs) below, both surfaces
glabrous or the costa beneath minutely and incon¬
spicuously puberulous, the cystoliths of both surfaces
inconspicuous under a lens. Flowers in short, dense
spikes forming a compound spicate inflorescence 3-
7 cm long and 2.5-3 cm wide including the corollas,
the flowers fascicled, the rachises and peduncles
rather densely and bifariously puberulous, the tri¬
chomes erect or ascending, about 0.2 mm long, the
peduncles 10-13.5 cm long, the nodes of the spike
bearing a pair of small leaves (their blades 13-22
mm long and 4-9 mm wide); bracts subtending the
flowers lanceolate, 5.5-6 mm long, 1 mm wide,
carinate, mucronulate at apex, firm, glabrous, the
margin ciliate; bracteoles similar but slightly larger;
calyx 8-9 mm long, deeply segmented, the five
segments lanceolate, 6.25-8.25 mm long, 1.75 mm
wide slightly above the middle, acuminate, the tip
curved, the costa prominent, bearing dorsally a few
glandular and eglandular trichomes, the margins
ciliate, the trichomes whitish, erect or ascending,
0.5 mm long; corolla rose-lavender, 16 mm long,
glabrous toward base, finely pilosulous toward tip,
the trichomes spreading, to 0.4 mm long, the tube
6 mm long, 3 mm broad at base, gradually enlarged
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
73
to 4 mm at mouth, the upper lip obovate to elliptic,
± erect, about 11 mm long, 6 mm wide at middle,
plicate, dorsally pilosulous, the trichomes glandular
or eglandular interspersed, hilobed at tip, the lobes
rounded, 0.5 mm long and 1 mm wide at base, the
lower lip striped white and lavender, obovate to
ovate, erect or reclinate, 12 mm long, 9 mm wide,
3-lobed, the lobes obovate, rounded, 3.5 mm long,
the middle lobe 3.5 mm wide, the lateral lobes 2.5
mm wide; stamens exserted 8 mm beyond the mouth
of the corolla, the filaments flattened and glabrous,
the anther thecae slightly superposed, the connec¬
tive 1 mm broad, the upper thecae 1.5 mm long,
0.5 mm broad, white-puberulous on the margin,
muticous and minutely spurred at base, the lower
thecae 1 mm long and 0.5 mm broad, white-pu¬
berulous on margin, terminating in a thecal ap¬
pendage about 1 mm long; pollen triporate, subpro-
late to perprolate, 36 x 20 /nm, the pores circular,
the membrane reticulate, trema area inconspicu¬
ously studded with 2 rows of ± circular insulae (Fig.
6A); style about 14 mm long, minutely puberulous,
the stigma minutely bilobed; ovary glabrous. Capsule
clavate, 9 mm long, 3 mm wide, brownish, puber¬
ulous; seeds not seen.
Distribution. In gallery forest, along banks of
rivulets and swale margins, Estado Bolivar, Vene¬
zuela, at 460-1,400 m.
Paratypes. VENEZUELA, bolivar: Gran Sabana, N of
Urirnan, 5°29'N, 62°36'W, 460 rn, Fernandez 2548
(MYF, US); Depto. Piar, Rio Karwai, around Irana-Wai-
mah, 20 kin from Kavanayen, 1,200 m, Ruiz-Teran <V
Lopez-Palacios 11311 (MERF, US); Rio Tarata, km
147-148 S of El Dorado, 1,220 m, Gentry, Morillo &
B. de Morillo 10554 (MO); ca. km 148 S of El Dorado,
headwaters of the Rio Aponguao, along Arauta-paru,
1,350-1,400 m, Steyermark & G. C. K & E. Dunster-
ville 104157 (US, VEN).
Justicia uvida superficially resembles /. ma-
guirei Wasshausen from Guyana. However, in J.
maguirei the bracts and bracteoles are narrowly
triangular, 2-3 mm long, the calyx segments 5.5
mm long, straight at tip and the corolla, probably
purple or red, is 18-22 mm long with the lohes of
the lower lip ovate, 3 mm long, the middle lobe 5
mm wide and the lateral lobes 3 mm wide. In con¬
trast, J. uvida has bracts and bracteoles lanceolate,
5.5-10 mm long, the calyx segments are 6.25-
8.25 mm long, the tips curved and the corolla is
rose-lavender, 16 mm long with the lobes of the
lower lip obovate, 3.5 mm long, the middle lobe 3.5
mm wide and the lateral lobes 2.5 mm wide.
9. Justicia delascioi Wasshausen, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas: Dep¬
to. Atabapo, Salto Yureba, Cano Yureba, Bajo
Ventuari, 4°3'N, 66°1'W, 120 150 m, 24
Oct.-4 Nov. 1981, Delascio & Guanchez
10896 (holotype, US). Figure 5A-C.
Frutex vel sufifrutex, caulibus subquandrangularibus,
glabris. Foliorum lamina late lanceolata vel ovata, acu¬
minata, basi angustata, firma, utrinque glabra vel parce
puberula, costa et venis lateralibus puberulis; cymae ax-
illares et terminales, pedunculis et pedicellis glabris; brac-
teae pedunculis subtentes foliaceae, ovatae, glabrae; brac-
teae floribus subtentes oblanceolatae, ciliolatae, glabrae;
calycis segmenta oblonga vel elliptica, glabra; corolla lil-
acina, glabra vel minute glanduloso-pubescens, labio su-
periore erecto, ovato, bilobato, labio inferiore trilobato,
lobis ellipticis, rotundatis; stamina exserta; lobis anther-
arum superpositis.
Shrub or subshrub, erect, 0.5 m tall; stem
subquadrangular, glabrous. Leaves on petioles 1.5
cm long, these puberulous in two rows on the edge
of channels, the trichomes retrorsely curved, the
blades broadly lanceolate to ovate, 6.5 8 cm long
and 2-2.7 cm wide, acuminate, narrowed at base,
rather firm, entire or undulate, both the upper and
lower surfaces glabrous or sparingly and inconspic¬
uously puberulous, the costa and lateral veins (4 or
5 pairs) bearing a few appressed-ascending tri¬
chomes, inconspicuous on both surfaces, the cys-
toliths conspicuous under a lens and scattered on
the upper surface, less conspicuous beneath. Flowers
(2 to 3) borne in short axillary and terminal cymes,
these 1.5-2 cm long, the peduncles 3.5-6 cm long,
the secondary peduncles 6-9 mm long, both primary
and secondary peduncles glabrous, subquadrangu¬
lar, the angles rounded; bracts subtending the sec¬
ondary peduncles leaflike, ovate, 8-15 cm long, 2
6 mm wide, acuminate at tip, glabrous without,
minutely puberulous within, the petioles 2-4 mm
long; bracts subtending the flowers oblanceolate, 10
mm long, 2 mm wide, ciliolate, glabrous, with scat¬
tered, brownish cystoliths visible under magnifica¬
tion; calyx deeply 5-segmented, the segments oblong
to elliptic, 14 mm long, 4 mm wide near the middle,
acuminate at apex, narrowed toward the base, co¬
riaceous, glabrous, drying yellowish; corolla lilac,
about 25 mm long, glabrous or minutely and in¬
conspicuously glandular pubescent, the tube 13 mm
long, 3 mm broad at base, 7 mm broad at mouth,
the upper lip erect, ovate, 12-13 mm long, 7 mm
wide, bilobed at apex, the lobes rounded, 0.5 mm
long and wide, cucullate, enclosing the stamens, the
lower lip 15 mm long and 1 cm wide at base of the
3 lobes, these elliptic, 7-8 mm long, 5-6 mm wide,
rounded; stamens exserted 11 mm beyond the mouth
of the corolla tube, the anthers 5.5 mm long, the
thecae superposed, vertically attached to a connec¬
tive which itself is 0.5 mm long, both thecae pu-
74
Novon
Figure 5. A-C. Justicia delascioi Wasshausen (Delascio & Guanchez 10896). —A. Habit. — B. Calyx. —C.
Corolla and stamens. D-F. Justicia breteleri Wasshausen (Breteler 4761). —D. Habit. —E. Bract and calyx. —
F. Bracteole, calyx segments, corolla, and stamens.
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
75
bescent, the lower thecae terminating in an ap¬
pendage 1.75 mm long; pollen 2-porate, prolate to
perprolate, 55 x 28 gm, the pores circular, the
equatorial outline of the pollen with a distinct in¬
dentation at the equator, the trema region micro-
reticulate (Fig. 6B); ovary puberulous; style 2 cm
long, glabrous; stigma minute, bilobed. Capsule
wanting.
Distribution. Known only from the type locality.
Justicia delascioi superficially resembles J. hoch-
reutineri Macbride from Venezuela. However, in J.
hochreutineri the flowers are borne in sessile or
short-pedunculate axillary spikes, the calyx seg¬
ments are oblong-lanceolate, setaceous-cuspidate,
scabrous pubescent and drying green, and the bracts
subtending the flowers are lanceolate and exceeding
the calyx. In contrast, in J. delascioi the flowers
are borne in axillary and terminal 2-3-flowered cymes
on peduncles 3.5-6 cm long, the calyx segments
are oblong to elliptic, acuminate, glabrous, and dry¬
ing yellowish, and the bracts subtending the flowers
are oblanceolate and shorter than the calyx.
10. Justicia breteleri Wasshausen, sp. nov.
TYPE: Venezuela. Territorio Federal Amazo¬
nas: left bank of Sipapo River near confluence
with Orinoco River, ±90 m, 20 Nov. 1965,
Breteler 4761 (holotype, US). Figure 5D-F.
Suffrutex, caulibus erectis, subquadrangularibus, gla-
bris vel minute puberulis. Foliorum lamina oblonga vel
elliptica, acuminata, basi obtusa, firma, supra glabra, sub-
tus minute puberula costa et venis lateralibus subtus prom-
inentibus; petioli puberuli; paniculae terminales, spicis par¬
vis compactis compositae, pedunculis et rachidibus
subquadrangularibus, puberulis; bracteae oblongae, cil-
iolatae; bracteolae oblongae vel oblanceolatae, ciliolatae;
calycis segmenta oblonga, glabra et ciliata; corolla nitida
aurantiaca, tubo luteo, sursurn gradatim ampliato, subtil-
iter pubescenti, labio superiore erecto, oblongo vel anguste
ovato, bilobato, labio inferiore trilobato, patulo, lobis ob-
longis, rotundatis; stamina exserta; lobis antherarum su-
perpositis.
Partly climbing, weak-stemmed shrub 2 4 m tall;
stem erect, subquadrangular, glabrous or minutely
and inconspicuously puberulous. Leaves opposite or
whorled, on petioles 1-1.5 cm long, these minutely
puberulous especially in two rows on the edge of
channels, the blades oblong to elliptic, 11-15 cm
long and 4-5.5 cm wide, acuminate, obtuse at base,
rather firm, entire, dark green and glabrous above,
dull paler green and minutely puberulous below,
especially the costa and lateral veins (7-9 pairs),
these inconspicuous above, raised and prominent
beneath, the trichomes erect, the cystoliths incon¬
spicuous under a lens above, scattered and con¬
spicuous under a lens below, the margins revolute;
inflorescence a terminal panicle composed of several
short, compact spikes 2-2.5 cm long (excluding
corollas), the peduncles 1-3 cm long, subquadran¬
gular, puberulous, the internodes of the inflorescence
6 mm long, somewhat flattened, 2 mm wide, pu¬
berulous; bracts oblong, 13-15 mm long, 2.25-3
mm wide, ciliolate, glabrous or minutely and incon¬
spicuously puberulous; bracteoles oblong to oblan¬
ceolate, 10 mm long, 1.4 mm wide, ciliolate, gla¬
brous or minutely and inconspicuously puberulous;
calyx deeply 5-segmented, the segments oblong, 8
9 mm long, 1.75 mm wide near the middle, acute
to acuminate at apex, slightly narrowed toward the
base, glabrous, the margins conspicuously ciliate;
corolla glossy orange, yellow at the base, about 50
mm long, the tube finely pubescent, 32 mm long,
2.5 mm broad at base, narrowed to 1.5 mm at 5
mm above base, thence gradually enlarged 1o 5.5
mm at mouth, the lips subequal, 17-18 mm long,
the upper lip oblong to narrowly ovate, finely pu¬
bescent, 8 mm wide near the base, 1 mm wide near
the tip, bilobed, the lobes 0.4 mm long, rounded,
the lower lip somewhat spreading, 3-lobed, the lobes
oblong, 4 mm long, 1.5 mm wide, rounded at tips,
densely pilose, the trichomes erect or spreading, 1-
1.3 mm long; stamens exserted 16 mm beyond the
mouth of the corolla tube, the filaments flat, gla¬
brous, the anthers 4.5-5 mm long, the thecae 2.5
mm long, superposed, somewhat obliquely attached
to the connective, this 0.5 mm wide, the upper
thecae muticous, the lower terminating in a short
blunt spur 0.5 mm long; pollen grains 2-porate,
bilateral, 60 x 25 pm, the pores circular, the trema
area microreticulate, traversed by 4 rows of 6-7
small smooth insulae (Fig. 6C, D); style 4.7 cm long,
glabrous. Capsule clavate (not seen at maturity).
Distribution. Rare and infrequent at edge of river
and along roadsides in the Departamento Atures,
Territorio Federal Amazonas, Venezuela, at eleva¬
tions between 80 and 200 m.
Paratypes. Venezuela. TERRITORIO FEDERAL AMAZONAS:
Depto. Atures, 2 km below Caserio San Pedro de Catan-
iapo, 50 km SE of Puerto Ayacucho, 5°38'N, 67°12'W,
100 m, Guanchez 179 (US); vicinity of “Las Pavas,”
25 km SE of Puerto Ayacucho on the road towards El
Gavilan, 5°35'N, 67°27'33"W, 80 m, Cuello 325 (US);
mountainous region of la Coromoto, sector “El Tobogan,”
37 km S of Puerto Ayacucho, 5°24’N, 67°35'W, 80-
200 m, Cuello 352 (US).
Justicia breteleri is not closely allied to the other
known species of neotropical Justicia. It is perhaps
nearest in relationship to J. sanchezioides Leonard
from the Comisaria Putumayo, Colombia, and the
76
Novon
Figure 6. Scanning electron (SEM) photomicrographs of Justicia pollen. —A. J. uvula , equatorial view, x 2,500
(Steyermark & Aristeguieta .35). —B. J. delascioi, equatorial view, x 1,600 (Delascio & Guanchez 1089 6). C,
D. J. bretcleri. —C. Equatorial view, x 1,600 (Breteler 4761). —D. Equatorial view, x 1,800 (C.uello 325).
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
77
Figure 7. A-E. Justicia parimensis Wasshausen (Huber & Colchester 8404). —A. Habit. —B. Bracts and calyx.
— C. Corolla and stamens. —D. Bracts, calyx segments, cupula, glands, and immature capsule. —E. Capsule and
retinaculum.
78
Novon
Figure 8. Scanning electron (SEM) photomicrographs of Justicia pollen. A, B. J. panarensis (Boom & Grillo 6315).
— A. Equatorial view, x 1,600. —B. Portion of surface, x 3,000. C, D. J. parimensis (Huber & Colchester 8404).
— C. Equatorial view, x 1,600. — D. Portion of surface, x 5,000.
Volume 2, Number 1
1992
Wasshausen
Justicia from Venezuelan Guayana
79
Provincia Napo, Ecuador; however, this species is
unique in that the inflorescence is a terminal panicle
composed of several unilateral spikes 5-12 cm long
(excluding the corollas), the corollas are pale purple
or bright rose, 65 mm long, with the subequal lips
37 mm long, and the subequal bracts and bracteoles
are lanceolate, 18 mm long, moderately hirsute with
the bracteoles slightly longer than the bracts. In
contrast, in J. breteleri the inflorescence is com¬
posed of short, terminal, compact spikes 2-2.5 cm
long (excluding the corollas), the corollas are glossy
orange, yellow at the base, 50 mm long, with the
subequal lips 17-18 mm long, and the bracts and
bracteoles are unequal, oblong to oblanceolate, gla¬
brous or inconspicuously puberulous with the bracts
13-15 mm long and the bracteoles 10 mm long,
respectively.
11. Justicia parimensis Wasshausen, sp. nov.
TYPE: Venezuela. Territorio Federal Amazo¬
nas: Depto. Atabapo, Sierra Parima, NE sector,
along the Rio Matacuni, approx. 20 km NNW
of Shimada-Wochi, 3°59'N, 64°41'W, 1,000-
1,500 m, 10 Nov. 1983, Huber & Colchester
8404 (holotype, US). Figure 7A-E.
Suffrutex, caulibus subquadrangularibus, sursuni pub-
erulis. Foliorum lamina elliptica vel ovata, acuminata, basi
angustata et in petiolum decurrens, firma, utrinque glabra
vel parce pilosa; petioli breves; spicae fasciculatae, in
aggregis terminalibus paniculam formans, fioribus secun-
dis; bracteae et bracteolae triangulares, glabrae et ciliatae;
calycis segmenta lanceolata, apiculata, puberula; corolla
rosea-lilacina, minute puberula, tubo anguste infundibu-
liformi, labio superiore erecto, ovato, minute bilobato,
labio inferiore oblongo, trilobato, lobis rotundatis; stamina
exserta; lobi antherarum superpositi; capsulae clavatae,
puberulae, acutae, unumquaeque valve bifida ad apicem.
Shrub or subshrub, erect, 1.5 m tall; stem
subquadrangular, glabrous below, becoming puber¬
ulous above, the trichomes retrorsely curved, sub-
appressed, arranged ± in two rows. Leaves on
short petioles 1-3 cm long, these glabrous or mi¬
nutely puberulous especially in two rows on the edge
of channels, the blades elliptic to ovate, 20-24 cm
long, 6.4-8.8 cm wide, acuminate and curved at
apex, narrowed at base and decurrerxt on the petiole,
firm, entire, the upper surface drying dark green,
glabrous or minutely and inconspicuously pilose, the
costa and lateral veins (about 1 1 pairs) prominent
but less so than those of the lower surface, the
cystoliths minute and scattered on both surfaces,
visible only under a lens, the lower surface drying
a lighter green, glabrous or minutely and incon¬
spicuously pilose especially along the venations; in¬
florescence terminal, the flowers secund in simple
spikes, these fascicled, the peduncles 9-12 cm long,
the spikes forming in aggregate a loose terminal
panicle 15-20 cm long and 5-8 cm wide (without
the corollas), the branches of the inflorescence slen¬
der, ± angular, puberulous, the trichomes short,
erect, sometimes multicellular intermixed with some
gland-tipped trichomes; bracts subtending the flow¬
ers triangular, barely 2 mm long and 1 mm wide
at base, strongly costate, glabrous and ciliolate; brac¬
teoles 1.5 mm long and 0.5 mm wide, in other
respects similar to the bracts; calyx 10-13 mm long,
light green, the segments narrowly lanceolate, 9-
1 1 mm long, about 1 mm wide at base, minutely
apiculate at apex, 3-nerved, puberulous, the tri¬
chomes erect, glandular and eglandular; corolla rose-
lilac, minutely and inconspicuously puberulous, some
of the trichomes gland-tipped, about 45 mm long,
the tube 3 mm wide at base, 8 mm wide at the
mouth, the upper lip erect, ovate, 20 mm long, 11
mm wide at base, bilobed at the tip, the lobes ro-
tundate, 0.5 mm long and 0.75 mm wide, lower lip
somewhat spreading, oblong, 24 mm long and 5 mm
wide, trilobate, the lobes rotundate, 2.5 mm long,
the middle lobe 2.5 mm wide, the lateral lobes 1
mm wide; stamens exserted, white, curved vertically
and exceeding the upper corolla lip, the anther the¬
cae oblique, superposed, the connective 1 mm long
and 0.5 mm wide; pollen 2-porate, prolate to per-
prolate, 51 x 27 jum, trema area not differentiated,
the whole surface of grain microreticulate, studded
with smooth obtuse-conical verrucae (Fig. 8C, D);
capsule clavate, 12 mm long, 4 mm wide, 3.5 mm
thick, acute with each valve bifid at apex, puberu¬
lous; seeds orbicular, flat, 3 mm wide and long,
glabrous.
Distribution. In rocky riverbed, between 1,000
and 1,500 m in Territorio Federal Amazonas, Ven¬
ezuela. Known only from the type locality.
Justicia parimensis superficially resembles J. ca-
lycina (Nees) V. A. W. Graham from the Guianas
and Venezuela. However, in J. calycina the flowers
are borne secund, in spikelike racemes, the latter
in a large, rather dense terminal panicle, the calyx
segments are linear and tapering with a long point,
15-17 mm long, the corolla is brick red, glabrous
and 50-60 mm long, and the valves of the capsule
are glabrous and entire at apex. In contrast, in J.
parimensis the flowers are borne in simple spikes,
these fascicled, forming in aggregate a loose terminal
panicle, the calyx segments are lanceolate, minutely
apiculate at apex, 9-11 mm long, the corolla is
rose-lilac, puberulous and 45 mm long, and the
valves of the capsule are puberulous and bifid at
apex.
Acknowledgments. My special thanks to Alice
Tangerini, who skillfully prepared the line drawings,
80
Novon
and the staff of the National Museum of Natural
History SEM Laboratory for their high-quality pollen
photomicrographs.
Literature Cited
Bremekamp, C. E. B. 1948. Notes on the Acanthaceae
of Java. Verh. Kon. Ned. Akad. Wetensch. Afd.
Natuurk., Tweede Sect. 45(2): 1-78.
Graham, V. A. W. 1988. Delimitation and infra-generic
classification of Justicia (Acanthaceae). Kew Bull.
43: 551-624.
Lindau, G. 1893. Beitrage zur Systematik der Acan-
thaceen. Bot. Jahrb. 18: 36 64, pis. 1, 2.
-. 1895. Acanthaceae. In: A. Engler & K. Prantl
(editors). Die Natiirlichen Pflanzenfamilien, IV(3b):
274-354.
Mabberley, D. 1987. The Plant-Book. Cambridge Univ.
Press, Cambridge.
Nees von Esenbeck, C. G. 1847. Acanthaceae. In: A.
P. de Candolle (editor), Prodromus Systematis N
turalis Regni Vegetabilis, 11: 46-519. Treuttel
Wiirtz, Paris.
Radlkofer, L. 1883. Ueber den systematischen Werth
der Pollenbeschaffenheit bei den Acanthaceen. Aka-
demie der Wissenschaften Munich Math.-Phy. Kl.
Sitzungsberichte 13: 256-314.
Stearn, W. T. 1971. Taxonomic and nomenclatural
notes on Jamaican gamopetalous plants. J. Arnold
Arbor. 52: 636-647.
i =3
Volume 2
Number 2
1992
NOVON vy
New Taxa and Nomenclatural Combinations of
Mesoamerican Grasses (Poaceae)
Gerrit Davidse
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Richard W. Pohl
Department of Botany, 353 Bessey Hall, Iowa State University,
Ames, Iowa 50011, U.S.A.
ABSTRACT. As a result of a review of the grass
family for Flora Mesoamericana, 13 new species
(Arthrostylidium judziewiczii, A. merostach-
yoides, Aulonemia clarkiae, Merostachys latifolia,
Rhipidocladum marlinezii, Guadua macclurei,
Arberella grayumii, Cryptochloa soderstromii,
Pariana argentea, Danthonia chiapasensis, Per-
eilema diandrum, Digitaria breedlovei, D. clavi-
tricha) and three new varieties (Eragrostis rufes-
cens var. mesoamericana, Panicum aquaticum var.
cartagoense, Andropogon gerardii var. honduren-
sis) are described, one new name (Elymus cordil-
leranus) is proposed, 17 new combinations (Guadua
longifolia, Chondrosum hirsutum var. glandulos-
um, Dichanthelium sphaerocarpon var. floridan-
um, D. sciurotoides, D. sciurotis, D. aciculare var.
ramosum, D. dichotomum var. unciphyllum, I).
umbonulatum, D. cordovense, D. pantrichum, Pas-
palidium chapmanii, P. distantiflora, P. leonis,
P. pradana, P. ophiticola, P. subtransiens, P.
utowanaea) are made, and one name (Bromus ex-
altatus) is lectotypified.
As a result of writing the account of the grass
family for volume 6 of Flora Mesoamericana, a
number of new taxa were recognized, and the need
for several new combinations became apparent, these
are published here under the authorship of the per-
son(s) primarily responsible for the account of the
taxon. The taxa are arranged in the taxonomic se¬
quence used in Flora Mesoamericana, which for a
large part follows the scheme of Clayton & Renvoize
(1986).
In the bamboo descriptions we refer to the two
main types of leaves found in bamboos as the culm
leaves and branch leaves. The latter leaf type has
been more commonly known in recent years as
foliage leaves. However, besides the redundancy
implicit in the term foliage leaves, the term branch
leaves, in parallel to culm leaves, is more precise in
describing the location of these leaves.
Bambuseae: Arthrostylidiinae
Arthrostylidium judziewiczii Davidse, sp. nov.
TYPE: Costa Rica. Heredia: primary forest
along Rio San Rafael, Atlantic slope of Volcan
Barva, 10°13'N, 84°05'W, 1,500 m, slender¬
stemmed, scandent and apparently epiphytic
bamboo on mossy trunk of large tree, 3-4 m
above ground, 12 Apr. 1986, M. H. Grayum
7024 (holotype, MO; isotypes, BM, CANB, CR,
F, ISC, K, PMA, SI, SP, US).
Bambusa lignosa caespitosa. Culmi cavi ad 4 m longi,
1-2 mm diametro, scandentes; nodis in medio culmi ramos
singulares gerentibus. Folia culmorum caduca. Folia ra-
morum laminis 6-13 cm longis, 1-2 cm latis, glabris.
Inflorescentiae bilaterales, spicatae terminales et axillares,
terminales 8-14 cm longae; rhachidi recta ca. 1 mm lata,
4-6 spiculas gerenti. Spiculae singulares, 3.5-6.5 cm
longae, sessiles; gluma inferna absenti; gluma superna
Novon 2: 81-110. 1992.
82
Novon
5.5-6.5 mm longa; flosculo infimo sterili; flosculis ferti-
libus 5-6, leinmate 8.0-11.5 mm longo, 9-nervi; callo
piloso trichomatibus 1.0-1.5 mm longis brunneis, lodiculis
3 ciliolatis, ovario apice puberulento, stylus 1, stigmatibus
2. Fructus non suppetens.
Delicate, caespitose, scandent bamboo to 4 m
long. Gulins 1-2 mm diam., hollow, cylindrical,
smooth, glabrous, the wall ca. l A as thick as the
culm diameter; nodal line evident but not expanded,
glabrous. Culm leaves (only 2 available from sec¬
ondary branches) soon stramineous, deciduous;
sheaths puberulent at the base and the lower mar¬
gins; oral setae similar to those of the branch leaves;
blades 2.0-2.5 cm long, 3-4 mm wide, lanceolate,
reflexed, earlier deciduous than the sheath, nar¬
rowed at the base to a pseudopetiole 0.2-0.5 mm
long, glabrous, marginally scaberulous in the upper
half. Branching intravaginal at the lowermost and
uppermost nodes; buds solitary; primary branches
at midculm nodes solitary, typically not rebranching
at the lower nodes and midculm internodes, sparingly
rebranched along their length, rarely with up to 3
branches at the uppermost nodes. Branch leaves 1-
6 per complement; sheaths slightly keeled, mostly
glabrous, puberulent near the base region and along
the overlapping margin; pseudopetiole 1.6-3.0 mm,
flattened, glabrous; external ligule a minute rim;
internal ligule 0.3-1.5 mm long, a glabrous mem¬
brane; oral setae 4.0-9.5 mm long, membranous,
whitish; blades 6-13 cm long, 1-2 cm wide, lan¬
ceolate, glabrous, basally obtuse, apically acuminate,
marginally scaberulous, the midnerve not evident,
the lower surface with evident tessellate venation.
Inflorescences bilateral, spicate, terminal and axil¬
lary from the main leaf complement and sometimes
from bladeless secondary branches; terminal spikes
8 14 cm long, with 4-6 spikelets; peduncle gla¬
brous, mostly not exserted from the subtending
sheaths; rachis ca. I mm wide, straight, glabrous,
strongly nerved, sulcate on the side bearing spike-
lets; axillary spikes similar to the terminal ones but
shorter and with fewer spikelets; uppermost 1 or 2
lateral spikelets subtended by solitary or paired
glumelike, obtuse, ciliolate bracts to 1 mm long;
lowest spikelet sometimes subtended by bracts to 9
mm. Spikelets 3.5-6.5 cm long, sessile, solitary,
linear, with 7-9 florets; lower glume absent; upper
glume 5.5-6.5 mm, asymmetric, 6-nerved, the mid¬
nerve prominently keeled on the back, excurrent as
an awn tip 0.2-1.8 mm long, glabrous or ciliolate
within the same inflorescence, the outer nerve on
the wider portion sometimes slightly keeled, the two
keels meeting the edge of the rachis groove; rachilla
segments 4-7 mm long, flattened, prominently his-
pidulous in the upper l A~H with brownish hairs on
the side opposite the subtending lemma; lowest floret
sterile, the lemma 9-nerved, abaxially tessellate-
nerved, the midnerve prominent, excurrent as an
awn tip 0.5-1.3 mm long, a palea absent or to Vi
as long as and much narrower than the lemma, the
callus glabrous; fertile, bisexual florets 5-6, the lem¬
ma 8.0-1 1.5 mm, 9-nerved with the midnerve ex¬
current as an awn point 0.5-2.0 mm long, glabrous,
marginally scaberulous in the upper Vi, the callus
prominently appressed pilose with hairs 1.0-1.5 mm
long, the hairs initially white, turning brown, the
palea nearly as long as the lemma, membranous, 2-
nerved, 2-keeled, the keels ciliolate; uppermost 1-
2 florets smaller, without flowers; lodicules 3, mem¬
branous, lanceolate, ca. 3-3.5 mm long, forming a
tube around the pistil, ciliolate at apex, sparsely
nerved; ovary with a puberulent apex; stamens 3,
the anthers 3.8-5.1 mm long, yellow; style 1; stig¬
mas 2, long-plumose. Fruit not available.
Paratype. Panama, bocas del TORO: along trail back
to Boquete, in cloud forest, scandent, no larger culms
seen, 6,000 ft., 13 Feb. 1979, Hummel 6173 (MO).
Although the type collection was apparently epi¬
phytic (M. H. Grayum, pers. comm.), this was prob¬
ably an isolated instance as happens sporadically
and accidently with other grasses when their seeds
happen to land on tree trunks and in branch crotches
with a large accumulation of epiphytic plant debris.
Just as at the lowermost and uppermost nodes,
the branching at midculm nodes is probably also
intravaginal, despite the fact that the mostly solitary
branches are predominantly divergent at the base
as in many other scandent bamboos. This habit with
widely spreading branches is doubtlessly important
in enabling these plants to climb into supporting
vegetation. Young culms with culm leaves are not
among the available collections, but it appears that
the culm leaves are quickly pushed away by the
expanding branch.
Arthrostylidium judziewiczii has three unusual
characters, one of which makes placement within
Arthrostylidium Rupr. somewhat problematical.
First, the prominently pilose callus is unusual.
The only other species with hair on the callus is A.
ecuadorense Judz. & L. G. Clark, but in this species
it is much less prominent.
Second, unusual in the genus is the production
of one branch per node that is not rebranched at
its basal nodes. However, at one midculm internode
of Grayum 7024 (MO), the primary branch was
immediately branched at the base. Furthermore, at
occasional nodes near the apex of the culm, the
primary branch may also be branched immediately,
thereby producing a complement of up to four lateral
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
83
branches. The occurrence of occasional basal
branching is judged to be an indication of funda¬
mental similarity to the typical branching pattern in
the genus. Among New World bamboos, basally
unbranched primary branches are characteristic of
Olmeca Soderstrom, Myriocladus Swallen, and most
species of Aulonemia Goudot. All these genera have
panicles in contrast to the unbranched spikes or
racemes in Arthrostylidium (except A. merostach-
yoides with a highly reduced panicle, see below).
Myriocladus is a totally distinct endemic genus from
the Guayana Highlands. Olmeca has fleshy fruits,
erect culm blades, and sympodial rhizomes. Aulo¬
nemia species usually have articulated, reflexed
blades, and in this regard A. judziewiczii appears
to be similar. However, only culm leaves from sec¬
ondary branches with moderately developed blades
were available for study, and it is not certain that
these have the same morphology as typical culm
leaves from lower to midculm nodes from the main
culm. The evidence therefore remains inconclusive.
The spike of A. judziewiczii is quite similar to that
of most species of Arthrostylidium, although most
species differ in having subsessile spikelets with a
small but evident pedicel, thus making the inflores¬
cence a spikelike raceme.
Third, also unusual is the occasional production
of small glumelike bracts at the base of the spikelets,
which, although very small, clearly embrace both
the spikelet base and part of the rachis. These are
therefore homologous to subtending leaves, and the
inflorescences are bracteate spikes. Furthermore,
this is one of the transitional stages postulated for
the origin of normal grass spikelets from pseudos-
pikelets as hypothesized by McClure (1934, 1966).
Additional comments on this topic will be presented
in a separate paper.
The specific epithet honors the outstanding con¬
tributions to agrostology of Emmet J. Judziewicz,
especially his contributions to our knowledge of bam¬
boos.
Arthrostylidium merostachyoides R. Pohl, sp.
nov. TYPE: Costa Rica. Puntarenas: Montev-
erde, 4 Jan. 1985, R. W. Pohl & L. G. Clark
14613 (holotype, ISC; isotypes, CR, F, K, MO,
US). Figure 1.
Merostachys glabra R. Pohl, nom. nud.
Bambusa lignosa caespitosa. Culmi cavi ad 10 m longi
et ad 1.5 cm diametro; nodis in medio culmi ramos 7-
40 gerentibus. Folia culmorum glabra; vaginis persisten-
tibus; laminis erectis, deciduis vel persistentibus. Folia
ramorum laminis 4-17 cm longis, 1.0-2.6 cm latis, adax-
ialiter glabris, abaxialiter pubescentibus secus marginem
unicum. Inflorescentia unilateralis spicata, terminalis, 2.5-
8.0 cm longa; rhachidi recta, 1.5-2.0 mm lata, spiculas
numerosas gerenti. Spiculae plerumque geminatae, 9.5-
14.0 mm longae, subsessiles, glabrae, supra lemma sterile
discedentes; gluma inferna 0.1-2.5 mm longa; gluma
superna 2.0-3.5 mm longa; flosculo infimo sterili, 3.5-
5.0 mm longo; flosculis fertilibus 2-3, lemmate 6.2 8.2
mm longo, 5-7-nervi, lodiculis 3, glabris, antheris 3, 3.2-
4.2 mm longis; ovario glabro, stylo 1, stigmatibus 2.
Fructus non suppetens.
Caespitose, ligneous bamboo. Rhizomes pachy-
morph. Culms 5-10 m long, 1.0-1.5 cm diam.,
arching, hollow, cylindrical, glabrous, yellow. Culm
leaves with the sheaths and blades glabrous; sheaths
persistent; blades soon deciduous or persistent in
older, larger culms, erect, tapering from the base
to the acuminate apex, longer than the sheaths,
lacking auricles; external ligule not evident; internal
ligule ca. 0.5 mm long, a stiff, minutely ciliolate
membrane. Midculm branch complement arising
from a distinct promontory, this ± truncate at the
apex; branches 7-40 per node, to 30 cm long, some
rebranched. Branch leaves 4-7 per complement,
the lower blades deciduous; sheaths truncate, pu-
berulent on the overlapping margin, bearing stiff
auricular bristles 7-16 mm long; pseudopetioles 1.5-
2.5 mm long, flattened, puberulent abaxially; inter¬
nal ligule 0.5-0.6 mm, a ciliolate membrane; blades
4-17 cm long, 1.0-2.6 cm wide, lanceolate-ovate,
pendent, flat, rounded to an asymmetric base, acu¬
minate, glabrous above, pubescent along one margin
below. Inflorescences 2.5-8.0 cm long, unilateral,
reduced racemelike panicles, terminal on leafy or
sometimes bladeless branches; rachis 1.5-2 mm wide,
obtusely angular, narrowly grooved above the point
of spikelet insertion, glabrous. Spikelets 9.5-14 mm
long, usually paired, pectinate in 4 rows along one
side of the rachis, sometimes solitary in 2 rows,
especially toward the tip of the raceme, laterally
compressed, with 4 or 5 florets, subsessile, the ped¬
icels 0.4 1.4 mm long; lower glume 0.1 2.5 mm
long, 1-3-nerved; upper glume 2.0-3.5 mm long,
3-5-nerved; lowest floret sterile, the lemma 3.5-
5.0 mm long, 3-7-nerved; disarticulation above the
sterile floret and between the fertile florets; fertile,
bisexual florets 2-3, the lemmas 6.2-8.2 mm long,
keeled, 5-7-nerved, acute, awnless, the palea sub¬
equal to the lemma; terminal sterile floret as long
as the fertile florets; lodicules 3, ovate, flat, vas-
culated, eciliate; stamens 3, the anthers 3.2-4.2
mm long; ovary glabrous; style 1, slightly thickened
at the base; stigmas 2. Fruit not available.
Paratypes. Costa Rica, alajuela: 17-20 km NNW
of San Ramon, 10°13'N, 84°32'W, 24 Apr. 1983, Lies-
ner & Judziewicz 14771 (MO). GUANACASTE: Parque
Rincon de la Vieja, 10°46'N, 85°49'W, 2 Dec. 1987,
84
Novon
Herrera 1488 (MO). LIm6n: Canton de Limon, cabeceras
de Rio Cariei, 09°47'20"N, 83°08T8"W, 26 Apr. 1989,
Herrera & Chacon 2801 (MO). PUNTARENAS: Monteverde
Forest Reserve, Cordillera de Tilaran, Dec. 1977, Dryer
1708 (ISC, MO), 12 Apr. 1983, Judziewicz 4628 (MO,
WIS), 24 Aug. 1985, Haber 2414 (MO), 2415 (MO),
2416 (CR, EAP, ISC, L, MEXU, MO, PMA, SI, US),
27 Aug. 1985, Haber & Bello 2481 (MO), 12 Feb.
1985, Grayum el al. 5126 (CR, INPA, ISC, K, MO,
US), 4 Jan. 1985, Pohl & Clark 14611 (ISC, MO).
Arthrostylidium merostachyoides is presently
known from cloud forests at elevations from 750 to
1,850 rn in the Cordillera de Tilaran, Cordillera de
Guanacaste, and one locality of the Atlantic slope
of the Cordillera de Talamanca. This distribution
suggests that it may also eventually be found in
Panama.
Seedlings were abundant on road embankments
and cliffs at the type locality in the Monteverde
Nature Reserve. Some had reached 1 m height and
had short rhizomes. They evidently grew from seeds
produced in previous seasons.
Although this species clearly belongs to the Ar-
throstylidiinae Soderstrom & Ellis (1987), its ge¬
neric disposition is less certain, a point also discussed
by Clark & Londoho (1991). The inflorescence
structure is unique in Arthrostylidium and initially
caused us to consider this a species of Merostachys
Sprengel. The epithet refers to this similarity. How¬
ever, A. merostachyoides possesses none of the
other diagnostic characters traditionally (McClure,
1973) ascribed to Merostachys. Specifically, its culm
blades are broad at the base and appressed to the
culm (vs. narrow, shortly pseudopetiolate and re¬
flexed); the midculm branching complement arises
from a truncate promontory (not a triangular plate);
the spikelets have a developed lower glume (vs.
absent or reduced to a rim); and the palea keels are
widely spaced and do not tightly enclose a long
rachilla internode with a spikelet rudiment.
The strongly secund inflorescence resembles that
of some Merostachys species in which the spikelets
are paired on each side of the rachis. However, the
grooves in the rachis above each point of spikelet
insertion are much narrower in A. merostachyoides
than in Merostachys species, and the raceme, there¬
fore, appears less flattened on the spikelet-bearing
side. In contrast, all previously described Arthros¬
tylidium species have bilateral spikes or racemes,
some of which are strongly zigzagged. Nevertheless,
the preponderance of diagnostic characters favors
a disposition outside of Merostachys.
Arthrostylidium and Rhipidocladum McClure
are very closely related but are distinguished pri¬
marily by their midculm branch complements, which
arise from a triangular plate in Rhipidocladum.
The inflorescence type of A. merostachyoides is
very similar to that of Rhipidocladum geminatum
(McClure) McClure, but the species seems otherwise
not closely related. Rhipidocladum as currently
circumscribed has three other inflorescence types,
including a zigzag raceme more common among
Arthrostylidium species (McClure, 1973; Clark &
Londoho, 1991).
It should also be noted that only the collections
from the Monteverde area show predominantly paired
spikelets on each side of the rachis, although toward
the tip there is often only one spikelet per node.
However, even in these collections, some of the
smaller inflorescences borne on leafless branches
may only have one spikelet per node on each side
of the rachis. Such inflorescences with solitary spike¬
lets at the rachis nodes are nearly identical to those
of Rhipidocladum maxonii (A. Hitchc.) McClure
in organization, but are much larger.
In contrast, in Herrera & Chacon 2801 and
Herrera 1488 all inflorescences bear one spikelet
per node on each side of the rachis. In occasional
inflorescences the lowest node of the inflorescence
bears an elongated branch with up to eight spikelets.
Since the pedicels of “typical'’ spikelet pairs in plants
from the Monteverde region are of slightly different
length, we interpret these to represent very reduced
branches and the inflorescence, therefore, to rep¬
resent a reduced secund panicle. Control of inflo¬
rescence development is evidently not stabilized in
this species.
We agree with Clark & Londoho (1991) that it
is likely that some characters traditionally employed
to define genera in the Arthrostylidiinae, especially
inflorescence type, may have evolved independently
several times in this subtribe and that generic de¬
limitations must be reconsidered.
Aulonemia clarkiae Davidse & R. Pohl, sp. nov.
TYPE: Mexico. Chiapas: Municipio Jitotol, 5
km SE of Jitotol along road to Bochil, in trees,
open forest with Pinus, Quercus, Nyssa , Liq-
uidambar, and Brunellia, culms to 8 m tall,
1,600 m, 8 Nov. 1981, D. E. Breedlove &
G. Davidse 55085 (holotype, MO; isotypes,
CAS, US). Figure 2.
Bambusa lignosa rhizomatibus 10-30 cm longis et 6-
15 mm crassis. Culmi 5-8 m longi, 1.0-1.6 cm diametro;
nodis in medio culmi ramos singulares gerentibus. Folia
culmorum vaginis persistentibus; setis oralibus abundan-
tibus et prominentibus; laminis reflexis, cauducis. Folia
ramorum laminis 10 36 cm longis, 9-15 mm latis, adax-
ialiter scabris, abaxialiter puberulis. Inflorescentia pani-
culata, terminalis, 10-33 cm longa. Spiculae 19~25 mm
longae; gluma inferna 3-5 mm longa arista 2.0-5.5 mm
longa munita; gluma superna 5.0-7.2 mm longa arista
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
85
Figure 1. Arthrostylidium merostachyoides R. Pohl. —A. Young culm with culm leaves, the blades erect. —B.
Flowering midculm branch complement. —C. Enlargement of a midculm branch complement showing the conspicuous
promontory. —D. Rachis segment with two pairs of spikelets. —E. Spikelet. (A, B based on Pohl & Clark 14613;
C based on Pohl Clark 14612; D, E based on Haber 2414.)
2.5-4.5 mm longa munita; flosculis 2; flosculo inferno
plerumque sterili interdum fertili, 9.5-10.5 mm longo
arista 3.0-5.5 mm longa munito; flosculo superno fertili
lemmate 13-15 mm longo, indimidio superiore pubescenti
arista 4-5 mm longa munito, lodiculis 3, ciliatis, antheris
3, 8.5-9.0 mm longis, ovario glabro, stigmatibus 2; rhach-
illa ultra flosculum terminalem producta. Fructus non
suppetens.
86
Novon
Figure 2. Aulonemia clarkiae Davidse & R. Pohl. —A. Flowering branch complement. —B. Midculm nodal
complex with a single primary branch. —C. Ligular region of two branch leaves showing the prominent oral setae.
— D. Spikelet. (A. C, D based on Breedlove 58512; B based on Breedlove & Davidse 55085.)
Caespitose, ligneous bamboo. Rhizomes 10-30
cm long, 6-15 mm thick, densely covered by scale
leaves. Culms 5-8 m long, 1.0-1.6 cm diam., erect,
solitary, hollow, cylindrical, glabrous, green, with
thin walls, the upper portion pendent, producing
branches in the upper 3 A. Culm leaves with long-
persistent, stramineous, glabrous to papillose-pilose
sheaths; auricles not developed; oral setae well-de¬
veloped, 8-15 mm long, the central ones flattened
at the base and borne on the back of the ligule, the
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
87
marginal ones cylindrical at the base and borne on
the sheath apex; external ligule a minute rim; in¬
ternal ligule a membrane 0.8-2.0 mm long, gla¬
brous; blades reflexed to spreading, soon deciduous,
narrower than the apex of the sheath, ciliolate.
Branches intravaginal, 1 per node, the primary
branches usually rebranching once or the larger
midculm primary branches forming a fascicle of
branches at the base. Branch leaves 4-8 per com¬
plement; sheaths of the lower leaves of a complement
usually prominently papillose-hirsute with reddish
brown, divergent hairs 1.5-3.0 mm long, ciliate with
finer hairs on the margins, the upper sheaths of a
complement puberulent or glabrous; external ligule
a rim 0.1-0.2 mm long, sometimes irregular and
to 0.8 mm long at the margin of the pseudopetiole;
internal ligule a membrane 0.5-1.0 mm long, mi¬
nutely ciliolate; auricles to 1 mm long on one side,
poorly or not developed or absent on the other side;
oral setae 7-15 mm long, prominent on each side
of the pseudopetiole; pseudopetiole hardly differ¬
entiated; blades 10-36 cm long, 9-15 mm wide,
linear-lanceolate, basally attenuate, apically long-
attenuate, retrorsely scabrid above, puberulent and
weakly tessellate below. Inflorescence a panicle 10
33 cm long, terminal on leafy branches; peduncle
not exserted; rachis ± triangular, puberulent and/
or scabrous; primary branches to 15 cm long, sol¬
itary, widely divergent at maturity, prominently pul-
vinate, the pulvini swollen, brown, puberulent, the
lowermost branch often subtended by a small bract;
secondary branches few, primarily in the lower part
of the panicle, pulvinate and divergent; pedicels
glabrous, appressed to the panicle branches, the
lateral ones 2-8 mm long. Spikelets 19-25 mm
long, linear-lanceolate, 2-flowered with a prominent
rachilla extension beyond the terminal floret; dis¬
articulation above the glumes and between the flo¬
rets; glumes 2, unequal, shortly awned, marginally
ciliate in the upper V 2 , the lower 3-5 mm long, 3-
nerved, lanceolate, the awn 2.0 5.5 mm long, the
upper 5.0-7.2 mm long, 5-7-nerved, lanceolate,
the awn 2.5-4.5 mm long; lower floret usually ster¬
ile, rarely fertile and then similar to the upper floret;
lower lemma 9.5-10.5 mm long, 1 1-13-nerved,
marginally ciliate on the upper V 2 , glabrous on the
back, awned, the awn 3.0-5.5 mm long; lower palea
usually 1.5-2.0 mm long, pubescent, subtending a
rudimentary bisexual flower with stamens to 0.5 mm
long and a minute pistil, rarely fully developed and
similar to the upper palea and subtending a lully
developed bisexual flower; rachilla internode be¬
tween the florets 2.5-6.0 mm long, thickened up¬
ward, puberulent; upper floret bisexual; upper lem¬
ma 13-15 mm long, 11-13-nerved, tessellate-nerved
in the upper V\, marginally ciliate in the upper V 2 ,
puberulent on the back and/or on the sides in the
upper V 2 and with the base shortly appressed-pu-
bescent, awned, the awn 4-5 mm long; upper palea
as long as the lemma, 4-nerved, 2-keeled and nar¬
rowly grooved between the keels, puberulent be¬
tween and on the keels, glabrous or pubescent in
the upper V 2 on the sides; rachilla extension 9.0-
12.5 mm long, tightly held in the groove between
the keels, without or with a minute, undifferentiated
rudimentary floret 0.1 0.2 mm long, puberulent in
the upper half and thickened toward the apex; lod-
icules 3, the 2 anterior ones 2.0-2.5 mm long,
lanceolate, flat, slightly thicker at the base, ciliate
in the upper V 2 , obscurely nerved, the posterior one
slightly smaller; stamens 3, the anthers 8.5-9.5 mm
long; ovary cylindrical-fusiform, glabrous; stigmas
2. Fruit not available.
Paratypes. MEXICO. CHIAPAS: Municipio of Jitotol, 5
km SE of Jitotol along road to Bochil, open forest with
Pinus, Quercus, Nyssa, Liquidambar, and Brunellia,
1,600 m, 0 Jan. 1981, Breedlove & Keller 49359 ( CAS),
10 Sep. 1981, Breedlove 52658 (CAS, MO), 24 Feb.
1982, Breedlove 5812 (CAS, MO, US).
This species is definitely known only from a single
population in Chiapas, Mexico. It was first observed
by Dennis E. Breedlove of the California Academy
of Sciences, who discovered it coming into flower
and who observed and collected it intermittently over
a period of 16 months. Davidse revisited the locality
in November 1984 and could not find a trace of
the plants, neither adult plants nor seedlings, from
which we tentatively conclude that this population
died after flowering synchronously and did not pro¬
duce viable seed.
This species was briefly discussed by Soderstrom
(1988) when he described Aulonemia fulgor Sod¬
erstrom from Oaxaca. Soderstrom (pers. comm.)
believed it to be a closely related, undescribed spe¬
cies, but his untimely death prevented him from
pursuing his study of these collections to completion.
Aulonemia clarkiae differs from A. fulgor in its
thicker and longer rhizomes, and, consequently, more
diffuse spacing of the culms; taller (5-8 m vs. 5-
6 m) and wider (1.0-1.6 cm vs. 0.5-1.0 cm), more
strictly erect culms; puberulent versus glabrous low¬
er leaf surfaces; usually sterile versus always fertile
lower floret; pubescent versus glabrous upper lem¬
ma; and longer (8.5-9.5 mm vs. 6.0-7.0 mm) an¬
thers.
Soderstrom (1988: 29, fig. 2) discussed and il¬
lustrated the rhizome system of the type collection
of A. fulgor in detail. However, as noted by him,
the description applies primarily to those decumbent
culms that root at the nodes and may at times form
88
Novon
rhizomes. At Soderstrom’s request, Davidse restu¬
died the main rhizome system of A. fulgor in the
field (Davidse et al. 30258) and found, as had been
supposed by Soderstrom, that it is a sympodial sys¬
tem. In A. fulgor the slender, pachymorph rhizomes
are generally 4-7 cm long before they turn upward
to produce a new culm. Well-developed plants pro¬
duce 20-30 loosely clumped culms. Those toward
the outside of the clump tend to he more decumbent
and more often root at the lower nodes. Although
the basic rhizome system is similar in A. clarkiae,
the rhizomes are much longer and thicker, and thus
the culms are widely spaced and not truly clump¬
forming.
Soderstrom (1988: 30) characterized the plants
that we are naming A. clarkiae as having three
glumes and also noted that Aulonemia , sensu Mc¬
Clure (1973), has species with two or three glumes.
The species pair here discussed clearly demonstrates
that it is better to interpret the so-called three-
glumed species as having two glumes and a sterile
lemma in which the enclosed palea and flower have
been totally lost through reduction. This becomes
apparent from the following comparison. In A. ful¬
gor all spikelets have two fully developed florets
above the two glumes (Soderstrom, 1988). In con¬
trast, in A. clarkiae two fully developed florets are
rare. Much more common are two florets in which
only the upper one is fully developed and in which
the lower one usually has a fully developed lemma
subtending a rudimentary palea and flower. By ho¬
mology, other species of Aulonemia with a third
empty bract represent a further step in this reduction
series, namely, one in which the lowermost floret
has been consistently reduced to a lemma.
For the reasons enumerated by Soderstrom (1988)
we follow him in tentatively classifying this species
in Aulonemia, even though the genus, as circum¬
scribed by McClure (1973) and accepted by Cal¬
deron & Soderstrom (1980), is probably hetero¬
geneous and is likely to be polyphyletic. As one of
the discordant elements in a widely circumscribed
Aulonemia, the A. fulgor-A. clarkiae species pair
may deserve to be segregated as a distinct genus.
It is another example of the need for an overall
reassessment of generic classification mentioned in
the discussion of Arthrostylidium merostachyoides.
The species is named for Lynn G. Clark, Iowa
State University, in recognition of her outstanding
contributions to agrostology, especially bamboos and
Chusquea in particular.
I he bamboo genus Merostachys is closely related
to Rhipidocladum, both genera having branch com¬
plements arising from a triangular, flat, adnate mer-
istem in a fan-shaped array. They are conventionally
distinguished by the reflexed culm blade in Meros¬
tachys and the erect blade in Rhipidocladum (Mc¬
Clure, 1973). While this structure is not known for
the species described below, from the form of the
inflorescences and nature of the spikelets, it is clearly
a new species of Merostachys.
Merostachys latifolia R. Pohl, sp. nov. TYPE:
Nicaragua. Jinotega: W. D. Stevens J. Hen-
rich 20418 (holotype, ISC; isotypes, BM, CAS,
COL, CR, EAP, F, HNMN, K, L, MEXU, MO,
NY, PMA, RB. SI, SP, US, VEN). Figure 3.
Bambusa lignosa caespitosa. Culmi 1.5-4 m longi, ad
8 mm diametro; nodis in medio culmi ramos ad 10 ger-
entibus. Folia culmorum ignota. Folia ramorum in com-
plemento 2-6, laminis 15-20 cm longis, 2.8-4.9 cm
latis, ovatis. Inflorescentia unilateralis, spicata, terminalis,
5-7 cm longa; rhachidi arcuata, ca. 1 mm lata, velutina.
Spiculae singulares vel geminatae, falcatae, 10-13 mm
longae; gluma inferna absenti; gluma superna 1.5-3.5
mm longa; flosculo inferno sterili, 4.7-7.0 mm longo ex
lemmate 7-nervi tantum constanti; flosculo superno fertilii
lemmate 9-10 mm mutico; palea superna anguste sulcata
inter carinas; rhachilla ultra flosculum terminale per 6.5-
10.0 mm producta, in flosculum abortivum minutum ter-
minanti. Flosculi sub anthesi etiam fructus non suppen-
tentes.
Caespitose, ligneous bamboo. Rhizomes pachy¬
morph. Culms 1.5-4.0 m long, to 8 mm diam.,
hollow, cylindrical, mottled with purple, asperous
below the nodes; nodes retrorsely bearded. Culm
leaves not known. Branches to 10 per node, to 45
cm long, including the inflorescence. Branch leaves
2-6 per complement; sheaths glabrous to puberu-
lent; auricular bristles prominent, 10-15 mm long,
also borne for several mm along the uppermost
portion of the overlapping sheath margin; internal
ligule 0.1-0.6 mm long, ciliolate; pseudopetiole 6-
10 mm long, flattened; blades 15-20 cm long, 2.8-
4.9 cm wide, ovate, flat, dark green, glabrous but
scabrid along one margin on the upper surface.
Inflorescences 5-7 cm long, spikelike, terminal on
leafy and bladeless lateral branches, slightly arcuate;
rachis ca. 1 mm wide, densely velutinous. Spikelets
10-13 mm long, diverging from the rachis, solitary
or paired, slightly falcate, slightly laterally com¬
pressed; lower glume absent; upper glume 1.5-3.5
mm long, triangular, 1-nerved; lower floret sterile,
consisting only of a lemma 4.7-7.0 mm long, gla¬
brous, 7-nerved; upper floret fertile, bisexual; upper
lemma 9.0-10.0 mm long, 11-nerved, awnless,
sparsely puberulent or shortly pubescent; upper pa¬
lea 9.2-10.5 mm long, slightly longer than the
lemma, narrowly grooved between the keels and
tightly clasping the slender rachilla extension; rach-
5 mm „ 5 cm
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
89
Figure 3. Merostachys latifolia R. Pohl. —A. Flowering midculm branch complement. —B. Rachis segment with
two spikelet pairs. —C. Fertile floret with rachilla extension bearing a minute rudimentary floret. (Based on Moreno
16367.)
90
Novon
ilia extension 6.5-10.0 mm long, bristlelike, tipped
with a minute rudimentary floret. Flowers and fruit
not available.
Paratypes. Honduras, olancho: Finca Santa Rosita,
a 5 km al S del Campamento, 1 June 1985, Tellez V.
<£ Martinez S. 8507 (MO). Nicaragua, matagalpa: NW
slope of Cerro El Picacho, ca. 13°00'N, 85°55'W, 25
May 1983, Stevens & Moreno 22117 (CR, HNMN, ISC,
MO, US), jinotega: km 146 carretera MatagalpaJino-
tega, 13°02'N, 85°56'W, 16 May 1982, Moreno 16367
(HNMN, MO, US). Costa Rica, puntarenas: Zapotel,
Montes de Oro, May 1961, Jimenez s.n. (F, ISC, MO).
ALAJUELA: La Palma de San Ramon, 17 Apr. 1927, Brenes
5433 (F).
Additional sterile collections. Guatemala. EL PRO¬
GRESO: Montana Canahui, between Finca San Miguel and
summit of mountain, near upper limits of Finca Caieta,
10 Feb. 1942, Steyermark 43772 (F). Nicaragua.
JINOTEGA: along Hwy. 3 from Jinotega to Matagalpa, ca.
5-8 mi. SW of Jinotega, 7 Aug. 1977, Croat 43057
(MO). Panama. cocl£: El Valle, 17 Jan. 1973, Soder-
strom 2011 (MO). DARlkN: along ascent of Serram'a de
Pirre, above Cana Gold Mine, between Rio Cana and Rio
Escucha Ruida, 27 July 1976, Croat 37779 (MO).
This species grows in the understory of cloud
forests at 1,100-2,300 m. The type collection and
all the paratypes are post-anthesis specimens. The
cited sterile collections are also tentatively identified
as this species, but we await the collection of fertile
specimens from these areas for confirmation. Other
sterile collections of Merostachys , in addition to the
cited ones, are known from Mesoamerica, but they
probably represent other species and are too frag¬
mentary for positive identification.
This is the species described by McClure in Swal-
len (1955: 207) under the name M. argyronerna
Lindman (1900). This Brazilian species differs in
having branch leaf blades 9-12 (not 15-20) cm
long and 1.2-1.6 (not 2.8-4.9) cm wide, up to 29
(not to 10) branches at the midcuhn nodes, 7-8
(not 2-6) branch leaves per complement, solitary
(not paired) spikelets, and lower lemma 5-nerved
(not 7-nerved). Lindman did not indicate either in
his description or illustration the presence of the
prominent elongated rachilla internode clasped by
the palea in spikelets of M. latifolia, but this is
characteristic of every known species in the genus.
Fhe genus is primarily Brazilian with more than
30 species, including a number of undescribed ones.
Among the broad-bladed species with strongly uni¬
lateral racemes with primarily paired spikelets, M.
latifolia resembles M. riedeliana Rupr. but has
much shorter spikelets (10-13 vs. ca. 18 mm), and
shorter, acute sterile lemmas (4.7-7.0 vs. 12-14
mm and short-awned). However, the genus is so
little understood that it is difficult to assess relation¬
ships.
Khipidocladum martinezii Davidse & R. Fold,
sp. nov. TYPE: Mexico. Chiapas: Mun. Union
Juarez, en el volcan Tacana, a 5 km al S de
Talquian, 1,550-1,700 m, bosque mesofilo de
montana, arbusto voluble, flor amarilla con es-
tambres morados y fruto, 8 Feb. 1987, E.
Martinez S., A. Marquez, G. Urquijo & M.
Ramirez 19767 (holotype, MO; isotype,
MEXU). Figure 4.
Bambusa lignosa caespitosa. Culmi 8 16 mm diametro;
nodis in medio culmi ramos 100-200 gerentibus. Folia
culmorum ignotae. Folia ramorum laminis 4-8 cm longis,
2.5-3.5 mm latis. Inflorescentia unilateralis racemosa,
terminalis, 2-4 cm longa, glabra, spiculas 2-4 gerens.
Spiculae 25-30 mm longae; gluma inferna 2.5-7.0 mm
longa arista 2-3 mm longa munita; gluma superna 6-11
mm longa arista 2.5-3.5 mm longa munita; floscufo infimo
sterili, lemmate 12 15 mm longo, arista 3-5 mm longa;
flosculis fertilibus 2-4, lemmate 12-19 mm longo, sca-
berulo, arista 3-6 mm longa, lodiculis 3, glabris, antheris
3, 6-7 mm longis, stylo 1, stigmatibus 2. Caryopsis 5.5-
6.1 mm longa, 1.0-1.2 mm lata; hilo caryopside aequi-
longo.
Caespitose, ligneous bamboo. Rhizome pachy-
morph. Culm length unknown; internodes 8-16 mm
diam., relatively thin-walled, hollow, cylindrical, gla¬
brous. Culm leaves unknown. Primary branch ele¬
ment flat, triangular, adnate to the main culm, bear¬
ing 100 200 flowering branches at midculm nodes;
flowering branches 10-30 cm long, glabrous, leafy
or leafless toward the tips of the main culm, branched
or unbranched, thin and delicate, the leafless branch¬
es mostly 4-5 cm long, bearing 2-4 spikelets. Branch
leaves 2 5 per complement; sheaths glabrous, trun¬
cate; oral setae 1.5-2.5 mm long, flexuous; external
ligule a rim; internal ligule 0.1-0.3 mm long, mem¬
branous, ciliolate; pseudopetiole 1.0-1.3 mm long,
0.2-0.3 mm wide, glabrous; blades 4-8 cm long,
2.5-3.5 mm wide, linear-lanceolate, flat, sparsely
puberulent at the base below, often scaberulous near
one margin above, otherwise glabrous. Inflorescence
a solitary, unilateral raceme bearing 2-4 spikelets;
peduncle scarcely exserted, puberulent; rachis 2 4
cm long, glabrous, the internodes 8-15 mm long;
pedicels to 0.5 mm long, appressed. Spikelets 25-
30 mm long, appressed, disarticulating above the
lowest, sterile floret and between the 2-4 fertile,
bisexual florets; florets 4-7, the lowest sterile, the
uppermost 1 or 2 staminate or sterile, the terminal
one usually rudimentary; lower glume sometimes
lacking entirely in terminal spikelets, when present
usually 2.5-7.0 mm long including a 2-3-inm-long
awn, narrowly lanceolate, 1 -nerved; upper glume
6-11 mm long including a 2.5-3.5-mm-long awn,
lanceolate, 5-7-nerved, glabrous; rachilla inter¬
nodes 2.7-4.5 mm long, glabrous; lower floret re-
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
91
Figure 4. Rhipidocladum martinezii Davidse & R. Pohl. —A. Leafy flowering midculm branch complement.
B. Leafless, flowering branch complement. —C. Spikelet. (A based on Martinez et al. 20286; B, C based on Martinez
et al. 19767.)
5 cm
92
Novon
duced to a lemma 12-15 mm long including a 3-
5-mm-long awn, lanceolate, 7-9-nerved, scaberu-
lous on the upper %; fertile lemmas 12-19 mm
long including a 3-6-mm-long awn, lanceolate, at¬
tenuate, 7-9-nerved, scaberulous in the upper 34;
fertile paleas 9.5-10 mm long, as long as or dis¬
tinctly shorter than the fertile lemmas, 4-6-nerved,
2-keeled, ciliolate in the upper 14 on the keels; callus
knoblike; lodicules 3, the two posterior ca. 2 mm
long, flat, somewhat thickened at the base, lanceolate
but with one side straight, glabrous, the upper mar¬
gin sparsely ciliolate, the anterior one narrower,
linear; stamens 3, the anthers 6-7 mm long; style
1; stigmas 2; ovary linear-lanceolate. Caryopsis 5.5-
6.1 mm long, 1.0-1.2 mm wide, flattened and slight¬
ly grooved on the hilar side, rounded on the embryo
side, dark brown; embryo ca. 1 mm long; hilum as
long as the caryopsis.
Paratypes. Mexico. CHIAPAS: Mun. Union Juarez, Vol¬
can Tacana, 5 km al S de Talquian, 8 Feb. 1987, Mar¬
tinez et al. 19813 (MEXU, MO); Mun. Union Juarez,
Volcan Tacana, 5 km al E de Talquian, 26 abr. 1987,
Martinez S. & Reyes 20286 (MEXU, MO).
This species is known only from the type locality
on Volcan Tacana, the large volcano straddling the
Mexican-Guatemalan border.
This species is most closely related to Rhipido-
cladum clarkiae R. Pohl, R. panamense R. Pohl,
and R. sibilans Davidse, Judz. & L. Clark, narrow
endemics of Costa Rica, Panama, and eastern Ven¬
ezuela and western Guyana, respectively. All four
species occur in montane forests. They may be more
widespread than present evidence indicates since
their infrequent flowering has no doubt discouraged
collectors. They may have arisen from a common
widespread ancestor that differentiated through iso¬
lation in widely separated mountain ranges or, per¬
haps more likely, they differentiated as new isolated
populations from a common ancestor as it spread
into previously uninhabited mountain ranges.
Rhipidocladum martinezii resembles R. clar¬
kiae, R. panamense, and especially R. sibilans in
its numerous branches per node, small leaf blades,
and obviously awned spikelets. The new species dif¬
fers from R. clarkiae in its smaller inflorescences
bearing fewer (2-4 vs. 7-8) spikelets per raceme
with glabrous versus puberulent rachises, more at¬
tenuate. less heavily scabrous lemmas, longer (6-7
vs. ca. 4.5 mm) anthers, and narrower (2.5-3.5 vs.
4-9 mm) branch leaves. Rhipidocladum marti¬
nezii differs from R. panamense in its smaller in¬
florescences bearing fewer (2-4 vs. 5-7) spikelets
per raceme, longer (9.5-12.0 vs. 5.0-6.5 mm)
much more attenuate lemma body, and longer an¬
thers. Rhipidocladum martinezii differs from R.
sibilans in its secund (vs. bilateral) raceme with the
spikelets more closely spaced (rachis internodes 0.8-
1.5 vs. 1.9-3.0 cm long) and glabrous (vs. densely
puberulent) rachilla internodes.
This species is named in honor of Esteban Mar¬
tinez S., Universidad Nacional Autonoma de Mexico,
a prolific plant collector and discoverer of many
interesting new plants for the flora of Mexico, in¬
cluding all known collections of this new species.
BaMBUSEAE: ClIADUINAE
We follow Soderstrom & Londono (1987) and
Soderstrom & Ellis (1987) in recognizing New World
Guadua Kunth as distinct from the Old World Bam-
busa Schreber, thus necessitating the following new
combination.
Guadua longifolia (Fourn.) R. Pohl, comb. nov.
Basionym: Arundinaria longifolia Fourn.,
Mexic. FI. 2: 131. 1886. Bambusa longifolia
(Fourn.) McClure, Smithsonian Contr. Bot. 9:
66. 1973. TYPE: Mexico. Jicaltepec, April,
Liebmann s.n. (holotype, C).
Arthrostylidium spinosum Swallen, J. Wash. Acad. Sci.
28: 6. 1938. Guadua spinosa (Swallen) McClure,
Phytologia 5: 82. 1954, non Bambusa spinosa Roxb.
ex Buch.-Ham., Trans. Linn. Soc. London 13: 480.
1822. Bambusa sivalleniana McClure, Smithsonian
Contr. Bot. 9: 68. 1973. TYPE: Belize. El Cayo
District: along Belize River at El Cayo, June Aug.
1936, Lundell 6939 (holotype, US; isotype, MO).
Guadua macclurei R. Pohl & Davidse, sp. nov.
TYPE: Honduras. Gracias a Dios: along Rio
Platano S of Ras, Moravian Mission settlement
called Las Marias or Batituk, elev. 50 m, culms
over 10 m long, arching, with numerous elon¬
gate spreading branches, this colony vegetative
except for one branch 1.5 m long, species is
abundant along lower Rio Platano, some col¬
onies blooming, 16 Dec. 1977, R. If’. Pohl &
M. Gabel 13340 (holotype, ISC; isotypes, F,
MO). Figure 5.
Bambusa lignosa spinosa. Culmi cylindrici cavi, 6-15
m longi, 3-4 cm diametro; nodis supra et infra cristam
nodalem annulos trichomatum alborum gerentibus, in me¬
dio culmi ramos singulares gerentibus. Folia culmorum
vaginis hispidis plerumque exauriculatis interdum auri-
culato-setosis, deciduis; laminis 7-14 cm longis adaxialiter
hispidis trichomatibis plerumque albis. Folia ramorum va¬
ginis auriculis 5.0-8.5 mm longis, lineari-falcatis pler¬
umque munitis; laminis 13-26 cm longis, 1.6-4.8 cm
latis, lanceolatis. Inflorescentia rhachidi gracili efoliata
usque ad 30 cm longa fasciculos densos numerosos pseu-
dospicularum sessilium gerenti. Pseudospiculae 10-15 mm
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
93
Figure 5. Guadua macclurei R. Pohl & Davidse. —A. Vegetative branch complement. —B. Culm leaf. —C.
Branch leaf detail showing prominent auricles. —D. Flowering branch with clusters of pseudospikelets. —E. Flowering
branch originating from a pseudospikelet. —F. Spikelet. (A-C based on McClure 21476; D-F based on Pohl &
Gabel 13340.)
94
Novon
longae, ca. 3 mm latae; flosculo fertili plerumque unico;
lemmate 8.5-9.0 mm longo; lodiculis 3, puberulentis;
antheris 6, ca. 5 mm longis; stigmatibus 3. Fructus non
suppentens.
Caespitose, ligneous, spiny bamboo. Rhizomes
short, pachymorph. Culms 6-15 m long, 3-4 cm
diam., arching, pendulous above; internodes to 55
cm long, green, sparsely appressed pubescent, hol¬
low, cylindrical, thin-walled; nodes not prominent,
with white pubescent bands above and below the
nodal ridge. Culm leaves tardily deciduous; sheaths
7-23 cm long, appressed-hispid with white hairs,
usually lacking auricles and auricular setae, rarely
with linear auricles to 9 mm long bearing prominent
setae; internal ligule 1.5-2 mm long, stiff, mem¬
branous, minutely ciliolate; blades 7-14 cm long,
as broad as the sheath apex, erect, broadly trian¬
gular, apiculate, ciliolate, densely appressed-hispid
above, sparsely hispid toward the margins below,
the hairs usually white or sometimes brown toward
the base, the base long-decurrent on one margin.
Branch complement consisting of a single primary
branch, the lowest nodes bearing 1 several hooked
thorns or secondary branches. Branch leaves with
the sheaths ciliate, glabrous to hispid on the back;
auricles 5-8.5 mm long, linear-falcate, sometimes
deciduous or absent; auricular setae 5-9 mm long;
external ligule a rim 0.1 -0.3 mm long; internal ligule
0.1-0.4 mm long, membranous; pseudopetiole 1.5-
2.5 mm long, pilose above; blades 13-26 cm long,
1.6-4.8 cm wide, flat, lanceolate, mostly glabrous,
ciliolate. Inflorescence axis to 30 cm, slender, leaf¬
less, bearing numerous dense fascicles of sessile
pseudospikelets. Pseudospikelets 10-15 mm long,
ca. 3 mm wide; basal bracts 2-4, 2-5.5 mm long,
similar to the glumes, often subtending axillary pseu¬
dospikelets in several orders; each pseudospikelet
terminating in a spikelet that disarticulates below its
glumes; lower glume 6.0-7.5 mm long, ovate, 11-
13-nerved, the apex with a stiff beak ca. 1 mm
long; upper glume 8.0-8.7 mm long, similar to the
lower glume; bisexual floret usually 1; lemma 8.5-
9.0 mm long, 11 — 15-nerved, broad, apiculate,
densely puberulent adaxially in the upper ] A; palea
broad, outwardly bowed, 8-10-nerved, about as long
as the lemma, 2-keeled, the margins each 3-nerved,
the keels broadly winged, the wings to 1.5 mm wide,
eciliate, completely enfolding the terminal rachilla
internode; rachilla stiff, 3-5 mm long, bearing a
cylindrical rudiment 3.5-5 mm long; lodicules 3,
flat, vaseulated, puberulent on the back; stamens 6,
the anthers ca. 5 mm long, yellow; stigmas 3. Fruit
not available. 2 n = 48.
Paratypes. Nicaragua, zelaya: Bluefields, El Recreo,
Agricultural Experiment Station, growing wild, forming
isolated colonies of limited extent on upland in quebradas
and second-growth jungle, branches all thorny, culms used
locally in construction of shade shelters for nursery plants,
known locally as “Cuajichote,” 23 Jan. 1946, McClure
21476 (ISC,' MO, US), 25 Jan. 1946, McClure 21479
(MO); road crossing of Rio Likus, 1.5 m SE of La Tron-
quera, S of Waspam, 6 Aug. 1971, Pohl & Erickson
12700 in part (MO-2613073). Honduras, gracias a
DIOS: Leymus, orilla del rio Segovia o Wanki, 100 km
SO de Puerto Lempira, 1 Feb. 1984, Nelson <£ Cruz
8696 (MO).
Additional sterile collections tentatively identified as this
species and not considered to be paratypes nor used in
writing the description are the following: NICARAGUA.
zelaya: ca. 0.5 km upstream from first suspension bridge
over Rio Sucio, also known as Rio Tunky, E of Rosita,
ca. 13°58'N, 84°28'W, 22 Feb. 1979, Stevens 12606
(MO); along new road between Rosita and Puerto Cabezas,
ca. 15.7 kmSWof Rio Kukalaya, ca. 13°58'N, 84°12'W,
30 Apr. 1978, Stevens 8464 (MO); ca. 3.9 km SE of
El Empalme on road to Limbaika, ca. 13°39'N, 84°24'W,
25 Apr. 1978, Stevens 8122 ( MO). Costa Rica, puntar-
ENAS: Piedras Blancas, edge of road in lowland forests
along Carretera Interamericana, 19 July 1966, Pohl &
Calderon 10103 (MO). Panama, barro Colorado island:
5 Mar. 1983, Judziewicz 4438 (MO, WIS), 30 Apr.
1970, Croat 10101 (MO). dariEn: along Pan-American
Highway between Pucro and Rio Punusa, 3 Aug. 1962,
Duke 5401 (MO).
This species is known with certainty only from
gallery forests along rivers in the Caribbean coastal
plain of Honduras and Nicaragua at an elevation
below 100 m. The additional sterile collections cited
from Costa Rica and Panama await flowering col¬
lections for confirmation. This species is named for
F. A. McClure, distinguished student of bamboos,
who recognized this species as distinct and gave it
a manuscript name, which is unfortunately a later
homonym. His detailed notes on vegetative speci¬
mens collected at Bluefields, Nicaragua, and portions
of his manuscript description have been incorporated
into this treatment. The one flowering specimen
available to McClure was immature, and some of
his statements about the pseudospikelet and spikelet
details are not correct.
This species is unusual in the genus in having
spikelets with only one or at most a few bisexual
florets. Vegetatively it most resembles G. amplex-
ifolia, but G. macclurei differs in the usually white
(not brown) hairs on the upper surface of the culm
leaf blades, the frequent occurrence of linear-falcate
(not triangular) setose auricles on the branch leaf
sheaths, the relatively thinner culm walls, and most
notably, in the much narrower pseudospikelets (ca.
3 mm vs. 4-5 mm wide).
OLYREAE
Arberella grayumii Davidse, sp. nov. TYPE: Cos¬
ta Rica. Puntarenas: along new road from Pie-
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
95
Figure 6. Arberella grayumii Davidse. —A. Habit. —B. Inflorescence with terminal female spikelets with only
the glumes remaining and the anthoecium dehisced. C, D. Female anthoecium. —C. Dorsal view showing the pubescence
around the germination flap. —D. Ventral view showing the prominently ciliate lemma margins. —E. Male inflorescence
with terminal male spikelet. (Based on Grayum et al. 3402.)
96
Novon
dras Blancas to Rincon de Osa, 8°45'N,
83°21'W, ca. 200 m, scattered on forest floor,
2 July 1984, M. H. Grayum, H. Jacobs, G.
Schatz, J. Kress & P. Sleeper 3402 (holotype,
MO; isotypes, CR, US). Figure 6.
Gramen perenne. Folia evoluta in quoque culmo 5-8;
vaginis ciliatis et pilosis; pseudopetiolo 1.0-1.5 mm longo;
laminis 5-10 cm longis, 1.6-3.0 cm latis, oblongo-ellip-
ticis, apice acuminatis, basi asymmetricis. Paniculae ad
quemque nodum medianum et supernum 1-5, 1.5-4.0
cm longae. Spiculae masculae 3.8-5.2 mm longae. Spic-
ulae femineae lanceolatae, 13.5-15 mm longae, ca. 2.5
mm latae; gluma inferna 7-nervi, superna 7-nervi; inter
gluma anthoeciumque internodio ca. 1 mm; lemmate ca.
8.7 mm longo, 2.3 mm lato, ciliato dense pubescenti
circum operculum germinationis; palea 4-nervi, glabra.
Caryopsis non suppetens.
Caespitose, perennial herb. Culms 35-50 cm long,
unbranched, erect or geniculate-ascending; inter-
nodes terete, shiny, glabrous, hollow, with a gland¬
like spot on one side toward the apex; exposed nodes
4-5, blackish, glabrous, formed from the thickened
basal rim of the sheath above and the thickened
upper rim of the internode below. Leaves of the
basal nodes consisting of only a sheath shorter than
the internode; fully developed leaves 5-8 per culm;
sheaths green-mottled, hirsute in the upper half,
ciliate; ligule a eiliolate membrane 0.2-0.4 mm long;
pseudopetiole 1.0-1.5 mm long, pilose with white
hairs on both surfaces; blades 5-10 cm long, 1.6-
3.0 cm wide, oblong-elliptic, sparsely to moderately
pilose below, glabrous to sparsely pilose above, cil¬
iate, acuminate, basally asymmetrical, the midrib
on both surfaces thickened at the base and termi¬
nating in a knoblike process. Panicles 1-5 from
each of the middle to uppermost nodes, 1.5-4.0 cm
long, narrow, erect, with a single large terminal
female spikelet and ca. 8-16 paired male spikelets
below, one of each pair long-pedicelled, the other
short-pedicelled, or the female spikelet lacking and
only male spikelets present; axis angular, glabrous;
pedicels glabrous, those of the female spikelets prom¬
inently thickened at the apex, those of the male
spikelets only slightly thickened. Male spikelets 3.8-
5.2 mm long, 0.5-0.6 mm wide, linear-lanceolate,
acuminate, glabrous, membranous, whitish; lemma
3-nerved, acuminate with an awn tip to 0.5 mm
long; palea as long as the lemma body, 2-nerved;
lodicules 3, 0.4-0.5 mm long, similar, oblanceolate;
stamens 3, the anthers 1.2-1.6 mm long, basifixed.
Female spikelets 13.5-15.0 mm long, ca. 2.5 mm
wide, lanceolate, long-attenuate, glabrous, whitish;
glumes subequal, membranous, longer than the an-
thoecium, 7-nerved, with a few transverse veinlets;
anthoecium ca. 8.7 mm long and 2.3 mm wide,
oblong-elliptic, the internode ca. 1.0 mm long; lem¬
ma smooth and shiny, whitish when immature,
densely and prominently ciliate, also densely pu¬
bescent on the germination flap and just above the
flap on the back; palea 4-nerved, glabrous; lodicules
3, ca. 1 mm long, vasculated, the posterior one and
one of the anterior ones marginally connate, the
free anterior one ± dumbbell-shaped; ovary fusi¬
form, ca. 2.8 mm long; style 1, as long as the
anthoecium; stigmas not seen. Caryopsis not avail¬
able.
Arberella Soderstrom & C. Calderon is a small
genus of five closely related species, plus the dis¬
tantly related A. bahiensis Soderstrom, the only
species that lacks the elongated internode between
the glumes and anthoecium (cf. Soderstrom & Zu-
loaga, 1985). Arberella grayumii differs from all
the glabrous-leaved species (A. bahiensis, A. flac-
cida (Doell) Soderstrom & Zuloaga, A. dressleri
Soderstrom & C. Calderon) in its much broader
blades, glabrous female palea, and different pubes¬
cence pattern on the back of the female lemma.
In Mesoamerica the four known species of Ar¬
berella occur only in tropical wet forests and have
very limited distributions: A. grayumii is only known
from the Osa Peninsula on the Pacific side of Costa
Rica; A. costaricensis (A. Hitchc.) Soderstrom &
C. Calderon is only known in the Rio Hondo area
on the Atlantic side of Costa Rica; A. dressleri is
known from eastern Colon and western San Bias,
Panama, on both sides of the continental divide but
is most common on the Caribbean side; and A.
lancifolia Soderstrom & Zuloaga is the most wide¬
spread Mesoamerican species, ranging from San Bias
to Bocas del Toro, Panama. Arberella lancifolia
occurs only along the Caribbean slope in the western
part of its range but crosses the lower continental
divide in the eastern part of its range.
This species is named for Michael H. Grayum,
Missouri Botanical Garden, in recognition of his out¬
standing work on the flora of Costa Rica. It is only
known from the type collection.
Cryptochloa sodersiromii Davidse, sp. nov.
TYPE: Panama. San Bias: Ailigandi, area along
trail from ocean to waterfall on river, 0-200
ft., culms clumped, to 1 m tall, leaves dark
green matte above, burgundy below, on slope
along river, 7 Oct. 1978, H. Hammel & W.
G. D’Arcy 4984 (holotype, MO; isotype, US).
Figure 7.
Gramen perenne. Culmi 60-100 cm longi. Folia in
complemento 7-9, scaberula; vaginis auriculatis margi-
nibus glabris; ligula 2-8 mm longa, asymmetrica; laminis
10-13 cm longis, 3-4 cm latis, oblongo-elliptico-ovatis.
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
97
Figure 7. Cryptochloa soderstromii Davidse. —A. Habit. —B. Portion of inflorescence with female spikelets. -
C. Portion of inflorescence with male spikelets. —D. Female spikelets. E, F. Female anthoecium (immature) with
prominent basal internode. —E. Ventral view. —F. Dorsal view. (Based on Hammel & D Arcy 4984.)
UiUJ £
98
Novon
Inflorescentiae paniculatae, ad quenque nodum foliarem
2-3, terminales et axillares, spiculas femineas apicem
versus et masculas basim versus gerentes. Spiculae mas-
culae 3.5-5.0 mm longae muticae; antheris 3, 3.3 3.6
mm longis. Spiculae femineae 9-13 mm longae; glumis
7-9-nervibus acutis vel aristatis ad 1 -3 mm; flosculo 7.8-
8.3 mm longo, 1.9-2.0 mm lato. Caryopsis non suppe-
tens.
Caespitose, perennial herb. Culms 60-100 cm
long, erect, with up to 6 elongated internodes bearing
sheaths without or with reduced blades preceding
the section ol the culm hearing the leaf complement;
internodes hollow, glabrous or sparsely scaberulous
between the veins at the apex, longer than the as¬
sociated sheath in the lower half, shorter in the leaf
complement; nodes glabrous to appressed scaberu¬
lous. Leaves with developed blades 7-9 per com¬
plement, scaberulous; sheaths prominently auricled,
the auricle adnate to the ligule, the margins glabrous;
ligule asymmetrical, 5-8 mm on the underlapping
side of the sheath, 2-3 mm on the overlapping side,
membranous, glabrous on the back, ciliolate; pseu¬
dopetiole ca. 1.5 mm long, scabrous; blades 10-13
cm long, 3-4 cm wide, oblong- to elliptic-ovate,
glabrous, basally rounded, slightly asymmetric, api-
cally abruptly acuminate. Panicles terminal and ax¬
illary, 2 -3 from each of the uppermost leafy nodes,
bearing female spikelets toward the tip and male
spikelets toward the base; peduncles not exserted
from the sheath; branches appressed, scaberulous,
ridged; female pedicels thickened at the tip, solitary;
male pedicels slender at the tip, solitary or in short-
and long-pedicelled pairs. Male spikelets 3.5-5.0
mm long, awnless, disarticulating immediately after
anthesis as one unit, glabrous; lemma and palea
subequal; lemma 3-nerved; palea 2-nerved; lodicules
3, truncate; stamens 3, the anthers 3.3-3.6 mm
long. Female spikelets 9-13 mm long including the
awn tips, fusiform, disarticulating above the glumes;
glumes as long as the spikelet, 7-9-nerved, glabrous
without, scabrous in the upper %, with 3 strong
nerves and 4 or 6 weak ones near the margin, with
transverse veinlets, the lower acute or with an awn
tip to 3 mm long, the upper acute or with an awn
tip to 1 mm long; internode between the glumes and
anthoecium 1.6-2.2 mm long; anthoecium 7.8-8.3
mm long, 1.9-2.0 mm wide, shiny; lemma 5-nerved;
palea 2-nerved, nearly as long as the lemma; lodi¬
cules 3, truncate, vasculated, eciliate; style 1, as
long as the anthoecium; stigmas 2; caryopsis not
available.
Cryptochloa soderstromii is very closely related
to the South American C. capillata (Trin.) Soder-
strom but differs in its smaller (3.5-5.0 mm vs.
6.0-9.0 mm), essentially awnless, male spikelets.
eciliate sheath margins, smaller female spikelets (9
13 mm vs. 20-23 mm) with essentially awnless
glumes or with the lower glume sometimes with an
awn point to 3 mm long, and inflorescence branches
more appressed, somewhat thicker, and not as highly
branched.
It is similar to C. capillata in its large size, similar
leaves with similar ligules, and paniculate terminal
inflorescences. In C. soderstromii the terminal in¬
florescence bears both male and female spikelets.
In C. capillata, as Soderstrom (1982) noted when
he transferred this species to Cryptochloa Swallen
from Olyra L., the terminal panicle may bear a
mixture of male and female spikelets, or it may be
uniformly unisexual, in which case it usually bears
male spikelets, or only rarely female spikelets. In
other species of Cryptochloa the terminal inflores¬
cences are nearly always male, and they are usually
much smaller. Whether C. soderstromii also usually
produces terminal male inflorescences awaits further
observations on new collections.
The species is named for the late Thomas R.
Soderstrom, who through his careful work and great
enthusiasm for the herbaceous bamboos, greatly in¬
creased our understanding of this group. It is known
only from the type collection.
Parianeae
The genus Pariana Aublet consists of approxi¬
mately 38 species distributed in lowland and lower
montane rainforests from Costa Rica (and probably
southern Nicaragua) south to Brazil and Bolivia. It
is especially abundant in the Amazon Basin. Victoria
C. Hollowell is currently monographing the genus.
It is not surprising that a new species needs to
be described since Costa Rica-Panama is one of the
major centers of bamboo diversity in the Neotropics,
with many species still awaiting description (Sod¬
erstrom et al., 1988).
Pariana argentea Hollowed & Davidse, sp. nov.
TYPE: Panama. Panama: El l.lano-Carti Road,
4 mi. beyond the (Pan-American) Highway,
500 m, in shade of forest, spikes green, the
stamens yellow, 5 Dec. 1979, T. B. Croat
49139 (holotype, MO).
Gramen perenne rhizomatibus brevibus. Culmi dimor-
phi; culmi steriles ad 40 cm longi, erecti vel geniculati,
ascendentes; culmi fertiles ad 20 cm longi, decumbentes.
Folia vaginis truncatis, fimbriatis, hirtellis; laminis linear-
ibus vel lineari-lanceolatis, adaxialiter viridibus, abaxialiter
manifeste argenteo-glaucis. Inflorescentiae spiciformes, ad
4.3 cm, terminales, graciles, gynecandrae, ex verticillis
spicularum dimorpharum constantes. Spiculae masculae
flosculum unum gerentes, lemmate anguste elliptico, stam-
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
99
inibus 9-15(-18). Spiculae femineae flosculum unum ger-
entes, glumis glabris, costatis, anthecio longioribus.
Caespitose, perennial herb. Culms dimorphic.
Sterile culms to 40 cm long, erect from decumbent
bases; internodes hollow, terete, the basal ones short,
the intermediate ones elongated, the apical ones
shortened and hidden within the leaf complement;
nodes vinaceous or stramineous, glabrous, linear to
dilated and biannular, the lower usually geniculate.
Fertile culms to 20 cm, basally decumbent, reduced
and often flexuous, bearing bladeless sheaths, the
sheaths basally imbricate, inflated and loose, stra¬
mineous or vinaceous-tinted. Leaves at lower and
midculm nodes of sterile culms reduced to bladeless
sheaths, often vinaceous-tinted, imbricate at culm
bases but equaling to half as long as the internodes
above. Upper leaves of sterile culms 7-10, forming
a terminal complement with fully developed blades;
sheaths truncate, dorsally keeled, hirtellous or gla-
brate, with 1-3 lunar marks at the summit; oral
setae to 8 mm long, numerous, spreading, antrorsely
scaberulous; ligules 0.6-0.7 mm long, truncate,
scarious, usually obscured by oral setae and lateral
callosities; pseudopetioles 1.5-2.5 mm long, gla-
brate, stramineous or vinaceous-tinted; blades 6.1-
12.0 cm long, 0.8-l. 1(-1.6) cm wide, linear or
linear-lanceolate, basally cuneate, apically acumi¬
nate or setaceous, glabrous, conspicuously blue-white
or silver glaucous below. Inflorescence to 4.4 cm
long, ca. 1.0 cm wide, slender; spikelets in ca. 7
whorls, these overlapping up to V 2 the length of the
staminate pedicels of the whorl above; elaiosomes
developed on the rachis segment at the base of the
pedicels. Staminate spikelets 7.2-8.3(-10.3) mm
long; pedicels 3.2-4.0 mm long, shorter than or
occasionally subequal to the lemma, basally canes-
cent, marginally ciliate; glumes 2.0-2.8 mm long,
stiffly triangular, apically setaceous, 2-3-nerved, to
Vi as long as the lemma, scaberulous or basally
ciliate; lemmas 3.5-4.0(-6.0) mm long, narrowly
elliptic, acuminate, glabrous; paleas subequal to the
lemmas, often 4(-5)-nerved; lodicules 2, to 0.5 mm
long, frontally directed at the palea margins, obo-
vate; stamens 9— 15(— 18), the anthers 2.6-2.8 mm
long, yellow, fully exserted at anthesis. Pistillate
spikelets 5.3-6.1 mm long, sessile; glumes 5.3 6.1
mm long, subequal, overtopping the anthoecium by
0.6-1.1 mm, costate with faint marginal nerves
occasionally evident, glabrous except scaberulous
along the costa and apical margins; anthoecium 4.8-
5.2 mm long, ovate, glabrous; lodicules 3, the an¬
terior pair to 1.5 mm long, obovate, acute with
apical microhairs, membranous, the posterior one
to 0.5 mm long, elliptic, lacking apical microhairs;
style 1, short, glabrous; stigmas plumose, subapically
exserted.
Paratypes. Panama. Panama: Card Road, 7 mi. from
turnoff at El Llano, in forest along road, 9°15'N, 78°58'W,
320 m, occasional on slopes in forest, leaves glaucous
below, 16-17 Apr. 1985, Hammel 13546 (MO); El Lla-
no-Carti Road, 10 km from Inter-American Highway,
forest grass, anthers yellow, 4 Oct. 1974, Mori & Knl-
lunki 2292 (MO); El Llano-Carti Road, 6-7 mi. from
Pan-American Highway, wet tropical forest, 300-350 ft.,
herb, leaves silvery below, inflorescence light green, sta¬
mens yellow, on a separate leafless scape, 28 Apr. 1981,
Sytsma 4151 (MO).
Pariana argentea is, as far as presently known,
endemic to a small area on the Pacific slope of east-
central Panama in the Province of Panama, where
it occurs in moist tropical forest.
Pariana argentea is distinguished by its narrow
leaf blades with silvery, glaucous abaxial surfaces
(the character from which the name is derived) and
its decumbent, flexuous fertile culms. It is most
closely related to P. strigosa Swallen and P. gracilis
Doell. From the first it differs most notably in its
smaller (6.1-12.0 x 0.8-1.1(-1.6) cm vs. 12.0-
1 5.0 x 4.5-5.5 cm), linear to linear-lanceolate (not
ovate to ovate-lanceolate) leaf blades, more numer¬
ous (not few or absent) and longer (to 8 mm vs. ca.
4 mm) oral setae, and pistillate glumes glabrous
except for the scaberulous costae and apical margins
(not puberulent and ciliate). It differs from P. grac¬
ilis in its glaucous (not green) lower leaf surfaces
and more numerous stamens.
One collection ( Hammel 13546) with the dis¬
tinctive foliage of P. argentea has atypically large
spikelets that resemble those of P. simulans Tutin
and P. strigosa. These two larger species have pu¬
berulent pistillate glumes in contrast to the glabrous
ones of P. argentea. This anomalous collection may
also have lightly puberulent pistillate glumes, an¬
other intermediate spikelet feature, which suggests
the possibility of hybridization and introgression
among these three sympatric species. Of interest
was the incidence of immature insects and insect
damage within the florets of this anomalous collec¬
tion.
Also notable is the differentiation of the rachis
internode at the base of each spikelet whorl into a
distinct callus. This callus is -noticeably swollen in
spikelet whorls containing mature pistillate spikelets,
i.e., those bearing fully developed caryopses. Mi¬
croscopic examination and staining with Sudan IV
(a lipid-specific stain) of this callus in mature spikelet
whorls of Hammel 13546 showed that the swelling
was caused by the accumulation of oil. Davidse
(1987) has reviewed the occurrence of similar struc-
100
Novon
tures in other genera of grasses and has interpreted
them as elaiosornes, which may be effective in ant
dispersal. This interpretation also seems to apply to
P. argentea and is probably also true of many other
species in the genus.
Bromeae
Bromus exaltatus Bernh., Linnaea 15(Litt. Ber.):
90. 1841. TYPE: Mexico. Anonymous s.n.
(lectotype, selected here, MO-2957788).
In the only major revision of Bromus in Mexico
and Central America, Soderstrom & Beaman (1968)
neotypified B. exaltatus Bernh., the earliest name
of section Bromopsis Dumort. (sect. Pnigma Du-
mort.) that applies to a Mesoamerican species. They
assumed that the type specimen from the Bernhardi
Herbarium (MO) was still missing, as both Wagnon
(1950), on the authority of George B. Van Schaack,
and Shear (1901), on the authority of W. Trelease,
had reported. Although we do not know what may
have happened to these specimens in the first part
of this century (perhaps misfiled or not recognized
as types), they were later found and annotated by
Van Schaack. Two sheets, both representing the
same species, exist. As is unfortunately character¬
istic of many specimens in the Bernhardi Herbarium,
both are poorly labeled with minimal information.
According to the annotation by Van Schaack, who
interpreted the handwriting, one of the specimens,
MO-2957788, has a label with "Bromus exaltatus,
Mexico," in pencil in Bernhardi’s own handwriting.
The second specimen only bears an unpublished
name. Since these are original specimens used by
Bernhardi to write his description, the earlier neo-
typification is superseded and the first specimen is
here chosen as the lectotype.
The lectotype has 3-nerved, pubescent glumes,
pubescent lemmas 9.5-11.5 mm long, rounded on
the back, auriclate sheaths, glabrous, obtuse ligules
0.8-1.1 mm long, a large nodding panicle, blades
7-9 mm wide, and anthers 2.7 mm long.
Both Wagnon (1950) and Soderstrom & Beaman
(1968) commented on Bernhardi’s description of the
lower glume as 4-nerved. This was a simple mistake
as all lower glumes in the lectotype are 3-nerved,
with the lateral nerves weak in a few spikelets.
Since Soderstrom & Beaman (1968) greatly em¬
phasized glume nervation as a key character, most
specimens that we recognize as B. exaltatus would
key out as B. porteri (J. Coulter) Nash in their
treatment. However, B. porteri is a nonauriculate
species of northern Mexico and the western United
States, and we have not seen any specimens from
Mesoamerica.
Triticeae
Agropyron attenuatum (Kunth) Koemer &
Schultes is the most widespread native species of
Triticeae in Latin America. It ranges along the main
cordilleras from Costa Rica to Argentina.
We, like Love (1984), place this species in the
genus Elymus L. However, while transferring this
species to Elymus, Love (1984) inexplicably created
an illegitimate later homonym while citing the le¬
gitimate earlier combination with this epithet. Thus,
a new name is required for this species in Elymus.
Elymus cordilleranus Davidse & R. Pohl, nom.
nov. Replaced name: Elymus attenuatus A.
Love, Feddes Repert. 95: 473. 1984, non (Gri-
seb.) K. Richter, 1890, based on Trilicum at¬
tenuatum Kunth in Humb., Bonpl. & Kunth,
Nov. Gen. Sp. 1: 180. 1816. Agropyron at¬
tenuatum (Kunth) Roemer & Schultes, Syst.
Veg. 2: 751. 1817. TYPE: Ecuador. Propter
Burropotrero et Chillo Quintensium, Humboldt
& Bonpland s.n. (P).
Arundineae
Danthonia chiapasensis Davidse, sp. nov. TYPE:
Mexico. Chiapas: steep slopes with montane
rainforest, evergreen cloud forest and elfin for¬
est, Drimys, Quercus, Ostrya, Liquidambar,
Styrax, and Ilex, on the SE side of Cerro Tres
Picos and the ridges near summit, elev. 2,100-
2,500 m, 11 Dec. 1972, D. E. Breedlove &
R. E. Thorne 30108 (holotype, DS).
Gramen perenne caespitosa. Culmi 40-60 cm longi.
Folia laminis 20 45 cm longis, plerumque involutis sed
ad 5 mm latis ubi complanatis. Panicula 8-13 cm longa,
ca. 1.5 cm lata, ramis glabris. Spiculae 17-21 mm longae;
flosculis 5-6; lemmate infimo usque ad aristae centralis
insertionem 4.0-4.5 mm longo, 9-nervi, dorsaliter pilosa
in dimidio inferiore, dentibus lateralibus 8.0-9.5 mm lon¬
gis aristam includentibus, aristis centralibus ad 25 mm
longis, callo piloso trichomatibus 0.5-1.0 mm longis. Car-
yopsis non suppetens.
Densely caespitose, perennial herb; innovations
intravaginal. Culms 40-60 cm long, erect, glabrous.
Leaves basal and cauline; sheaths rounded on the
hack, glabrous except for tufts of hairs 0.7-1.1 mm
long in the auricular area; ligule a dense row of
hairs 0.5-1.5 mm long; blades 20-45 cm long,
mostly involute, to 5 mm wide when flat, glabrous
to sparsely pilose toward the base above and below,
the upper surface deeply and closely furrowed. Pan¬
icle 8-13 cm long, ca. 1.5 cm wide, linear; branches
to 7 cm long, ascending, glabrous. Spikelets 17-21
mm long; glumes subequal, as long as the spikelet,
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
101
lanceolate, long-acuminate, 3-5-nerved, purplish in
the center, translucent whitish toward the margins;
florets 5-6, smaller upwards, the uppermost rudi¬
mentary; lowermost lemma 4.0—4.5 mm to the point
of insertion of the central awn, 9-nerved, pilose on
the back and along the margins in the lower half,
the lateral teeth prominent, awned, 8.0-9.5 mm
long including the awn, the central awn to 25 mm
long, the basal part flattened and twisted; lowermost
palea ca. 6 mm long, prominently 2-keeled, acute,
sulcate between the scaberulous keels; callus linear,
pilose with hairs 0.5-1.0 mm; lodicules 2, obovate,
obtuse, vasculated, sparsely ciliate with hairs to 0.5
mm long; stamens 3, the anthers 1.7-3.4 mm; styles
2, separate, the stigmas plumose; caryopsis not
available.
This species is only known from the type locality
in the Sierra Madre in the State of Chiapas (after
which it is named), at ca. 16°12'N, 93°37'W.
Only one other native species of Danthonia DC.
is known from Mesoamerica, D. filifolia Hubb.,
which ranges from Puebla, Mexico, to Guatemala.
Danthonia chiapasensis is easily distinguished from
that species by the lemma, which is prominently
long-pilose on the back (vs. glabrous), and the short-
pilose callus (0.5-1.0-mm vs. 3-5-mm-long hairs).
Although it may not necessarily be particularly
closely related to either, it is morphologically most
similar to D. parryi Scribner from the Kocky Moun¬
tains in the north, and D. cirrata Hackel & Arechav.
from southern Brazil, Uruguay, and Argentina in
the south.
Danthonia parryi differs from D. chiapasensis
in its larger spikelets, smaller inflorescences, pu-
berulent panicle branches, broader and less promi¬
nently nerved glumes and lemmas, and broader cal¬
luses. Danthonia cirrata differs in its narrower leaf
blades, florets smaller in all parts, and denser lemma
pubescence.
Conert (1987) has pointed out that Danthonia
(sens, lat.) in the New World has a large gap in its
distributional range in that seven species occur pri¬
marily in the United States and 18 species in south¬
ern South America. In the north only D. spicata
(L.) P. Beauv., primarily of the United States, ranges
as far south as Puebla, Mexico, but it becomes quite
rare in the southernmost part of its range. In the
south the widely ranging D. secundiflora J. S. Presl
reaches as far north as Colombia. Danthonia chia¬
pasensis and D. filifolia partially bridge the gap
along the probable Cordilleran migration route for
the genus. Interestingly, they each represent one of
the major morphological groups of Danthonia (sens,
str.) in the Americas, i.e., those with lemmas long-
pilose all over the back, and those ciliate only along
the margins.
Eragrostideae: Eleusininae
Eragrostis rufescens Schrader ex Schultes var.
mesoamericana Davidse, var. nov. TYPE:
Honduras. Morazan: 2.5 km N of Zambrano
on road to San Francisco de Soroguara, pine
forest along a small stream, on dry tuff out¬
crops, elev. 1,370 m, 1 July 1970. R. If. Pohl
& G. Davidse 12125 (holotype, MO; isotype,
ISC).
A var. rufescenti inflorescentiis ramorum axillis pilosis
et pedicellis divergentibus differt.
Caespitose, eglandular, annual herb. Culms 5-35
cm tall, erect to decumbent, usually unbranched,
glabrous. Leaves mostly basal; sheaths rounded, gla¬
brous to papillose-pilose, the apex pilose; ligule 0.1-
0.2 mm long, a row of hairs; blades 5-20 cm long,
2.0-4.5 mm wide, flat to rolled, papillose-pilose to
glabrous. Panicle 2-13 cm long, 1.5-6 cm wide,
ellipsoid to ovoid, moderately dense; axils densely
pilose, rarely sparsely pilose or nearly glabrous;
branches spikelet-bearing to the base; pedicels much
shorter than the spikelets, slightly divergent. Spike-
lets 8-15 mm long, 1.6-2.3 mm wide, oblong to
oblong-lanceolate, purplish, disarticulating from the
base; glumes 1-nerved, acute, the lower 12 mm
long, the upper 1.2-1.8 mm long; rachilla persis¬
tent; florets 13-45; lemmas 1.4-2.1 mm long, acute,
sharply keeled, scaberulous on the keels, overlap¬
ping for Vi- Z A of their length, the lateral nerves
evident; paleas V\ to nearly as long as the lemmas,
the keels ciliolate; stamens 2, the anthers 0.2-0.3
mm, purple. Caryopsis 0.4-0.5 mm long, 0.3 mm
wide, ovate in outline, roundly triangular in cross
section, apically obtuse.
Paratypes. Mexico. VERACRUZ: Coatzacoalcos, 21 Mar.
1910, Orcutt 3103 (MO). TABASCO: Mun. Huimanguillo,
Laguna de los Limones, 18 Nov. 1981, Magana & Za-
mudio 111 (MO). CHIAPAS: Mun. La Trinitaria, 9 km S
of Comitan, 6 Nov. 1981, Breedlove & Davidse 54951
(CAS, MO); Mun. La Trinitaria, 10 km E of La Trinitaria,
7 Nov. 1981, Breedlove & Davidse 54977 (CAS, MO).
Belize. BELIZE: mi. 7 on Northern Highway, 23 Jan.
1974, Dwyer & Liesner 12147 (MO); from Belize City
to 5 mi. beyond Ladyville Estates, 22 Mar. 1973, Dwyer
10403 (MEXU, MO); Sibun River, 25 Feb. 1936, Gentle
1812 (MO); Belize, 29 Jan. 1931, Bartlett 11257 (MO).
CAYO: road S to Augustine and San Luis to Camp Six, 19
Mar. 1967, Divyer et al. 300 (MO). ORANGE walk: 5
km N of August Pine Ridge, 19 Mar. 1987, Davidse &
Brant 32778 (MO). TOLEDO: lower part of Richardson
Creek, 2-3, 11 Mar. 1987, Davidse & Brant 32326
(MO). Guatemala, chiquimula: 10 km ESE of Esqui-
pulas, 12 Aug. 1970, Harmon & Dwyer 3746 (MO).
102
Novon
Honduras. col6n: 4.5 mi. NE of Trujillo, 28 Jan. 1981,
Saunders 959 (MO). COMAYACUA: Siguatepeque, 26 May
1972, Burch 6144 (MO). EL paraIso: road to Yuscaran,
21 Dec. 1975, Williams et al. 31196 (MO); 2 mi. W
of Giiinope, 5 Jan. 1947, Williams & Molina R. 11518
(MO); 3 km NW of Giiinope, 27 Feb. 1949, Williams
& Merrill 15704 (MO); 25 km W of Danli, 20 July
1971, Pohl 12612 (MO), cracias a dios: Ahuas, 12-
14 die. 1972, Clewell 3606 (MEXU, MO), lempira:
cerca la ciudad de Gracias, 7-9 die. 1971, Nelson et al.
176 (MO). FRANCISCO MORAZAN: 8 km E of El Zamorano,
14 June 1970, Pohl & Davidse 11911 (MO); El Za¬
morano, 2 Nov. 1981, Swollen 11266 (MO); 14 km
NW of Lepaterique, 2 Jan. 1979, Pohl <£ Gabel 13865
(MO); 4 km SE of Rio Agua Galiente, 1 Jan. 1979, Pohl
& Gabel 13848 (MO), OCOTEPEQUE: 8 km SW of Santa
Fe, 16 Jan. 1976, W illiams et al. 31263 (MO). OLANCHO:
11 km NE of Pataste, 23 Dec. 1978, Pohl & Gabel
13779 (MO). Nicaragua. COMARCO del cabo: near Bil-
waskartna, 12 Mar. 1971, Seymour 4579{ MO), zelaya:
junction of road to Alarnikanba with road between El
Empalme and Limbaika, 24 Feb. 1979, Stevens 12780
(MO). Costa Rica, guanacaste: 5 km S of Liberia, 5
Dec. 1968, Pohl <6 Davidse 11553 (MO), san JOSfc: vie.
of El General, Dec. 1936, Skutch 3068 (MO).
This variety has been collected in Trachypogon
savannas, open areas in Pinus forests, and in Pinus
savannas at 0-1,600 m elevation from Oaxaca,
Mexico, to Costa Rica.
This distinctive suite of specimens from Oaxaca
and Veracruz to Costa Rica has an acute lemma
apex. They have almost invariably been identified
as E. maypurensis (Kunth) Steudel, a widespread
species ranging from Durango, Mexico, to Brazil
and Bolivia. However, true E. maypurensis has an
acuminate, somewhat divergent lemma apex. These
Mesoamerican specimens agree well in general mor¬
phology and spikelet characters with E. rufescens ,
a South American species, previously unreported
from Mesoamerica. The Mesoamerican plants differ
somewhat from the South American plants in having
the main axils of the inflorescence branches usually
densely pilose (rarely sparsely pilose or nearly gla¬
brous) versus glabrous or sparsely pilose and in
having the spikelets more divergent from the inflo¬
rescence branches versus virtually appressed in typ¬
ical E. rufescens. For these reasons the Mesoamer¬
ican plants are recognized as a new variety. The
epithet reflects its Mesoamerican distribution.
Cynodonteae: Boutelouinae
Chondrosum Desv. has most recently been rec¬
ognized by Clayton (1982) and Clayton & Renvoize
(1986). The species included have usually been
placed in Bouteloua Lagasca (e.g., Griffiths, 1912;
Gould, 1980a), from which they differ in having
usually fewer pectinate spikes whose spikelets dis¬
articulate above the glumes. Species retained in Bou¬
teloua usually have numerous spikes that disartic¬
ulate as a unit from the rachis. The recognition of
both genera for Flora Mesoamericana requires one
new combination.
Chondrosum hirsutum (Lagasca) Sweet, Hort. Brit.
455. 1827. Bouteloua hirsuta Lagasca, Var¬
ied. Ci. 2: 141. 1805. TYPE: Cult. Madrid
Bot. Gard., Lagasca s.n. (lectotype, selected
by Gould (1979), MA).
Chondrosium hirtum Kunth in Humb., Bonpl. & Kunth,
Nov. Gen. Sp. 1: 176. 1816. TYPE: Crescit in
temperatis, apricis Novae Hispaniae, inter Mexico
et Gasave, alt. 1180 hex., Humboldt & Bonpland
s.n. (P).
The treatment in Sweet consists only of the state¬
ment, “4. hirsutum Hairy.” This appears to be a
very deficient description, hut Sweet referred in his
naming of the genus to “K.S.” This cryptic refer¬
ence appears to be Kurt Sprengel’s sixteenth edition
of Systerna Vegetabilium. Page 293 of this work
contains a portion of Sprengel’s treatment of Ath-
eropogon Muhlenb., a synonym of Bouteloua.
Number 3 of Sprengel’s listing is A. hirtus, indicated
by him as a reference to Chondrosium hirtum Kunth
(1816) or the earlier Bouteloua hirsuta Lagasca
(1805).
Chondrosum hirsutum (Lagasca) Sweet var.
glandulosum (Cerv.) R. Pohl, comb. nov.
Basionym: Erucaria glandulosa Cerv., Natur-
aleza (Mexico City) 1: 347. 1870. TYPE: Mex¬
ico. In collibus de Guadelupe, et de Moctezuma
prope Mexicum.
Eragrostideae: Sporobolinae
Pereilema diandrum R. Pohl, sp. nov. TYPE:
Costa Rica. Heredia: Puente Mulas, S of San
Antonio, canyon of Rio Virilla, 850 m, 28 Nov.
1968, R. W. Pohl & G. Davidse 11482 (ho-
lotype, ISC, fragment, MO; isotype, US).
Gramen annuum caespitosum a P. beyrichiano (Kunth)
A. Hitchc. panicula densiore ramis ascendentibus super-
positis, arista lemmatis 10-24 mm longa, antheris duabus
et chromosomatum numero 2a = 80 recedit.
Caespitose, annual herb. Culms 40-80 cm long,
sometimes decumbent and rooting from lower nodes,
usually simple; internodes 1-2 mm diam., slightly
scabrid, often reddish; nodes glabrous. Leaves with
the sheaths shorter than the internodes, scabrid;
prophylls prominent, 2-3 cm long, bifid; ligule a
thick membrane 0.7-1 mm long; blades 10-20 cm
long, 5-8 mm wide, scabrid; auricles prominent,
clasping, ciliate. Panicle solitary, terminal on the
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
103
main culm or leafy branches, 10-20 cm long, 1-
2 cm wide, cylindrical; peduncle included or exserted
to 6 cm; rachis angular, ciliate-scabrous; branches
ascending, overlapping, solitary, 1-3 cm long,
densely covered with dense fascicles of spikelets.
Sterile spikelets reduced to bristles 3-5 mm long;
functional spikelets terete; glumes subequal, 0.7-
1.5 mm long, oblong to broadly ovate, 1-nerved,
with an awn 1.5-4 mm long; lemma 1.9 2.5 mm
long, 3-nerved, scahrid, with a straight awn 10 24
mm long; callus hairs 0.6-0.8 mm, scant; palea
bidentate, slightly longer than the lemma; stamens
2, the anthers 0.8-1.0 mm long, often purple. 2 n
= 80 from type.
Representative paratypes. CosTA RlCA. SAN JOSE: Rio
Tiliri, Tonduz 3121 (ISC); San Francisco de Guadalupe,
Tonduz 9817 (ISC, US); Llano Grande de Puriscal, Ji¬
menez 890 (US), alajuela: Carrillos de Poas, Hrenes
14606 (US). Numerous other specimens from the San
Jose area are in US.
This species is only known from the Meseta Cen¬
tral and Canton de Dota, Costa Rica, at elevations
of 500-1,200 m.
This species is similar to P. beyrichianum (Kunth)
A. Hitchc. (syn. P. brasilianum Trin.) of South
America, differing in the denser panicle with as¬
cending appressed branches, longer lemma awn, pos¬
session of 2 anthers (basis of the epithet) rather than
3, and in the octoploid chromosome number, n =
40. The chromosome number of P. diandrum was
determined from the type hut was mistakenly re¬
ported under the name P. beyrichianum (Pohl &
Davidse, 1971). The only known count for true P.
beyrichianum is n = 20 (Davidse & Pohl, 1978,
as P. brasilianum Trin.). It differs from Meso¬
american P. crinitum J. S. Presl in its straight rather
than flexuous awn, broader leaf blades, and in chro¬
mosome number. The chromosome number for P.
crinitum has been reported as n = 10 (Could &
Soderstrom. 1970; Pohl & Davidse, 1971; Davidse
& Pohl, 1974; Reeder, 1984).
Paniceae: Setariinae
Panicum aquaticum Poiret var. cartagoense
Davidse, var. nov. TYPE: Costa Rica. Cartago:
0.5 km E of Planta Radiografica along railroad,
2 km W of Paraiso, 1,300 m, open grass marsh,
plants 1-2 m tall, rooted in dense tangle ol
herbage, chromosome number determined to
be n = 36, 6 Feb. 1969, R. IV. Pohl & G.
Davidse 11700 (holotype, ISC).
A var. aquatico culmis longioribus simplicibus foliorum
vaginis pubescentibus paniculis grandioribus et foliorum
laminis erectis differt.
Perennial, rhizomatous herb. Culms 1-2 m long,
erect, unbranched; internodes and nodes glabrous.
Leaves with sheaths papillose-pubescent to glabres-
cent; ligule with the membrane 0.4-0.6 mm long
and the cilia 0.7-1.0 mm long; blades to 35 cm
long, 7-10 mm wide, sparingly pilose. Panicle ca.
27 cm long, terminal; branches 10-14 cm long,
scaberulous. Spikelets 3.0-3.1 mm long, basally in
unequal pedicellate pairs, terminally solitary, lan-
ceoloid-ellipsoid, glabrous; lower glume 0.8-1.0 mm
long, !/ s -Xo as long as the spikelet, 1 -nerved, obtuse;
upper glume as long as the lower lemma, 7-nerved,
acute; lower floret staminate; lower lemma as long
as the spikelet, 7-9-nerved, acute; lower palea as
long as the lower lemma; stamens 3, the anthers
0.7-1.4 mm long; upper floret 1.9-2.0 mm long,
O. 9-1.0 mm wide, smooth and shiny, glabrous, acute;
stamens 3, the anthers 1.3 1.4 mm long.
This distinctive middle-elevation population is only
known from the type collection. It is perennial, has
spikelets like P. aquaticum, but differs from the
typical variety of the species by its much larger
stature, unbranched culms, pubescent leaf sheaths,
larger panicles, and upright leaf blades.
Panicum sect. Dichotomifiora A. Hitchc. & Chase
ex Honda contains a widespread, annual-perennial
species pair(P. dichotomiflorum Michaux P. aqua¬
ticum) that has often been treated as a single species
(e.g., Reeder & Reeder, 1971; Renvoize, 1984).
We recognize the two taxa at the species level be¬
cause of the difference in life form and because P.
aquaticum has a functional male flower in the lower
floret. This male flower is lacking in P. dichotom¬
iflorum, even in plants in which the lower palea is
fully developed. As Hitchcock & Chase (1910) noted
many years ago, the lower palea of P. dichotomi¬
florum may be completely absent to fully developed.
The new variety seems to have a relationship with
P. aquaticum var. aquaticum equivalent to that of
P. dichotomiflorum var. bartowense (Scribner &
Merr.) Fern, with variety dichotomiflorum, with the
outstanding differences of both of the nontypical
varieties being the unbranched culms and pubescent
sheaths.
Variety cartagoense is an octoploid with n = 36,
as is variety aquaticum (Davidse & Pohl, 1974,
reported as P. aquaticum). The varietal name is
based on the name of the province in which it was
collected.
The recognition of Dichanthelium (A. Hitchc. &
Chase) Gould as a segregate genus from Panicum
L. continues to be controversial. Nevertheless, it
seems likely that, despite a relatively uniform spike¬
let structure, Panicum as recognized in the broad
104
Novon
sense by Zuloaga (1987) and Clayton & Renvoize
(1986) is polyphyletic, since elements included in
Panicum sens. lat. are probably more closely related
to other segregate genera now generally recognized.
Under these circumstances narrow generic concepts
seem prudent. Most of the combinations for Me-
soamerican species of Dichanthelium are already
available from the work of Hansen & Wunderlin
(1988), Gould (1974, 1980b), and Gould & Clark
(1978). The few additional ones not yet available
are made here:
Dichanthelium sphaerocarpon (Elliott) Gould
var. floridanum (Vasey) Davidse, comb. nov.
Basionym: Panicum sphaerocarpon Elliott var.
floridanum Vasey, U.S.D.A. Div. Bot. Bull. 8:
33. 1889. P. erectifolium Nash, Bull. Torrey
Bot. Club 23: 148. 1896, nom. nov. for P.
floridanum (Vasey) Chapman, FI. South. U.S.
(ed. 3) 585. 1897, non Trin. 1835. Dichan¬
thelium erectifolium (Nash) Gould & C. A.
Clark, Ann. Missouri Bot. Card. 65: 1105.
1979. TYPE: United States. Florida: Mosquito
Inlet, moist pine barrens, 5 May 1879, Curtiss
3599 (lectotype, selected by Hitchcock & Chase
(1910), US; isolectotype, MO).
Dichanthelium erectifolium was distinguished
from D. sphaerocarpon by Gould & Clark (1978),
following Hitchcock & Chase (1910), by its smaller
spikelets (1.0-1.1 vs. 1.4-2.0 mm long). Hitchcock
& Chase (1910) additionally noted differences in
growth habit, blade nervation, and panicle size. How¬
ever, reexamination of specimens throughout the
range (including D. sphaerocarpon var. isophyllum
(Scribner) Gould & C. A. Clark, P. polyanthes
Schultes) shows a complete intergradation in habit,
inflorescence size, and spikelet size. However, plants
traditionally referred to D. erectifolium have mod-
ally smaller spikelets, 1.2 versus 1.5 mm, suggesting
that varietal status is appropriate. As suggested by
the specific epithet, the leaves of variety floridanum
tend to be erect and appressed to the culm, and
they also have a more clearly defined ligule, a cil-
iolate rim 0.3-0.4 mm long. The characters that
distinguish variety floridanum are nearly equivalent
to those that distinguish variety isophyllum from
variety sphaerocarpon, as was done by Gould &
Clark (1978).
Dichanthelium sciurotoides (Zuloaga & Mor-
rone) Davidse, comb. nov. Basionym: Panicum
sciurotoides Zuloaga & Morrone, Novon 1:1.
1991. TYPE: Brazil. Minas Gerais: Districto
Diamantina, Christais, near Corriga Duas Pon¬
tes, on damp earth in shade, common, 1,160
m, 13 May 1931, Mexia 5819 (holotype, MO;
isotypes, G, K, M, P, R).
Dichanthelium sciurotis (Trin.) Davidse, comb,
nov. Panicum sciurotis Trin., Gram. Panic.
228. 1826. TYPE: Brazil. Chamisso s.n. (ho¬
lotype, LE).
Zuloaga & Morrone (1991) recently clarified the
status of the above two species. Mesoamerican plants
of D. sciurotoides have previously been misidentified
as D. viscidellum (Scribner) Gould and as Dichan¬
thelium sciurotis Trin. The latter is a rare, narrowly
restricted species from northeastern Brazil (Zuloaga
& Morrone, 1991). Dichanthelium sciurotoides is
so far known only from Belize, Bolivia, Brazil, Ec¬
uador, French Guiana, Guyana, Panama, and Ven¬
ezuela. Dichanthelium viscidellum ranges from
southern Mexico throughout Mesoamerica to Co¬
lombia and Venezuela. The three species may be
distinguished as follows:
la. Lower glume nerveless, truncate; lower lemma
inflated at the base . D. sciurotis
lb. Lower glume l-3(-7)-nerved, acute to sub¬
acute.
2a. Ligule with the membrane always longer
than the minute cilia; spikelets glabrous
. D. sciurotoides
2b. Ligule with the cilia always longer than the
minute rudimentary membrane; spikelets
usually at least sparsely, marginally pu-
berulent, rarely glabrous, often densely pu-
berulent. D. viscidellum
Dichanthelium aciculare (Desv. ex Poiret) Gould
& C. A. Clark var. ramosum (Griseb.) Dav¬
idse, comb. nov. Basionym: Panicum neuran-
thum Griseb. var. ramosum Griseb., Cat. PI.
Cub. 232. 1866. TYPE: Cuba occ., Wright
3454 (holotype, GOET).
The element of D. aciculare traditionally known
as Panicum fusiforme A. Hitchc. (Hitchcock &
Chase, 1910), which has large, acute spikelets with
a long-attenuate base, seems modally distinct from
the rest of the species, especially in Mesoamerica,
although there is some intergradation. This kind of
variation is best recognized at the varietal rank.
Dichanthelium dichotomum (L.) Gould var.
unciphyllum (Trin.) Davidse, comb. nov.
Basionym: Panicum unciphyllum Trin., Gram.
Panic. 242. 1826, and the autonym created
by P. unciphyllum Trin. var. implicatum
(Scribner) Scribner & Merr., Rhodora 3: 123.
1901. TYPE: United States. Trattinick s.n.
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
105
(lectotype, selected by Hitchcock & Chase
(1910), LE).
Panicum tenue Muhlenb., Descr. Gram. 118. 1817. Di-
chanthelium dichotomum (L.) Gould var. tenue
(Muhlenb.) Gould & C. A. Clark, Ann. Missouri Bot.
Gard. 65: 1119. 1979. TYPE: United States. Muhl¬
enberg Herb. 192 (holotype, PH-M, fragment, US).
As pointed out by Hansen & Wunderlin (1988),
unciphyllum is an earlier varietal epithet. Although
they include this variety in Dichanthelium ensifol-
ium (Baldwin ex Elliott) Gould, I agree with Gould
& Clark (1978) that because of more numerous
elongated culm internodes, it is best considered a
part of D. dichotomum.
Dichanthelium umbonulatum (Swallen) Dav¬
idse, comb. nov. Basionym: Panicum umbon¬
ulatum Swallen, Contr. U.S. Natl. Herb. 29:
420. 1950. TYPE: Guatemala. Zacapa: upper
slopes of Sierra de Las Minas, along Rio Re-
pollal, 2,100-2,400 m, 12-13 Jan. 1942,
Steyermark 42469 (holotype, US; isotype, F).
Panicum ramiparum Swallen, Contr. U.S. Natl. Herb.
29: 423. 1950, syn. nov. TYPE: Guatemala. Quetz-
altenango: mountains near Santa Maria, S of Quetz-
altenango, 25 Mar. 1932, Weatherwax 160 (ho¬
lotype, US).
Panicum alcobense Swallen, Contr. U.S. Natl. Herb. 29:
423. 1950, syn. nov. TYPE: Guatemala. Jalapa:
oak wood around top of Cerro Alcoba, just E of
Jalapa, 1,700 m, 2 Dec. 1939, Steyermark 32515
(holotype, US; isotype, F).
Panicum alsophilum Swallen, Contr. U.S. Natl. Herb.
29: 422. 1950, syn. nov. TYPE: Guatemala. Jalapa:
in oak woods around the top of Cerro Alcoba, just
E of Jalapa, 1,700 m, 2 Dec. 1939, Steyermark
32513 (holotype, F).
Dichanthelium umbonulatum is closely related
to D. commutatum (Schultes) Gould sens. lat., in¬
cluding D. albomaculatum (Scribner) Gould, syn.
nov., but differs in its glabrous spikelets and leaves
with a prominent line of hairs at the base of the
blade near the throat of the sheath; it also tends to
have smaller leaf blades. The tip of the upper lemma
is umbonate with a distinct, nearly herbaceous, pro¬
tuberance. This is also true in D. commutatum in
Mexico, Mesoamerica, and the southern United
States but gradually becomes less pronounced north¬
ward in the United States. Panicum ramiparum is
a form of D. umbonulatum with broad, pubescent
leaves, and P. alcobense is a form with slightly
smaller leaf blades. Although Swallen (1950) also
noted that P. alcobense has the upper glume and
lower lemma shorter than the upper floret, D. um¬
bonulatum is variable in this regard and this char¬
acter is not useful for separating these taxa. Finally,
Panicum alsophilum is a form with the spikelets
less densely arranged in the panicle.
Dichanthelium cordovense (Fourn.) Davidse,
comb. nov. Basionym: Panicum cordovense
Fourn., Mexic. PI. 2: 26. 1886. TYPE: Mex¬
ico. Cordova, Schaffner 293 (holotype, P, frag¬
ments, BAA, US).
Dichanthelium pantrichum (Hackel) Davidse,
comb. nov. Basionym: Panicum pantrichum
Hackel, Verh. K.K. Zool.-Bot . Ges. Wien 65:
72. 1915. TYPE: Brazil. Rio Grande do Sul:
Mun. Rio Pardo, Fazenda Leitao, 70 m, Feb.
1909, Jurgens s.n. (holotype, W, fragment,
US).
Typical species of Setaria P. Beauv. are well
characterized by the possession of dense, cylindrical
panicles with the spikelets borne in disordered fas¬
cicles on the main rachis or on the primary branches
and flat, nonplicate leaf blades. In the majority of
cases, each spikelet is subtended by one to several
sterile branches or bristles. Another group of species
has but a single bristle borne below the terminal
spikelet of each branch. Such species were assigned
by Hitchcock & Chase (1910) to Panicum suhg.
Paurochaetium A. Hitchc. & Chase and more re¬
cently by Pilger (1940: 72) to Setaria sect. Pau¬
rochaetium (A. Hitchc. & Chase) Pilger. Rominger
(1962) noted that these species do not constitute a
natural group, and his observation is substantiated
by the anomalous structure of the inflorescence and
spikelets in ,S. chapmanii (Vasey) Pilger. In this
species, the slender inflorescence is made up of a
number of unilateral spikes. The spikelets are bis-
eriate and turned with the blunt first glume turned
away from the rachis and the hack of the fertile
lemma toward it. The end of the rachis of the spike
is a triquetrous extension beyond the terminal spike¬
let. This has led to the inclusion of this species in
Setaria sect. Paurochaetium , hut the neat arrange¬
ment of the spikelets in unilateral spikes is highly
anomalous in that genus. The nature of the inflo¬
rescence indicates that this species is better accom¬
modated in Paspalidium Stapf, characterized by
spikelets borne in unilateral spikes and oriented with
the back of the upper lemma turned regularly toward
the rachis. Paspalidium with about 30 species is
pantropical with its greatest center of speciation in
Australia. The other Mesoamerican species of Pas¬
palidium, P. geminatum (Forsskal) Stapf. is aquatic
and has thick, spongy culms, while P. chapmanii
is an inhabitant of more temporary pools and marsh¬
es on limestone, coral, and sand, with more slender
culms.
106
Novon
As already noted by Rominger (1962), the re¬
maining species in subgenus Paurochaetium fall into
a primarily West Indian-Floridian group of six spe¬
cies, and a mainland group of three species centered
in Texas and northern Mexico, with a fourth species
from the Yucatan Peninsula. Although the arrange¬
ment of the spikelets in the West Indian species is
less neat than in P. chapmanii, it nevertheless cor¬
responds quite closely to the type of inflorescence
structure found in certain Australian species of Pas-
palidium, such as P. clementii (Domin) C. E. Hubb.,
P. constrictum (Domin) C. E. Hubb., P. criniforme
S. T. Blake, and P. grandispiculatum B. Simon.
For this reason we take the opportunity here to
transfer these species to Paspalidium. This rela¬
tionship was also hinted at by Clayton & Renvoize
(1986). We prefer to maintain the mainland group
in Setaria.
Paspalidium chapmanii (Vasey) R. Pohl, comb,
nov. Basionym: Panicum chapmanii Vasey,
Bull. Torrey Bot. Club 11: 61. 1884, as P.
“ chapmani . ” Setaria chapmanii (Vasey) Pil-
ger, Nat. Pflanzenfam. (ed. 2) l ie: 72. 1940.
TYPE: United States. Florida: Chapman s.n.
(lectotype, selected by Hitchcock & Chase
(1910), US).
Paspalidium distantiflorum (A. Rich.) Davidse
& R. Pohl, comb. nov. Basionym: Panicum
distantiflorum A. Rich, in Sagra, Hist. Fis.
Cuba, Bot. 11: 302. 1850. Setaria distanti-
flora (A. Rich.) Pilger, Nat. Pflanzenfam. (ed.
2) 14e: 72. 1940. TYPE: In montosis insulae
Cubae, Sagra s.n. (bolotype, P).
Paspalidium leonis (E. Ekman) Davidse & R.
Pohl, comb. nov. Basionym: Panicum leonis
E. Ekman in A. Hitchc., Man. Grasses W. Ind.
295. 1936. TYPE: Cuba. Cojimar, near Ha-
bana, in fruticetis littoralibus, 9 Sep. 1921,
Ekman 13155 (holotype, US).
Paspalidium ophiticola (A. Hitchc. & E. Ek¬
man) Davidse & R. Pohl, comb. nov. Basionym:
Panicum ophiticola A. Hitchc. & E. Ekman
in A. Hitchc., Man. Grasses W. Ind. 293. 1936.
Setaria ophiticola (A. Hitchc. & E. Ekman)
Sauget-Barb., Contr. Ocas. Mus. Hist. Nat. Col-
egio “De La Salle” 8: 163. 1946. TYPE: Cuba.
Pinar del Rio: Loma Cajalbana, in cuables, on
serpentine, 2 Jan. 1921, Ekman 12712 (ho¬
lotype, US).
Paspalidium pradanum (Sauget-Barb.) Davidse
& R. Pohl, comb. nov. Basionym: Panicum
pradanum Sauget-Barb. in A. Hitchc., Man.
Grasses W. Ind. 294. 1936. Setaria pradana
(Sauget-Barb.) Sauget-Barb., Contr. Ocas. Mus.
Hist. Nat. Colegio “De La Salle” 8: 164. 1946.
TYPE: Cuba. Oriente: in open thickets on slope
of Mesa de Prada, lower Jauco Valley, Jauco,
17 July-4 Aug. 1924, Leon 11710 (holotype,
US).
Paspalidium subtransiens (A. Hitchc. & E. Ek¬
man) Davidse & R. Pohl, comb. nov. Basionym:
Setaria subtransiens A. Hitchc. & E. Ekman
in A. Hitchc., Man. Grasses W. Ind. 351. 1936.
TYPE: Cuba. Santa Clara: in thickets on ser¬
pentine, Motembo, 27 June 1923, Ekman
16828 (holotype, LJS).
Paspalidium utowanaeum (Scribner) Davidse &
R. Pohl, comb. nov. Basionym: Panicum uto¬
wanaeum Scribner, Publ. Field Columbian Mus.,
Bot. Ser. 2: 25. 1900. Setaria utowanaea
(Scribner) Pilger, Nat. Pflanzenfam. (ed. 2) 14e:
72. 1940. TYPE: Puerto Rico. From a dry
hillside near Guanica, 22 Jan. 1899, Mills-
paugh 702 (holotype, F).
Paniceae: Digitariinae
Digitaria breedlovei R. Pohl & Davidse, sp. nov.
TYPE: Mexico. Chiapas: Municipio de Villa
Corzo, 65 km S of Mexican Highway 190 on
road from Tuxtla Gutierrez to Nueva Concor¬
dia, steep canyon with seasonal evergreen forest
and slopes of Quercus, elev. 850 m, 12 Sep.
1974, D. E. Breedlove 37709 (holotype, MO;
isotypes, CAS, MEXU). Figure 8.
Digitaria spiculis ternatis gluma prima absenti, gluma
secunda obsoleta vel minuta, usque ad 0.4 mm longa,
lemmate fertili brunneolo vel castaneo, rhachidi triquetra,
hirsuta.
Caespitose, annual herb. Culms 25-50 cm long,
branching from the base; internodes glabrous. Leaves
with the sheaths and blades densely papillose-hirsute;
ligule 0.7-1 mm long, membranous, truncate; blades
4-10 cm long, 2-3 mm wide, linear, flat. Inflores¬
cences 612 cm long, terminal; peduncles exserted
for 6 8 cm; rachis of the inflorescence short; ra¬
cemes 4-5, 6-8 cm long, slender, ascending; rachis
of the racemes 0.2-0.3 mm wide, triquetrous, bear¬
ing conspicuous hairs 1-2 mm long; pedicels sca¬
brous, unequal, the longest 1.3-1.5 mm long. Spike-
lets 1.3-1.5 mm long, in triads, acute; lower glume
absent; upper glume 0-0.4 mm long, a hyaline scale
when present; lower lemma narrower and shorter
than the upper, 5-nerved, sparsely appressed-pu-
bescent, the hairs with dilated tips; upper lemma
tan to light brown, minutely striate; stamens 3, the
anthers ca. 0.5 mm; pollen and caryopses devel¬
oping.
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
107
Figure 8. Digitaria breedlovei R. Pohl & Davidse. —A. Habit. —B, C. Two views of a rachis segment with a
spikelet triad. D, E. Spikelet. —D. Ventral view showing upper lemma. —E. Dorsal view showing small hyaline lower
upper glume and back of upper floret. (Based on Breedlove 37709.)
This species is known only from the type collec¬
tion. It is distinguishable from other American spe¬
cies with ternate spikelets by the minute, hyaline
upper glume and the conspicuously hirsute rachis.
This species is named in honor of Dennis E.
Breedlove, California Academy of Science, in rec¬
ognition of his outstanding contributions to Mexican
botany, especially his work on the Flora of Chiapas.
108
Novon
Digilaria clavitricha R. Pohl, sp. nov. TYPE:
Panama. Colon: Chorrera, prairie, especially
on dry or gravelly places, 16 Sep. 1911, A.
S. Hitchcock, Amer. Grasses Natl. Herb. 1564
(holotype, MO; isotype, US) [also distributed
as A. S. Hitchcock 8149 (US)].
Digitaria ad sectionem Corynotrichas Henrard per-
tinens. Gramen perenne. Inflorescencia ex racemis 1-2
constans. Spiculae ternatae, late ellipticae, 1.8-1.9 mm
longae, 0.9-1.1 mm latae; gluma prima minuta, truncata;
gluma secunda truncata, 1.5-1.7 mm longa; lemmate
infero spiculam aequanti; gluma secunda et lemmate in-
fero pilis minutis clavatis obsitis; lemmate supero badio.
Caespitose, perennial herb. Culms 30-90 cm long;
internodes glabrous; nodes papillose-hirsute, es¬
pecially the lower ones. Leaves with the sheaths
papillose-hirsute; ligules ca. 0.5 mm long, membra¬
nous; blades 6-24 cm long, 2-3 mm wide, glabrous
except lor a few elongated trichomes near the ad-
axial base. Inflorescence 2-7 cm long, of 1-2 ra¬
cemes 2-4 cm long. Spikelets 1.8-1.9 mm long,
0.9-1.1 mm wide, broadly elliptic, in triads; lower
glume a minute, nerveless, hyaline cuff 0.05-0.15
mm long; upper glume 1.5-1.7 mm long, broad,
blunt, 3-nerved; lower lemma as long as the spikelet,
5-7-nerved; upper glume and lower lemma beset
with clavate hairs ca. 0.3 mm long and dilated nearly
to their bases; upper lemma dark brown; stamens
3, ihe anthers ca. 0.9 mm long; styles purple.
Paratype. Panama. CANAL ZONE: between Panama and
Corozal, prairie, 20-50 m, 17 Dec. 1911, Hitchcock
9/96 (US).
I bis species is only known from grasslands in
central Panama at low elevations. It belongs to Hen-
rard’s (1950) section Corynotrichae, which is char¬
acterized by minute balloon-shaped hairs. The epi¬
thet of the new species refers to this type of hair.
All previously known species of this section are from
South America. It differs from its closest ally, D.
venezuelae Henrard, in the presence of a lower
glume and in the shorter upper glume. Swallen (1943:
170) identified the two cited specimens as D. sin-
gularis Mez, a species described from Paraguay and
also known from Argentina (Rugolo de Agrasar,
1974). However, D. clavitricha differs from D.
singularis, in its much smaller spikelets and non¬
protruding fertile floret.
Andropogoneae
Andropogon gerardii Vitman var. hondurensis
R. Pohl, var. nov. TYPE: Honduras. Road to
Teupasenti, open pine forest, 1,350 m, 22 June
1980, R. W. Pohl & L. G. Clark 14011 (ho¬
lotype, ISC).
A var. gerardii culmis gracilibus brevibusque, spiculis
sessilibus brevioribus ca. 5-5.3 mm longis, gluma inferiore
plana esulcata recedit.
Caespitose, perennial herb with short, thick rhi¬
zomes. Culms 70-200 cm long. Leaves mostly basal,
glabrous except for a tuft of hairs in the ligular area;
ligule 1.0-2.5 mm, a ciliolate membrane; blades to
50 cm long, 3.0-5.5 mm wide, folded or flat, the
tip acute. Inflorescences several, scarcely overlap¬
ping; spathes narrow; racemes 1-3 per peduncle,
3-8 cm long, ascending, sparsely pubescent with
hairs 2.0-3.5(-4.0) mm long; rachis internodes cla¬
vate, flat. Sessile spikelets 5.0-5.3 mm long; callus
oblique, pilose with hairs 0.5-2.5 mm long; glumes
awnless, the lower flat, several-nerved between the
keels, coriaceous; awn of the upper lemma exserted
to 1 1 mm, twisted in the lower X A, geniculate; sta¬
mens 3, the anthers 1.0-1.2 mm long; pedicellate
spikelets 5-7 mm long, acute, the flower absent or
staminate; stamens 3 when present, the anthers to
2.4 mm long. 2 n = 80.
Representative paratypes. Belize. EL CAYO: Mountain
Pine Ridge, July-Aug. 1936, Lundell 6791 (F).
Honduras, comayagua: vie. of Siguatepeque, 4 July 1936,
Yuncker et al. 5641 (MO); 10 km E of Siguatepeque,
17 June 1980, Pohl & Clark 13988 (ISC), distrito
CENTRAL: El Hatillo, Pohl & Gabel 13794 (ISC). EL
paraIso: Guinope, 25 Oct. 1951, Swallen 11105 (ISC);
22 kin W of Danli, 21 July 1970, Pohl & Davidse
12400 (MO), francisco morazan: 4 km N of La Venta,
20 June 1970, Pohl & Davidse 12031 (ISC); Las Mesas,
15 Oct. 1951, Swallen 10762 (ISC). INTIBUCA: La Es-
peranza, 12 Sep. 1981, Segovia 101 (MO); 8 km SW
de Masaguara, 6 June 1985, Tellez V. & Martinez S.
8675 (MO). Costa Rica, guanacaste: SE slopes of Volcan
Santa Maria, 27-28 Jan. 1983, Davidse et al. 23331
(MO).
Plants of variety hondurensis are especially com¬
mon, but scattered, in the open [line forests of central
Honduras, and this is the basis for tbe epithet. They
have been collected at elevations of 800-2,000 m.
Andropogon gerardii var. hondurensis in Me-
soamerica is smaller and more gracile than the typ¬
ical variety encountered in the United States. The
Mesoamerican plants also differ from the North
American plants in having shorter sessile spikelets,
averaging ca. 5 mm long, whereas plants in the
temperate United States usually have spikelets 7-
10 mm long. Lower glumes of the sessile spikelets
of the Mesoamerican plants are flat, while those
from the United States are sulcate. Although various
ploidy levels for this species are known, the Hon¬
duran plants, so far as known, are octoploid, 2 n =
80, (Davidse & Pohl, 1972, as A. gerardii).
Acknowledgments. Work on the production of
Flora Mesoamericana has been supported by a se-
Volume 2, Number 2
1992
Davidse & Pohl
Mesoamerican Grasses
109
ries of grants from the National Science Foundation
to Davidse. We gratefully acknowledge the contri¬
bution of Pariana by Victoria C. Hollowell. We
thank Linda G. Ellis for the illustrations. We ap¬
preciate the useful discussions by Lynn G. Clark
and Emmet J. Judziewicz about some of the new
bamboos, as well as the very helpful reviews of the
manuscript by Roy E. Gereau and Emmet J. Jud¬
ziewicz. We thank the U.S. National Herbarium for
access to the late F. A. McClure’s manuscript notes
on Guadua macclurei and the illustrations of this
species drawn by Agnes Chase. We are grateful to
the curators of the cited herbaria for making their
facilities and herbarium specimens available to us.
Davidse thanks Dennis E. Breedlove for the oppor¬
tunity to conduct fieldwork with him in Chiapas in
1981.
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Love, A. 1984. Conspectus of the Triticeae. Feddes
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McClure, F. A. 1934. The inflorescence in Schizos-
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Pohl, R. W. & G. Davidse. 1971. Chromosome num¬
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-& C. G. Reeder. 1971. Gramineae. Pp. 823-
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199 209.
-. 1988. Aulonemia fulgor (Poaceae: Bambu¬
soideae), a new species from Mexico. Brittonia 40:
22-31.
- & J. H. Beaman. 1968. The genus Bromus
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- & X. Londono. 1987. Two new genera of
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157 in W. R. Heyer & P. E. Vanzolini (editors).
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Proceedings of a Workshop on Neotropical Distri¬
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de Janeiro.
Swallen, J. R. 1943. Gramineae. In: Flora of Panama.
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Merostachys burmanii (Poaceae: Bambusoideae: Bambuseae),
a New Species from Brazil
Tatiana Sendulsky
Herbario, Instituto <le Botanica, Caixa Postal 4005,
01061 Sao Paulo, Sao Paulo, Brazil
ABSTRACT. Merostachys burmannii, a new spe¬
cies from Brazil, is described and illustrated.
The South American genus Merostachys (Po¬
aceae: Bambusoideae: Bambuseae) comprises ap¬
proximately 40 species and has a center of diversity
in Brazil. The first attempt to recognize and order
the species in this genus was made by McClure
(1973), who listed ca. 25 species. During fieldwork
and herbarium studies aimed at completing a revi¬
sion of this genus, several new species have been
discovered, one of which is described and illustrated
here.
Taxonomic understanding of genus Merostachys
has grown slowly. This is due to the long interval
between flowering cycles—some 30-33 years—and
to the difficulties in bringing together the rare col¬
lections. Because of long-standing diversity of opin¬
ion regarding interpretation of the spikelet mor¬
phology in the genus, a well-founded and adequate
terminology for bamboo descriptions is under in¬
vestigation. For now, the most suitable terms from
standard grass taxonomy are used.
Merostachys burmanii Sendulsky, sp. nov. 1’YPE:
Brazil. Sao Paulo, Jardim Botanico, 25 Mar.
1944 (fl), Kuhlmann s.n. (holotype, SP sheet
no. 79757; isotype, US sheet no 2678539).
Figure 1.
Culmi praeter apices flexuosos erecti, 10-15 m alti,
ca. 4 cm diametro; internodiis fistulosis, luteis vel fusco-
viridibus plerumque laevibus, nonnurnquam scabris, acu-
leis retrorsis, acutisque; nodis prominentibus, fuscis; culmo
sub nodo pilis albidis adpressis interdum cera mixtis ves-
tito. Folia culmi vagina ca. 30 cm longa, 10 cm lata, ad
apicem asymmetrica, rotundata, fimbriata, setis ca. 0.5
mm longis; lamina reflexa, ca. 17 cm longa, 18-20 mm
lata, acuminatissima. Rami verticillati, in complemento
100-130, usque ad 65 cm longi, 1-2 mm diametro,
teretes, laeves, glabri. Folia ramorum in complemento 7-
9; vagina glabra, setis oralibus caducis, 2-3 mm longis;
lamina 15-17 cm longa, 2-3 cm lata, lineari-lanceolata,
glabra, apice acuminata. Inflorescentia spiciformis, non
pectinata, 8-11 cm longa, erecta, rami ad apicem dis-
posita; spiculis saepe deflexis, plerumque binatis, interdum
solitariis, rarissime ternatis. Caryopsis oblonga, ovoidea,
asymmetrica, rostrata, luteo-castanea, laevis, 6-7 mm
longa, 2-2.5 mm lata.
Culms 10-15 m long, ca. 4 cm diam., erect
except at flexuous tips, the slender young culms
strongly and retrorsely scabrous. Internodes 38-40
cm long, terete, fistulose, dark yellow or greenish,
mostly smooth and shiny, though otherwise smooth
culms may display strongly scabrous areas with re-
trorse, sharp, hooklike prickle hairs; culm walls weak,
1-3 mm thick. I\odes salient, dark, with a layer of
velvety whitish hairs below, hairs and wax sometimes
mixed together. Culm leaves caducous: sheaths ca.
28-30 cm long, 10 cm wide, rounded and asym¬
metrical at the summit, fringed at the apex only
with fine hairs, these ca. 0.5 mm long; margins
smooth and shiny along the adaxial surface, mem¬
branous, without cilia; abaxial surface nearly gla¬
brous at the base, retrorsely scabrous at the apex,
the adaxial surface shiny; inner ligule a fine mem¬
brane, ciliolate on margin, 1-2 mm long, wider in
the middle; outer ligule a salient rim; blades reflexed,
ca. 17 cm long, 18-20 mm wide, strongly nerved,
gradually attenuate to a fine point. Branch com¬
plement with 100-130 branches (200-300 fide
Soderstrom), these up to 50 65 cm long, 1-2 mm
thick, yellowish, smooth, glabrous, shiny, the inter¬
nodes ca. 17 cm long, 2-3(-5)-noded before the
leaves, nodes usually black, sometimes dark yellow,
prominent. Foliage leaves 7~9 per complement,
the lower well developed, distant below; sheaths tight,
glabrous, adaxial surface smooth, densely dark-spot¬
ted; oral setae caducous, 2-3 mm long, whitish,
diminishing in size downward along the upper margin
of the sheath; inner ligule arcuate, ca. 1 mm long,
coriaceous, minutely ciliolate on the margin; outer
ligule a salient, glabrous rim; pseudopetiole flat, dark,
glabrous, 4 mm long; blades 15-17 cm long, 2-3
cm wide, linear-lanceolate, asymmetrical, gradually
attenuate toward the apex, glabrous on both sur¬
faces, the adaxial surface sometimes with 2-3 mar¬
ginal nerves on one side with rows of very fine prickle
hairs, the margins and the apex finely scabrous.
Inflorescence terminal on a leafy branch, spicate,
not pectinate, erect, 8-1 1 cm long, ca. 2 cm wide,
bearing secund, mostly deflexed, laterally outspread,
paired spikelets. Peduncle of the inflorescence densely
and retrorsely scabrid. Rachis sinuous, longitudinally
Novon 2: 111-113. 1992.
112
Novon
Figure 1. Merostachys burmanii Sendulsky. —a. Flowering branches and part of a branch complement. —b. Old,
empty anthers. —c. Gynoecium. —d. Lodicules. —e. Part of the inflorescence: rachis and paired spikelets. —f.
Pedicels. —g. Lower glume. —h. Upper glume. —i. Lemma. —j. Palea. —k. Prolongation of rachilla. —1. Culm
leaf. —m. Branch bud in development. —n. Caryopsis, lateral view. —o. Pair of spikelets with mature caryopses.
— p. Caryopsis, embryo view. —q. Caryopsis, hilum view. —r. Upper portion of the sheath, ligule, and base of a
foliage leaf. All illustrations based on Kuhlmann s.n. (SP sheet no. 79757) except “1” and “m,” based on Toledo
s.n. (SP sheet no. 238492).
Volume 2, Number 2
1992
Sendulsky
Merostachys burmanii
113
deeply ridged, puberulous to villous, the hairs white.
Spikelets in pairs, sometimes solitary, very rarely
ternate, frequently one of a pair of spikelets depau¬
perate or rudimentary. Normal spikelet 1 -flowered,
hermaphrodite, 8-11 mm long, 2 mm wide, nar¬
rowly fusiform when young, becoming thicker and
shorter when mature. Pedicels adnate, thick, pu¬
bescent. Glumes 2, unequal. Lower glume trian¬
gular, 2 mm long, 1.5 mm wide at the base, acu¬
minate toward the apex, 1-nerved, abaxially sparsely
hirsute along the nerve, adaxially puberulous near
the apex, the hairs white. Upper glume 5 mm long,
3.5 mm wide, broadly lanceolate, slightly asym¬
metrical, carinate, marginally ciliate toward the apex,
5-7-nerved, the nerves anastomosing, the adaxial
surface densely dark-spotted. Lemma 8 mm long,
3.5-4 mm wide, carinate, broadly lanceolate, slight¬
ly curved, fully embracing the palea at the base,
slightly asymmetrical, 10-1 1-nerved, the nerves
anastomosing, the adaxial surface dark-spotted. Pa¬
lea fusiform, curved, bifid, 2-keeled, narrowly sul-
cate abaxially, 6-nerved, the nerves anastomosing
in upper part, the keels finely and sparsely dentic¬
ulate. Prolongation of the rachilla bristlelike, as long
as the palea, finely pubescent and bearing a minute
rudiment. Lodicules 3, subequal, ca. 2 mm long,
finely ciliate at the apex. Stamens 3, the anthers 6
mm long, 0.5 mm wide, somewhat twisted in later
stages. Stigmas 2, ca. 4 mm long, plumose. Mature
caryopsis 6-7 mm long, 2-2.5 mm wide, oblong,
ovoid, asymmetrical, rostrate, dark yellow or brown,
smooth, shiny, with thin, coriaceous pericarp, the
embryo and the hilum not manifest externally.
Phenology ■ This species has been collected in
flower in 1915, in the periods between 1944 and
1948 and between 1972 and 1978(-1980).
Distribution. Brazil, states of Bahia, Rio de Ja¬
neiro and Sao Paulo, at (400-)600-1,600 m above
sea level.
Paratypes. BRAZIL. BAHIA: Jaguaquara, km 30 da ro-
dovia Jaguaquara-Apuerema, 25 Oct. 1972 (fl), Pinheiro
s.n. (US). RIO DE JANEIRO: Serra dos Orgaos, Parque
Nacional, km 4, 7 July 1947 (fl), Occhisni 1046 (US);
Serra dos Orgaos, 16 km S of Itaipara, 950 m, 26 Mar.
1976 (fr), Davidse et al. 11419 (NY, US); Serra dos
Orgaos, Fazenda Dr. Portugal, 1,400 m, 27 Nov. 1948
(fl), tirade 19457 (SP); Organ Mountains, 12 Aug. 1915
(fr), Rose & Russel 20768 (NY, RB, US); Orgelgeberge,
1,400-1,600 in. May 1915, Luetzelburg 6255 (NY);
Serra dos Orgaos, Nova Friburgo, road to television tower,
1,350 m, 14 Mar. 1980 (fl—abnormal late growth),
Burman & Soderstrom 525 (US); Campo Grande, Serra
do Mendanha, 600-700 m, 4 Apr. 1978 (fl), Martinelli
et al. 4129 (RB,SP); Tijuca, Parque Nacional, Mesa do
Imperador, ca. 20°50'S, 43°20'W, 400-650 m, 21 Oct.
1977 (fl), Landrum 2031 (RB). SAO PAULO: Jardim Bo-
tanico, 19 Jan. 1944 (fl), Toledo s.n. (SP sheet 50362,
US sheets 2678540, 2900180, 2921684), 19 May 1959
(st), Toledo s.n. (SP sheet 238492), 23 Mar. 1944 (fr),
Gert & Kuhlmann s.n. (SP sheet 50462).
Only three of the 40 or so known species of
Merostachys, described and undescribed, bear
spikelets in clusters on the inflorescence spikes: in
Merostachys sp. indet. (Pohl, 1980) and M. bur¬
manii, the spikelets are paired, and in M. ternata
Nees they are grouped in threes, thus the specific
name. Merostachys ternata is further distinguished
by its large spikelets (up to 14 mm long) and the
size of its leaves (35-50 cm long, up to 10 cm
broad). Thus, M. burmanii bears a close resem¬
blance only to the species described, but not named,
by Pohl from Costa Rica: it may be distinguished
from the latter by its inflorescence, 8-1 1 cm long,
with deflexed, nonpectinate, laterally outspread
spikelets. The inflorescence of Pohl’s species is 4.5-
6.5 cm long, with spikelets appearing to form a
single pectinate rank.
Burman & Soderstrom 525 is an unusual form
of M. burmanii, with occasionally 2 or 3 florets
(rather than 1) per spikelet. Exceptionally, under
the stress of late growth, when other plants have
already flowered and died in the region, a few plants
may tardily produce new shoots bearing late spike¬
lets. Some of these spikelets may be abnormal, with
2 or 3 florets each, sustained one above the other
by a continuous upward development of the rachilla.
As it represents an abnormal form, the collection
cannot be chosen as the holotype and has been
replaced by another, more representative collection.
This species is named for Alasdair G. Burman,
colleague and friend, who dedicates much of his time
to studying South American bamboos. It was he,
who, in 1980, in the Serra dos Orgaos, State of Rio
de Janeiro, Brazil, collected this species in the com¬
pany of the late Thomas R. Soderstrom, who on
that occasion had the intention of naming it for him.
Acknowledgments. I thank the Conselho Na¬
cional de Desenvolvimento Cientifico e Tecnologico
(CNPq) for a grant that has made this work possible.
I also thank the staff of the U.S. National Herbar¬
ium, Smithsonian Institution, for assistance and for
making their collections available for study, and the
anonymous reviewers for critically reading the
manuscript.
Literature Cited
McClure, F. A. 1973. Genera of bamboos native to the
New World (Gramineae, Bambusoideae). Smithson¬
ian Contr. Bot. 9: i-xii, 1-148 (edited by T. R.
Soderstrom).
Pohl, R. W. 1980. Gramineae. In: W. C. Burger (editor),
Flora Costaricensis. Fieldiana, Bot. n.s. 4: 321-322.
Muhlenbergia cualensis and M. michisensis (Poaceae: Eragrostideae):
Two New Species from Mexico
Yolanda Herrera A.
CIIDIR Unidad Durango, Instituto Politecnico Nacional,
Apartado Postal 738, Durango, Dgo., 34000 Mexico
Paul M. Peterson
Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, U.S.A.
ABSTRACT. Muhlenbergia cualensis from Jalisco,
and M. michisensis from Durango, Mexico, are
described and illustrated. A table distinguishing
among the new species and related taxa is given.
The genus Muhlenbergia Schreber is represented
in Mexico by at least 1 15 species, ol which 47
percent are endemic (Beetle, 1987). Morphological
characters that delimit the genus are spikelets with
a single perfect floret and lemma usually with three
prominent nerves. While collecting for a systematic
study of Muhlenbergia rnontana (Nutt.) A. Hitchc.
and related species in the Sierra Madre of Durango
(Herrera, 1991), a morphologically distinct taxon
was discovered. Later, during an examination of
herbarium specimens, a second new taxon was found.
The Muhlenbergia rnontana complex is a loose
assemblage of caespitose, nonrhizomatous perennials
of perhaps 12 species that usually have a dentate
or 3-toothed second glume and an awned lemma.
Members of this complex generally occur above
1,000 in in pine and oak forests from Montana to
California and Texas, and extend southward into
Mexico and Guatemala. These two new species and
an accompanying table are presented prior to com¬
pletion of the overall treatment of the Muhlenbergia
rnontana complex, so others in Mexico may become
aware of their existence and search for additional
locations. The specific epithet of M. cualensis is
derived from the Sierra el Cuale, the type locality,
and the epithet of M. michisensis refers to the small
village of San Juan de Michis where it was first
collected.
Muhlenbergia cualensis Herrera & P. Peterson,
sp. nov. TYPE: Mexico. Jalisco: SSE of Puerto
Vallarta, 14 km from the opening of the Zi-
inapan Mine, 1 km W of Providencia boundary
El Tuito-El Cuale, 20°15'N, 105°15'W, 31
Jan. 1983, H. Guzman 6090 (holotype, COCA;
isotypes, IBUG, US). Figure 1.
A Muhlenbergia eriophylla culmis altis 50-70 cm,
ligulis 10 12 mm longis; laminis planis ad involuta supra
hirsutula; paniculis 15-25 cm longis; lemmatibus
(2.6-)2.8-3.0 mm longis, aristatis luteolis, recedit.
Densely caespitose perennial without rhizomes.
Culms 50-70 cm tall, erect, rounded and branching
near the base, puberulent just above the mostly basal
nodes; internodes mostly glabrous. Sheaths 7-34
cm long, mostly longer than the lower internode,
glabrous to scaberulous, stiff and papery below, usu¬
ally folded; margins membranous. Ligules 10-12
mm long, membranous, hyaline, decurrent; apex
acuminate, often lacerate. Blades (ll-)20-25 cm
long, 0.5-2 mm wide, flat to involute, apically acu¬
minate, somewhat stiff, hirsutulous above and gla¬
brous to scaberulous below. Panicles 15 25 cm long,
0.7-4 cm wide, narrow to somewhat open, loosely
flowered, ascending branches mostly loosely ap-
pressed or spreading up to 35° from the culm axis;
pedicels 0.6-3 mm long, flattened, ascending, sca¬
brous to scaberulous; inflorescence branches 0.6-7
cm long; central axis with 4-6 ribs near base. Spike-
lets 3-4 mm long, erect, 1 -flowered. Glumes (3-)3.5-
4 mm long, oblong, longer than the lemma, usually
equal in length, 1-nerved, olivaceous, short pilose
near base and along nerve; apex acute to obtuse,
occasionally minutely erose. Lemma (2.6-)2.8-3.0
mm long, oblong-lanceolate, awned, olivaceous; mid¬
nerve, margins, and proximal !A to V\ loosely to
densely appressed pubescent to pilose, the hairs up
to 0.4 mm long; apex acute to acuminate; the awn
16-20 mm long, yellow, flexuous to straight. Palea
(2.6-)2.8-3.0 mm long, oblong, the proximal %
loosely to densely appressed pubescent to pilose be¬
tween the nerves; apex obtuse with an abruptly
mucronate tip. Anthers 1.6-2.2 mm long, purple.
Caryopsis not seen.
Novon 2: 114-118. 1992.
Volume 2, Number 2
1992
Herrera & Peterson
Muhlenbergia cualensis and
M. michisensis
115
Figure 1. Muhlenbergia cualensis Herrera & P. Peterson, Jalisco, Mexico (R. Guzman 6090). —A. Habit. —
B. Ligule. — C. Inflorescence. —D. Spikelet. —E. Glumes. —F. Lower glume, dorsal view. — G. Floret. —H.
Lemma, dorsal view. —I. Lemma, ventral view. —J. Palea, dorsal view. —K. Palea enclosing the pistil and lodicules,
ventral view. —L. Palea and stamens, lateral view.
116
Novon
Figure 2. Muhlenbergia michisensis Herrera & P. Peterson, Durango, Mexico (Herrera <& Acevedo 986). —A.
Habit. —B. Ligule. —C. Inflorescence. —D. Spikelet. —E. Glumes. —F. Lower glume, dorsal view. -—G. Floret.
— H. Lemma. —I. Palea, dorsal view. —J. Palea enclosing the stamens, pistil, and lodicules.
Volume 2, Number 2
1992
Herrera & Peterson
Muhlenbergia cualensis and
M. michisensis
117
Table 1. Salient characters distinguishing Muhlenbergia c
iserisis, and M. watsoniana.
ualensis, M. durangensis, M. eriophylla, M. micli-
Characters
M.
cualensis
M.
durangensis
M.
eriophylla
M.
michisensis
M.
watsoniana
Rhizomes
absent
present
absent
absent
absent
Ligule length (mm)
10 12
0.5-0.7(-l)
8 10
4-7
0.5 1
Adaxial blade surface
hirsutulous
scabrous
long villous
hirsute-glabrous
hirsutulous
Panicle length (cm)
15-25
10-25
4-13
12-17
4-12(15)
Spikelet length (mm)
3-4
(5-)6-7(-7.5)
3.5-4
4.5-5
3-3.8
Glume length (mm)
(3-)3.5-4
(5)6—7(—7.5)
3-4
4.5-5
3-4
Second glume shape
entire to erose
entire
dentate
entire
erose
Lemma length (mm)
(2.6-)2.8 -3
5-6.7(-7)
3-3.5
3.5-4
2.5-3.3
Lemma awn
flexuous-straight
flexuous
straight
flexuous
flexuous
Lemma awn color
yellow
yellow
olive-green
purplish yellow
yellow
Anther length (mm)
1.6-2.2
(2.5-)3-3.5
1.8-2
2-2.4
1.7-2
Distribution and habitat. Muhlenbergia cu¬
alensis is known only from the area surrounding
the Zimapan Mine at approximately 1,000 m in the
mountains southeast of Puerto Vallarta. At this lo¬
cation M. cualensis occurs on sandy to sandy clay
loam slopes with granitic stones in open forests of
Quercus and Pinus ayacahuite Ehrenb.
Additional specimens examined. Mexico. JALISCO: E
of the Zimapan Mine, 31 Jan. 1983, R. Guzman 6100,
6101, 6102, 6104 (all COCA).
Muhlenbergia cualensis is morphologically very
similar to M. eriophylla Swallen, which can be
distinguished from the former by possessing culms
17-40 cm tall, ligules generally 8-10 mm long,
leaf blades with large, silvery, densely appressed-
villous, unicellular macrohairs that measure 0.1-
0.2 mm wide and up to 4 mm long, panicles 4 13
cm long, and lemmas 3-3.5 mm long with olive-
green awns (see Table 1). The leaf blade in trans¬
verse section of M. cualensis differs from M. er¬
iophylla by having elliptical or vertically elongated
primary (1°) vascular bundles that alternate between
a single, smaller tertiary (111°) vascular bundle. In
M. eriophylla the 1° vascular bundles are circular
or round in outline, and there are two vascular
bundles, either secondary (11°), 111°, or in combi¬
nation, between each 1° vascular bundle.
Muhlenbergia cualensis resembles M. watson¬
iana A. Hitchc., although the latter differs in a few
morphological characteristics. Short ligules (0.5-1
mm long), folded leal blades, and short, narrow
panicles (4-12 cm long x 1-2 cm wide) are the
most prominent features of M. watsoniana.
The flavonoid chemistry of M. cualensis is most
similar to M. flaviseta Scribner and M. virescens
(Kunth) Kunth by sharing the following compounds:
quercetin 3-0 glucoside with M. virescens; and lu-
teolin 5-0 glucoside. luteolin 6 arabinoside, luteolin
7-0 galactoside, apigenin 6. 8 diglucoside (vitexin),
apigenin 7-0 arabinoside, tricin 5-0 glucoside, tricin
7-0 glucuronide, and 4'hydroxyflavone 7-0 gluco¬
side with both species (Herrera & Bain, 1991).
Muhlenbergia michisensis Herrera & P. Pe¬
terson, sp. nov. TYPE: Mexico. Durango: Mun-
icipio de Suchil, Las Escobas, San Juan de
Michis, 23°24'N, 104°8'W, 17 Sep. 1989,
Herrera & Acevedo 086 (holotype, CIIDIR;
isotypes, MTMG, US). Figure 2.
A Muhlenbergia durangensis rhizoinate nullo; ligulis
4-7 mm longis; spiculis 4.5-5.0 mm longis; glumis 4.5-
5.0 mm longis, glumis secundis 1-nervis vel 2-nervis;
lemmatibus 3.5-4 mm longis; paleis 2.8-3.5 mm longis,
apice 3-lobis; antheris 2-2.4 mm longis, recedit.
Densely caespitose perennial without rhizomes.
Culms 68-80 cm tall, branching and rounded near
base, glabrous just below the mostly basal nodes;
internodes glabrous. Sheaths 8-14 cm long, mostly
longer than the lower internode, scaberulous, be¬
coming flattened and conspicuously spirally coiled
below, loose and papery. Ligules 4-7 mm long,
membranous, hyaline, decurrent; apex acuminate.
Blades (8.5-)10-20 cm long, 1-3 mm wide, flat to
folded or involute, scabrous below and glabrous to
sparsely hirsute above. Panicles 12-17 cm long, 2-
4 cm wide, open, loosely flowered, ascending branch¬
es mostly spreading 20-50° from the culm axis;
pedicels 0.2-1 mm long, short and stout, scabrous
to hispid; inflorescence branches 0.4-3 cm long;
central axis triquetrous, 3-6-ribbed, scabrous.
Spikelets 4.5-5.0 mm long, erect, tightly clustered
or paired along each branch, 1-flowered. Glumes
4.5-5.0 mm long, oblaneeolate, entire, longer than
the lemma, usually equal in length, 1 -nerved, oc-
118
Novon
casionally the second 2-nerved, olivaceous, scabrous
along the entire length and usually short pilose near
base and along midnerve; apex acute. Lemma 3.5-
4.0 mm long, oblongdanceolate, hyaline to yellow¬
ish, awned; midnerve and margins on the proximal
Vi appressed pubescent to pilose, the hairs to 0.3
mm long; apex acute, minutely bifid, the lobes to
0.2 mm long, the awn 10-15 mm long, flexuous,
yellowish distally and purplish near base. Palea 2.8-
3.5 mm long, oblanceolate, loosely appressed pu¬
bescent between the nerves on the proximal %; apex
3-lobed to tridentate, the lateral lobes obtuse and
the central lobe acute. Anthers 2-2.4 mm long,
reddish purple. Caryopsis not seen.
Distribution and habitat. Muhlenbergia mich-
isensis is known from southeastern Durango, south¬
west of Vicente Guerrero between 2,450 and 2,650
m. It occurs on rocky slopes in forests composed ol
Quercus and Pinus.
Additional specimens examined. MEXICO. DURANGO:
Cerro Blanco, Reserva de la Biosfera “La Michilia,” 17
Apr. 1986, S. Gonzalez 3740 (CIID1R); San Juan de
Michis (Potrero de Escobas), 23 Jan. 1986, J. Alvarado,
s.n. (C11D1R).
Muhlenbergia michisensis is morphologically
similar to M. durangensis Herrera, which can be
distinguished from the former by having short rhi¬
zomes; ligules 0.5-0.7(-l) mm long; spikelets (5-)6-
7(-7.5) and glumes (5-)6-7(-7.5) mm long, the
second 3-nerved; lemmas 5-6.5(-7) cm long with
awns (10-)15-20(-25) mm long; paleas 5-6 mm
long; and anthers (2.5-)3-3.5 cm long (Table 1).
The flavonoid chemistry of M. michisensis is most
similar to M. eriophylla by sharing: quercitin 3-0
rhamnosilxyloside, luteolin 6 galactoside, luteolin 6
glucoside, luteolin 8 glucoside, apigenin 6, 8 dig-
lucoside (vitexin), apigenin 7-0 arahinoside, apigenin
7-0 diarabinoside, apigenin 7-0 glucoside, tricin 5-
0 glucoside, tricin 7-0 glucuronide, and
4'hydroxyflavone 7-0 glucoside (Herrera & Bain,
1991).
Acknowledgments. This study was partially sup¬
ported by the Banco de Mexico, COFAA-IPN, Sig-
ma-Xi, and an NSERC operating grant. We thank
William F. Grant and Susan Aiken for critically
reading an earlier version of the manuscript and
Alice Tangerini for providing the illustrations.
Literature Cited
Beetle, A. A. 1987. Noteworthy grasses from Mexico
XIII. Phytologia 63: 209 297.
Herrera, Y. A. 1991. A biosystematic study of Muh¬
lenbergia montana complex (Poaceae, Eragrosti-
deae). Master’s Thesis, McGill University, Montreal.
- & J. F. Bain. 1991. Flavonoid profiles in the
Muhlenbergia montana complex (Poaceae). Bio-
chem. Syst. Ecol. 19: 665-672.
Leucoloma crosbyi (Dicranaceae), a New Species
Endemic to Northern Madagascar
Ca t her in e IL a Fa rge-Engl and
Cryptogamic Herbarium, Department of Botany, University ol Alberta,
Edmonton, Alberta T6G 2E9, Canada
Abstract. Leucoloma crosbyi, a new species from
northern Madagascar, is described, resulting from
a revision of the pantropical genus. Diagnostic char¬
acters include: robust habit; narrow, opaque, jux-
tacostal bands tapering to basal region; costal-lam¬
ina] transition zone with 2-9 multi- to histratose
rows; interior cells forming broad scarious region;
narrow hyaline margin; and longitudinally thick-
walled alar cells. It is most closely related to L.
grandidicri Ren. & Card, and has been confused
with L. talazaccii Ren. & Card.
Ferdinand Renauld published Essai sur les Leu-
colomas in 1909, the first and only comprehensive
treatment of Leucoloma, enumerating 131 species
for the world. Renauld & Cardot (1915) produced
Les Mousses dc Madagascar, which described 38
species of Leucoloma for Madagascar. Crosby et al.
(1983) listed 51 species for Madagascar and nearby
islands, based on available literature. From current
revisionary work on the genus, collections from
Montagne d’Ambre by Crosby in 1972 have resulted
in the description of a distinct taxon, Leucoloma
crosbyi LaFarge-Kngland.
Leucoloma is a large pantropical genus that has
a strong center of diversity in the rainforests of east
Africa, Madagascar, and surrounding islands. Spe¬
cific endemism for this region is 94 percent. The
revision of the genus is expected to modify this
figure, though rates of endemism should remain high
for this region. The species described below supports
this view.
Leucoloma crosbyi LaFarge-England, sp. nov.
TYPE: Madagascar. Diego Suarez: Montagne
d’Ambre, Parc National, 10 km along trail be¬
tween Petit Lac and Grand Lac, 12°34'S,
49°12'E, 13 Nov. 1972, 1,200 m, Crosby &
Crosby 7167 (holotype, MO; isotype, ALTA).
Figures 1-12.
Dioicous. Plantae robustae, pallide vel glauco-virides
ad 5 cm altae. Folia longa et flexuosa. Cellulae juxtacos-
tales papillosae densissimae obscurae in vittis utrinque
% distalis costae latere dispositae. Latibasis scariosa
sensim angustata versus marginem hyalinam. Cellulae
alares planae, rufo-fuscae, scalariformes. Setae 1.0-2.0
mm longae. Capsulae 0.9-1.5 mm longae, ovales ad ob-
longae, erectae, immersae. Dentes peristomii quasi ad
basim divisi. Calyptrae initratae. Species haec ab Leu¬
coloma grandidieri Ren. et Card, differt vix rugosis lam-
inis et foliis longioribus, costa latiore et vittis cellularum
juxtacostalium papillosarum versus basim angustioribus.
Dioicous. Plants robust, erect to spreading, pale
to glaucous green or tan to light-brown, forming
loose tufts up to 5 cm tall. Stems red, nontomentous,
elliptic in transverse section (0.36 0.32 mm long
by 0.31-0.24 mm wide), central strand lacking,
outer 3-5 layers of red-brown, thick-walled cortical
cells with smaller lumen than the homogenous inner,
yellow to tan walled cortical cells; stems densely
leaved, unbranched or sympodially branched, fertile
branches distally ramulose from subapical innova¬
tions below perichaetia or perigonia, 1 to several.
Rhizoids smooth, red-brown, several to numerous
at the base of stem or potentially deciduous branches
encircling supporting axis. Juvenile leaves 0.3-3.0
mm long, ovate to ovate-lanceolate, apices shortly
acuminate to acuminate, at base of each branch and
stem. Leaves erect to wide spreading wet or dry,
4.0-6.5 mm long, 0.2-1.2 mm wide, narrowly ovate-
lanceolate to lanceolate, plane to slightly concave
below, not to slightly transversely rugose in medial
region, tapering gradually to a long, subulate, sub-
tubulous, nonrugose acumen; apices flexuose to
slightly incurved when dry. Mean acumina to shoul¬
der, shoulder to base ratio is 1 : 1.65. Margins plane,
entire, with a distinct, narrow, hyaline border rang¬
ing from 1 to 2 cells at the base of the leaf, reaching
a maximum width of 3-6 cells (9.6-16.8 gm wide)
in the median region of the leaf, narrowing to a
single cell in the upper region and ending below the
apex; apex rounded or acute, serrulate to multis-
errate. Costa 52.8-106.0 gm wide at base, sub-
percurrent, ending 3-6 cells below the apex, shiny,
translucent; transverse section plano-convex to el¬
liptic, 24.0-33.6 gm diam., with abaxial and adaxial
stereids in 3-5 rows, guide cells 4-6 in a single
row, sometimes double. Juxtacostal cells 4.8-12.0
fim long, 4.8-7.2 jxm wide, irregularly quadrate
Novon 2: 119-122. 1992.
120
Novon
figs. 1,2:-1mm
figs. 3,4,8,9:-1mm
figs. 5,6,7,10,11,12:- 0.01mm
Figures 1-12. Leucoloma crosbyi LaFarge-England. —1. Mature capsule, Crosby & Crosby 7167 (ALTA).
2. Operculum with fringed calyptra, 7167 (MO). —3. Calyptrae, 7167 (MO). —4. Polysetous perichaetium, immature
capsules, , 161 (MO). —5. Mature leaf apicies, 7167 (MO, ALTA). —6. Medial interior and juxtacostal cells, 7167
Volume 2, Number 2
1992
LaFarge-England
Leucoloma crosbyi
121
rounded to oblong, pluripapillose, with low multifid
papillae on abaxial and adaxial surfaces, filling acu¬
men and extending as narrow, opaque bands on
either side of the costa, gradually tapering as a
narrow V-shaped wedge in the lower l A of lamina,
bi- to multistratose in 2-6(-9) rows on either side
of the costa, becoming unistratose toward the in¬
terior cells, occasionally with isolated, irregular, nar¬
row bistratose bands, sharply delimited from smooth
interior cells. Interior cells smooth, hyaline to
brownish, nonchlorophyllose, thick-walled, forming
a shiny, scarious membrane, extending from the base
to the base of the acumen; medial to upper 24-
72(-106) gmt long, 6-7 gm wide, elongate to linear,
nonporose to porose; basal (24-)36-132 gm long,
7-11 gm wide, elongate to linear, strongly porose.
Marginal cells 168-204 gun long, 2-3 gun wide,
narrowly linear, hyaline, smooth. Alar cells quad¬
rate to rectangular, 12 72 gun long, 12—34 gun
wide, in 11-17 seriate columns, cell lumen con¬
spicuously granulose, longitudinal walls thick, ± no¬
dose, scalariform, red-brown (sometimes hyaline),
flat, sharply differentiated into a triangular to trap¬
ezoidal region, separated from the costa by a band
of strongly nodose, elongate, yellow-brown cells.
Perichaetia terminal, sessile. Perichaetial leaves
2-6 mm long, 0.75-1.1 mm wide, broadly ovate-
lanceolate, sharply contracted to long, narrow, acu¬
minate, aristate, flexuose apex; base clasping con¬
volute, alar cells not differentiated; basal cells 36-
60 gun long, 7-12 gun wide, elongate, strongly po¬
rose, yellow-brown to orange-brown; medial cells
elongate, elliptic, somewhat inflated (enlarged), or¬
ange-brown in basal region; margins subentire at
base, plane; costa 72-48 /urn wide at base, single,
excurrent. Archegonia 0.8-1.4 mm long, with necks
0.7-1.3 mm long, red-brown, ca. 20 28 per per-
ichaetium; with numerous hyaline paraphyses 0.7-
0.9 mm long. Perigonia terminal, sessile with 14
short subapical branehlets, each producing a ter¬
minal perigonium. Perigonial leaves 0.9-1.6 mm
long, ovate with abruptly narrowing, short, acumi¬
nate apices, convolute; basal cells 36 84 gun long,
7- 1 2 n m wide, elongate, yellow to yellow-orange,
extending up into 2 A of leaf; medial cells 24-6 i gtm
long, 7-24 gun wide; antheridia 0.70 0.95 mm
long, 0.10-0.15 mm wide, red-brown to yellow-
brown, with numerous hyaline paraphyses, 0.60
0.95 mm long. Sporophyte single or polysetous (to
Figure 13. Distribution of Leucoloma crosbyi.
4 per perichaetium observed), immersed. Seta stout,
short, 1.0-2.0 mm long, slightly twisted toward the
right, somewhat flexuose, orange-tan, smooth. Cap¬
sules oval to oblong 0.9-1.5 mm long, 0.5 0.6 mm
wide, stomates lacking, light brown to tan, annulus
not differentiated; exothecial cells 19-60 gmi long,
9.6 36.0 gim wide, rectangular to quadrate, becom¬
ing oblate just below rim. Operculum conic with
flanged base. Peristome teeth 16, asymmetrical,
bifid almost to 2 A of length, bases commonly reflexed,
distally erect, filiform (some capsules have incurved
peristome), inserted well below rim, red-brown, dif¬
ferentially thickened on inner surface; primary per¬
istomal layer with external surface smooth at base,
medial portion roughened to papillose, papillose to
striate-papillose distally; inner peristomal layer with
internal surface smooth at base, papillose to papillose
striate distally. Calyptra mitrate, broadly fringed to
lobate (7-20 lobes), hyaline, pale yellow to tan be-
(MO, ALTA). —7. Basal interior cells, Crosby & Crosby 7170 (MO). —8. Mature stem leaf, 7170 (MO). —9.
Juvenile leaf, Crosby & Crosby 7034 (L). —10. Alar cells, 7170 (MO). —11. Costal cross section, 7304 (L), 7167
(MO). —12. Exothecial cells just below mouth, 7167 (ALTA).
122
Novon
low, orange-brown above, smooth to slightly rough¬
ened above. Spores 24-31 /am, finely granulate.
Paratypes. Madagascar, diego suarez: Montagne
d'Ambre, Parc National, 10 km along trail between Petit
Lac and Grand Lac, 12°34'S, 49°12'E, 13 Nov. 1972,
1,200 m, Crosby & Crosby 7170 (MO); 0-5 kin along
trail between Petit Lac and Grand Lac, 12°34'S, 49°12'E,
12 Nov. 1972, 900-1,100 m, Crosby <£ Crosby 7304
(L).
Habitat. Corticolous, ramicolous, 900-1,200 m
above sea level.
Leucoloma crosbyi is distinguished by its robust
habit, with erect to wide spreading leaves with trans¬
parent, shiny laminae sharply differentiated from a
narrow band of opaque, densely papillose juxtacostal
cells gradually tapering to costa in the basal region,
which are not to hardly rugose, with long flexuose
acutnina. The leaves are bordered by a very narrow,
hyaline margin and have abruptly differentiated alar
cells, which are longitudinally thick-walled, red-
brown, arranged in 11-17 columns, forming a tri¬
angular to trapezoidal region. Leucoloma crosbyi is
distinguished from L. grandidieri Ren. & Card, by
its longer, stiffer, nonrugose leaves; opaque juxta¬
costal bands extending farther into the basal region
of the leaf and gradually narrowed; laminal trans¬
verse section showing a greater number of bi- or
multistratose rows between the costal and unistratose
laminal cells. Leucoloma grandidieri has 1 — 2(—3)
rows, and L. crosbyi has (2-)3-6( 9).
I he known distributions of these two species are
allopatric, L. crosbyi endemic to Montagne d’Ambre
at the northern tip of Madagascar and L. grandi¬
dieri confined to the eastern central region. Leu¬
coloma crosbyi is distinguished from L. talazaccii
by its more robust habit, lack of caducous lower
stems, larger leaves, longer interior cells, and well-
developed, granulose, scalariform alar cells.
The leaves of Leucoloma crosbyi vary from
smooth to slightly rugose in the medial opaque re¬
gion, with flexuose to slightly incurved acumina. The
apices vary from acute to rounded and toothed. The
number of rows of bi- to multistratose cells between
the costa and the unistratose lamina varies from 2
to 9. Isolated bistratose strips have been observed
within the opaque bands, as well as the more com¬
mon bistratose to multistratose transition between
the costa and unistratose lamina.
Distribution. Leucoloma crosbyi is known only
from Montagne d'Ambre, Province de Diego Suarez,
Madagascar (Fig. 13). I he distribution is based on
four specimens, which form the type material for
the new species.
Etymology. The species has been named L. cros¬
byi after Marshall R. Crosby, the collector of all
the known specimens. His contribution of recent
material from the 1970s, as well as the critical
compilation of the bryological literature, has helped
to establish a foundation for future bryological re¬
search in Madagascar.
Acknowledgments. Special gratitude is expressed
to Marshall R. Crosby for encouragement and sup¬
port in the initial part of this project and for his
rigorous collecting in the early 1970s, which has
provided the essential data for this taxon. Further
support and helpful discussions have been provided
by Dale H. Vitt.
Literature Cited
Crosby, M. R., U. Schultze-Motel & W. Schultze-Motel.
1983. Katalog der Laubmoose von Madagaskar und
den umliegenden Inseln. Willendowia 13: 187-255.
Renauld, F. 1909. Essai sur les Leucoloma. Prodome
de la Flore Bryologique de Madagascar, des Mas-
carenes et des Comores, Imprimerie de Monaco.
- & J. Cardot. 1915. Mousses de Madagascar.
In: A. & G. Grandidier, Histoire Physique, Naturelle
et Politique de Madagascar 39: 1-562.
Porotrichum cavifolium comb. nov. (Musci: Neckeraceae)
Robert E. Magill and Bruce Allen
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 62166, U.S.A.
ABSTRACT. Recent collections and new morpho¬
logical information justify the transfer of Pireella
cavifolia to Porotrichum and its treatment within
Neckeraceae in the Moss Flora of Mexico.
New morphological evidence and a review of re¬
cent collections of Pireella cavifolia (Card. & Herz.)
Card, have led to the following reevaluation of its
relationships.
Pireella cavifolia is a small, dendroid moss known
from the West Indies and Mexico south through
Guatemala (San Marcos, Croat 40961A, MO) to
Panama (Darien, Allen 8917C, MO) and Colombia
(Risaralda, I Volf 1435, MO). The species has un¬
differentiated alar cells and clearly differs from other
members of Pireella in having obovate leaves that
are broader above and lack a distinct flattened apical
portion. The sporophyte of P. cavifolia is unknown.
Allen (1987), in a broad survey of pseudopara -
phyIlia in the Pterobryaceae, found that the family
is characterized by the presence of filamentous pseu-
doparaphyllia. In his survey, the only two species
that lacked such structures were considered mis¬
placed in the Pterobryaceae. Pireella cavifolia does
not have filamentous pseudoparaphyllia; instead, it
appears to have either folious pseudoparaphyllia or
rudimentary branch leaves around its branch pri-
mordia. The absence of sporophytic evidence for P.
cavifolia makes its placement problematical, but on
the basis of its lack of filamentous pseudoparaphyllia
we exclude the species from the Pterobryaceae.
The presence in this species of folious structures
around the branch primordia, a dendroid habit, un¬
differentiated alar cells, and especially the occasional
production of flagelliform gemmiferous branches all
indicate the Neckeraceae as a likely placement for
it. Within the Neckeraceae, Porotrichum could ac¬
commodate it. This transfer is therefore made so
that the species can be treated within the Necker¬
aceae for the Moss Flora of Mexico.
Porotrichum cavifolium (Card. & Herz.) Magill
& Allen, comb. nov. Basionym: Pirea cavifolia
Card. & Herz. in Card., Rev. Bryol. 38: 39.
1911. Pireella cavifolia (Card. & Herz.) Card.,
Rev. Bryol. 40: 17. 1913. TYPE: Mexico.
Hidalgo: Honey-station, Pringle 15668 pro
parte (holotype, PC). Illustrations: Theriot
(1940, pi. 54, 5a-5f as Jaegerinopsis ramo-
sa); Arzeni (1954, pis. 22-23).
Acknowledgment. We thank the anonymous re¬
viewer of the manuscript for useful comments.
Literature Cited
Allen, B. 1987. On distinguishing Pterobryaceae and
Meteoriaceae by means of pseudoparaphyllia. Bryol.
Times 42: 1-3.
Arzeni, C. B. 1954. The Pterobryaceae of the southern
United States, Mexico, Central America and the West
Indies. Amer. Midi. Naturalist 52: 1-67.
Theriot, I. 1940. Complement au catalogue des mousses
de Cuba III. Mem. Soc. Cub. Hist. Nat. “Felipe
Poey” 14: 349-372.
Novon 2: 123. 1992.
Notes on the Solanaceae of China and Neighboring Areas
U illiam G. D'Arcy
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Zhang Zhi-yun
Institute of Botany, Academia Siniea, 141 Xizhimenwai Dajie, Beijing 100044, China
ABSTRACT. Nomenclatural changes are made for
the Flora of China, affecting Anisodus, Archi-
physalis, Leucophysalis, Physaliastrum, and Sco-
polia.
Anisodus
the nomenclature relating to the Asian genus
Anisodus and its type species differs in various floras
and revisions. The generic name and its type species
are clarified below, and another species is transferred
from Scopolia to Anisodus.
Since Anisodus was described in 1824, the tax¬
onomic validity of its separation from Scopolia Jacq.
(1764) has been disputed. Some workers have sim¬
ply placed species of Anisodus in Scopolia (I)unal,
1 852); others have recognized Anisodus as a section
or series in Scopolia (Wettstein, 1895; Weinert,
1972); and others have considered it a distinct genus
(Pascher, 1909; Chen & Chen, 1977; Wu & Chen,
1978; Sandina & Tarasevich, 1982). It will be
treated as a distinct genus in the Flora of China
(Zhang et al., in press).
Scopolia and Anisodus are members of the so-
lanaceous tribe Hyoscyameae, characterized by cap¬
sular fruits that are partly or completely covered
by the persistent fruiting calyx and strongly curved
embryos contained in compressed seeds. Both gen¬
era include herbaceous plants with subactinomorphic
flowers and five stamens situated at the level of the
corolla mouth. However, in addition to grossly dif¬
ferent overall appearance, the groups differ in the
conspicuous details shown in Table 1.
Sandina & Tarasevich (1982) studied the pollen
of Anisodus (Whitleya ) and Scopolia and found
differences, particularly in the structure of the pores
and the pattern of the exine, that they considered
to be of generic significance. Differences in the pollen
of Scopolia ( S. carniolica) (Punt & Monna-Brands,
1980) and Anisodus (A. carniolicoides (as S. car-
niolicoides), A. tanguticus (as A. mairei ), and A.
acutangulus (Zhang & Lu, 1984)) are contrasted
in fable 1.
Anisodus and Scopolia are widespread in the Old
World; their geography was reviewed by Weinert
(1972) and by Lu & Zhang (1986). When sepa¬
rated, the two genera have distinct geographical
ranges. Anisodus occurs in the Himalayan region
from western Nepal to Yunnan, China, and north¬
ward from Assam along the central Yangtze-Me-
kong-Salwin watersheds (Hengduan Mountains) of
China to nearly 39°N latitude. The two species of
Scopolia show a disjunction between Japan, ,S. ja-
ponica Maxim., and eastern Europe, S. carniolica
Jacq.
Weinert (1972) referred to an unpublished dis¬
sertation by Semenowa (1955), which we have not
seen, but names he attributed to her in his publi¬
cation were not effectively published by her nor
validly published by him.
Nomenclature of Anisodus and A. luridus
Anisodus Link in Sprengel, Syst. Veg. 1: 699.
1825 [1823 fide Flora 9: 2, 495. 1826, late
1824 fide Stafleu & Cowan 5: 813]. Scopolia
sect. Anisodus Dunal in DC., Prodr. 13(1):
555. 1852. TYPE: A. luridus Link ex Spren¬
gel.
If hit leya Sweet, News of Lit. & Fashion 3: 108. 13 Aug.
1925; Brit. Flower Gard. 2: 125. 1 Oct. 1825 [dates
fide Airy Shaw, J. Bot. 85: 192-193. 1937], TYPE:
If hitleya stramonifolia Sweet.
Anisodus was described by Link in Sprengel’s
edition of Caroli Linnaei . . . Systema vegetabi-
liurn, edition 16, which actually appeared before its
title page date of 1825. The original publication,
which included the description of A. luridus, cited
as a synonym Nicandra anomala Link, which was
not validly described until the following year by Link
and Otto (see below). The material on which Ani¬
sodus was based was grown in Berlin from seed
obtained from England that was noted as being from
Nepal. About a year later, the genus was again
described under the name Whitleya from different
material that also had come into cultivation in Eu¬
rope from seed obtained in Nepal.
Novon 2; 124-128. 1992.
Volume 2, Number 2
1992
D’Arcy & Zhang
Solanaceae of China
125
The plant known as Anisodus luridus was de¬
scribed four different times with three different ep¬
ithets in publications that bear the frontispiece date
1824 or 1825; confusion about its correct name
persists. The following reviews the history of naming
this plant and presents the correct name.
Anisodus luridus Link in Sprengel. Syst. Veg. 1:
699. 1825 [late 1824 fide Stafleu & Cowan
5; 813]. Scopolia lurida (Link) Dunal in DC.,
Prodr. 13(1): 555. 1852. TYPE: same as Ni-
candra anomala.
Physalis stramonifolia Wall, in Roxburgh, FI. India, ed.
Carey, 2: 242. 1824. [Mar.-Apr. 1824 fide Stafleu
& Cowan 4: 957]. Scopolina stramonifolia (Wall.)
Kuntze, Rev. Gen. PI. 2: 452. 1891. Scopolia stra¬
monifolia (Wall.) Shrestha, Bull. Dept. Med. PI.
Nepal 2: 81. 1969. Scopolia stemonifolia Chen
Cheih & Chen Chung-lien, Acta Phytotax. Sin. 15(2):
61. 1977, orthographic variant. Scopolia stramon¬
ifolia (Wall.) N. P. Balakrishnan, Bull. Bot. Surv.
India 22: 176. 1980 [ 1982] redundant combination.
TYPE: cultivated, London, from seed obtained from
Robert Henry Jenkinson, perhaps through Reginald
Whitley, noted as being from Nepal, specimen not
known. Nepal, herb. Wallich 2632 not seen, IDC
microfiche 7394: 295.
Nicandra anomala Link & Otto, Icon. pi. select. 77. t.
35. July-Dee. 1825 [fide Stafleu & Cowan 3: 68].
TYPE: cultivated, Berlin, from seed obtained from
England, noted as being from Nepal, specimen not
known (lectotype, Link & Otto plate 35; same as
Anisodus luridus). Scopolia anomala (Link & Otto)
Airy Shaw, J. Bot. 75: 195. 1937.
Whitleya stramonifolia Sweet, News of Lit. & Fashion
3: 108. 13 Aug. 1825; Brit. Flower Card. 2: 125.
1825 [1 Oct. 1825 date fide Airy Shaw, J. Bot. 85:
192-193. 1937]. Anisodus stramonifolius (Sweet)
G. Don in Louden, Hort. Brit. 61. 1830. Based on
Whitleya stramonifolia Sweet. Physalis stramon-
ifera Chen Cheih & Chen Chung-lien, Acta Phyto¬
tax. Sin. 15(2): 61. 1977. orthographic variant.
Anisodus stemonifolius Wu Cheng-yih & Chen
Cheih, FI. Reipubl. Popularis Sin. 67(1): 23. 1978,
orthographic variant.
Anisodusfischerianus Pascher, Repert. Spec. Nov. Regni
Veg. 7: 226. 1909. Anisodus luridus var. fischer¬
ianus (Pascher) C. Y. Wu & C. Chen, Acta Phy¬
totax. Sin. 15(2): 62. 1977. TYPE: cultivated in
Leningrad, seed from Sikkim-Tibet, specimen not
seen.
Scopolia mairei Leveille, Bull. Geogr. Bot. 25: 37. 1915
[Annee 24 (4° Serie)], fide Lauener, Notes Roy. Bot.
Gard. Edinburgh 37: 146. 1978. Anisodus mairei
(Leveille) C. Y. Wu & C. Chen, Acta Phytotax. Sin.
15(2): 64. 1977. TYPE: China. Yunnan (Yun-Nan):
Ma-Kong, rocks, 2,800 m, E. E. Maire s.n. not
seen.
The first description of a plant of Anisodus was
of Physalis stramonifolia by Nathaniel Wallich in
a compilation of his plant descriptions published by
Carey early in 1824. The description of Anisodus
Table 1. Differences between Scopolia and Aniso¬
dus.
Scopolia
Anisodus
Calyx and corolla conical
with straight lobes and
limb
Flower and fruits twice as
large
Calyx and corolla broadly
campanulate with re¬
curved lobes and limb
Corolla conspicuously ex-
serted, at least twice
as long as the calyx
Corolla shorter relative to
calyx, less than twice as
long or only slightly ex-
serted
Flowering calyx of five,
equal, membranaceous
lobes
Flowering calyx splitting
irregularly into 1-5 un¬
equal, fleshy or subcor-
iaceous lobes
Corollas reddish purple,
drying blue-purple
Pollen grains elliptical in
equatorial view, aper-
turate with scabrate
exine and 3 4-colpate
Corollas brownish purple
or yellow, usually dry¬
ing yellow
Pollen grains consistently
subspheroidal, nonaper-
turate with verrucate
exine and indistinct
stratification
luridus followed later in the year: late in the fol¬
lowing year, the same species was described as Whi¬
tley a stramonifolia by Robert Sweet. One might
suspect that Sweet’s name was in some way based
on that chosen by Wallich, but there is no evidence
to substantiate this. Sweet noted that his plant was
“nearest related” to Link's Anisodus, which was
first cultivated in Berlin in 1821, but he did not
mention Wallich’s work: his description is in differ¬
ent terms and differs in some details from that of
Wallich, for example, “peduncle short, densely vil¬
lous” compared with Wallich’s “peduncle . . . spar¬
ingly villous.”
Soon thereafter, George Don (1830) accepted
Link’s generic concept and transferred Sweet’s name
to Anisodus. Thus, the name A. stramonifolius is
occupied by A. stramonifolius (Sweet) G. Don, pre¬
venting the transfer of the earlier Physalis stra¬
monifolius Wall, into Anisodus and supplanting A.
luridus Link.
We recognize Anisodus as a genus, but for those
who place Anisodus in Scopolia, the correct name
in Scopolia is S. stramonifolia (Wall.) Shrestha.
We saw no material of Anisodus fischerianus
and place it in synonomy based on the original
description, which notes a funnelform-campanulate
flowering calyx. Wu & Ghen (1978), in making the
combination A. luridus var. fischerianus, noted
126
Novon
leaves with 1-3 pairs of irregular coarse teeth and
purple eyes inside the base of the corolla tube, but
apparently they did not see the type. Pascher sus¬
pected this species to be a hybrid of A. luridus and
A. tanguticus. Its provenance from Xizang (Tibet),
China, and Sikkim, India, is within the range of A.
lurida. Anisodus luridus is recorded from Sikkim,
Bhutan, Nepal, and in China from Sichuan, Xizang,
and northwestern Yunnan, occurring from 3,000
to 4,450 m.
Anisodus carniolicoides (C. Y. Wu & C. Chen)
D’Arcy & Zhang Zhi-yun, comb. nov. Basion-
ym: Scopolia carniolicoides C. Y. Wu & C.
Chen, Acta Phytotax. Sin. 15(2): 59. 1977.
TYPE: China. Yunnan: Deqen Xian, Baima
Mountain (Pei-ma-shan), 7. 7. Yu 8773 (ho-
lotype, HY). Paratypes: K. M. Fang 392, T.
T. Yu 9451, 1139 6, 73692 (all A).
Scopolia carniolicoides C. Y. Wu & C. Chen var. den-
tata C. Y. Wu & C. Chen, syn. nov. Acta Phytotax.
Sin. 15(2): 60. 1977. TYPE: China. Sichuan: Mu-
li Xian, T. T. Yu 6 291 (holotype, HY not seen;
isotype, A). Paratype: T. T. Yu 7180(A).
Following our review of material from the full
geographical range of the tribe Hyoscyameae—Eu¬
rope to Japan and the former U.S.S.H. to India
we conclude that Scopolia carniolicoides should be
placed in Anisodus and not with the European and
Japanese species that have been called Scopolia.
Variety dentata is based on specimens with toothed
leaves, but such leaves sometimes occur in other
specimens that are otherwise not different from more
typical plants. It should be noted that during the
period when Wu and Chen described these taxa,
difficulties beyond their control prevented consul¬
tation of collections from places outside of China.
I. YCIAA'THES
Lycianthes neesiana (Nees) D'Arcy & Zhang Zhi-
yun, comb. nov. Basionym: Solarium neesian-
um Wall, ex Nees, Trans. Linn. Soc. London
17: 42. 1837. Solarium subtruncatum Wall,
ex Dunal in DC., Prodr. 13(1): 180. 1852.
Lycianthes subtruncata (Wall, ex Dunal) Bit¬
ter, Abh. Naturwiss. Vereine Bremen 25: 478.
1919 [1920], TYPE: Wallich. Cat. supp.
2620, year 1828 [or later?].
Lycianthes neesiana was treated as Solanum sub¬
truncatum in the handwritten catalog of specimens
collected by Nathaniel Wallich that was prepared
by Wallich and George Bentham, and as Solanum
neesianum in the supplement to the catalog. The
catalog and supplement are handwritten lists without
descriptions, and they do not meet requirements for
effective, much less valid, publication. Up to the
present, this species has gone under the name So¬
lanum subtruncatum, probably because this name
was used first in the 1828 catalog; Solanum nee¬
sianum first appeared in the supplement, presum¬
ably a later publication. The names were not validly
published until later, the first being Solanum nee¬
sianum by Nees in 1837, who attributed the name
to Wallich. Lhe name Solanum subtruncatum was
not validated until 1852 by Dunal and is considered
to be a synonym of L. neesiana.
This species has a calyx with an entire, truncate
margin and 1-10 teeth arising below the apex, a
feature placing it in Lycianthes rather than Sola¬
num.
PHYSALIASTRUM
Solanaceae subtribe Physalidinae Miers includes
about a dozen genera characterized by having ac¬
crescent calyces, longitudinally dehiscent anthers,
and mostly rotate corollas. The subtribe is centered
in northern Mexico ( Chamaesaracha, Jaltomata,
Margaranthus, Physalis, Quincula), but it is also
represented in temperate Asia (Physaliastrum,
Physalis) and elsewhere. Generic lines have been
unclear, leading to diverse nomenclature. A series
of Asian species that has been variously known in
Archiphysalis, Chamaesaracha, Leucophysalis,
Physaliastrum, and Physalis appear to belong to
a single genus, which should be called Physalias¬
trum.
Physaliastrum was described by Makino (1914)
with two Japanese species, P. echinatum (Yatabe)
Makino and P. savatieri (Makino) Makino, which
were transferred from the American genus, Cha¬
maesaracha. In a revision of Chamaesaracha, Av-
erett (1973) concurred in excluding these species
from Chamaesaracha. Kuang & Lu (1965) revised
Physaliastrum, recognizing seven species, all in Asia.
Kuang (1966) described the genus Archiphysalis,
basing it on one element from Japan and another
from western China. These species had been known
as Physalis, Physaliastrum, or Chamaesaracha.
Averett (1977), extending his studies on Chamae¬
saracha, reviewed a series of the Chinese species
hitherto placed in Physaliastrum and concluded
that they and other enigmatic North American spe¬
cies are congeneric with Leucophysalis; he had
transferred these species into Leucophysalis earlier
(Averett, 1970, 1973). Following Averett’s view,
Grierson & Long (1978) described a new variety in
Leucophysalis from Bhutan.
Volume 2, Number 2
1992
D’Arcy & Zhang
Solanaceae of China
127
After studying a range of material for preparation
of the Solanaceae treatment for the Flora of Chirm
and for a study of leucophysalis grandiflora
(D’Arcy et al., 1990; D'Arcy & Keating, in prep.),
the type species of Leucophysalis, we are able to
separate the Asian from the American groups at the
generic level on the basis of aspects of their fruiting
calyces. In some of the Asian species the walls of
the fruiting calyx are elaborated by emergences that
give the entire living fruit a bristly appearance. This
is illustrated for Physaliastrum japonicum by Ya-
saka (1983: 80, figs. 4, 5) and by Yoshisuke (1985:
199). When dried, the emergences are sometimes
difficult to see, resembling flattened, near-hyaline
scales or giving the calyx wall a muricate or rough¬
ened appearance. In two other cases, which had
been segregated in the genus Archiphysalis, the
calyx modification is somewhat different. In one of
these, Physaliastrum sinense , the ribs are greatly
thickened. In the other, Physaliastrum chamae-
sarachoides (Makino) Makino, illustrated by Kuang
& Lu (1978: 51, plate 14, fig. 7, as Archiphysalis
kwangsiensis Kuang), the calyx ribs are thickened
and bumpy, suggestive of incipient teeth. Such emer¬
gences or elaborations are lacking on American spe¬
cies of Leucophysalis and related groups, e.g.,
Ph ysa lis, Chamaesarach a.
The separation of Physaliastrum from Physalis
rests largely on the elaboration of the calyx in Phys¬
aliastrum. Flowering material of most species of
Physaliastrum appears to be scarce in herbaria and
was not available for our study. However, the lit¬
erature notes that in most species of Physaliastrum
the corolla is lobed, while in most species of Phys¬
alis —all those from the Americas—the corolla is
apically subentire. (Although Waterfall's (1958,
1967) revision of the North American species in¬
cluded some species with lobed corollas, these have
since been removed to other genera.) Physalis al-
kekengi, perhaps the only species of Physalis native
to the Old W orld, does have shallowly lobed corolla
lobes, but it bas no sign of calyx elaborations typical
of Physaliastrum.
One of the main characters that was used to
justify separation of Archiphysalis is the degree to
which the berry fills the fruiting calyx. In the species
that were left in Physaliastrum, the berry fills the
fruiting calyx, which is appressed to the berry wall.
In Archiphysalis, the fruiting calyx greatly exceeds
the berry in diameter and length, and the berry is
free within the bladdery calyx. In Physalis, the fruit
is usually like that of Archiphysalis, but sometimes
it is like that in Physaliastrum. For example, in
Physalis philadelphica, which is widely marketed
for food in North America and Mexico, within a
single lot of fruits, some will have the berry filling
the calyx and tightly appressed to it, and others will
have a small berry that is free from the bladdery
calyx. This variability seems to be related to degree
of overall development of the fruit: larger berries
tend to fill the calyx, while younger or “poorer”
fruits tend to have loose calyces. Thus, the degree
to which the berry fills the calyx varies within a
single population or species, and because the species
in Physaliastrum and Archiphysalis are closely
related to Physalis, we do not think this is an ap¬
propriate character for generic separation of Ar¬
chiphysalis from Physaliastrum.
In Physaliastrum ( Archiphysalis) chamaesar-
achoides, there are no emergences on the fruiting
calyx walls, but the greatly thickened calyx ribs
have erect teeth, presenting a somewhat bristly ap¬
pearance. In Physaliastrum ( Archiphysalis ) si¬
nense, the calyx ribs are thickened, resembling those
of P. chamaesarachoides, but there are no teeth.
This species is more like Physalis than any of the
others in Physaliastrum, but differs in the thickened
calyx ribs, which we have not seen in any species
of Physalis or in any species of Physalidinae from
North America. We consider the Asian species to
be distinct from the American species and to form
the single genus Physaliastrum on the basis of the
elaborations of the fruiting calyx.
The differences noted between the species from
Archiphysalis and those previously part of Phys¬
aliastrum may warrant recognition of the group at
the sectional level or perhaps as a subgenus.
Physaliastrum yunnanense subsp. hhutani-
cum (Grierson & Long) D’Arcy & Zhang Zhi-
yun, comb. nov. Basionym: Leucophysalis
yunnanensis subsp. bhutanica Grierson &
Long, Notes Hoy. Bot. Card. Edinburgh 36:
141. 1978. TYPE: Bhutan: Rinchu-Kancham,
5,000 ft.. Cooper 3943 (holotype, E not seen;
isotype, BM not seen).
Although no material of this taxon was seen, the
description by Grierson and Long notes "the ac¬
crescent acutely muricate calyx,” which is char¬
acteristic of the Asian Physaliastrum and not the
American Leucophysalis.
Physaliastrum sinense (Hemsley) D’Arcy &
Zhang Zhi-yun, comb. nov. Basionym: Cha-
rnaesaracha sinensis Hemsley, J. Linn. Soc.,
Bot. 26: 174. 1890. Archiphysalis sinensis
(Hemsley) Kuang, x\cta Phytotax. Sin. 11(1):
62, pi. 8, 9, figs. 1 -6. 1966. Physalis sinensis
128
Novon
(Hemsley) Averett, Ann. Missouri Bot. Card.
57: 380. 1970. TYPE: China. Hubei (Hupeh):
Yichang (Ichang) Xian and immediate neigh¬
borhood, Henry 2902 (holotype, K not seen).
Acknowledgment. We are grateful to Armen L.
Takhtajan, V.L. Komarov Botanical Institute, Len¬
ingrad, for assistance relating to the unpublished
dissertation of Semenowa (1955).
Literature Cited
Averett, J. E. 1970 (1971). New combinations in the
Solaneae (Solanaceae) and comments regarding the
taxonomic status of Leucophysalis. Ann. Missouri
Bot. Card. 57: 380-381.
-. 1973. Biosystematics of Chnmaesaracha (So¬
lanaceae). Rhodora 75: 335-365.
-. 1977. Taxonomic notes and new combinations
in Leucophysalis (Solanaceae). Ann. Missouri Bot.
Card. 64: 141-143.
Chen, C. & Chen, C-l. 1977. On the Chinese genera
Scopolia Jacq., Anisodus Link et Otto and Atro-
panthe Pascher. Acta Phytotax. Sin. 15(2): 57-68.
D’Arcy, W. G., K. Pickett & R. C. Keating. 1990.
Investigation into Leucophysalis graruliflora. Wild-
flower 3: 20-26.
Don, G. 1830. Anisodus. P. 61 in J.C. Louden, Hurt.
Brit.
Dunal, M. F. 1852. Solanaceae. In; A. de Candolle,
Prodroinus 13( 1 b): 4-690.
Grierson, A. J. C. & D. G. Long. 1978. Notes relating
to the flora of Bhutan. Notes Roy. Bot. Card. Ed¬
inburgh 36: 139-144.
Jacquin, N. V. 1764. Observationum Botanicarum. Vi¬
enna.
Jaeger, P.-M. L. & F. N. Hepper. 1986. A review of
the genus Solanum in Africa. Pp. 41-55 in W. G.
D'Arcy (editor), Solanaceae: Biology and Systemat-
ics. Columbia Univ. Press, New York.
Kuang Ko-zen. 1966. Archiphysalis Kuang genus no¬
vum Solanacearum Asiae Orientalis. Acta Phytotax.
Sin. 11: 59-63.
- & Lu An-ming. 1965. Revisio Physaliastro-
rum Makino. Acta Phytotax. Sin. 10: 347-355.
- & -. 1978. FI. Reipubl. Popularis Sin.
Linnaeus, C. 1753. Species Plantarum. Stockholm.
Lu An-ming & Zhang Zhi-yun. 1986. Studies of the
subtribe Hyoscyaminae in China. Pp. 56-78 in W.
G. D’Arcy (editor), Solanaceae: Biology and System-
atics. Columbia Univ. Press, New York.
Makino, T. 1914. Observations on the flora of Japan.
Bot. Mag. Tokyo 28: 20-22.
Nees von Esenbeck, C. G. 1837. Monograph of the
East Indian Solaneae. Trans. Linn. Soc. London 17:
37-78.
Pascher, A. 1909. Zwei neue Arten der Gattung An¬
isodus (Solanaceae). Repert. Spec. Nov. Regni Veg.
7: 226-227.
Punt, W. & M. Monna-Brands. 1980. Solanaceae. Pp.
1-30 in W. Punt & G. C. S. Clarke (editors), The
Northwest European Pollen Flora, 8. Elsevier, Am¬
sterdam. Reprint from Rev. Palaeobot. & Palynol.
Supp. “NEPF” 1-30. 1977.
Sandina, L. B. & V. F. Tarasevich. 1982. Some pal-
ynological data on the study of the genera If hitleya,
Atropanthe, and Scopolia s. str. (Solanaceae). Bot.
Zhurn. (Moscow & Leningrad) 67: 146-154.
Semenowa, M. I.. 1955. Scopolia and its economic
significance. Dissertation, Library of Komarov Bo¬
tanical Institute, St. Petersburg. [Not seen.]
Waterfall, U. T. 1958. A taxonomic study of the genus
Phvsalis in North America north of Mexico. Rhodora
60: 107-173.
-. 1967. Physalis in Mexico, Central America
and the West Indies. Rhodora 69: 82-329.
Weinert, E. 1972. Zur Taxonomie und Chorologie der
Gattung Scopolia Jacq. Feddes Repert. 82: 617-
628.
Wettstein, R. von. 1895. Solanaceae. In: Engler &
Prantl, Natiirlichen Pflanzenfamilien 4(3b): 4-38.
Wu, C. Y. & C. Chen. 1978. Anisodus. Pp. 22-27
in Kuang Ko-zen & Lu An-ming, FI. Reipubl. Po¬
pularis Sin. 67: 158.
Yasaka, H. 1983. Wild Flowers of Japan. Yama to
Keikokusha, Tokyo.
Yoshisuke, S. 1985. Wild Flowers of Japan. Heibonsha,
Tokyo.
Zhang Zhi-yun & Lu An-ming. 1984. Pollen morphol¬
ogy of the subtribe Hyoscyaminae (Solanaceae). Acta
Phytotax. Sin. 22: 175-180.
-, - & W. G. D’Arcy. Solanaceae. Flora
of China, volume 17. (In press.)
67: 158.
Two New Combinations in the Endemic Hawaiian Genus
Cyanea (Campanulaceae: Lobelioideae)
Thomas G. hammers
Department of Botany, Field Museum of Natural History,
Chicago, Illinois 60605-2496, U.S.A.
Abstract. Examination of types for 74 names
published in Delissea by the late Harold St. John
showed that most represent previously described
species that are currently treated as members of the
related genus Cyanea. Two of the types, however,
represent morphologically distinctive species, which
are referable to Cyanea sect. Hirtellae. New com¬
binations in Cyanea are proposed, the two species
are described in greater detail, and a key to the
species of the section is provided.
Cyanea Gaudich. (Campanulaceae: Lobelioideae)
is a genus of trees and shrubs endemic to the Ha¬
waiian Islands. St. John (1987a) merged the genus
with its close relative Delissea Gaudich. and de¬
scribed 74 new taxa under the latter name (St. John,
1987b, 1988). Altogether, St. John recognized 166
species and 50 nonautonymic varieties and forms
within his concept of Delissea. In the treatment of
Campanulaceae (hammers, 1990) for the Manual
of the Flowering Plants of Hawaii, I argued lor
the retention of the traditional circumscription of
these two genera, recognizing 52 species plus nine
nonautonymic subspecies in Cyanea and just nine
species plus three nonautonymic subspecies in De¬
lissea. Publishing deadlines did not permit me to
examine types and account for all the new names
proposed by St. John in Delissea. However, it was
suggested that most would fall within my concept
of Cyanea and prove to be referable to previously
described species, particularly C. coriacea (A. Gray)
Hillebrand, C. fissa (H. Mann) Hillebrand, and C.
sylvestris A. Heller.
In May and June 1991, I visited BISH in order
to examine the relevant types and determine the
best disposition for each ol these names. This ex¬
amination confirmed the original suggestion that most
of St. John’s names should be treated as synonyms
of various previously described species. Two of the
types, however, did represent morphologically dis¬
tinctive species that were new to science and refer¬
able to Cyanea as circumscribed in the Manual.
These two species are here transferred to that genus
and described more fully. The remaining names will
be accounted for in detail in a forthcoming mono¬
graph of the genus (Lammers, in prep.).
Cyanea eleeleensis (H. St. John) Lammers, comb,
nov. Basionym: Delissea eleeleensis H. St. John,
Phytologia 63: 341. 1987. TYPE: Hawaiian
Islands. Kaua‘i: Wainiha Valley, on side of
intermittent stream below Pali ‘Ele’ele, shaded
gulch in wet forest, 700 ft., 19 July 1977,
Christensen 261 (holotype, BISH; isotype,
BISH). [The holotype consists of a standard
herbarium sheet bearing two detached leaves
plus a jar of liquid-preserved floral material.]
Shrub, 1.8 in tall; stem glabrous. Lamina oblan-
ceolate, 38.5-40.5 cm long, 11.5 cm wide, gla¬
brous; upper surface green; lower surface pale green;
margin minutely denticulate; apex acute; base cu-
neate. Petiole 7.8 8.5 cm long, 4 mm diam., gla¬
brous. Inflorescence 11 -20-flowered, densely short-
pubescent; peduncle deflexed, 8-9 cm long, 5-7
mm diam.; rachis 2.5-4 cm long; bracts linear or
narrowly triangular, 3 mm long, 1 mm wide, the
apex acute; pedicels 10-17 mm long, bibracteolate
in the lower Vz; bracteoles linear or narrowly tri¬
angular, 2 mm long. Hypanthium obovoid, deeply
10-sulcate, 10-11 mm long, 11-12 mm diam.,
densely short-pubescent. Calyx lobes triangular, 4-
5 mm long, 3 mm wide; apex acute. Corolla bilabiate,
46-47 mm long, purple with lighter longitudinal
stripes, short-pubescent; tube curved, 19 mm long,
7 mm diam., cleft dorsally for Vz its length; dorsal
lobes linear, 32 mm long, 3 mm wide, spreading
horizontally; ventral lip 19 mm long, cut to its middle
into three linear lobes 3-4 mm wide (the middle one
a little wider than the others) and acute at apex.
Staminal column slightly exserted, glabrous; fila¬
ments 35 mm long; anther tube 9 mm long, 3 mm
diam., the lower 2 anthers with tufts of white hairs
at apex. Berry (immature) subglobose, 12 mm long,
14 mm diam., purple, deeply 10-sulcate, crowned
by the persistent calyx lobes. Seeds unknown.
Known only from the type locality in Wainiha
Valley, Kaua’i, where it was found growing in wet
forest at an elevation of 213 m.
Novon 2: 129-131. 1992.
130
Novon
Cyanea kolekoleensis (H. St. John) Lammers,
comb. nov. Basionym: Delissea kolekoleensis
H. St. John, Phytologia 63: 344. 1987. TYPE:
Hawaiian Islands. Kaua‘i: Wahiawa Valley, left
side of stream, 765 m, 23 Sep. 1979, S. Perl¬
man 498 (holotype, BISH; isotypes, BISH—
2 sheets). [The holotype consists of a standard
herbarium sheet plus a jar of liquid-preserved
floral material.]
Shrub; stem unhranched, 1.5-1.9 m tall, gla¬
brous. Lamina narrowly elliptic, 16.5-27.5 cm long,
2.7-5.7 cm wide; upper surface green, glabrous;
lower surface greenish white, glabrous or the midrib
minutely and sparsely pubescent; margin minutely
serrulate; apex acuminate; base cuneate. Petiole
terete, 3.5-9 cm long, 2 mm diam., glabrous. In¬
florescence 4-8-flowered, glabrous; peduncle de-
flexed, 13.5-16 cm long, 2-4 mm diam.; rachis 3-
6.5 cm long; pedicels sharply recurved, 18 27 mm
long, reduced in length toward apex of rachis. IIy-
panthium obconic or ohovoid, 6-13 mm long, 6-
11 mm diam., densely short-pubescent. Calyx lobes
triangular or deltoid, 1.5-3 mm long, 1.5-3.5 mm
wide, the apex acute. Corolla bilabiate, white shading
to purple on the lobes, 50-52 mm long, densely
short-pubescent; tube curved, 30-39 mm long, 5.5-
9 mm diam., cleft dorsally for Vi its length; dorsal
lobes linear, 13-19 mm long, 1.5-3 mm wide, acute
at apex; ventral lobes linear, 10-15 mm long, 1.5-
3 mm wide, acute at apex. Staminal column ex-
serted; filaments 3.7-4.9 cm long, purple, glabrous;
anther tube dark purple, 9-11 mm long, 2.5-4 mm
diam., the lower 2 anthers with tufts ol white hairs
at apex. Berries unknown.
Endemic to the Wahiawa drainage of southern
Kaua'i, in wet Metrosideros forest at 650-765 m
elevation.
Additional specimens examined. Hawaiian Islands.
Kaua'i: Koloa District, Lihu'e-Koloa Forest Reserve, NW
of Wahiawa Bog, along tributary of Wahiawa Stream,
NW of stream and SE of Hulua, 7 Dec. 1988, Flynn &
If ood 3229 (F, PTBG); Wahiawa Mts., NE of Hulua,
near Waiinea-Koloa district boundary, 6 Sep. 1991, Perl¬
man et al. 12235 (F, PTBG).
Cyanea eleeleensis and C. kolekoleensis are both
referred here to section Hirtellae Hock, a group of
species endemic to Kaua'i (Rock, 1919). This sec¬
tion is characterized by shrubby habit; relatively
slender, unarmed stems; entire or minutely toothed
leaves; densely pubescent flowers; hypanthium 6-
13 mm long; triangular calyx lobes 1-5 mm long;
and corollas 25-52 mm long. Both are distinct from
the five previously known members of the group.
Their long deflexed peduncles are unique to the
section, as is the deeply sulcate hypanthium of C.
eleeleensis. They and the other members of this
section may be distinguished by the following key.
Key to the species of Cyanea sect. Hirtellae
la. Peduncle deflexed, 8-16 cm long, with an elon¬
gate rachis 2.5-6.5 cm long.
2a. Lamina oblanceolate, 38.5-40.5 cm long,
11.5 cm wide; peduncle 8-9 cm long, 5-
7 mm diam., 11-20-flowered, pubescent;
hypanthium deeply 10-sulcate, 11-12 mm
diam.; dorsal corolla lobes longer than the
corolla tube .
. C. eleeleensis (H. St. John) Lammers
2b. Lamina narrowly elliptic, 16.5-27.5 cm
long, 2.7-5.7 cm wide; peduncle 13.5-16
cm long, 2-4 mm diam., 4-8-flowered,
glabrous; hypanthium smooth, 6 11 mm
diam.; dorsal corolla lobes shorter than the
corolla tube .
. C. kolekoleensis (H. St. John) Lammers
lb. Peduncle horizontal or ascending, 0.7-6 cm
long, the rachis condensed or obsolete, less than
2 cm long.
3a. Lamina narrowly elliptic.
4a. Lamina 28-40 cm long, the margin
undulate; peduncle 43 60 mm long;
berries yellow . . . . C. undulata C. Forbes
4b. Lamina 12-28 cm long, the margin
flat; peduncle 7-10 mm long; berries
purple .... C. recta (Wawra) Hillebrand
3b. Lamina obovate, oblanceolate, or oblong.
5a. Lamina 32 80 cm long; petiole 0.5-
4 cm long, stout, winged; latex yellow;
berries broadly obovoid, 10-15 mm
diam., red-orange .
. C. Jissa (H. Mann) Hillebrand
5b. Lamina 10-37 cm long; petiole 2.5-
8 cm long, slender, terete; latex white;
berries globose or obovoid, 6-10 mm
diam., yellow or orange.
6a. Lamina 2-6 cm wide; calyx lobes
0.5-1 mm wide; corolla 30-40
mm long, the tube 3-4 mm diam.,
suberect .
. . . C. hirtella (H. Mann) Hillebrand
6b. Lamina 5-13 cm wide; calyx
lobes 1-2 mm wide; corolla 40-
48 mm long, the tube 4-5 mm
diam., gently curved .
. C. sylvestris A. Heller
Acknowledgments. The staffs of BISH and PTBG
are acknowledged for the many courtesies extended
during visits to their institutions.
Literature Cited
Lammers, T. G. 1990. Campanulaceae. Pp. 420-489
in W. L. Wagner, D. R. Herbst & S. H. Sohmer,
Manual of the Flowering Plants of Hawai'i. Univ. of
Hawai'i Press, Honolulu.
Rock, J. F. 1919. A monographic study of the Hawaiian
species of the tribe Lobelioideae family Campanula¬
ceae. Mem. Bernice Pauahi Bishop Mus. 7(2): i-xvi,
1-395.
Volume 2, Number 2
1992
Lammers
Cyanea
131
St. John, H. 1987a. Enlargement of Delissea (Lobeli-
aceae). Hawaiian Plant Studies 138. Phytologia 63:
79-90.
-. 1987b. Diagnoses of Delissea species (Lob-
eliaceae) from Kauai. Hawaiian Plant Studies 145.
Phytologia 63: 339-349.
-. 1988. Diagnoses of new Delissea species (Lob-
eliaceae). Hawaiian Plant Studies 162. Phytologia
64: 165-168.
Five New Species of Ferns from the American Tropics
Robbin C. Moran
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT. Five new species of ferns, Blechnum
anthracinum, B. werffii, Saccoloma squamosum,
Polypodium pinnatissimum, and Tectaria pubens,
are described as a result of work on general fern
identification at MO and for the pteridophyte volume
of Flora Mesoamericana.
Blechnum anthracinum R. C. Moran, sp. nov.
TYPE: Bolivia. La Paz: Provincia de Nor Yun-
gas, Cotapata, roadside behind gas station,
16°15'S, 67°50'W, 3,225 m, 27 July 1989,
Fay & Fay 2446 (holotype, MO; isotypes, LPB
not seen, UC, US). Figure 1.
Plantae epiphyticae. Rhizoma 5-12 mm latum, an¬
thracinum, reptans, internodiis 5-15 mm longis, stolo-
nifera, stolonibus ca. 1 mm longis, anthracinis, squamis
rhizomatis 3-6 mm longis, 1-2 mm latis, lanceolatis,
anthracinis, denticulatis. Folia sterilia et fertilia mono-
morphica, petiolis 20-70 cm longis, ad basim anthracinis,
ad apicein strainineis vel brunneis, omnino dense squa-
mulosis, squamis 0.5-1 mm longis, adpressis, ciliatis rub-
robrunneis, caducis. Laminae 20-55 cm longae, 6-13
cm latae, anguste lanceolatae, 30-55-jugatae, omnino 1-
pinnatae, pinnis 3-6.5 cm longis, 0.5-1 cm latis, falcatis,
basi valde cordatis auriculis rhachidi superpositis, ad basim
foliorum valde reflexis, supra sparse et infra inodice squa-
mulosis, squamis valde denticulatis vel dissectis. Rhachis
dense squamulosa, squamis rubrobrunneis, denticulatis,
denticulis apice interdum bifurcatis.
Plants epiphytic; rhizome 5-12 mm wide, black,
shiny, creeping, with internodes 5-15 mm long,
stoloniferous, the stolons ca. 1 mm wide, black,
shiny; rhizome scales 3-6 x 12 mm, lanceolate,
blackish, denticulate; sterile and fertile leaves mon¬
omorphous; petioles 20-70 cm long, blackish ba-
sally, stramineous to brown distally, densely scaly
throughout, with the scales 0.5-1 mm long, ap-
pressed, ciliate, reddish brown, caducous; lamina
20-55 x 6-13 cm, narrowly lanceolate, 1-pinnate
throughout; pinnae 3-6.5 x 0.5-1 cm, 30-55
pairs, falcate, with the base strongly cordate-auric -
ulate, the auricles sometimes overlapping the rachis,
the basal ones strongly reflexed, sparsely scaly above,
moderately scaly below, with the scales strongly
denticulate or highly dissected; rachis densely scaly,
with the scales reddish brown, denticulate, the apex
ol the denticulae sometimes bifurcate.
Paratype. Bolivia, la paz: Province of Nor Yungas,
1 km W of Chuspipata, elfin forest with Clusia, Wein-
mannia, and Myrica , 16°17'S, 67°49'W, 3,140 m, So¬
lomon 7259 (MO).
Blechnum anthracinum grows in cloud forests
in Bolivia, from 3,140 to 3,225 m. No other species
of Blechnum with monomorphic sterile and fertile
leaves grows at such high elevations or is epiphytic.
Several characteristics of this species readily distin¬
guish it from other monomorphic Blechnum species.
The rhizome and stolons are black and shiny (thus
the specific epithet). In leaf cutting, the pinnae are
short-stalked or sessile throughout the length of the
lamina (rather than adnate and confluent distally).
The basal pinnae are strongly reflexed. The pinnae
are more numerous (30-55 pairs) compared to
closely related species such as B. occidental L.
(12-25 pairs) or B. glandulosum Kaulf. (19-40
pairs). The scales on all parts of B. anthracinum
are strongly denticulate, and those on the rachis
and costae are so dense they obscure the surface.
The strongly toothed scales suggest a relationship
with B. stipitellatum (Sodiro) C. Chr., a species
known from Venezuela, Ecuador, and Peru. That
species, however, is terrestrial, has dimorphic sterile
and fertile leaves, larger rhizome scales, dull brown
rhizomes, and lacks stolons.
Blechnum werffii R. C. Moran, sp. nov. TYPE:
Panama. Chiriqui: Distrito Bugaba, Cerro Pun-
ta, around STRI house, cloud forest, rich in
epiphytes, 8°52'N, 82°33'E, 2,220 m, 27 Jan.
1985, t>an der Werff & Herrera 6500 (holo¬
type, MO). Figure 2.
Plantae terrestres. Rhizoma erectum nec subarbores-
cens, nec stoloniferum, squamis 1-1.5 cm longis, 0.2-
0.3 cm latis, ovatis vel lanceolatis, concoloribus, pallide
flavobrunneis, integris. Folia sterilia et fertilia dimorpha.
Folia sterilia 17-41 cm longa, petiolis 8-20 cm longis,
omnino atropurpureis vel pallide brunneis et maculis ob-
scurioribus conspersis. Laminae steriles 9-21 cm longae,
3-6 cm latae, 11-25-jugatae, utrinque glabrae, 1-pin-
natae, versus apicem pinnatifidae et gradatim angustatae,
gemmis carentibus; pinnis 1.5-3 cm longis, 0.6-1 cm
latis, anguste involutis, coriaceis, serratis, sessilibus, basi
cordatis, rhachidi superpositis; rhachis et costae steriles
pallide brunneae (vel rhachis atropurpurea), non papil-
losae, glabrae vel sparsim squamosae, squamis ovatis,
pallide aurantiaco brunneis, adpressis, integris; aerophora
basi pinnarum praesentia, minus quam 1 mm longa, ob-
longa. Folia fertilia 25-57 cm longa, quam foliis sterilibus
Novon 2: 132-138. 1992.
Volume 2, Number 2
1992
Moran
New Ferns from the American Tropics
133
Figure 1. Blechnum anthracinum R. C. Moran. —A. Fertile leaf. —B. Rachis and pinna juncture. Note the
slightly raised, circular aerophore on the rachis at the juncture. (Solomon 7259, MO.)
altiora, petiolis 17-35 cm longis, pallide brunneis et ina-
culis obscurioribus conspersis, pinnis 1.2-2.5 cm longis,
0.2-0.3 cm latis.
Plants terrestrial; rhizomes erect but not subar-
borescent, not stoloniferous, the scales 1-1.5 x
0.2-0.3 cm, ovate to lanceolate, concolorous, light
yellow-brown, entire; sterile and fertile leaves di¬
morphic; petioles 8-20 cm long, tan with brown
spots or atropurpureous; lamina 9-21 x 3-6 cm,
1-pinnate, buds absent, the apex tapered, pinnatifid;
pinnae 1.5-3 x 0.6-1 cm, pairs 11 25, narrowly
involute, coriaceous, serrate, sessile, with the base
cordate, overlapping the rachis; rachis and costae
light tan or the rachis atropurpureous, not papillose,
glabrous or sparsely scaly, the scales ovate, light
orange-brown, appressed, entire; aerophores present
at the pinna bases, less than 1 mm, oblong; laminar
tissue glabrous on both surfaces; fertile leaves 25-
57 cm long, erect, taller than the sterile leaves;
petiole 17-35 cm long, tan with brown spots or
atropurpureous; pinnae 1.2-2.5 x 0.2-0.3 cm.
Paratypes. COSTA RlC.A. HEREDIA: on the road N of
San Rafael, ca. 6 mi. above Monte de la Cruz, cloud
forest, 2,000 m, Moran 2269 (MO). Panama. chiriquI:
Guadalupe, Cerro Punta, Finca Alfaro, 2,000 m, Cab¬
allero 141 (MO, PMA).
Blechnum werjffii is endemic to Costa Rica and
Panama, where it grows in cloud forests from 2,000
to 2,200 m. The pinnae are distinctive by being
short, obtuse, coriaceous, and glabrous on both sur-
134
Novon
cn
o
3
i-
1 cm
Figure 2. Blechnum werffii R. C. Moran. —A. Entire plant. Note the long petiole of the fertile leaf. —B. Petiole
scales. —C. Pinna bases overlapping the rachis. (van der Werff & Herrera 6500 , MO.)
faces. The petioles of the fertile leaves are extremely
long relative to other species in the genus, being
about twice the length of the fertile lamina (Fig.
2A). I cannot suggest another species of Blechnum
that is closely related to B. werffii.
Saccoloma squamosum R. C. Moran, sp. nov.
TYPE: Ecuador. Zamora-Chinchipe: Nangar-
itza Canton, Valle del Rio Nangaritza, Miazi,
bosque sobre pendientes fuertes de roca caliza
o de pizarra, 78°40'W, 04°18'S, 1,200 m,
Palacios 6721 (holotype, MO; isotypes, F, NY,
QCNE, UC, US). Figure 3.
Plantae rheophyticae. Rhizoma erectum, squamis 0.5-
1 mm latis, 0.3-0.6 mm longis, linearibus, lucidis, cas-
taneis, denticulatis. Laminae 2-pinnato-pinnatisectae,
abaxialiter dense squamosae. Sporae flavidae.
Plants rheophytic; rhizome erect; scales 0.5-1
x 0.3-0.6 mm, linear, shiny, castaneous, dentic¬
ulate; petiole ca. as long as the lamina, brown, dense¬
ly scaly basally, lacking epipetiolar buds; lamina 15-
45 x 6-15 cm, lanceolate; pinna pairs 10-18,
alternate, stalked, the stalks 0.5-1 cm long; pinnules
1.5-4 cm long, anadromically arranged throughout,
slightly falcate and ascending, stalked, the stalks 1-
3 mm long; rachis, costules, and veins strongly scaly
abaxially, much less so adaxially, the scales 0.5-2
mm long, spreading, often tortuous, brownish, lucid,
denticulate; sori ending 0.3-0.5 mm before the mar¬
gin, embossed on the adaxial surface of the leaf;
spores yellowish.
Paratype. Ecuador. ZAMORA-CHINCHIPE: Nangaritza
Canton, Valle del Rio Nangaritza, Miazi, bosque sobre
pendientes fuertes de roca caliza o de pizarra, 78°40'W,
04°18'S, 1,200 m, Palacios 6737 (AAV, F, MO, QCNE).
Saccoloma squamosum most resembles S. in-
aequale (Kunze) Mett., which probably represents
a complex of several species. Saccoloma squamo¬
sum differs from all its congeners by its densely
scaly leaves, smaller leaf length, and linear, dentic-
Volume 2, Number 2
1992
Moran
New Ferns from the American Tropics
135
Figure 3. Saccoloma squamosum R. C. Moran. —A. Fertile leaf. —B. Pinnule base showing the strongly scaly
axes and veins. —C. Rhizome scale. (Palacios 6721, UC.)
ulate, castaneous rhizome scales. Whereas all other
neotropical species of Saccoloma are terrestrial, S.
squamosum is a rheophyte. The valley where the
specimens were collected is unusual geologically be¬
cause of its vast limestone and slate outcrops. It
may eventually be shown that S. squamosum is
restricted to these rock types.
Saccoloma was reviewed by Tryon (1962), who
recognized three species in the American tropics
(i.e., S. domingense (Sprengel) C. Chr., S. elegans
Kaulf., S. inaequale (Kunze) Mett.). Since then, the
number of taxa has doubled from the addition of S.
membranaceum Mickel, S. elegans subsp. char-
taceum Nair ex Cremers & Kramer, and S. squa¬
mosum.
Polypodium pinnatissimum R. C. Moran, sp.
nov. TYPE: Panama. Panama-San Bias bor¬
der: along El Llano-Carti road, ridge trail along
the divide, wet forest, 09°20'N, 78°58'W, 250-
300 m, Moran 4098 (holotype, MO; isotype,
UC). Figure 4.
Plantae epiphyticae. Rhizoma 5-8 mm latum, repens,
squamis 2-5 mm longis, 1-1.5 mm latis, ovatis, conco-
loribus, brunneis, non clathratis, integris, adpressis, pu-
bescentibus. Laminae (15-)25-40 cm longae, (6-)20-
30 cm latae, 1-pinnatae, apice conforme vel hastato,
pinnis (3 )10—15 cm longis, 0.8-1.9 cm latis, integris,
8-16 utroque rhachidis latere; rhachis adaxialiter pubes-
cens; venae liberae; sporangia setulosa.
Plants epiphytic; rhizome 5-8 mm wide, not pru-
inose, creeping, the scales 2-5 x 1-1.5 mm, ovate,
brown, concolorous, not clathrate, entire, oppressed,
usually comose basally with brown hairs at the point
of attachment; petiole ca. l A the length of the lamina,
brown, sparsely puberulent abaxially, exalate; lam¬
inae (15-)25-40 x (6-)20-30 cm, debate, 1-pin¬
nate throughout, not reduced basally, abruptly re¬
duced distally to a hastate or subconform terminal
segment; pinnae (3-) 10-15 x 0.8-1.9 cm, entire,
pairs 8-16, opposite or subopposite and sessile in
the basal part of the lamina, becoming adnate and
alternate distally; rachis brown, pubescent adaxially,
the hairs 0.2-0.3 mm long, 2-4-celled, antrorse;
136
Novon
Figure 4. —A. Fertile leaf of Polypodium pinnatissimum R. C. Moran (van der Werff 7027, MO). —B. Fertile
leaf of P. dulce Poiret in Lain. (Moran 3198, MO).
costae mostly whitish or brown basally, sparsely
pubescent abaxially, the hairs ca. 0.1 mm long,
occasionally branched, very pale reddish or brown¬
ish, moderately pubescent adaxially, the hairs ca.
0.2 mm long, antrorse, 2 4-celled; veins free; lam¬
inar tissue between the veins glabrous on both sur¬
faces; sori round to elliptic, in one row between the
costa and margin; sporangia setulose, the setulae
ca. 0.1 mm.
Paratypes. Costa Rica. ALAJUELA: Finca Los Ensayos,
ca. 11 mi. NW of Zarcero, 900 m, Croat 43509 (MO,
UC), 43512 (MO). SAN Jos£: Reserva Biologica Carara,
W part of Montanas Jamaica, ca. 3.5 km NE of Bijagual
de Turrubares, 9°45.5'N, 84°33.5'W, 500-600 m, Gra-
yum 8582 (MO); Zona Protectora La Cangreja, along
Quebrada Grande and on adjacent ridges, ca. 2 km NNE
of Mastatal de Puriscal, 9°42'N, 84°22'W, 400-540 m,
Grayum 8636 (MO, UC). PANAMA. COCLfe: along road
between Llano Grande and Coclesito (N of Pintada), 4
mi. N of Llano Grande, 600 m, Antonio 3577 (MO); La
Mesa, above El Valle, 800 m, Croat 25340 (MO); road
from La Pintada to Coclesito, 80°30'W, 8°45'N, 600 m,
Hamilton & Davidse 2845 (MO, UC); S of Cascajal
along Continental Divide, 8°45'N, 80°25'W, 800-900
m, Knapp 1989 (CR, MO, UC), 1991 (MO); ca. 3 km
NE of El Valle, Mori & Kallunki 2973 (MO); foot of
Cerro Pilon, above El Valle de Anton, ca. 600 m, Porter
et al. 4616 (MO); W slope and summit of Cerro Valle
Chiquito, 700-800 m, Seibert 510 (MO). col6n: Santa
Rita Ridge, end of road from Transisthmian Hwy., ca.
10 mi. from hwy., Porter et al. 4829 (MO). HERRERA:
18 km W of Las Minas, trail to top of Alto Higo, 800
m, Hammel 4220 (CR, MO). PANAMA: Cerro Campana,
near summit, 1,000 m, Croat 22805 (MO), 12149 (MO),
Kennedy et al. 2055 (MO), Porter et al. 4880 (MO),
Sytsma 3528 (MO), van der If erjf & Herrera 6204
(MO, UC). panama/san HI,AS: Nusagandi, van der ff erff
7027 (MO, UC). SAN blas: road from El Llano to Carti,
Pacific side, 79°00'W, 9°20'N, 200-300 m, Hamilton
& Stockwell 2936 (MO, UC); Campamento Nusagandi,
sendero Ina Igar en el Llano-Carti, 19 km de la Via
Interamericana, 350-400 m, Herrera & Irvine 352 ( MO,
UC, US). VERAGUAS: 5 mi. W of Santa Fe on road past
Escuela Agricola, Alto Piedra, on Pacific side of divide,
800-1,200 m, Croat 23058 (MO).
Polypodium pinnatissimum is closely related to
P. dulce Poiret in Lam. (a species that has previously
been called P. sororium Humb., Bonpl. & Willd.;
Volume 2, Number 2
1992
Moran
New Ferns from the American Tropics
137
Figure 5. Tectaria pubens R. C. Moran. —A. close-up of the abaxial surface of the leaf, showing dense pubescence
of long, septate hairs. —B. Sterile leaf. (Moran 3647, MO.)
Proctor & Lourteig, 1990). Both have free veins,
similarly cut laminae, and round to elliptic sori.
Polypodium pinnatissimum differs by its complete¬
ly 1-pinnate lamina (Fig. 4A) and setulose sporangia.
In contrast, P. dulce has mostly pinnatisect laminae
(or 1-pinnate basally; Fig. 4B) and glabrous spo¬
rangia. Three other differences occur between the
species, but these are not constant: P. pinnatissi¬
mum tends to have less visible veins, narrower (0.8-
1.9 cm wide) pinnae, and hastate or suhconform
apices. In contrast, P. dulce often has darkened
and easily visible veins (especially adaxially), wider
(0.9-2.5 cm wide) pinnae, and more or less evenly
tapered, pinnatifid apices. The two species also differ
in range: P. pinnatissimum occurs in Costa Rica
and Panama, whereas P. dulce occurs from southern
Mexico to western Panama (Chiriqui), Venezuela,
Colombia, Ecuador, and Peru. Most of the specimens
of P. pinnatissimum are from Panama, where P.
dulce is rare.
138
Novon
Tectaria pubens R. C. Moran, sp. nov. TYPE:
Peru. Loreto: Prov. Maynas, ca. 50 mi. down¬
river from Iquitos, at Peter Jensen’s Explorama
Lodge, virgin forest, clay soils, ca. 120 m,
Moran 3647 (holotype, MO; isotype, UC). Fig¬
ure 5.
Plantae terrestres. Rhizoma erectum vel decumbens,
squamis 5-10 mm longis, lanceolatis, brunneis, integris.
Petioli 40-60 cm longi, pubentes. Laminae 35-50 cm
longae, late ovatae, 1-pinnatae, gemmis carentes, pub¬
entes, pilis 0.4-1 mm longis, 5-8-cellularibus rufis prae-
ditae; pinnae basales 10-16 cm latae, falcatae, acumi-
natae, ad basim basiscopice lobatae. Indusia reniformia
vel circularia, pubescentia.
Plants terrestrial; rhizome erect or decumbent,
the scales 5-10 mm long, lanceolate, dull brown,
entire; lamina 35-50 cm long, broadly ovate, 1-
pinnate, lacking buds; basal pinnae 10-16 cm wide,
free with a short stalk up to 3 mm long, falcate,
acuminate, entire except for a single basal basiscopic
lobe; medial segment 20-30 x 8-12 cm, opposite,
falcate, entire, acuminate, decurrent; terminal seg¬
ment ovate, with one or two shallow lobes; rachis
and costae abaxially densely pubescent (the surface
often obscured); costules and veins pubescent abax¬
ially, the hairs 0.4-1 mm long, 5-8-celled, pale
reddish with most of the color aggregated in the
septae, the cells often flattened, adaxially glabrous
to sparsely pubescent; laminar tissue between the
veins pubescent abaxially, glabrous adaxially; sori
in 2(-4) rows between the main veins; indusia ren-
iform to circular, persistent, brown, concolorous,
pubescent on the surface and margins, the hairs ca.
0.1 mm long.
Paratypes. Ecuador. NAPO: Jatun Sacha Biological
Station, 8 km E of Misahualli, 1°04'S, 77°36'W, 400 m.
Fay & Fay 2714 (F, MO, QCNE). pastaza: Via Auca,
115 km al S de Coca, 10 km al S de la frontera Napo -
Pastaza, cerca del Rio Tiguino, carretera de PETRO-
CANADA en construccion, bosque humedo tropical, lomas
del suelo rojo, 01°15'S, 76°55'W, 320 m, Hurtado 3c
Neill 1486 (MO). Peru. LORETO: Prov. Maynas, Yana-
mono Explorama Tourist Camp, halfway between Indiana
and mouth of the Rio Napo, nonflooded forest on rather
fertile soil, 130 m, van der If erjff et al. 9886 (MO),
Gentry 29142 (MO, UC).
Tectaria pubens most closely resembles T. pilosa
because of its pubescent laminae and similar leaf
cutting. However, 7. pubens has pinnae 10-16 cm
wide, long-decurrent apical segments, and one pair
of free basal pinnae. In contrast, 7. pilosa has pinnae
2-5 cm wide, cuneate or short-decurrent apical
segments, and 1-4 pairs of free pinnae.
Tectaria pilosa (Fee) R. C. Moran, comb. nov.
Basionym: Cardiochlaena pilosa Fee, Mem.
Foug. 10: 45, t. 40, f. 4. 1866. TYPE: Brazil.
Rio de Janeiro: Weddell 656 (P? not seen).
This species has been previously identified in her¬
baria as Tectaria incisa Cav. var. pilosa (Fee) C.
Morton. It is here recognized at the specific level
because the plants are morphologically distinct and
do not intergrade with related species. In Costa Rica
and Panama, T. pilosa grows commonly in lowland
forests with T. incisa Cav. and 7. vivipara Jermy
& T. Walker. I have not seen intermediates that
might be interpreted as hybrids between these spe¬
cies. From a distance, T. pilosa can be distinguished
from the other two by its smaller leaves, lighter
green laminae, and fewer (1-4) pairs of pinnae. The
other species have leaves about twice as long, darker
green laminae, and more numerous (4-12) pairs of
pinnae. Closer examination of 7. pilosa reveals that
both surfaces of the lamina are pilose with spreading,
septate hairs 0.2-0.8 mm long. In contrast, both
7. incisa and T. vivipara are glabrous (except on
the axes). Fee’s illustration of the type appears to
match the Mesoamerican plants in cutting of the
lamina and abundance of long hairs.
Acknowledgments. I thank Alan R. Smith (UC)
and David B. Lellinger (US) for helpful comments
on the manuscript.
Literature Cited
Proctor, G. R. & A. Lourteig. 1990. Nomenclatura
plantarum Americanum. XIII. Pteridophyta. Llradea
5: 384-387.
Tryon, R. M. 1962. Taxonomic fern notes. III. Contr.
Gray Herb. 191: 91-107.
New Species of Ruellia (Acanthaceae) from the Venezuelan Guayana
Dieter C. Wasshausen
Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, U.S.A.
ABSTRACT. Recent expeditions in the Venezuelan
Guayana have resulted in additional new taxa in the
genus Ruellia (Acanthaceae). The following places
these novelties on record for the convenience of
other taxonomists prior to the publication of the
Flora of the Venezuelan Guayana. Five new Ven¬
ezuelan species of Ruellia, R. wurdackii, R. stey-
ermarkii, R. delascioi, R. liesneri, and R. boli-
varensis, are described, illustrated, and compared
with their closest relatives.
Ruellia L. is the second largest genus of Acan¬
thaceae, with approximately 250 species of peren¬
nial herbs and shrubs with a pantropical distribution.
The genus, named for Jean de la Ruelle (1474-
1537), a French physician and herbalist, exhibits
large and usually showy flowers borne either singly
or in various inflorescence forms, which are either
Key to the Venezuelan Guayana Species of Ruellia
la. Inflorescence paniculate, flowers borne on simple or dichotomously branched penduncles or flowers in clusters
of 2-4 at the leaf axil.
2a. Flowers axillary, sessile, verticillate and clustered.
3a. Corolla narrowly infundibular, 25 mm long, white with violet veins; leaves petiolate, the blades
ovate-oblong . R. merit hoides
3b. Corolla funnelform, 35-50 mm long, light pink to pale mauve or purple; leaves sessile to subsessile,
the blades linear to narrowly lanceolate . R. geminiflora var. angustifolia
2b. Flowers terminal and axillary, pedicellate, pedunculate, the peduncles 25-100 mm long.
4a. Flowers terminating conspicuously long peduncles, these 70-100 mm long.
5a. Corolla tube white, the limb 15 mm broad, the lobes obovate, ca. 6 mm long and wide;
peduncles slender, curved .1. R. wurdackii
5b. Corolla tube scarlet or crimson, the limb 25-30 mm broad, the lobes oblong, 12 mm long,
5-6 mm wide; peduncles rather stout, erect-ascending . R. macrophylla
4b. Flowers not terminating on conspicuously long peduncles, these 25-60 mm long (if peduncle is
more than 40 mm long then inflorescence is a terminal panicle 60-130 mm long).
6a. Corolla yellow, greenish yellow, or pale green, the limb oblique.
7a. Corolla pale green, 30-33 mm long, the lobes revolute-twisted with age; calyx segments
narrowly ovate, 14-17 mm long, winged on the angles; leaf blades elliptic to oblong, 7-
13 cm long and 3.5-5 cm wide .2. R. steyermarkii
7b. Corolla pale yellow or greenish yellow, 35-60 mm long, the lobes spreading with age;
calyx segments linear-lanceolate, 25-30 mm long, membranous; leaf blades ovate or
oblong-ovate, 18-30 cm long and 8.5-12 cm wide.
8a. Corolla pale yellow, 50-60 mm long, 20 mm wide near the mouth; stamens exserted,
not reflexed; leaf blades oblong-ovate, 26-30 cm long, 10.5-12 cm wide, narrowed
at the base . R. rnalaca
8b. Corolla greenish yellow, 35 mm long, 10 mm wide near the mouth; stamens exserted,
reflexed; leaf blades ovate, 18-20 cm long, 8.5-9.3 cm wide, obtuse at the base
.3. R. delascioi
6b. Corolla mauve, the limb regular . R. tuberosa
lb. Inflorescence spicate or a 2-3-flowered terminal fascicle.
axillary or terminal or both. The funnelform or sal-
verform corollas are either red, yellow, white, or
purple (mauve) with a narrow tube and an often
campanulate limb with five spreading lobes. The
genus is characterized by the four didynamous sta¬
mens each with bilocular anthers, which are usually
muticous at the base. The capsules are either oblong
or clavate, with the dissepiment remaining attached
to the capsule wall at maturity. Each capsule con¬
tains between 4 and 20 piano-compressed, subor-
bicular seeds. These are provided with a mucilagi¬
nous coat that exhibits, after moistening, tapered,
short trichomes with walls marked internally by rings
or spirals (Long, 1970). In an attempt to identify
recent collections of Ruellia from the Venezuelan
Guayana, I have found five undescribed species ol
the genus.
Novon 2: 139-148. 1992.
140
Novon
9a. Flowers borne in terminal, 2 3-flowered fascicles or terminal, few-flowered, congested spikes.
10a. Corolla scarlet or red, 45-50 mm long; petioles 5-12 mm long, pilose and puberulous .
.4. R. liesneri
10b. Corolla white, 60-80 mm long; petioles 3-5 mm long, puberulous . R. sprucei
9b. Flowers borne opposite in pairs on lax, axillary and terminal spikes .5. R. bolivarensis
1 . Ruellia wurdackii Wasshausen, sp. nov. TY PE:
Venezuela. Bolivar: Bio Parguaza, at base of
Piedra Mapollos, E bank of river about 10 km
above El Carmen (about 60 river kin lrorn
mouth), 110 m, 2 Jan. 1956, J. J. Wurdack
& J. V. Monachino 41082 (holotype, US; iso¬
type, NY). Figure 1E-G.
Suffrutex caulibus quadrangularibus, angulis rotunda-
tis. Folium lamina oblongo-elliptica vel ovata, graciliter
acuminata, basi angustata in petiolum decurrenti, mem-
branacea, supra atroviridi, subtus pallide viridi costa et
venis prominentibus, utrinque glabra; petiolo glabro. In-
florescentiae ex cymis dichotomis constantes; pedunculis
gracilibus, glabris vel parce puberulis; bracteolis anguste
oblongis glabris vel puberulis et parce glanduloso-pilosis;
calycis segmentis inaequalibus, anguste linearibus, glan¬
duloso-pilosis; corollae tubo albo, leviter curvato, limbo
suberecto, purpureo, ad basim pallide luteo lobis obovatis,
rotundatis; staminibus inclusis.
Shrub or suffrutescent herb; stems unbranched,
25-40 cm tall, quadrangular, 1-1.5 mm wide, the
angles rounded. Leaves petiolate, petioles about 10
mm long, glabrous, narrowly winged, the blades
oblong-elliptic to ovate, 8 11 cm long and 2.5-4
cm wide, long, slenderly acuminate at apex, atten¬
uate, membranous, entire or shallowly crenate, rath¬
er thin, the upper surface dark green, glabrous, the
lower surface paler green, glabrous, the costa and
lateral veins (about 7 pairs) somewhat more con¬
spicuous underneath, the cystoliths numerous and
conspicuous under a lens. Flowers borne in lax di-
chotomously branched cymes, the peduncles slen¬
der, curved, subquadrangular, about 70 mm long,
glabrous or sparingly and inconspicuously puberu¬
lous, the primary branches 1.5-2.5 cm long, pu¬
berulous, the trichomes erect, occasionally gland-
tipped; bracteoles narrowly oblong, 3.5-4 mm long,
0.75 mm wide, glabrous to puberulous and sparingly
glandular-pilose, the pedicels 2 mm long, subquad¬
rangular, puberulous, the trichomes minute and erect;
calyx segments unequal, narrowly linear, 11-14
mm long, 1 mm wide, glandular-pilose, the trichomes
spreading, 0.5-2 mm long; corolla 46 mm long, the
tube white, slightly curved, 32 mm long, 1.6 mm
broad at base, narrowed to 1 mm at 1 nun above
base, the throat turned up from the tube, 8 mm
wide, the limb suberect, purple with darker spots on
the lower portion, pale yellow at base, the lobes
obovate, about 6 mm long and wide, rotundate;
stamens included, about 8 mm long; pollen sphe¬
roidal, 92 /am diam., the reticulum homobrochate,
membrane granular (Fig. 3A, B). Capsule clavate,
1.2 cm, 2.5 mm wide and thick, narrowed at the
base, the solid portion 4.5 mm long, apically obtuse,
minutely and sparingly puberulous, upward the tri¬
chomes are intermixed with a few glands. Mature
seeds wanting.
Distribution. Occasional in forests and understo¬
ry in Amazonian Venezuela and Brazil at elevations
between 70 and 250 m.
Paratypes. VENEZUELA. TERRITORIO FEDERAL AMAZONAS:
Depto. Atures, road Puente Cataniapo-Gavilan, 20-25
km SE of Puerto Ayacucho, 90 m, P. J. M. Maas & O.
Huber 5094 (U, US); carretera via a Gavilan puente de
Paso del Diablo hasta 2 km mas adelante, sur-este de
Puerto Ayacucho, 5°30'N, 67°27'W, 70 80 m, Fran¬
cisco Gudnchez 274 (US). BRAZIL. PARA: Serra dos Ca-
rajas, 8.5 km W of AMZA camp N-l on road to Rio
Itacaiunas, 5°59'S, 50°20'W, 250 m, C. R. Sperling ,
R. S. Secco, M. Condon, .4. L. Mesquita, II. G. S. Ribeiro
& L. R. Marinho 5795 (NY).
Ruellia wurdackii is related to another Vene¬
zuelan species, R. pterocaulon Leonard. In R. pter-
ocaulon the corolla is 25 mm long, the corolla tube
is lavender, erect, 15 mm long, the leaf blades are
firm and rigid, nitid, acuminate with an obtuse tip,
and the stem is about 100 cm tall and 3 mm wide.
In contrast, the corolla of R. wurdackii is 46 mm
long, the corolla tube is white, slightly curved, 32
mm long, the leaf blades are membranous, dark
green above, paler green below, long, slenderly acu¬
minate, and the stem is 25-40 cm tall and 1-1.5
mm wide.
2. Ruellia steyermarkii Wasshausen, sp. nov.
TYPE: Venezuela. Bolivar: Altiplanicie de Nu¬
ria, cloud forest on summit of SE-facing es¬
carpment, E of Cerro El Picacho, N of Las
Nieves and Las Chicharras, 45 km N of Tum-
eremo, vicinity of Deborah, 600-650 m, 5-8
Feb. 1961, Julian A. Steyermark 89113 (ho-
lotype, US; isotypes, NY, VEN). Figure 2A-G.
Frutex volubilis caulibus subteretibus, simplicibus vel
ramosis, glabris. Folium lamina elliptica vel oblonga, brev-
iter acuminata, basi angustata, subcoriacea vel membra-
nacea, supra atroviridi, subtus pallide viridi costa et venis
prominentibus, utrinque glabra; petiolo glabro. Inflores-
centiae ex paniculis terminalibus, puberulis constantes;
pedunculis puberulis; bracteolis oblanceolatis, glabris, cil-
iolatis; calycis segmentis atrovirentibus, anguste ovatis,
Volume 2, Number 2
1992
Wasshausen
Ruellia
141
Figure 1. A-D. Ruellia menthoides (Nees) Hiern (F. Guanchez 1954). - —A. Habit. —B. Bract. —C. Bracteole
and calyx segments. —D. Corolla. EG. Ruellia wurdackii Wasshausen (Wurdack & Monachino 41082). —E.
Habit. —F. Bracteoles and calyx segments. —G. Calyx segments and corolla.
142
Novon
Figure 2. A-G. Ruellia steyermarkii Wasshausen (J. A. Steyermark 88995). —A. Habit. —B. Bracteoles, calyx
segments, and pistil. —C. Bracteoles, calyx segments, and ovary. —D. Corolla, stamens, and pistil. —E. Expanded
corolla showing attachment of filaments. —F. Calyx segments and capsule. —G. Seed.
Volume 2, Number 2
1992
Wasshausen
Ruellia
143
glabris, ciliatis; corolla pallide viridi, parce puberula, tubo
leviter curvato, limbo obliquo, lobis vetustate revoluto-
tortis, suborbicularibus; staminibus exsertis. Capsulae ovo-
ideae, obtusae et apiculatae.
Vining subsprawling; stem erect, simple or
branching, glabrous, brownish, subterete. Leaves
petiolate, the petioles 0.5-1.5 cm long, canaliculate,
glabrous; the blades elliptic to oblong, 7-13 cm long
and 3.5-5 cm wide, short-acuminate at apex, nar¬
rowed at base, subcoriaceous or membranous, entire
or undulate, both the upper and lower surface gla¬
brous, dark green above, paler green below, the
costa and lateral veins (4-6 pairs) conspicuous be¬
low, less so above, the cystoliths barely visible under
a lens on the upper surface. Panicle terminal, 6 9
cm long and 6-7 cm wide, puberulous, the trichomes
brownish, eglandular, the peduncles 2.5 3.5 cm
long, puberulous; bracteoles oblanceolate, 12-14
mm long, 2-3 mm wide, glabrous and ciliolate; calyx
grass green, winged on the angles, the segments
narrowly ovate, 14-17 mm long, 7-8 mm wide,
acuminate, glabrous and ciliolate, the cystoliths con¬
spicuous under a lens; corolla pale green without,
buff-green within on the lobes, sparingly puberulous
with both glandular and eglandular trichomes, 30-
33 mm long, the tube sometimes slightly curved,
hasally 4 mm wide, near mouth 15 mm wide, the
limb oblique, 22-25 mm wide, the lobes revolute¬
twisting with age, suborhiculate, 9 mm long, 1 1 mm
wide; stamens exserted, to 45 mm long; filaments
greenish white, glabrous; anthers buff-brown, ob¬
long, 8 mm long, 2 mm wide, glabrous, basal lobe
7 mm long; pollen spheroidal, 91 /am diam., the
reticulum homobrochate, the membrane granular
(Fig. 3C, D); ovary soft tomentose, the trichomes
brownish; style greenish white, slender, exserted about
30 mm beyond mouth of the corolla. Capsule ovoid,
20 mm long, 1 0 mm wide, obtuse and apiculate at
apex, puberulous, the trichomes appressed; seeds
flat, 6 mm long and wide, minutely moss-like-pu-
berulous when moistened; retinacula subcarinate,
curved, 5 mm long, 0.5 mm wide, erose at tip.
Distribution. Steep moist forested slopes over
boulders just below summit and on ridge on summit
at elevations between 200 and 650 m.
Paratypes. VENEZUELA. BOLfVAR: Rio Toro (Rio Gran¬
de), between Rio La Reforma and Puerto Rico, N of El
Palmar, 200-250 m, Julian A. Steyermark 88139 (VEN);
Cerro El Picacho and vicinity, N of Las Chicharras, 45
km N of Tumeremo Altiplanicie de Nuria, 300-620 m,
Julian A. Steyermark 88995 (NY, US, VEN); Camp “El
Paraiso” and “La Yugua,” 48 km NE of village Los
Rosos, E end of 17 km of Upata, Carlos Blanco 447
(US).
Ruellia steyermarkii is perhaps most closely re¬
lated to another Venezuelan Guayana species, R.
rnalaca Leonard, which differs markedly by having
linear-lanceolate calyx segments; pale yellow corol¬
las 50 60 mm long; and oblong-ovate, 26-30-cm-
long, 10.5-12-cm-wide leaf blades. In contrast, R.
steyermarkii has narrowly ovate, winged on the
angles calyx segments; pale green without, buff-
green within corollas, 30-33 mm long; and elliptic
to oblong, 7-13 cm long, and 3.5-5 cm wide leaf
blades.
3. Ruellia delascioi Wasshausen, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas: Dep-
to. Atabapo, Salto Yureba, Bajo Ventuari, 4°3'N,
66°1'W, 120-150 m, 24 Oct.-4 Nov. 1981,
Francisco Delascio & Francisco Gudnchez
10693 (holotype, IJS). Figure 4A-C.
Suffrutex caulibus erectis, simplicibus vel ramosis, to-
mentosis. Folium lamina ovata, breviacuminata, basi ob-
tusa, aliquanto firma, utrinque molliter hirtella; petiolo
dense tomentoso vel minute puberulo. Inflorescentiae ex
paniculis terminalibus, dense puberulis constantes; pe-
dunculis dense et molliter tomentosis; bracteolis persis-
tentibus, dense puberulis. Flores persistentes; calycis seg-
mentis firmis, lineari-lanceolatis, longo-acuminatis, dense
puberulis, costa et venis obscuris; corolla minute glari-
duloso-puberula, viridi-flava, tubo curvato, limbo obliquo,
lobis rotundatis; staminibus reflexis.
Shrub or subshrub; stem erect, simple or branch¬
ing, grayish, subquadrangular, densely and softly
tomentose. Leaves petiolate, the petioles 3.5-5 cm
long, densely tomentose or minutely puberulous, the
blades ovate, 18-20 cm long and 8.5-9.3 cm wide,
short-acuminate at apex, obtuse at base and briefly
decurrent on the petiole, firm, entire or undulate,
the upper surface softly hirtellous, drying dark green,
the trichomes erect, the lower surface paler green,
softly hirtellous, the trichomes erect, grayish, the
costa and lateral veins (6-8 pairs) rather prominent,
especially below, the cystoliths inconspicuous under
a lens. Panicle terminal, 6 cm long and wide, densely
puberulent, the trichomes eglandular; peduncle 60
mm long, densely and softly tomentose; bracteoles
persistent, oblong, 3.7 cm long, 8 9 mm wide,
densely puberulent; flowers persistent; calyx seg¬
ments firm, linear-lanceolate, about 3 cm long, 4
mm wide, long-acuminate, densely puberulent, the
costa and veins obscure; corolla minutely glandular
puberulent, greenish yellow, 35 mm long, the tube
slightly curved, 3 mm wide at base, 10 mm wide
near the mouth, the limb oblique, 20 mm wide, the
lobes oblong, 7 mm wide, rotundate; stamens ex¬
serted and reflexed; filaments puberulous; anthers
144
Novon
Figure 3. Scanning electron (SEM) photomicrographs of Ruellia pollen. AB. R. wurdackii (F. Guanchez 274).
— A. Equatorial view, x700. —B. Portion of surface x 2,000. C-D. R. steyermarkii (J. A. Steyermark 89113).
— C. Equatorial view, x800. —D. Portion of surface x 3,000.
oblong, 10 mm long, 2 mm wide, glabrous, basal
thecae 6 mm long. Capsule wanting.
Distribution. Known only from the type locality.
Ruellia delascioi superficially resembles R. rna-
laea Leonard from the Venezuelan Guayana. How¬
ever, in R. malaca the pale yellow corolla is 50
60 mm long, 20 mm wide near the mouth, the
stamens are exserted but not reflexed, and the leaf
blades are oblong-ovate, 26-30 cm long, 10.5-12
cm wide, and narrowed at base. In contrast, in R.
delascioi the greenish yellow corolla is 35 mm long,
10 mm wide near the mouth, the stamens are both
exserted and reflexed, and the leaf blades are ovate,
18-20 cm long, 8.5-9.3 cm wide, and obtuse at
base.
Volume 2, Number 2
1992
Wasshausen
Ruellia
145
Figure 4. A-C. Ruellia delascioi Wasshausen (Delascio A' Guanchez 10693). —A. Habit. —B. (.alyx segments.
— C. Corolla and stamens. DF. Ruellia liesneri Wasshausen (R. Liesner 2S638). —D. Habit. —E. Leafy bracts
and calyx segments. —F. Corolla and stamens.
146
Novon
Figure 5. Scanning electron (SEM) photomicrographs of Ruellia pollen. A-B. R. liesneri (/V. Cuello 400). —A.
Equatorial view, x 1,000. —B. Portion of surface x 3,000.
4. Kiiellia liesneri Wasshausen, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas: Dep-
to. Atabapo, E slope of Cerro Huachamacari,
3°49'N, 65°42'W, 600-700 m, 2 Nov. 1988,
Ronald Liesner 25638 (holotype, US; isotype,
MO). Figure 4D F.
Herba volubilis caulibus ascendentibus, subquadran-
gularibus, parce pilosis et puberulis. Folium lamina ob-
longa vel anguste ovata, acuminata, basi rotundata, inem-
branacea, atrovirenti, utrinque pilosa. Inflorescentia ex
fasciculi) terminali 2-3-floro bracteis foliaceis subtento
constans; pedicellis strobilaceis, molliter et dense pilosis;
bracteolis foliaceis, anguste lanceolatis, dense pilosis. Flos
calycis segmentis lanceolatis, pilosis et glanduloso-punc-
tatis; corolla coccinea, parce pilosa, glanduloso-punctata,
bypocrateriformi, tubo curvato et ventricoso, lobis oblon-
go-ovatis, emarginatis; staminibus exsertis. Capsulae cla-
vatae, puberulae, glanduloso-punctatae, apice subobtusae.
Clambering herbs 0.3 2 m high; stems ascending,
subquadrangular, sparingly pilose and puberulous,
the pilose trichomes erect, 1.5 mm long, puberulous
trichomes (inconspicuous) appressed. Leaves petio-
late, the petioles 5-12 mm long, pilose and puber¬
ulous, the pilose trichomes whitish, erect, 2 mm
long, the blades oblong to narrowly ovate, 4.5 10
cm long and 2.5-4 cm wide, acuminate at apex,
rounded at base, entire, undulate or shallowly cre-
nate, rich green, membranous, both surfaces pilose,
the trichomes ± erect, soft, prominent on the costa
and lateral veins (7-9 pairs), the cystoliths incon¬
spicuous under a lens. Fascicle terminal, 2-3-flow-
ered, subtended by leafy bracts 2.5 3 cm long and
0.8- 1.2 cm wide; pedicels conelike, 1-3 mm long,
softly and densely pilose; bracteoles leaflike, nar¬
rowly lanceolate, 10 mm long, densely pilose; calyx
5 9 mm long, pilose and gland-dotted, the segments
lanceolate, 4.5-8 mm long, 0.8 1 mm wide, pilose
and gland-dotted; corolla scarlet or red, sparingly
pilose, gland-dotted, 45-50 mm long, salverform,
the tube slightly curved and ventricose, 2-3 mm
broad at base, slightly narrowed above the ovary,
thence gradually enlarged to about 7 mm at mouth,
the limb 25-35 mm broad, the lobes oblong-ovate,
13 mm long and 10 mm wide near the tip, emar-
ginate; stamens exserted 8-10 mm beyond mouth
of the corolla tube; filaments sparingly and incon¬
spicuously pilose; anthers oblong, 3 mm long, 1.5
mm wide, sparingly and inconspicuously pilose; pol¬
len spheroidal, 65 /xm diam., the reticulum hom-
obrochate, the membrane mostly smooth (Fig. 5A,
B); the pislil exceeding the stamens, glabrous. Cap-
Volume 2, Number 2
1992
Wasshausen
Ruellia
147
Figure 6. A-F. Ruellia bolivarensis Wasshausen (F. Cardona 49 ?). —A. Habit. —B. Enlargement of lower leaf
surface. —C. Bracts, bracteoles, and calyx segments. —D. Calyx s jgments. —E. Expanded corolla, stamens, and
pistil. —F. Bract, bracteoles, calyx segments, and capsule.
sule clavate, puberulous, gland-dotted, 15 mm long,
5 mm broad, rather bluntly pointed, narrowed to a
slender, solid base about 3 mm long; seeds brown,
appressed-puberulous, about 5 mm long and 4 mm
wide.
Distribution. Open and forested areas around
waterfall and on forested areas and igneous outcrops
at elevations between 80 and 750 m.
Parat rpes. VENEZUELA. TERRITORR) FEDERAL AMAZONAS:
Depto. ktabapo, slope of Huachamacari, 3°39 / N,
65°42'V , 750 m, Ronald L. Liesner 18378 (MO, US);
Depto. t tures, 35 km SE of Puerto Ayacucho, along Rio
Coromot), at Tobogan de la Selva, 5°22'N, 67°33'W,
100 m,. teyermark , Davidse & Guanchez 122508 (MO,
NY, USg forest 1 km below Tobogan de la Selva, 80 m,
Steyermark, Holst & Manara 131550 (MO, US); Ser-
rania de la Coromoto, sector “El Tobogan,” 37 km S of
Puerto Ayacucho, 5°24'N, 67°35'W, 80 200 m, \ idia
Cuello 400 (US).
148
Novon
Ruellia liesneri is related to the widespread Ven¬
ezuelan and Colombian species R. humboldtiana
(Nees) Lindau. In R. humboldtiana the inflores¬
cence is a thyrsoid panicle, with the branches each
bearing from several to 20 or more flowers, these
not subtended by any conspicuous, leafy bracts, the
calyx 9-12 mm long, and with petioles 10-1 5 mm
long, puberulous. In contrast, the inflorescence of
R. liesneri is a 2-3-flowered fascicle, subtended by
conspicuous, large, leafy bracts, the calyx 5-9 mm
long, and the petioles are 5-12 mm long and both
pilose and puberulous.
5. Ituellin bolivarensis Wasshausen, sp. nov.
TYPE: Venezuela. Bolivar: Sitio Parupa, rio
Paragua, Apr. 1943, Felix Cardona 492 (ho-
lotype, US). Figure 6A-F.
Herba vel suffrutex caulibus quadrangularibus, sulcatis,
parce pilosis. Folium lamina oblonga vel ovata, graciliter
acuminata, basi rotundata, firma, praeter costain et venas
pilosas glabra. Inflorescentiae ex spicis terminalibus laxis,
flores in paribus oppositis gerentibus constantes, inter-
nodiis glabris vel parce pubescentibus; bracteis foliaceis
quam foliis multo minoribus, oblongis; bracteolis anguste
triangularibus. Flos calycis segmentis subulatis, pilosis;
corolla anguste infundibuliformi, tubo curvato, lobis su-
borbicularibus, ad apicem irregulariter lobulatis; stami-
nibus inclusis, antheris mucronulatis, glabris. Capsulae
clavatae, puberulae, ad apicem acutae.
Plants herbaceous or suffrutescent; stems qua¬
drangular, shallowly sulcate, the angles narrowly
winged, the wings pilose, the trichomes barely 1 mm
long, the cystoliths minute. Leaves petiolate, the
petioles 5-15 mm long, minutely pilose, the blades
oblong to ovate, 9-14 cm long, 3.3 6.4 cm wide,
rather slenderly acuminate, distinctly rounded at
base, firm, shallowly crenate, glabrous except the
costa and lateral veins (5 or 6 pairs), these pilose,
the trichomes about 0.5 mm long, spreading or
subappressed, the cystoliths conspicuous under a
lens on the upper surface. Flowers borne opposite
in pairs on lax, nearly straight, rigid, ascending
axillary and terminal spikes 2.5-8.5 cm long, the
lowermost internodes 3 cm long, the others succes¬
sively shorter, somewhat flattened near the nodes,
glabrous or sparingly pubescent, the trichomes as¬
cending to subappressed, about 0.2 mm long; bracts
leaflike but much smaller, oblong, narrowed at both
ends, 1 1 mm long and 2 mm wide; bracteoles nar¬
rowly triangular, 1 mm long, sparingly pilose; calyx
6-7 mm long, the segments subulate, 4-5 mm long,
0.75-1 mm wide at base, pilose, consisting of both
glandular and eglandular trichomes, these erect or
spreading; corolla 12-14 mm long, inconspicuously
pilose, narrowly infundibular, the narrow portion of
the tube slightly curved, 6-7 mm long, 1 mm broad
at base, the limb about 4.5 mm wide, the lobes
suborbicular, about 2 mm diam., irregularly lobulate
at apex; stamens barely reaching the base of the
corolla lobes; anther oblong, mucronulate, the upper
pair 1.5 mm long, 0.5 mm wide, glabrous, the lower
pair 1 mm long, 0.3 mm wide. Capsule slenderly
clavate, about 13 mm long, puberulous, pointed at
apex, the slender sterile basal portion about 6 mm
long and 1 mm broad at base, enlarging to 1.25
mm at base of seed-bearing cavity, this flattened
and about 2 mm broad; retinacula slender, curved,
about 1.75 mm long, the slender tip ± erose.
Distribution. Known only from the type locality.
Ruellia bolivarensis superficially resembles R.
menthoides (Nees) Hiern from Amazonian Vene¬
zuela and Brazil. However, in R. menthoides (Fig.
1A-D) the flowers are borne axillary, sessile, clus¬
tered, and verticillate; the corollas are 25 mm long,
and the upper leaf blades are narrowed and acute
at base. In contrast, in R. bolivarensis the flowers
are borne opposite in pairs on lax, axillary and
terminal spikes; the corollas are 12-14 mm long,
and the upper leaf blades are distinctly rounded at
base. This new taxon is unique from the other known
species of neotropical Ruellia. It possesses mu¬
cronulate anther lohes, which thus far have not been
reported for this genus in the literature, and a rather
long, pointed capsule. Unfortunately, the single Car¬
dona 492 collection is somewhat scanty; additional
specimens should be collected for a proper diagnosis
of these character states.
Acknowledgments. My special thanks to Alice
Tangerini, who skillfully prepared the line drawings,
and to Stephen F. Smith and the staff of the National
Museum of Natural History SEM Laboratory for
their high-quality pollen photomicrographs.
Literature Cited
Long, R. W. 1970. The genera of Acanthaceae in the
southeastern United States. J. Arnold Arbor. 51:
257-309.
Nomenclatural Changes in the Acanthaceae and Two New Species of
Odontonema from Venezuela and Guyana
Dieter C. Wasshausen
Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, U.S.A.
ABSTRACT. The recent completion of the Acan¬
thaceae treatment for the Flora of the l enezuelan
Guayana has necessitated a new name in the genus
Staurogyne , a new combination in Anisacanthus,
and the recognition of two undescribed species of
Odontonema , based on relatively recent collections.
Staurogyne spraguei Wasshausen, norm nov. Re¬
placed name: Staurogyne leptocaulis Leonard,
Contr. U.S. Natl. Herb. 31: 671. 1958. not
S. leptocaulis Bremek. 1957. TYPE: Colom¬
bia. Meta: T. A. Sprague 27 (holotype, K).
Anisacanthus boliviensis (Nees) Wasshausen,
comb. nov. Basionym: Drejera boliviensis Nees
in DC., Prodr. 1 1: 334. 1847. TYPE: Bolivia.
Orbigny 786 (lectotype, designated here, G;
isolectotype, GZU). [The syntypes, Otto 1006
(Venezuela) and Hostmann 1123 (Surinam),
are Anisacanthus secundus Leonard.]
Schaueria caduciflora Griseb. in Symb. FI. Argent. 261.
1879. A. caduciflorus (Griseb.) Ariza in Bob Soc.
Argent. Bot. 22(1/4): 255. 1983. TYPE: Argen¬
tina. Gran Chaco: Laguna del Palmar, Lorentz &
Hieronymus 554 (holotype, GOET).
Jacobinia caducifolia Griseb. in Symb. FI. Argent. 261.
1879. Anisacanthus caducifolius (Griseb.) Lindau
in Bot. Jahr. 19, Beibl. 48: 18. 1894. TYPE: Ar¬
gentina. Oran, Tabacal, Lorentz & Hieronymus 926
(holotype, GOET).
Odontonema liesneri Wasshausen, sp. nov.
TYPE: Venezuela. Territorio Federal Amazo¬
nas: Depto. Rio Negro, Cerro de La Neblina
Camp V, valley N base of Pico Cardona, 0°49'N,
66°0'W, 1,250 m, 21-24 Mar. 1984, Ronald
Liesner & Brian Stannard 16610 (holotype,
US; isotype, MO). Figure 1A-F.
Frutex caulibus juvenilibus subquadrangularibus, gla-
bris. Folium lamina oblonga vel elliptica, acuta vel acu¬
minata, basi angustata et in petiolum decurrenti, mem-
branacea, utrinque minute et parce puberula; petiolo parce
puberulo. Racemi terminales, graciles, rhachi sicut ped-
icello puberula; bracteis lineari-lanceolatis; bracteolis an-
guste triangulatis, minute puberulis. Flos calycis segmentis
anguste triangularibus; corolla rubra, glabra vel minute
glanduloso-pubescenti, labiis subaequalibus, superiore bi-
lobo lobis ellipticis vel late oblongis, inferiore prope basim
trilobo lobis loborum superiorum similibus; staminibus vix
exsertis.
Plants fruticose, 1-1.5 m tall, the stems terete
m older portions to subquadrangular in younger
portions, glabrate. Leaves petiolate, the petioles 10-
20 mm long, minutely and inconspicuously puber-
ulous with appressed trichomes, the blades oblong-
elliptic, 16-30 x 6.5-11 cm, acute to acuminate
at apex, narrowed at base and decurrent on the
petiole, thin, membranous, minutely and inconspic¬
uously puberulous on both surfaces especially along
the midrib and lateral veins of the upper leaves, the
margins entire to crenate, the cystoliths minute and
scattered on the upper surface, visible only under
a lens; inflorescence a loose, slender, terminal ra¬
ceme 10-22 cm long with 1 or 2 flowers per node,
the rachis puberulous, the pedicels 2-4 mm long,
minutely puberulous; the bracts subtending the ra¬
chis linear-lanceolate, the lowermost pair 5.5-9 x
0.8 mm, the others successively reduced upwardly,
the bracts subtending the pedicels narrowly trian¬
gular, 2 x 0.6 mm; bracteoles 2 per flower, nar¬
rowly triangular, 1 x 0.5 mm, all minutely puber¬
ulous; calyx segments narrowly triangular, 2 x 0.8
mm, acute, sparingly to moderately puberulous; co¬
rolla red, narrowly urceolate, 15-25 mm long, 1-
2 mm wide at base, 3-4 mm wide at throat, glabrous
to inconspicuously glandular-puberulent without, pu-
berulent within, the lips similar, the upper lip with
elliptic to broadly oblong lobes, 3x2 mm, the lower
lip with similar lobes 2.5-3 x 2 mm; stamens ex¬
tended to the edge of the corolla lips, the filaments
glabrous, 12 mm long (heterostyly is known to occur
in many species of Odontonema; however, only one
morph is currently known from this taxon), the
anthers 3.5 mm long; staminodes 1 mm long, gla¬
brous; pistil exserted, the stigma minutely hilobed,
the style 20 mm long, glabrous; ovary glabrous.
Capsule not seen.
Distribution. Noted as locally common, but known
only from the type locality, Cerro Neblina in south¬
ern Venezuela.
Novon 2: 149-152. 1992.
150
Novon
Figure 1. A-F. Odontonema liesneri (Liesner & Stannard 16910). —A. Habit. — B. Bract, bracteole, pedicel,
and calyx. C. Calyx and juvenile corolla. —D. Corolla. —E. Corolla tube and stamens. —F. Bract, bracteoles,
pedicel, calyx, and juvenile corolla.
Odontonema liesneri resembles O. fuchsioides
(Nees) Kuntze from the coastal regions of northern
Colombia. However, in O. fuchsioides the leaf blades
are rather firm, 9-12.5 X 2-6 cm, the corollas are
25-30 x 5-6 mm at the throat, and the calyx
segments are 5x1 rnm. In contrast, (). liesneri
has leaf blades that are thin and 16-30 X 6.5 -1 1
cm, the corollas are 15-25 x 3-4 mm at the throat,
and the calyx segments are 2 X 0.8 mm.
Odontonema mazarunensis Wasshausen, sp.
nov. TYPE: Guyana. Upper Mazaruni River
region, Karowtipu Mountains, 5°45'N,
60°35'W, ca. 460 m, 26 Apr. 1987, Brian
Volume 2, Number 2
1992
Wasshausen 151
Changes and New Species in Acanthaceae
Figure 2. A-D. Odontonema mazarunensis (Boom & Gopaul 7748). —A. Habit. —B. Bract subtending the
rachis. —C. Bracteoles, pedicel, and calyx. —D. Corolla, stamens, and pistil.
152
Novon
Boom & Doorjoohan Gopaul 7748 (holotype,
NY). Figure 2A-I).
Suffrutex caulibus juvenilibus subquadrangularibus,
glabris. Folium lamina elliptica vel lanceolata, acuminata,
basi angustata, firma, utrinque glabra vel in costa et venis
lateralibus parce hirtella; petiolo brevi, glabro. Racemi
terminales, breves, rhachi sicut pedicello rubra, puberula;
bracteis rubris, lineari-lanceolatis, pilosis; bracteolis sub-
ulatis, minute pilosis. Flos calycis segmentis rubris, an-
guste lanceolatis, glanduloso-puberulis; corolla rubra, tub-
ulosa, glanduloso-pubescenti, labiis subaequalibus, superiore
bilobo, lobis ellipticis vel ovatis, inferiore prope basim
trilobo lobis loborum superiorum simiiibus; staminibus
exsertis, filarnentis subroseis, antheris rubris.
Shrub 1 m tall, the stems terete in older portions
to subquadrangular in younger portions, glabrate.
Leaves short-petiolate, the petiole (unwinged por¬
tion) 5 mm long, glabrous, the blades elliptic-lan¬
ceolate, 23-24 x 6.2-8.2 cm, acuminate at apex,
narrowed at base, entire or slightly crenulate, firm,
glabrous above, glabrous below except for the costa
and lateral veins (6-8 pairs), these sparingly hir-
tellous, the trichomes appressed. Inflorescence a
short terminal raceme 3 cm long with 2 flowers per
node, the rachis red, puberulous, the pedicel red,
4.5-6 mm long, glandular-puberulous; bracts sub¬
tending the rachis red, linear-lanceolate, the low¬
ermost pair 14 X 1 mm, pilose, the bracts sub¬
tending the pedicel red, linear, 12 x 0.5 mm,
glandular-puberulous; bracteoles subulate, 3 x 0.3
mm, minutely and inconspicuously pilose; calyx seg¬
ments reddish, narrowly lanceolate, 6.5-7.5 x 1
mm, glandular-puberulous; corolla red, glandular-
puberulent without, glabrous within, tubular, 40-50
mm long, the tube 3 mm wide at the base, 6.5-7
mm wide at the throat, the upper lip with elliptic-
ovate lobes 5x3 mm, the lower lip with elliptic-
ovate lobes 6x4 mm; stamens exceeding the
corolla lobes by 5 mm, the filaments pinkish, 27
mm long, pubescent basally, tbe anthers red, 2.5
mm long; staminodes 2 mm long, retrorsely puber¬
ulous, apically capitate with a rudimentary anther;
pistil exserted, the stigma minutely bilobed, glabrous,
the style 45 mm long (heterostyly is known to occur
in many species of Odontonema; however, only one
morph is currently known from this taxon), sparsely
pilose; ovary glabrous. Capsule not seen.
Distribution. Humid forest at the base of moun¬
tain. Known only from the Karowtipu Mountains,
Guyana.
Odontonema mazarunensis is perhaps most
closely related to the lowland Venezuelan species O.
album V. M. Baum, which differs markedly by hav¬
ing a white corolla to 30 mm long, bracts subtending
the pedicels lanceolate, 1-1.5 mm long, and the
stamens extended to the edge of the corolla lobes.
In contrast, 0. mazarunensis has a red corolla 40-
50 mm long, bracts subtending the pedicels red,
linear, 12 mm long, and the stamens exceeding the
corolla lobes by 5 mm.
Acknowledgment. My special thanks to Alice
Tangerini for the skillfully prepared line drawings.
A New Species of Aptandra (Olacaceae) from Amazonian Peru
Alwyn //. Gentry and Rosa Ortiz
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, LJ.S.A.
ABSTRACT. A new species of Olacaceae from Am¬
azonian Peru is described as Aptandra caudata.
Aptandra caudata A. Gentry & Ortiz, sp. nov.
TYPE: Peru. Loreto: Provincia Maynas, Que-
brada Sucusari, N bank of Rio Napo below
Mazan, mature upland forest on lateritic soil,
elev. 130 m, 3°15'S, 72°55'W, 7 July 1983
(fl), Gentry , Vasquez & Jaramillo 42718 (ho-
lotype, MO; isotypes, AMAZ, USM). Figure 1.
Arbor parva, ramulis teretibus, glabris. Folia anguste
elliptica vel elliptico-oblonga, apice abrupte per 1.5-3.5
cm caudata, glabra, membranacea. Inflorescentia pauci-
fiora, pedicellis filiformibus, 1-1.5 mm longis. Flores in
alabastro cylindrici, ca. 5 mm longi; calyce obtuse 4-
lobulatos, 0.5 mm longo. Fructus globosus, ca. 2.5 cm
diametro, calyce subpatelliformi expanso subtensus.
Treelet 3-5 m tall; branchlets slender, subterete,
smooth, glabrous. Leaves narrowly elliptic or elliptic-
oblong, the apex abruptly caudate-acuminate for
1.5-3.5 cm, the base ± cuneate, membranaceous,
olive green when dry, glabrous, 3.5-15 cm long
(excluding drip tip), 1.2-4.5 cm wide, the lateral
nerves 6-14 pairs; petiole slender, 4-8 mm long,
ca. 1 mm thick. Inflorescence an open axillary pan¬
icle, few-flowered, the branches slender, the pedun¬
cle ca. 1 cm long, the flowers arranged in 2-3-
flowered umbels at ends of the inflorescence branch¬
es, the filiform pedicels ca. 1-1.5 cm long. Flower
bud a cylindrical tube with globose apex; calyx shal¬
lowly and obtusely 4-lobed, ca. 5 mm long at an-
thesis, much enlarged in fruit; petals 4, linear below,
somewhat broadened at apex, greenish cream, ca.
5 X 1 mm, forming a strictly cylindrical tube with
a globose apex; stamens 4, the filaments connate
into 4-mm-long tube, the anthers ca. 0.5 mm long,
fused to an annulus, opening by a thin reflexing
valve; pistil ca. 4 mm long, the slender style apicallv
thickened at level of anthers. Fruit globose, single-
seeded, green, ca. 2.5 cm diam., the fruiting calyx
coriaceous, becoming broadly cup-shaped to almost
patelliform, enveloping less than the basal quarter
of drupe; seed globose, ca. 7 mm across.
Distribution. Known only from the type locality
at the Explornapo Camp on the Rio Sucusari in
Loreto Department, Peru, 116-130 m elevation.
Additional collections examined. PERU. LORETO: (all
from type locality), 5 July 1983 (fl), Gentry et al. 42628
(AMAZ, MO, USM), 14 June 1986 (st), Gentry et al.
54321 (AMAZ, MO), 7 Oct. 1986 (st), Vasquez & Jar¬
amillo 8156 (AMAZ, MO), 7 Nov. 1989 (fr), Vasquez
& Jaramillo 13103 (AMAZ, MO, USM).
Only three species of Aptandra were recognized
by Sleumer (1984), two in the Neotropics and one
in tropical Africa. Of these, A. caudata is closest
to A. tubicina (Poeppig) Benth. ex Miers, which
also occurs in Amazonian Peru, on account of the
membranaceous, cuneate leaves and strictly cylin-
dric pre-anthesis corolla tube. 1 he most striking
differentiating feature of A. caudata is the con¬
spicuously caudate leaf apex, especially pronounced
in small leaves where a 3-cm-long leaf can have a
1.5-cm-long drip tip. In contrast, A. tubicina is
acuminate with the drip tip 0.5 1.5 cm long. Ihe
leaves of A. caudata are also narrower and more
elliptic than in A. tubicina ; although the latter some¬
times has some leaves as narrow as those of A.
caudata , the narrower leaves are mostly lanceolate.
The flower of A. caudata is also noticeably longer
(5 mm vs. 3 mm) than that of A. tubicina and has
a more prominently globose bud apex and more
shallowly and obtusely lobed calyx. Another differ¬
ence is the almost patelliform fruiting calyx of A.
caudata , which covers only the extreme base of the
drupe and is very different from the broadly cupular
calyx of A. tubicina, which covers most of the lower
half of the drupe.
It is possible that two species are included m
Sleumer’s (1984) concept of A. tubicina: one with
membranaceous to chartaceous, strongly acuminate
leaves, which includes the type; the other with more
subcoriaceous, subacuminate leaves and smaller,
more numerous flowers, which is much more com¬
mon, at least in Amazonian Peru. Aptandra cau¬
data is vegetatively more similar to the Brazilian
type material of A. tubicina than to the rest of the
Peruvian material. However, A. caudata cannot be
conspecific with either variant of A. tubicina be¬
cause of the differences in size and shape of flowers
and buds and the patelliform fruiting calyx.
Aptandra caudata is the largest-flowered species
in the genus and approaches Chaunochiton in flower
size and in the almost patelliformly expanded fruiting
Novon 2: 153-154. 1992.
154
Novon
Figure 1. Aptandra caudata A. Gentry & Ortiz. —A. Habit (Gentry et al. 42718). — B. Variation in leaf size
and shape (Vasquez & Jaramillo 8156). —C. Fruit ( Vasquez & Jaramillo 13103). Scale bar = 1 cm.
calyx. It is locally one of the most common under¬
story species at the type locality, with seven treelets
>2.5 cm DBH in a 0.1 ha. sample.
Literature Cited
Sleuiner, H. O. 1984. Olacaceae. FI. Neotrop. Monogr.
38: 1-139.
Four New Species of Meliosma (Sabiaceae) from Peru
Alwyti //. Gentry
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
ABSTRACT. Four new species of Meliosma are de¬
scribed from the Peruvian Andes as M. pumila, M.
simiarum, M. sirensis , and M. youngii.
Meliosma is emblematic of Peruvian cloud forest
plants in its tendency to local endemism and in the
incompleteness of taxonomic knowledge about it.
Not a single species of Sabiaceae was known from
Peru when Macbride (1956) treated related families
in the Flora of Peru, but at least 17 Meliosma
species (and three of lowland Ophiocaryon) are now
known from the country (Gentry, 1986, in press).
Four of the most distinctive of these are described
here, all from the Peruvian Andes. These four spe¬
cies are all shrubs or small trees or treelets and are
the smallest in stature of any neotropical Meliosma.
Three of them are also among the smallest-leaved
species of the genus; the fourth is a large-leaved
pachycaul treelet.
Meliosma pumila A. Gentry, sp. nov. TYPE:
Peru. Amazonas: Bongara Prov., Shillac, N of
Pedro Ruiz, 5°49'S, 78°01'W, 2,300 m, moist
primary montane rainforest, 31 Aug.-2 Sep.
1983 (fr), /). Smith & S. Vasquez 4894 (ho-
lotype, MO; isotype, USM).
Frutex 0.5-2 m altus. Folia oblanceolata, 4-17 cm
longa, 1-5 cm lata, acute serrata, plus minusve glabra.
Inflorescentia terminalis, sparsim puberula, pedicellis 0-
0.5 mm longis, sepalis 0.3-0.4 mm longis, petalis 0.7-
1 mm longis. Fructus globosus, 1-1.3 cm diametro.
Shrub or treelet 0.5-2 m tall; branchlets terete,
glabrous or sparsely and glabrescently pubescent
with appressed trichomes when young, usually with
a few small, inconspicuous lenticels. Leaves alter¬
nate, oblanceolate, 4-17 cm long, 1-5 cm wide,
acuminate, cuneate at base, chartaceous, sharply
serrate, the midvein strongly impressed above, raised
below, secondary veins plane and inconspicuous
above, raised below, each terminating in a conspic¬
uous tooth, the tertiary venation inconspicuous and
plane above, plane to subprominulous below, gla¬
brous above, below glabrous except for a few in¬
conspicuous appressed trichomes along main veins
and sometimes very sparsely on surface; petiole 0.5-
2.5 cm long. Inflorescence terminal, subpaniculate
with a well-developed slender central axis and 2 4
widely separated lateral branches, sparsely puber-
ulous, the pedicels lacking to 0.5 mm long. Flowers
(seen only in bud) greenish, the sepals 5, broadly
ovate, rounded, 0.3-0.4 mm long, three slightly
larger than the other two, the margin minutely fim¬
briate; outer petals broadly ovate, ca. 1 mm long,
inner petals narrow, ca. 0.7 mm long and 0.2 mm
wide; fertile stamens 2, the anthers thick and su-
borbicular, each ca. 3 mm long, only slightly sep¬
arated by the thickened connective, including con¬
nective ca. 0.6 mm across, the slender filament ca.
0.5 mm long, completely fused to the narrow inner
petal; ovary ovoid, ca. 0.5 mm long. Fruit cream-
colored when fresh, asymmetrically globose, ca. 1-
1.3 cm diam., slightly contracted into a broad and
poorly demarcated basal stipe, the surface glabrous.
Distribution. Endemic to mid-elevation cloud for¬
ests of northern Peru, south of the Huancabamba
Depression, 1,850-2,300 m elevation.
Additional specimens examined. Peru. AMAZONAS:
Prov. Bongara, Dist. Sipabamba, Shilla, montane forest,
1,850-1,900 m, 6 May 1981 (fr). Young & Eisenberg
378 (MO). CAJAMARCA: Prov. Cutervo, Distr. San Andres
de Cutervo, carretera entre San Andres y Santo Tomas,
km 12, 2,320 m, 7 Aug. 1987 (fl), Diaz & Osores 2536
(MO); lower edge of Cutervo National Park, 10-15 km
N of San Andres de Cutervo, montane cloud forest, 6°10'S,
78°40'W, 2,200-2,250 in, 11 Feb. 1988 (st). Gentry
et al. 61511 (MO, USM).
Meliosma pumila is the smallest species in the
genus and can reportedly be reproductive as a 0.5-
m-tall subshrub. It differs from M. solomonii A.
Gentry, which has similarly small leaves, in the
chartaceous leaf texture and consistently and sharp¬
ly serrate leaf margin. It differs from M. simiarum ,
which also has small chartaceous leaves, in the cream-
colored (rather than black) fruits at maturity, in the
narrower leaves with conspicuously spinose serrate
margins, and in the twigs and leaves either glabrous
or with small appressed trichomes.
Meliosma simiarum A. Gentry, sp. nov. TYPE:
Peru. San Martin: Venceremos near Amazonas
border, km 291 on Rioja-Pomacocha road,
5°45'S, 77°40' W, 1.850 m, wet lower montane
forest, 9 Feb. 1984 (fl, fr). Gentry & Smith
Novon 2: 155-158. 1992.
156
Novon
Figure 1. Meliosma simiarum. —A. Habit showing in¬
florescence, leaves, and fruit (Gentry & Smith 45225).
B. Close-up of inflorescence and fruit (Gentry & Smith
45225). —C. Close-up of flowering inflorescence (Gentry
<£- Smith 45335).
45225 (holotype, MO 3517422; isotypes, MO
3517423, USM). Figure 1.
Arbor parva 6-10 rn alta. Folia anguste ovata vel
elliptica vel oblanceolata, 5-14 cm longa, 1.5-4.5 cm
lata, puberula pro parte maxima infra in costa. Inflores-
centia pauci-ramosa; plerumque axillaris, puberula, ped-
icellis ca. 1 mm longis, sepalis ca. 1 mm longis, petalis
ca. 1 mm longis. Fructus obpyriformis, 1.4 1.7 cm lon-
gus, 1 1.5 cm diametro.
Small tree 6-10 rn tall; branchlets terete, with
laxly spreading reddish trichomes, lenticels large but
very few and scattered. Leaves alternate, narrowly
ovate to narrowly elliptic or oblanceolate, 5-14 cm
long, 1.5-4.5 cm wide (-22 x 6 cm in juveniles),
acuminate, cuneate to obtuse at base, chartaceous,
entire or with 2-3 shallow teeth near apex at ma¬
turity, rather remotely and coarsely serrate in ju¬
veniles, the midvein impressed above, secondary
veins plane above, raised below, tertiary veins plane
above, finely prominulous below, puberulous with
suberect trichomes along main vein below, also
sparsely on secondary veins below and even more
sparsely with widely scattered trichomes on surface
above and below; petiole 0.3-0.7(-l) cm long, pu¬
bescent with suberect flexuous trichomes. Inflores¬
cence a few-branched panicle, mostly axillary or
borne below the leaves, with a straight rachis and
5-10 widely separated short lateral branches, pu¬
bescent with 2 kinds of trichomes, some longer and
suberect, others short and gland-tipped (or the gland
sessile with the stalk obsolete), the pedicels ca. 1
mm long. Flowers white, the sepals 5, broadly ovate,
ca. 1 mm long, the margin erose-fimbriate, otherwise
glabrous; outer petals broadly ovate, ca. 1 mm long,
inner petals thinner, obtriangular, 0.5 mm long, 0.5
mm broad at apex, the filament ca. 0.8 mm long,
the lower half fused to inner petal, the anthers
subglobose, 0.3 mm long, widely separated by the
thickened connective, sometimes only 1 developed;
pistil 0.5 mm long. Fruit obpyriform 1.4-1.7 cm
long, 1-1.5 cm diam., turning black at maturity,
the poorly demarcated thick basal stipe having an
asymmetrically squarish corner at base, glabrous,
drying black.
Distribution. Endemic to lower elevation cloud
forests of San Martin Department, 1,850 2,040 m
elevation.
Additional specimens examined. Peru, san martIn:
type locality, 10 Feb. 1984 (st), Gentry et al. 45318
(MO, USM), 10 Feb. 1984 (fl), Gentry et al. 45335
(MO, USM), 12 Feb. 1984 (fl, fr), Gentry et al. 45497
(MO, USM); Rioja Prov., Pedro Ruiz Moyobamba road,
km 390 Venceremos, 2-3 Aug. 1983 (fr), D. Smith <£-
S. Vasquez 4573 (MO, USM); Province of Rioja, sin.
loc., M. Leo 110 (MO).
This species is most similar to M. pumila, the
only other species of Peruvian Meliosma with small
noncoriaceous leaves, but differs in the more con¬
spicuous vegetative pubescence of suberect flexuous
trichomes, the entire or subentire mature leaves,
the black (rather than cream-colored) fresh fruits,
and the apically much broader inner petal and broad¬
er connective. Although the two species grow in the
same general area, M. pumila occurs in the cloud
forests of the Marar'ion Valley, while M. simiarum
occurs on the Amazon-facing slopes.
Volume 2, Number 2
1992
Gentry
Meliosma from Peru
157
Meliosma simiarum is named for its association
with the nearly extinct yellow-tailed woolly monkey,
Lagothrix flavicaudus, which shares the same cloud
forest habitat. It is known only from the Venceremos
forest, one of the last habitats of that threatened
primate, where it is one of the most common un¬
derstory species, averaging 40 plants >2.5 cm DBH/
ha.
Meliosma sirensis A. Gentry, sp. nov. TYPE:
Peru. Huanuco: Prov. Pachitea, Serrania de
Siri, 26-28 km ESE of Puerto Inca, 1,750 m,
elfin forest, 9°25'S, 74°43'W, 23 July 1988
(fl), B. Wallnoffer 11-23788 (holotype, MO;
isotypes, USM, W).
Arbor parva 7 m alta. Folia anguste elliptica, 5-9.5
cm longa, 1.1-2.5 cm lata, plus minusve glabra. Inflo-
rescentia axillaris vel subterminalis, glabra, pedicellis 2-
3 mm longis, sepalis ca. 1 mm longis, petalis exterioribus
2 mm longis. Fructus ignotus.
Small tree 7 m tall and 4 cm DBH; branchlets
terete, glabrous or with very few scattered tri-
chomes, the few lenticels large and tannish. Leaves
alternate, narrowly elliptic, 5 9.5 cm long, 1.1-2.5
cm wide, gradually sharply acuminate, narrowly cu-
neate at base, coriaceous, entire, the midvein im¬
pressed above, strongly raised below, the secondary
veins 6-8 on a side, brochidodromous, making near¬
ly a 90° angle with midvein, plane and inconspicuous
above, prominulous below, the tertiary venation fine¬
ly and intricately prominulous-reticulate above and
below, glabrous except for very few small appressed
trichomes widely scattered along midvein below, both
surfaces minutely lepidote-glandular, drying dark
brown above, gray-brown below with contrasting
brown midrib and whitish finer venation; petiole 0.6-
1.4 cm long, glabrous. Inflorescence axillary or leaf-
opposed to subterminal, paniculate with a well-de¬
veloped central rachis, glabrous, the pedicels 2-3
mm long. Flowers white, the sepals broadly and
obtusely ovate, ca. 1 mm long, with a few short,
gland-tipped trichomes along margin, otherwise gla¬
brous; outer petals broadly ovate, 2 mm long, inner
petals rectangular, 1 mm long, 0.5 mm wide, not
bifid, fused only at base with the filament of fertile
stamen; fertile stamens 2, the slender filament 1.5
mm long, apically thickened into connective, the
anthers small, oblong, each ca. 0.3 mm long, held
horizontally at ends of the 0.5-mm-thick connective;
ovary ovoid, 1 mm long and wide, glabrous, tapering
to a short narrow style. Fruit unknown.
Distribution. Known only from the type locality
in elfin forest along the crest of the isolated Serrania
de Siri.
This species is closest to M. solomonii , the only
other Andean Meliosma with small, narrow, coria¬
ceous leaves. Meliosma solomonii has mostly larger
leaves with more numerous, more strongly ascending
secondary veins, a larger inflorescence with sessile
or subsessile flowers and scattered appressed tri¬
chomes, and larger sepals (ca. 2 mm long). That
species is known only from higher altitudes (2,450
2,850 m) in the main Andean Cordillera.
Meliosma youngii A. Gentry, sp. nov. TYPE:
Peru. San Martin: Prov. Mariscal Caceres, Rio
Abiseo National Park, 7°S, 77°W, Las Palmas,
2,350 m, 17 Aug. 1986 (fr). Young 4070
(holotype, MO; isotype, USM not seen).
Arbor parva 3 m alta. Folia anguste obovata, ca. 21-
50 cm longa, ca. 8-16 cm lata, adpresso-puberula pler-
umque in costis, petiolis ca. 7 cm longis. Inflorescentia
folio opposita, pyramido-paniculata, puberula. Fructus
globosus, 1.4-1.7 cm diarnetro.
Small pachycaul tree 3 m tall, branches thick
and somewhat angled, glabrous or glabrate, the hark
rather wrinkled, grayish, with contrasting reddish
brown raised lenticels. Leaves (only 2 seen, both
broken) clustered near apex (fide K. Young), alter¬
nate, narrowly obovate, ca. 21-50 cm long, ca. 8-
16 cm wide, apex not seen, base obtusely asym¬
metrically cuneate, membranaceous to chartaceous,
apparently entire (badly eaten), the midvein im¬
pressed above, secondary veins plane above, prom¬
inent below, tertiary venation plane and not apparent
above, prominulous below, appressed-puberulous
along main veins above and below and sparsely over
surface below; petiole ca. 7 cm long, tannish from
the dense parallel-appressed short trichomes. Inflo¬
rescence (seen only in fruit) leaf-opposed, pyramidal-
paniculate with a well-developed central rachis, ca.
14-24 cm long, puberulous. Flowers not seen. Fruits
brownish when fresh, drying black, globose, essen¬
tially estipitate, 1.4-1.7 cm diam.
Distribution. Known only from the type specimen
from montane rainforest in Rio Abiseo National Park
at 2,350 m.
Although the type collection has only two leaves,
which are not in very good condition, there is no
doubt that this represents an undescribed species.
The species is characterized by the pachycaul treelet
growth form and by the large thin-textured leaves
(larger than any other high-altitude Meliosma) with
conspicuously tannish, almost sericeous, petioles and
midveins, and by the very smooth upper leaf surface.
Acknowledgments. Support for this study of Pe¬
ruvian plants has been provided by a grant from
158
Novon
the Mellon Foundation and by a lellowship from the
Few Scholars program.
Literature Cited
Gentry, A. H. 1986. New neotropical species of Me-
liosma (Sabiaceae). Ann. Missouri Bot. Card. 73:
820-824.
-. Diversity and floristic composition of Andean
cloud forests of Peru and adjacent countries: impli¬
cations for their conservation. In K. Young & N.
Valencia (editors), Biogeografia, Ecologia y Conser-
vacion del Bosque Montano en el Peru. Mem. Mus.
Hist. Nat. Univ. San Marcos 21 (in press).
Macbride, J. F. 1956. Flora of Peru. Publ. Field Mus.
Nat. Hist., Bot. Ser. 13(3A): 291-744.
Six New Species of Bignoniaceae from Upper Amazonia
Alwyn H. Gentry
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
ABSTRACT. Six new species of Bignoniaceae from
upper Amazonia are described as Arrabidaea ([(fin¬
is, Cuspidaria emmonsii, Distictis occidentalis,
Haplolophium nunezii, Schlegelia cauliflora, and
Schlegelin hirsuta. In addition, merger of Urban-
olophium with Haplolophium is proposed, neces¬
sitating the new combination H. glaziovii.
Names are needed for these six undescribed spe¬
cies of Bignoniaceae lianas from upper Amazonia
for the familial treatment in Flora de Colombia
and/or the Catalogue of the Flowering Plants and
Gymnosperrns of Peru. Four of the six are wide¬
spread, and one is documented to have ethnobotan-
ical uses. In addition, discovery of Haplolophium
nunezii makes unavoidable the reduction of l r-
banolophium to Haplolophium , necessitating a new
combination for its only species.
Arrabidaea affinis A. Gentry, sp. nov. 1 4 PE:
Ecuador. Napo: Lago Agrio, forest on banks ot
lake, 250 m, 0°7'N, 76°55'W, 1 Apr. 1980
(fl), J. Rrandbyge & E. Asanza 30393 (ho-
lotype, MO; isotype, AAU).
Frutex scandens, ramulis teretibus, minute puberulis,
consociebus glandularum in nodis inter petioles. Folia 2-
3-foliolata, interdum cirrho simplici, foliolis ellipticis vel
anguste ovato-ellipticis, minute puberulis non nisi in nervis
principalibus. Inflorescentia floribus in panicula terminali
dispositis. Calyx campanulatus, valde bilabiatus, 5-7 mm
longus, eglandulosus; corolla alba, tubulo-campanulata,
extus sparsim puberula. Capsula ignota.
Liana to at least 16 cm diam., the branchlets
terete, lepidote and minutely puberulous or papil-
lose-puberulous, longitudinally striate, drying brown¬
ish with inconspicuous lenticels, lacking pseudosti¬
pules, interpetiolar glandular fields large and
conspicuous, usually with notched upper margin.
Leaves 3-foliolate or 2-foliolate with a simple tendril
or tendril scar; leaflets elliptic to narrowly ovate-
elliptic, acute to acuminate, obtuse to truncate at
base, often the extreme base abruptly short-decur¬
rent onto petiole apex, 5-17 cm long, 3.5-9.5 cm
wide, chartaceous, secondary veins 4-6 on a side,
rather conspicuously minutely lepidote above and
below, also minutely subpuberulous on midvein above
and basal main veins below, drying gray to dark
brown above, olive-brown or olive-gray below with
conspicuously darker main veins; petiolules 0.5-6
cm long, petiole 0.6-8 cm long, lepidote and mi¬
nutely inconspicuously puberulous. Inflorescence a
terminal panicle with well-developed central axis,
the flowers mostly arranged in small ± flat-topped,
corymbose clusters, each with 1-4-cm-long pedun¬
cle (cf. A. florida), tannish puberulous and lepidote,
bracts and bracteoles minute or absent. Flowers with
the calyx campanulate, strongly bilabiate, 5-7 mm
long, 4 5 mm wide, the indument mostly of glan-
dular-lepidote and stalked-lepidote trichomes, with¬
out plate-shaped glands; corolla white, broadly cam¬
panulate above the narrowly tubular base, 1.6-2
cm long, 0.8-1 cm wide at mouth of tube, the tube
1-1.5 cm long, the lobes ca. 0.5 cm long, rather
sparsely pubescent with flexuous multicelled tri¬
chomes outside, the lobes inside puberulous mostly
with stalked lepidote glands, tube inside glabrous
except for short glandular trichomes at filaments’
insertion; stamens didynamous, the thecae divari¬
cate, 2 mm long, the connective slightly extended;
ovary oblong, 1.5 mm long, 0.8 mm wide, drying
blackish, densely lepidote; disk cylindric, 0.6 mm
long, 1 mm wide. Fruit unknown.
Distribution. Andean foothill region of upper
Amazonia from Colombia to Peru, mostly on rich-
soil forest; 130-500 m elevation.
Additional specimens examined. COLOMBIA. META:
alrededores de La Macarena, Sabanas de San Juan de
Arama, Laguna de Plaza Bonita, 500 m, 22 Apr. 1957
(fl), J. Idrobo 2606 (COL, fragment MO). Ecuador, napo:
15 km W de Coca, via Los Zorros, 0°35'S, 77°2'W, 250
m, 18-20 Apr. 1985 (fl), J. Zaruma, D. Neill, M. Baker
& W. Palacios 108 (MO). PERU. LORETO: Rio Corrientes
at Ecuador border, between Teniente Lopez and Puesto
Avanzado, 280-350 in, 4 Apr. 1977 (st), Gentry, Revilla
& Daly 19087 (AMAZ, MO); Explorama Inn, 1 km S
of Indiana, 3°30'S, 73°1’W, 130 m, 18 June 1986 (st).
Gentry, Vasquez & Jaramillo 54672 (AMAZ, MO), 13
Feb. 1987 (st), Gentry, Vasquez & Jaramillo 55764
(MO); Allpahuayo, 39 km SW of Iquitos on road to Nauta,
3°50'S, 73°25'W, 130 m, 24 Feb. 1988 (st). Gentry,
Vasquez, Blaney & Jaramillo 61884 (AMAZ, MO).
PASCO: Prov. Oxapampa, Iscozacin, Rio Iscozacin, 10°12'S,
75°13'W, 350 m, 11 July 1986 (st), 5. Knapp, R. Perez
& O. Sharehua 7828 (MO). MADRE DE Dios: Tambopata.
alluvial soil along Rio Tambopata, 12°49'S, 89°18'W,
280 m, 19 Feb. 1984 (st), Gentry, Vasquez & Jaramillo
45698 (MO).
Novon 2: 159-166. 1992.
160
Novon
This nondescript species is similar to A. fiorida
DC. except for tlie strongly bilabiate calyx and large
and conspicuous interpetiolar glandular field. The
small, white, broadly campanulate corollas are es¬
pecially reminiscent of A. fiorida. The Colombian
specimen was originally determined with a query as
A. fiorida by Dugand and later (Dugand, 1969:
260) cited as that species. However, 1 think that
the coincident combination of well-developed inter¬
petiolar glandular fields and bilabiate calyces with
the rich-soil ecology is adequate for specific rec¬
ognition.
Cuspidaria emmonsii A. Gentry, sp. nov. TYPE:
Peru. Ucayali: Provincia Coronel Portillo, Bos¬
que von Humboldt, km 86, Pucallpa-Tingo
Maria road, alt. 270 m, 8°40'S, 75°0'W, 9
Feb. 1981 (fl), A. Gentry, K. Young, S. Li-
benson, M. Olsen & T. Trucios 31126 (holo-
type, MO; isotypes, AMAZ, USM).
Frutex scandens, ramulis teretibus, consociebus glan-
dularum in nodis inter petiolos. Folia 2-foliolata, interdum
cirrho simplici, foliolis ovatis, infra valde puberulis. Inflo-
rescentia floribus in panicula terminali dispositis. Calyx
cupulatus, minute 5-denticulatus, 4-5 mm longus; corolla
rosea, tubulo-campanulata, extus puberula. Capsula li¬
nearis, teres, valde sulcata, puberula, seminibus velutinis.
Liana, the branchlets terete, the nodes with con¬
spicuous interpetiolar glandular fields, with tiny tri¬
angular pseudostipules. Leaves 2-foliolate, the leaf¬
lets ovate, acute to obtuse, usually apiculate or
abruptly short-acuminate, rounded to truncate at
base, 7-13 cm long, 4.5-10 cm wide, above sparse¬
ly puberulous mostly along main veins, below densely
finely puberulous with suberect trichomes, the upper
surface drying dark brown, lower surface light brown
or grayish. Inflorescence corymbose-paniculate, pu¬
berulous. Flowers with the calyx cylindric-cupular,
4-5 mm long, 2-3 mm wide, minutely 5-denticu-
late, the teeth extended as ridges on upper part of
calyx, puberulous; corolla magenta, sometimes with
purple dots deep in throat and purple striations in
throat, tubular-campanulate above a narrowly cy¬
lindrical base, 2-4 cm long, 0.6-1.2 cm wide at
mouth of tube, the tube 1.5—3 cm long including
the ca. 1-cm-long constricted basal part, the lobes
0.5-1 cm long, puberulous outside and on lobes and
at level of stamen insertion inside; stamens didyn-
amous, the anther thecae bent and reflexed sharply
forward near middle, ca. 1 mm long, glabrous, the
connective extremely thick, extended, minutely pa¬
pillose, puberulous apically; ovary ellipsoid, densely
lepidote, 2 mm long, 1 mm wide. Capsule linear,
terete, with a conspicuous median furrow on each
valve, 45-53 cm long, 1.8-2 cm wide, puberulous,
drying grayish, with scattered raised lenticels; seeds
bialate, flattened, 1.3-1.7 cm long, 5-6 cm wide,
the hyaline-membranaceous wings sharply demar¬
cated from the seed body, the seed body densely
velutinous.
Distribution. Southwestern Amazonia in the sea¬
sonally dry part of the southern half of Amazonian
Peru and adjacent Rondonia, 200 500 m elevation.
Additional specimens examined. Peru, madre de Dios:
Cocha Cashu, Manu National Park, Rio Manu, Oct. 1979
(fl), L. Emmons s.n. (MO), 31 Dec. 1983 (fl), Emmons
230 (MO), 1 Jan. 1984 (fl), Emmons 233 (MO). Brazil.
RONDONIA: Municipio de Santa Barbara, Rodovia BR364,
km 120, 9°10'S, 63°7'W, 10 km da Sede da Mineragao,
26 May 1982 (fr), L. Teixeira et al. 782 (MO).
This species is close to Cuspidaria subincana A.
Gentry of central and northern Amazonia, from which
it differs most strikingly in the densely velutinous
rather than glabrous seed body. The fruit of C.
subincana is also much smaller (23-28 x 1 -1.3
cm) and more lenticellate. 1 cannot reliably differ¬
entiate the two species in flower, although the tannish
pubescence of the leaflet underside of the Peruvian
material, at least, is quite distinct from the whitish
canescent leaf undersurface of T. subincana. The
tannish leaf pubescence of the Peruvian collections
of C. emmonsii is more similar to that of C. we-
berbaueri (Sprague) A. Gentry of the dry Huallaga
and Ghanohamayo valleys of Peru, which also has
velutinous seed bodies, but that species has a much
narrower white flower and distinctive grayish ma¬
genta young leaves subtending the inflorescence.
The vegetative differences between C. emmonsii
and C. subincana break down in Rondonia where
the Texeira et al. collection has the leaf undersur¬
face densely canescent as in C. subincana but the
strongly velutinous seed body of C. emmonsii. This
collection differs from other material of the complex
in a less strongly grooved capsule valve. A flowering
collection from Rondonia ( Cordeiro 613, not in¬
cluded in the description) may also be conspecific;
although it has been identified as C. subincana on
account of the densely canescent leaf undersurface,
it vegetatively matches the Texeira et al. specimen
equally well. Cuspidaria subincana is otherwise un¬
known in Rondonia. That geographically interme¬
diate material apparently is morphologically inter¬
mediate between C. subincana and C. emmonsii,
with the leaves of the former but the fruits of the
latter, might be interpreted to indicate that these
two taxa represent extremes of a cline if the differ¬
ence between glabrous and densely velutinous seeds
were not so striking.
Volume 2, Number 2
1992
Gentry
6 New Bignoniaceae Species
161
Figure 1. New species of Distictis and Schlegelin. A, B. Type of Distictis occidentals (Gentry & L. Forero
15409). —A. Inflorescence with side view of flower. —B. Front view of corolla. C, D. Schlegelia cauliflora. C.
Cauliflorous inflorescence of type (Gentry & Revilla 20656). — D. Close-up of flowers of variant with more ventricose
corolla (Gentry 42364).
Distictis occidentalis A. Gentry, sp. nov. TYPE:
Colombia. Santander: 3 km N of Yarima on
road to Albania SE of Barranca Bermeja, 25
July 1975 (fl). Gentry & L. Forero 15409
(holotype, MO; isotype, COL). Figure 1A, B.
Frutex scandens, ramulis hexagonis, stellato-puberulis.
Folia 2-foliolata, interdum cirrho trifurcato, foliolis ellip-
ticis vel late ovatis, trichoinatibus dendroideis saltern infra
in nervorum secundarium axillis. Inflorescentia floribus in
racemo terminali dispositis, pedicellis bracteis linearibus
subtends. Calyx cupulatus, minute 5-denticulatus, subtil-
iter stellato-puberulus; corolla alba, tubulo-campanulata,
extus minute stellato-puberula. Capsula elliptica, dense
stellato-tomentosa.
Liana, the branchlets hexagonal, the angles ribbed,
sparsely lepidote and thick-stellate puberulous, dry¬
ing brownish or reddish brown, the ribs usually darker,
pseudostipules absent or very early caducous, when
present on youngest branchlets subfoliaceous, in-
terpetiolar glandular fields lacking. Leaves 2-foliol-
ate, sometimes with a trifid tendril, the leaflets elliptic
to broadly ovate or oblong-obovate, obtuse to short-
acuminate or cuspidate, the base rounded or asym¬
metrically truncate (to subcordate on one side), 5-
17 cm long, 3-9 cm wide, membranaceous to char-
taceous, above scattered lepidote or lepidote-punc-
tate, otherwise glabrous except for small, few-
branched trichomes at base of midvein, below rather
densely lepidote over surface, also puberulous with
small, mostly few-branched, stellate and dendroid
trichomes at least in lower vein axils and sometimes
sparsely along main veins, drying brown to dark
olive above, paler olive or brownish olive below;
petiolules 0.8-4 cm long, petiole 1-4 cm long, mi¬
nutely puberulous with a mixture of lepidote scales
and simple and thick-stellate trichomes. Inflores¬
cence a terminal raceme, the rachis stellate-puber-
ulous, a pair of conspicuous linear bracts subtending
each pair of pedicels, these 5-12 mm long and ca.
1-2 mm wide. Flowers with the calyx cupular, 5
6 mm long, 5-8 mm wide, truncate, minutely 5-
denticulate, finely tannish stellate-puherulous, with
conspicuous, linear, glandular fields below the mar¬
gin; corolla white, tubular-campanulate above the
narrow base, 5.5-9.5 cm long, 1.4-2.2 cm wide at
mouth of tube, the tube 4.5-7.5 cm long, the lobes
1-2 cm long, dendroid- or stellate-pubescent out¬
side, with linear glandular fields below the lobes.
162
Novon
7WW$&/CK?
Z.OC.M
Figure 2. Haplolophium nunezii A. Gentry. —A. Habit (Nunez & Phillips 10412). — B. Close-up of tendril.
inside glandulardepidote and somewhat stellate, the
tube inside mostly glabrous, villous with multicellular
trichomes at base of stamens; stamens didynamous,
the thecae divaricate, 4 mm long; ovary ellipsoidal,
3-4 mm long, 3 mm wide, densely stellate tomen-
tose; disk annular-pulvinate, 1.5 mm long, ca. 4
mm wide. Capsule (immature) elliptic, 5 x 1.7 cm,
drying dark brown, finely and densely stellate-to-
mentose and lepidote.
Distribution. Upper Amazonia near the base of
the Andes from southern Peru to northern Ecuador,
also the Magdalena Valley, and northern Choco.
Distictis occidentalis is the westernmost South
American species of the genus.
Additional specimens examined. Colombia, choco:
hills behind Bahia Solano, 0-250 m, 5 Jan. 1973 (st),
Gentry <& Forero 724 IB (COL, MO); upper Rio Truando,
La Teresita, 100-200 m, 19 Jan. 1974 (st). Gentry 9361
(COL, MO); trail from Unguia to Cerro Mali, Serrania
del Darien, 300-500 m, 20 Jan. 1975 (st), Gentry &
Mori 13737 (MO). Santander: road from Barranca Ber-
rneja to El Centro, 180 m, 25 July 1975 (fl), Gentry &
horero 15398 (COL, MO); El Centro, vie. Barranca Ber-
meja, 100 m, 19 July 1938 (fl), llaught 2209 (MO).
Ecuador, napo: Rio Cuyabeno below Laguna de Cuya-
beno, 250 m, 0°3-6'S, 76°10-13'W, 14 Apr. 1980 (fl),
Volume 2, Number 2
1992
Gentry
6 New Bignoniaceae Species
163
J. Brandbyge, E. Asanza & L. Reib 30541 (AAU, MO).
Peru, madre de nios: Lago Tres Chimbadas, 15 km NW
mouth of Rio La Torre, Rio Tambopata, 260 in, 12°49'S,
69°17'W, 16 June 1980 (fl, fr), Barbour 5739 (MO);
tributary of Rio Madre de Dios below Puerto Maldonado,
250 m, 22 Apr. 1977 (st). Gentry et al. 19658 (MO).
UCAYALI: Bosque Nacional von Humboldt, km 86 Pu-
callpa-Tingo Maria, 300 m, 27 Mar. 1977 (st), Gentry
et al. 18737 (AMAZ, MO).
This species is closely related to I), granulosa
Bur. & Schumann, and most of the collections were
originally referred to that species. These two species
share such important taxonomic features as a finely
tannish tomentose calyx, lack of pseudostipules, and
reduced leaf and stem pubescence. Distictis gran¬
ulosa differs in a lavender rather than white corolla,
in lacking the conspicuous inflorescence bracts, and
especially in the more coriaceous leaflets that are
more evenly elliptic, often with a broadly cuneate
base, and that dry a different grayish olive color
below, typically with contrasting, darker-drying,
parallel tertiary veins. As thus interpreted, I>. gran¬
ulosa occurs mostly in the Guayana area, ranging
sparsely southwest to Madre de Dios, Peru. At Tam¬
bopata, Peru, the two species overlap and are clearly
distinct, with white-flowered D. occidentalis occur¬
ring in swampy areas around lake margins and lav¬
ender-flowered l). granulosa occurring in sandy
upland forest.
The other most closely related Distictis species
is D. scabriuscula (Mart, ex DC.) A. Gentry of
coastal Brazil, which differs in its larger (7 9 mm
long), more coarsely stellate-puberulous calyx. An¬
other coastal Brazilian species, D. stipularis (Mart,
ex DC.) A. Gentry, has leaflets closely resembling
I). occidentalis in texture and color but differs in
its prominent, persistent leafy pseudostipules.
There may be yet another undescribed Distictis
in Amazonian Peru, differing from I), occidentalis
in that the leaflet venation is intricately prominulous
below and the undersurface much more densely
puberulous. Included here are Croat 18576 (MO)
from 7 km southwest of Iquitos and Killip & Smith
29163 (collected by Dennis) (US) from Rio Morona,
Loreto, Peru, the latter a fruiting specimen discussed
by Sandwith (1965) in his notes on Anornocteniurn
and related genera. Sandwith suggested that this
probably represents a new taxon, which he refrained
from describing in the absence of flowers. Huashikat
1912 (MO) from Quebrada Caterpiza, Rio Santiago,
200 m, represents buds and calyces of this same
taxon, which has bracteate inflorescences like D.
occidentalis and may be only a strikingly more
pubescent form of the new species.
Haplolophium nunezii A. Gentry, sp. nov. 1 YPE:
Peru. Madre de Dios: Tambopata Province,
Cuzco Amazonica Tourist Lodge, 18 May 1989,
P. I\uhez & 0. Phillips 10412 (holotype, MO;
isotypes, CUZ, NY, USM). Figure 2.
Frutex scandens; ramuli subsexangulati sine consocie-
bus glandularum in nodis inter petiolos. Folia bifoliolata,
interdum cirrhis fissis vel simplicibus apicibus discifor-
mibus, foliolis ellipticis, coriaceis, subtus stellato-puberulis.
Inflorescentia floribus in racemo vel panicula angusta dis-
positis, bracteis et bracteolis linearibus; calyx cupulatus,
margine expanso crispato, stellato-tomentosus; corolla alba,
tubulo-campanulata, ad medium flexa; stamina didynama,
thecis divaricatis; ovarium subglobosum, dense tomento-
sum; discus patelliformis. Capsula ignota.
Liana, the branchlets inconspicuously hexagonal,
stellate-puberulous, without interpetiolar glandular
fields, with obovate foliaceous pseudostipules to 1
cm long; with a dimorphic juvenile phase growing
appressed to tree trunks. Leaves dimorphic, 2-fo-
liolate, sometimes with a short trifid tendril, one or
more tendril ends becoming disk-tipped, the others
becoming vestigial; juvenile leaflets elliptic, 1 -6 cm
long, 0.6-4 cm wide, membranaceous, puberulous
below with mostly simple, in part forked or stellate,
trichomes mostly along main veins, petioles 0.2-
0.5 cm long, the tendril apparently simple and disk-
tipped, occasionally with 1-2 short, non-disk-tipped
branches; mature leaflets elliptic, obtuse to rounded
at base and apex, 2.5-1 1 cm long, 1.5-8 cm wide,
coriaceous, above glabrescently puberulous with
simple and forked trichomes mostly near base of
midvein, below ± densely tannish puberulous with
stellate trichomes, drying shiny and olive-gray above,
tannish below; petiolules 0.3-1 cm long, the petiole
0.2-1.4 cm long, stellate-puberulous. Inflorescence
an axillary raceme or narrow panicle with short,
few-flowered lateral branches, tannish puberulous
with stellate trichomes, each flower or flower cluster
subtended by a pair of ca. 5-mm-long linear bracts
or bracteoles, the pedicels ca. 1 2 mm long, flowers
with the calyx having a cupular base and expanded
frilly margin, the base 7-10 mm long and 7-8 mm
wide, the frilly membranaceous margin irregularly
expanded into 5 lobes ca. 10-15 mm long, rather
sparsely stellate-puberulous, also with small peltate
scales; corolla white, tubular-campanulate, bent at
90° angle above base, 4-5 cm long, 1.3-1.6 cm
wide at mouth of tube, the tube cylindrical and ca.
1 cm to the bend, 2.5-3 cm long above the bend,
the lobes 0.6-1.0 cm long, outside puberulous with
thick-stellate trichomes, also with scattered stalked-
lepidote glands mostly near base of lobes, the lobes
inside stellate-puberulous, tube inside mostly gla¬
brous, glandular villous at stamen insertion; stamens
164
Novon
didynamous, the filaments 1.3-1.5 cm long, the
anther thecae divaricate, 3 mm long, the staminode
5 mm long; pistil ca. 3 cm long, style sharply bent
near middle, stellate-puberulous toward base, the
ovary subglobose, 3 mm long, 3 mm wide, densely
stellate-tomentose; disk patelliform, 1 mm long, 3
mm wide. Capsule unknown.
Distribution. Known only from lowland terra firme
forest in southwestern Amazonia from Madre de Dios
in southern Peru to Santa Cruz, Bolivia.
Additional specimens examined. Peru. MADRE de dios:
Cocha Cashu camp, Parque Nacional de Manu, 380 m,
17 Oct. 1979 (st), Gentry et al. 26802 (MO, USM).
Bolivia, santa cruz: Santa Cruz Botanical Garden, 12
km E of Santa Cruz, 17°46'S, 63°04'W, 375 m, 18 May
1991 (st), Gentry & Limpias 74083 (LPB, MO, NY,
USZ).
Haplolophium heretofore consisted of two spe¬
cies, //. rodriguesii A. Gentry, widespread in Ama¬
zonia but poorly collected, and II. bracteatum Cham.,
endemic to coastal Brazil. The fundamental char¬
acters of the genus, which belongs to subtribe Pi-
thecocteniinae and is closely related to Amphilo-
phium and Pithecoctenium, include 6-angled
branchlets, trifid tendrils, and most notably, the un¬
usual calyx with a cupular base and frilly expanded
margin. The new species differs from H. bracteatum
in its multiflowered inflorescence with linear bracts
and bracteoles, white flowers, and densely puberu-
lous lower leaflet surface, and from //. rodriguesii
in the denser inflorescence and much larger calyx
lobes. It differs from both species in its rounded
leaflet apices, more densely pubescent leaf under¬
surfaces, shorter petioles and petiolules, less con¬
spicuously 6-angled branchlets and the short disk-
tipped tendril.
In the latter features, H. nunezii more nearly
approximates Urbanolophium, a poorly known
monotypic genus of coastal Brazil, which was sep¬
arated lrom Haplolophium by Melchior (1927) on
account of terete rather than hexagonal branchlets,
disk-tipped tendrils, and nonechinate capsules. In
fact, //. nunezii is more closely related to U. gla-
ziovii (Bur. ex Schumann) Melch. than it is to either
species of Haplolophium and might well be regarded
as conspecific with U. glaziovii on vegetative
grounds. The most striking difference is that U.
glaziovii has magenta or wine-colored flowers borne
in axillary clusters of (1 —)2—3 rather than the well-
developed inflorescence of white flowers of //. nu¬
nezii; the frilly calyx margin of U. glaziovii is also
much less accentuated than in II. nunezii. Even
though specifically distinct, II. nunezii and U. gla¬
ziovii are each other’s closest relatives, and con¬
stitute another example of the remarkable disjunc¬
tion between southeastern Brazil and the Bolivia-
Peru border area of Amazonia (cf. Prance, 1973;
see also Clytostoma campanulatum (Cham.) Bur.
& Schumann).
The discovery of morphologically intermediate II.
nunezii erodes the grounds for generic recognition
of Urbanolophium , which is better considered con¬
generic with Haplolophium. Echinate and none¬
chinate fruits are found widely within the same gen¬
era in subtribe Pithecocteniineae (e.g., in
Pithecoctenium (including Neves-armondia), Am-
philophium, and Haplolophium sens, str., where
H. rodriguesii turns out to have a nonechinate fruit),
fhe branchlets of U. glaziovii are often more or
less hexagonal, although less so than in related taxa;
moreover, the tendril apex enlarged into an adhesive
disk turns out to be of much less taxonomic signif¬
icance than supposed by Melchior (1927), and is
not even constant within a species. In fact, in Pi¬
thecoctenium crucigerum (L.) A. Gentry, devel¬
opment of adhesive disks at the ends of some tendril
branches (but not others) is clearly no more than a
phenotypic response. There is also an interesting
parallel with Mansoa, another genus with typically
branched tendrils like those of most of subtribe Pi-
thecocteniinae. Like Pithecocteniinae, Mansoa has
a few species that have disk-tipped tendrils, the most
striking of which is M. parvifolia (A. Gentry) A.
Gentry, anomalous in the genus in the simple ten¬
drils, just as //. glaziovii and juvenile //. nunezii
are in Haplolophium. The similarity is so strong
that the first collection of H. nunezii, of a sterile
juvenile plant, was identified as M. parvifolia. Ap¬
parently, there is a correlation between single and
disk-tipped tendrils in otherwise trifid-tendrillate
genera.
Another close relative of H. nunezii is monotypic
Gla ziovia, which is vegetatively similar to both U.
glaziovii and //. nunezii, especially since the form¬
er’s supposedly definitive dimorphic juvenile form is
now seen to be shared with Haplolophium. Gla-
ziovia has the strongly bilabiate “pseudocleistoga-
mous” corolla of Amphilophium and seems better
retained as generically distinct. However, generic
merging of Urbanolophium and Haplolophium
seems unavoidable, necessitating the following new
combination.
Haplolophium glaziovii (Bur. ex Schumann) A.
Gentry, comb. nov. Pithecoctenium glaziovii
Bur. ex Schumann, in Engler & Prantl, Nat.
Pflanzenf. 4(3b): 218. 1894. Distictis gla¬
ziovii (Bur. ex Schumann) Bur. & Schumann,
Volume 2, Number 2
1992
Gentry
6 New Bignoniaceae Species
165
in Mart., FI. Bras. 8(2b): 180. 1896. Urban-
olophium glaziovii (Bur. ex Schumann) Mel¬
chior, Feddes Repert. Beih. 46: 80. 1927.
Bothriopodium glaziovii (Bur. & Schumann)
Rizzini, Arch. Jard. Bot. Rio de Janeiro Jan. 9:
71. 1949. TYPE: Brazil. Rio de Janeiro, Sal-
danha 7445 (B possibly destroyed, P).
Haplolophium dusenianum Kriinzl., Feddes Repert. 17:
118. 1921. Urbaniella duseniana (Kranzl.) Dusen
ex Melch., Feddes Repert. Beih. 46: 81. 1927.
TYPE: Brazil. Parana: Dusen 8015 (lectotype, BM;
isolectotypes, MO, S).
Bothriopodium glaziovii (Bur. & Schumann) Rizzini var.
symmetricum J. C. Gomes, Rev. Bras. Biol. 11: 51.
1951. TYPE: Brazil. Santa Catarina: Reitz 6 11
(RB).
Schlegelia cauliflora A. Gentry, sp. nov. TYPE:
Peru. Loreto: forest behind Mishana, Rio Na¬
nay between Santa Maria and Iquitos, 150 m
alt., 13 Nov. 1977, Gentry & Revilla 2065b
(holotype, MO; isotypes, AMAZ, F, NY, USM).
Figure 1C, D.
Frutex scandens hemiepiphyticus, ramulis plus minusve
teretis, lenticellatis. Folia elliptica vel anguste elliptica,
basim rotundata vel cuneata, coriacea, glabra. Inflores-
centia cauliflora, racemosa, pedicellis puberulis. Calyx
cupulatus, subtruncatus, 3-5(-6) mm longus; corolla ru¬
bra, anguste tubularis, 1.8—2(—2.2) cm longa, extus gla¬
bra; stamina subexserta. Fructus ignotus.
Hemiepiphytic liana growing appressed to tree
trunks, the stem without phloem arms in cross sec¬
tion, the branchlets terete to somewhat angulate and
flattened, lenticellate when older, without interpe-
tiolar ridges or glandular fields. Leaves opposite,
elliptic to narrowly elliptic (more than twice as long
as wide), rounded to apiculate at apex, rounded or
cuneate at base, without a conspicuously inrolled
basal auricle, 9-16(-19) cm long, 4-8(-9) cm wide,
coriaceous, venation plane above, midvein promi¬
nent and secondary veins prominulous below, gla¬
brous except for a few scattered lepidote scales, with
a few large plate-shaped glands near base of midvein
below and a few smaller persistent glands scattered
on undersurface; petiole woody, 5-10 mm long.
Inflorescence a cauliflorous cluster of several ±
contracted racemes with a woody short-shoot rachis
and minute inconspicuous bracts, the pedicels pu-
berulous, 3-5 mm long. Flowers with the calyx
cupular, subtruncate, 3-5(-6) mm long, 3-4(-5)
mm wide, drying brown, slightly puberulous or gla-
brescent, with a few scattered, large, dark-drying
plate-shaped glands; corolla red, narrowly tubular,
sometimes slightly ventricose toward apex, 1.8-2(-
2.2) cm long, 4-5 mm wide at mouth of tube, the
lobes 1-2 mm long, the tube glabrous outside, inside
lepidote-glandular and glandular puberulous at sta¬
men insertion; stamens about equal, subexserted,
inserted 5 mm from base of corolla, the thecae
parallel, l(-2) mm long; pistil ca. 5 mm long, the
ovary ovoid, 1.5 mm long, 1.5 mm diam., glabrous;
disk fused with and not clearly differentiated from
base of ovary. Fruit unknown.
Distribution. Known only from northwestern
Amazonia in forests on white sand; elevation below
200 m.
Additional specimens examined. COLOMBIA. AMAZONAS:
Araracuara, 2 Aug. 1986 (st), Battjes 331 (U); Rio
Apaporis, raudal Jirijirimo, (fl), Schultes 13468 (COL).
CAQUETA: Santa Isabel, quebrada El Chururuco, 5 Nov.
1984 (st), La Rotta & Mirana 534 (HA), vaupes:
Inambu, 18 Nov. 1952 (bud), Romero-Castaneda 3627
(COL). Brazil. AMAZONAS: Rio Javari behind Palmeiras
Army Post, 5°8'S, 72°49'W, 31 July 1973 (fl), Lleras
et al. P16963 (MO, NY). PERU. LORETO: Mishana (type
locality), 14 July 1977 (fl), Solomon 3620 (MO); Mis-
huyacu, 100 m, Oct.-Nov. 1929 (fl), Klug 466 (US);
SW of Iquitos at Pena Negra, 600 ft., 21 Nov. 1975
(fl), C. Davidson 3657 (MO); Quistacocha, 200 m, 27
May 1978 (fl), Gentry & Jaramillo 22312 (AMAZ, MO);
Mishana, 140 m, 6 jan. 1983 (fl), Gentry et al. 39185
(AMAZ, MO); Rio Vainilla below Rio Maniti, Caseria
Catalan, 3°35'S, 72°45'W, 29 June 1983 (fl). Gentry
42364 (AMAZ, MO); Allpahuayo, 130 in, 35 km SW
of Iquitos, 3°50'S, 73°25'W, 16 Feb. 1987 (fl). Gentry
et al. 56024 (AMAZ, MO).
Local names. “Lengua de picon” (Caqueta), “te-
nekoniyanejo” (Mirana).
Uses. The Mirana Indians use this plant as one
of the ingredients of their curare (fide La Rotta 534).
This species is closely related to S. spruceana
Schumann of the Guayana area of southern Vene¬
zuela and adjacent Brazil and Guyana, but is oth¬
erwise isolated in the genus. That species has a larger
(5-7 x 4-6 mm) calyx and longer (2-2.5 cm long)
corolla tube and generally larger (15-26 cm long),
less coriaceous leaves with the tertiary venation typ¬
ically more prominulous beneath and the leaf base
with conspicuously incurved basal auricles. The in¬
florescence of S. spruceana usually has fewer flow¬
ers and is a single raceme rather than branched at
the base. It is possible that the western Amazonian
plant should be regarded as part of a variable S.
spruceana, but the uniformity of the available col¬
lections and the different geographic distributions
support species recognition. The new Schlegelia
species from Amazonian Colombia here described as
S. hirsuta is also part of this complex; it differs
from S. cauliflora primarily in the same vegetative
characters (subbullate leaves with veins hirsute be¬
low) that distinguish it from S. spruceana (see be¬
low).
166
Novon
Schlegelia hirsuta A. Gentry, sp. nov. TYPE:
Colombia. Vaupes: margenes del Rio en Pu-
caron, 240 m, 2 Oct. 1939, Cuatrecasas 7128
(holotype, COL 35587; isotypes, COL 35589,
fragment MO).
Frutex scandens (hemiepiphyticus?), ramulis subtere-
tibus. Folia elliptica, basim rotundata sed auriculis basa-
libus, coriacea, plus minusve bullata, infra hirsuta. Inflo-
rescentia cauliflora, contracto-racemosa, pedicellis
puberulis. Calyx campanulatus, truncatus, 3-5 mm lon-
gus; corolla rubra, anguste tubularis, aliquantum ventri-
cosa, 2-2.2 cm longa. Fructus ignotus.
Liana (probably bemiepiphytic; one collection in¬
dicated as treelet), branchlets subterete to irregularly
subangled, glabrous, without interpetiolar ridges or
glandular fields; pseudostipules apparently lacking.
Leaves opposite, simple, elliptic, acute or round and
shortly cuspidate apically, basally rounded except
lor a conspicuously auriculate 5-mm-long basal at¬
tenuation, 11-24 cm long, 4-16 cm wide, coria¬
ceous, ± bullate, the secondary veins 6 9 on a side,
impressed above and prominent below, glabrous ex¬
cept for a few, scattered, long trichomes above,
below hirsute with 1 -2-mm-long stiff trichomes along
main veins and widely scattered over surface; petiole
woody, 5 10 mm long to level of base of infolded
auricles, an additional 5 mm between the auricles.
Inflorescence a contracted raceme, borne on older
stems, the branches somewhat puberulous. Flowers
with the calyx campanulate, truncate, 3-5 mm long,
4-5 mm wide, slightly puberulous, corolla red, nar¬
rowly tubular, 20-22 mm long, to 5 mm wide,
slightly ventricose in upper half, the lobes minute
(1-1.5 mm long), minutely glandular-lepidote out¬
side, slightly puberulous at level of stamen insertion;
stamens inserted 6 mm from base of corolla tube,
subexserted, the thecae parallel, 1 mm long, the
longer filaments ca. 12 mm long, shorter filaments
8 9 mm long, staminode 5 mm long; pistil 3-4 mm
long, the ovary spherical, 1.5 mm long, 2 mm wide,
minutely lepidote, the ovules multiseriate in each
locule on a single central placenta; disk indistin¬
guishable from base of ovary. Fruit unknown.
Distribution. Known only from the Rio Vaupes
region of Amazonian Colombia.
Additional specimens examined. COLOMBIA. VAUPfe:
Yurupari (orilla Vaupes) 350 km arriba de Mitu, 220 in,
24 Sep. 1939 (H), Cuatrecasas 6994 (COL); entre el
Vaupes y el Apaporis, selva del Cano Popore, afluente
del Apaporis, 240 m, 30 Sep. 1939 (fl), Cuatrecasas
7085 (COL).
Schlegelia hirsuta is closely related to S. spru-
ceana Schumann of the Guayana region of Guyana,
Venezuela, and Brazil. The two species share cau-
lifiorous tubular red flowers and rather large cori¬
aceous leaves that are usually auriculate at the base.
The Colombian plants differ in the ventricose upper
part of the corolla, impressed leaf venation above,
hirsute midrib and secondary veins below, and short¬
er calyx. Schlegelia spruceana has an evenly tu¬
bular corolla, glabrous leaves with the main veins
slightly raised or plane above, and a calyx 5-7 mm
long. While these differences could prove inter¬
pretable as varietal or subspecific variation, the strik¬
ingly long trichomes along the mid vein below are
unique in the genus, which otherwise has glabrous
or lepidote leaves, and strongly favor specific rec¬
ognition.
Acknowledgments. The fieldwork leading to the
discovery of these species was supported by grants
from the National Science Foundation (most recently
BSR-8607113) and the Mellon Foundation. The
paper was written with the support of a fellowship
from the Pew Scholars Program.
Literature Cited
Dugand, A. 1969. Notas sobre la flora de Colombia y
paises vecinos, III. Caldasia 10: 247-286.
Melchior, H. 1927. Der natiirliche Formenkreis der
Pithecocteniinae innerhalb der Familie der Bignoni-
aceae. Feddes Repert. Beih. 46: 71-82.
Prance, G. 1973. Phytogeographic support for the the¬
ory of Pleistocene forest refuges in the Amazon Basin,
based on evidence from distribution patterns in Car-
yocaraceae, Chrysobalanaceae, Dichapetalaceae and
Lecythidaceae. Acta Amazon. 3: 5-26.
A New Species of Aegiphila (Verbenaceae) from Colombia
Ricardo M. Rueda
Departamento de Biologia, Universidad Nacional Autonoma de Nicaragua, Leon, Nicaragua
Current address: Missouri Botanical Garden, P.O. Box 299,
St. Louis, Missouri 63166, U.S.A.
ABSTRACT. A new species, Aegiphila narinensis
(Verbenaceae), from La Planada Reserve, Colombia,
is described, and its affinities are discussed. The new
species has mucronate calyx lobes, and the corolla
is larger than in any other species in the genus and
has a curved tube. This name is needed to complete
the botanical list of plants collected at La Planada
Reserve being compiled by Olga de Benavides and
Alwyn Gentry.
Aegiphila is a genus of about 160 species of
shrubs and trees found from Cuba and Mexico
through the West Indies and Central America to
Peru and northern Argentina. About 40 species are
found in Colombia (Moldenke, 1980), 10 of which
are endemic, including this new one.
Aegiphila narinensis Rueda, sp. nov. TYPE: Co¬
lombia. Narino: La Planada Reserve, near Ri-
caurte, 1.800 m, 01°O5'N, 78°01'W, 21 Dec.
1987 (fl), Gentry, Benavides & Keating 59648
(holotype, MO; isotype, PSO not seen). Fig¬
ure 1.
Suffrutex 0.5-1 m altus, glabrus. Folia petiolata; lam-
inis subtus punctatis, ellipticis, 14-18 cm longis x 8-10
cm latis, chartaceis, 6-8 venis secundariis. Inflorescentiae
3.5-6 cm longae x 3-5 cm latae; bracteis 2-4 mm
longis; pedicellis 1.5-3 cm longis. Flos lobis calycinis 1-
3 mm longis mucronatis; corolla hypocrateriformi alba,
glabra, tubo arcuato, 2.5-3 cm longo, lobis 1.5-2 cm
longis, inaequalibus.
Subshrub 0.5-1 m tall; branches tetragonal or
subterete, sparsely pilose, glabrescent. Leaves de¬
cussate-opposite; blade elliptic, acute to acuminate
at apex, acute to cuneate at base, 14-18 x 8-10
cm, chartaceous, sparsely hirsute, both laminar sur¬
faces glabrous, lower surface punctate; venation eu-
camptodromous to weakly brochidodromous, the
midrib slightly elevated adaxially, prominently ele¬
vated abaxially, secondary veins 6-8 on each side
of costa, at an angle of 45° with the costa, impressed
adaxially and elevated abaxially, arcuate-ascending,
with well-marked minor veins; margins entire to
slightly sinuate; petioles 0.8-1 cm long. Inflores¬
cences cymose, axillary, few-flowered, 3.5-6 x 3-
5 cm; peduncles 1.5-2.5 cm long, bracts linear, 2-
4 mm long; pedicels 1.5 3 mm long; peduncle,
bracts, and pedicels glabrous. Calyx glabrous, the
lobes triangular, mucronate, 1-3 mm long; corolla
white, glabrous, salverform, the tube curved, 2.5-
3 cm long, limb lobed, 1.5-2.5 cm wide, the lobes
obovate, 1.5-2 cm long, unequal; stamens 4, at¬
tached to the throat of the corolla, equal, the fila¬
ments filiform, ca. 2 mm long, slightly surpassing
the corolla mouth, the anthers oblong; ovary globose,
the style filiform, reaching to 3-5 mm below stamen
insertion, 1.5 cm long, stigma bifid, the branches 1
mm long. Fruit not seen.
Aegiphila narinensis is distinguished from other
species in the genus by having mucronate calyx
lobes and a curved corolla tube about 3 cm long.
The corolla is larger than in any of the other species,
with unequal lobes 1.5-2 cm long. This species
resembles the complex of closely related and large-
flowered species formed by Aegiphila costaricensis
Mold., from Central America, and A. haughtii Mold.,
Clerodendrum tessmanni Mold., and C. ulei Hayek
from South America, as discussed by Nash & Nee
(1984). However, elements of this group have straight
corolla tubes of less than 25 mm long, with lobes
more or less equal, whereas A. narinensis has a
curved corolla tube of more than 25 mm in length,
and unequal lobes. In addition, the species of Cler¬
odendrum in this group have stamens that are long
exserted, whereas A narensis has stamens that only
slightly surpass the corolla mouth.
The type was collected at 1,800 m elevation, in
the cloud forest of the La Planada Reserve, in the
state of Narino, after which the species is named.
This area is famous for its high biological endemism
(Terborgh & Winter, 1982), to which this distinctive
new species adds yet another example.
Acknowledgments. I thank John Myers for the
excellent illustration, and William G. D’Arcy, Alwyn
Novon 2: 167-168. 1992.
168
Novon
Figure 1. Aegiphila narinensis Rueda (Gentry, Benavides <£- Keating 59648). —A. Habit. —B. Flower. —C.
Flower, partial longitudinal section.
H. Gentry, and P. Mick Richardson for comments
on the manuscript.
Literature Cited
Moldenke, H. N. 1980. Phytologia Memoirs II. A Sixth
Summary of the Verbenaceae, Avicenniaceae, Stil-
baceae, Chloanthaceae, Symphoremaceae, Nyctan-
thaceae, and Eriocaulaceae of the World as to V alid
Taxa, Geographic Distribution and Synonymy. H. N.
& A. L. Moldenke, Plainfield, New Jersey.
Nash, D. L. & M. Nee. 1984. Verbenaceae. Flora de
Veracruz 41: 1-54.
Terborgh, J. & B. Winter. 1982. Evolutionary circum¬
stances of species with small ranges. Pp. 587-600
in G. T. Prance (editor), Biological Diversification in
the Tropics. Columbia Univ. Press, New York.
Mesanthophora , a New Genus of Vernonieae
(Asteraceae) from Paraguay
Ha rold Robinson
Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, U.S.A.
ABSTRACT. The new genus Mesanthophora is from
a limestone area of central Paraguay. It looks similar
to Lepidaploa, but has decurrent auriculate-based
leaves, pedunculate heads from the middle of inter¬
nodes, nearly equal numbers of involucral bracts
and florets, and triporate, totally lophate, strongly
columellate pollen lacking a perforated tectum.
A Paraguayan specimen belonging to the Aster-
aceous tribe Vernonieae that was sent by the Mis¬
souri Botanical Garden for identification has proven
to represent a distinctive, previously undescribed
genus. A superficial resemblance to Lepidaploa
proves misleading. Both the genus and the species
seem undescribed in spite of the many members of
the Vernonieae that have previously been described
from Paraguay by such authors as Chodat (1901-
1902).
The type specimen of the new genus has a general
habit of the common neotropical Vernonian genus
Lepidaploa (Cass.) Cass. (Robinson, 1990), with
series of heads among large foliose bracts along the
branches of the inflorescence. However, initial ob¬
servation shows that each head is pedunculate, un¬
like the heads of Lepidaploa. The new genus also
differs by the position of the pedunculate heads on
the middle of the internodes rather than at the nodes.
The supra-axillary position seems rigorously main¬
tained throughout the inflorescence and is a char¬
acteristic that distinguishes the plant from all other
Vernonieae. In a final distinction, the leaf bases are
sessile, decurrent for up to 5 mm, and are auriculate
below the decurrency. The decurrency holds the
basal auricles of the leaf at a downward angle com¬
pared to the rest of the leaf blade. Lepidaploa does
not have sessile auriculate leaf bases. Other South
American Vernonieae with sessile, markedly auric¬
ulate leaf bases, such as Vernonia auriculata Gri-
seb., V. jubifera Rusby, and V. prenanthoides Glea¬
son, all of Bolivia, have no decurrency, have more
densely pyramidal inflorescences without large fo-
liiform bracts, and have smaller heads bearing type
A pollen (Jones, 1979).
In the reproductive parts of the plant, other than
pollen, it is the nature of involucral bracts that most
sets the new genus apart from Lepidaploa and its
relatives. Mesanthophora has nearly equal numbers
of bracts and florets of the heads, which is similar
to the Vernonia of the eastern United States. Lep¬
idaploa and its close relative Lessingianthus H.
Robinson have one and a half to three times as many
bracts as florets in a head. The only examples among
the Lepidaploa relatives that have fewer bracts than
florets are some species of Chrysolaena H. Rob¬
inson. which have reduced numbers of both bracts
and florets. The involucre of the new genus differs
from those of most Lepidaploa because it lacks
strong structural differentiation between inner and
outer bracts.
The pollen of the new genus shares a lophate
condition with Lepidaploa and its relatives, but the
detailed form is totally different. The pollen lacks
evident colpi and has numerous, rather uniform
areoles covering the poles and sides equally. The
areolation of the grains does not seem rigorously
consistent in form; generally, an areole seems to be
centered at each pole, surrounded by six subpolar
areoles. The areoles containing the pores are some¬
times directly below an areole of the subpolar series
and sometimes below a wall between two subpolar
areoles. Lateral areoles between the pores always
seem staggered. Basal columellae are concentrated
toward the junctures of the muri of the exine and
often seem to radiate from the curved lower edge
of the crests of the muri. The muri lack a perforated
tectum of the type found in Lepidaploa and its
relatives, and the grains lack the spines and contin¬
uous perforated tectum of the type A pollen found
in Vernonia and many other members of the tribe.
The general pollen form with many polar areoles
and no evident colpi is rare among neotropical Ver¬
nonieae, but is more common in the Faleotropics.
In South America only Acilepidopsis H. Robinson
has been found to have precisely the same type of
pollen with columellae concentrated near the junc¬
tures; a relationship to paleotropical Vernonieae has
been suggested for that genus (Robinson, 1989).
The inflorescence of Acilepidopsis has sessile heads
Novon 2: 169-172. 1992.
170
Novon
Figures 1-7. Mesanthophora brunneri H. Robinson. —1. Top of plant with inflorescence. —2. Head. —3. Corolla
showing tips of style and anthers. —4. Style showing basal node. —5, 6. Young achenes with part or all of inner
pappus attached. —7. Older achene with inner pappus lacking.
Volume 2, Number 2
1992
Robinson
Mesanthophora
171
Figures 8-11. Mesanthophora pollen. —8. Polar view. —9. Lateral view with pore. —10, 11. Details of exine
with lintels of crests broadened in middle and columellae concentrated at corners of areoles.
containing only ca. 30 oblong, short-tipped involu-
cral bracts and 8-13 florets, and the heads are not
positioned in the middle of the internodes. As in the
case of Acilepidopsis, the chemistry and chromo¬
some number of the new genus, which might indicate
hemispheric relationships, are not known. The neo¬
tropical aquatic Pacourina Aubl., which is outside
of the traditional generic concept of Vernon in with
172
Novon
its large achenes and short pappus, has pollen similar
to that of Mesanthophora and Acilepidopsis, and,
in spite of the great difference in appearance, it
could be a comparatively close relative.
The new genus is named Mesanthophora in ref¬
erence to the position of the heads in the inflores¬
cence at the middle of the internodes. The species
is named for the collector, David R. Brunner.
Mesanthophora brunneri H. Robinson, gen. el
sp. nov. TYPE: Paraguay. Concepcion: Arroyo
Tagatiya-nd, 22°37'S, 57°32'W, 140 m, 4 abr.
1986, David R. Brunner 1720 (holotype, US;
isotypes, MO, PY). Figures 1-11.
A Lepidaploa foliis base decurrentibus et auriculatis,
capitulis ex rnediis internodiorum pedunculatis, bracteis
involucralibus et floribus ca. 100 in quoque capitulo, et
granis pollinis triporatis isodiametrice multiareolatis in par-
tibus polaribus et lateralibus tecto imperforato distincta.
Perennial herbs to 80 cm high. Stems slender,
smooth, glabrous, nonfistulose. Leaves alternate,
sessile, oblong-ovate, mostly 1.5—7.5 cm long, 1.0-
3.2 cm wide, upper leaves smaller, base of blade
bending downward and distinctly decurrent, ex¬
panded outward into broad auricles, margins sub-
denticulate, apex shortly acute and apiculate, sur¬
faces glabrous, glandular-punctate below.
Inflorescence narrowly thyrsoid-paniculate, with se-
riately cymose apex and branches, lower heads ma¬
turing first; peduncles arising from middle of inter¬
nodes without closely subtending bract, 4-10 mm
long, glabrous. Heads broadly campanulate; invo-
lucral bracts ca. 100, graduated in ca. 5 series,
linearly lanceolate, 3-6 mm long, base to 1 mm
wide, apex narrowly acute, outside glabrous. Florets
90-100; corollas bluish lavender, ca. 6.5 mm long,
tube narrowly funnelform, ca. 3.5 mm long, throat
ca. 1 mm long, lobes narrowly oblong, ca. 2 mm
long, ca. 0.4 mm wide, with glandular dots outside
more dense distally; anther thecae ca. 1.5 mm long;
apical appendages oblong-ovate, ca. 0.2 mm long,
glabrous, with thin-walled cells; style base with dis¬
tinct narrow node. Achenes ca. 2 mm long, 8-10-
costate, with many short, subappressed setulae most
persistent between costae; pappus bristles white, ca.
25, ca. 3 mm long, fragile, narrowed at base, not
broadened at apex, squamae of outer series ca. 2
mm long. Pollen grains ca. 55 /am diam., triporate.
lophate, with numerous subisodiametric areoles in
polar and lateral parts, without perforated tectum,
columellae broadly attached to base, concentrated
at junctures of muri, lintels of muri broadened in
middle with convex lower margins.
Additional label data is, “Bosque ribereno de 20
35 m de altura cambiando a savanas y campos fuera
del arroyo. Suelo calcareo de poco profundidad sobre
pedra caliza.” The description of the type locality
is interpreted as a wooded streamside over limestone
in a savanna and campo area. The collection area
is on the southern edge of a large limestone and
dolomite region extending northward into adjacent
westernmost Mato Grosso do Sul in Brazil. The latter
is an area of low hills in the Pantanal adjacent to
seasonally flooded lowlands. Additional collections of
the new genus and many other distinctive plants
should be found in the area.
Acknowledgments. The drawings of Mesantho¬
phora were prepared by Alice Tangerini of the De¬
partment of Botany, National Museum of Natural
History, Smithsonian Institution. The material of the
pollen was prepared by Sherry Pittam and Carol
Annable of the Department of Botany, under the
supervision of Joan Nowicke. The photographs were
taken by Suzanne Braden of the National Museum
of Natural History SEM Laboratory using a Hitachi
S-570 scanning electron microscope. Prints were
prepared by Victor E. Krantz, staff photographer.
National Museum of Natural History.
Literature Cited
Chodat, R. 1901-1902. Plantae Hasslerianae, soit en¬
umeration des plantes recoltees au Paraguay par le
Dr. Emile Hassler, D'Aarau (Suisse) de 1885-1895
et de 1898- 1900. Compositae. Bulletin de l.'Herbier
Boissier, 2me serie 1: 408 419; 2: 297-312, 382-
400.
Jones, S. B. 1979. Synopsis and pollen morphology of
Vernonia (Compositae: Vernonieae) in the New World.
Rhodora 81: 425-447.
Robinson, H. 1989. Acilepidopsis, a new genus of
Vernonieae from South America (Asteraceae). Phy-
tologia 67: 289-292.
-. 1990. Studies in the Lepidaploa complex
(Vernonieae: Asteraceae) VII. The genus Lepida¬
ploa. Proceedings of the Biological Society of Wash¬
ington 103: 464 498.
Recognition of Brownea coccinea Jacq. subspecies
capitella (Leguminosae: Caesalpinioideae)
Dilia Velazquez
Departamento de Biologia de Organismos, Universidad Simon Bolivar,
Apartado 89000, Caracas 1081, Venezuela
ABSTRACT. A new subspecies, Brownea coccinea
subsp. capitella, is recognized based on geographical
separation and morphological differences. A key to
the subspecies is provided.
Brownea is a neotropical genus of about 12 spe¬
cies occurring between sea level and 1,300 m, usu¬
ally in humid microenvironments. Following an ini¬
tial study of the genus in Venezuela (Velasquez &
Agostini, 1981), a need to revise the classification
of B. coccinea became clear. Study of numerous
specimens revealed a wide range of phenetic vari¬
ation, which had previously led to descriptions of
several new species, here considered as synonyms.
Two groups are now being recognized, correspond¬
ing to different geographical regions in Venezuela:
B. coccinea subsp. coccinea, from the western part
of the Cordillera de la Costa, Zulia, and the Cordillera
de los Andes; and B. coccinea subsp. capitella,
occurring in the central-eastern part of the Cordil¬
lera de la Costa south to the Territorio Federal
Amazonas. Subspecies capitella is the more com¬
mon of the two, also occurring in Cuyana, Trinidad-
Tobago, and northern Brazil. Klitgaard (1991) re¬
duced Brownea angustifolia Little to a subspecies
of B. coccinea, but it is here considered to he a
distinct species restricted to Ecuador and south¬
western Colombia. Besides the geographical sepa¬
ration, B. coccinea has five equal petals, whereas
B. angustifolia has three large, spathulate petals
and two smaller, narrower ones. The two species
also differ in the shape of the sepals and the form
and apex of the bracteoles.
Brownea coccinea Jacq., Select. Stirp. Amer. Hist.
194, t. 121. 1763. Hermesias coccinea (Jacq.)
Kuntze Rev. Gen. 1: 191. 1891. TYPE: tabula
121 in Jacq., Select. Stirp. Amer. Hist. 1763.
Trees 3-8 m high. Leaves petiolate; petioles 5-30
mm long; rachis 4-28 cm long; stipules deciduous;
petiolules 1-4 mm long; leaflets in (4 )6 pairs, 4-
20.5 X 2.2-7.2 cm, glabrous or puberulous at the
base and on the costa, opposite or subopposite, el¬
liptic, with a gland at the base; base cordate, oblique
or acute; apex long acuminate; costa flat or slightly
hollow at the abaxial surfaces. Inflorescence with
ca. 10 30 flowers; peduncle glabrous or puberulous,
2-10 mm long; the floral rachis 2-15 cm long;
bracts widely obovate, puberulous, deciduous; brac-
tlets filiform, deciduous, 12-42 mm long; the con¬
nate bractlets glabrous or puberulous, red, 13-35
mm long, 2-lobulate, the lobule apex acute or round¬
ed. Flowers 5-7.8 cm long; pedicel 2-18 mm long;
hypanthium 6-25 mm long; sepals 4, 14-25 x 2-
8 mm, red; petals 5, 22-48 x 7-19 mm, red,
spathulate, the apex rounded, apiculate or retuse.
Stamens 11, 33-48 min long; the staminal sheath
14-35 mm long, irregularly connate; anthers 1.5-
4 x 1-1.5 mm. Ovary 7-12 mm long, the stipe
5-1 7 mm long; style glabrous or puberulous at the
base, 25-38 mm long. Fruit puberulous, villous or
glabrate, 13-24 x 2.6-4.4 cm, the surface usually
smooth and shiny when mature, the stipe 15-40
mm long, the apex acute, curved. Seeds dark brown,
20-30 x 15-20 mm.
Key to the subspecies of Brownea coccinea
la. Connate bractlets longer than the hypanthium
(ratio of connate bractlets : hypanthium > 1.5:
1); ovary stipe 5-9(-ll) mm long; anthers 3
4 mm long; leaflets glabrous or puberulous at
the base and on the costa, the margins occa¬
sionally slightly revolute; style glabrous; inflo¬
rescences with 25-30 flowers; pedicels 2-12
mm long .1. B. coccinea subsp. capitella
lb. Connate bractlets as long as or shorter than the
hypanthium (connate bractlets : hypanthium ra¬
tio ca. 1:1); ovary stipe 14-17 mm long; an¬
thers 1.5-3.5 mm long; leaflets glabrous, the
margins usually slightly revolute; style puber¬
ulous at the base; inflorescences with 15-20
flowers; pedicels 5-17 mm long .
.2. B. coccinea subsp. coccinea
1. Brownea coccinea Jacq. subsp. capitella
(Jacq.) D. Velazquez & Agostini, stat. et comb,
nov. Brownea capitella Jacq., Fragm. Bot.
Illustr.: 26, t. 18, 19. 1809. Hermesias cap¬
itella (Jacq.) Kuntze, Rev. Gen. 1: 191. 1891.
TYPE: tabulae 18. 19 in Jacq., Fragm. Bot.
Illustr. 1809.
Novon 2: 173-175. 1992.
174
Novon
Hrownea latifolia Jacq. Fragm. Bot. Illustr.: 25, t. 17.
1809. Hermesias latifolia (Jacq.) Kuntze, Rev. Gen.
1: 191. 1891. TYPE: tabula 17 in Jacq., Fragm.
Bot. Illustr. 1809.
Hrownea racemosa Jacq., Fragm. Bot. Illustr.: 25, t. 16.
1809. Hermesias racemosa (Jacq.) Kuntze, Rev.
Gen. 1: 191. 1891. TYPE: tabula 16 in Jacq.,
Fragm. Bot. Illustr. 1809.
Hrownea guianensis Klotzsch in R. M. Schomburgk,
Reis. Br.-Guiana 3: 1209. 1848, nom. nud.
Distribution. Widely distributed through eastern,
southern, and north-central Venezuela, Trinidad-
Tobago, Guyana, and extreme northern Brazil, from
50 to 950 m.
Specimens examined. Brazil, roraima: vicinity of
Auaris, 4°3'N, 64°22'W, Prance et al. 9845 (K). Guyana.
Cuyuni River, Sandwith 626 (K); Essequibo River, Mor-
aballi Creek, Forest Dept. 2418 (K); Baramita, Hardy
75 (K); Barina River, Jenman 6933 (K); Mazaruni River,
Jenman 7621, 7595 (both K); Pomeroon District, Im
Thurn 1946 (K), Schomburgk 90 (K); Russell District,
Jenman 2082 (K); Mazaruni River, Hohenkok 904 (K).
Tobago. Eggers 5670 (GH); near Charlotesville, F. IF.
II. 20106 (GH); Mason Hall area, Howard 10468 (BM,
GH). Trinidad. Arima Valley, near Simla, Smith 10063
(GH); heights of Aripo, Broadway 9794 (GH); Blanchis-
seuse road, Broadway 6003 (BM, MO); near Jabaquito,
Broadway s.n. (GH). VENEZUELA. TERRITORIO FEDERAL
AMAZONAS: alrededores de San Juan de Manapiare, Agos¬
tini 1553 (VEN); Sierra Parima, vecindades de Simara-
wochi, Rio Matacuni, Steyermark 107439 (MO, VEN);
vecindades del Rio Coro Coro y del Aeropuerto de Yutaje,
Steyermark 117903 (MO, VEN); selva de galena de El
Gristo, Williams 12565 (K); El Palmar, Williams 12897
(K); Cacuri, 4°50'N, 65°20'W, Lister & Colchester 2377
(K); Cano Marieta, 5°10'N, 66°30’W, Lister & Col¬
chester 2294 (K). ANZOATEGUI: La Mata de Cafe, NE de
Bergantin, Morillo 163 (VEN); a lo largo del rio Leon,
500 m cerca de la Quinta Danta, Steyermark 61007
(MO, VEN). ARAGUA: Parque Nacional Rancho Grande,
Garcia 34 (VEN), Lasser 2338 (VEN), Fittier 15348
(VEN); carretera Cumboto-Turiamo, Trujillo 5682 (MY);
El Limon, Williams 10230 (VEN). bolivar: NE de Ca-
naima, S de Cerro Venado, Agostini 306 (VEN); Depto.
Caroni, Ciudad Guayana, Puerto Ordaz, Parque Loefling,
Agostini 2629 (VEN); Ciudad Piar, laderas no ferrugi-
nosas del Cerro Bolivar, Aristeguieta 2206 (Y EN); Puerto
Ordaz, San Felix, Aristeguieta 5303 (MO, VEN); pie de
Sierra Imataca, estacion Magdalena, El Palmar, Bernardi
7051 (MER); carretera Upata-San Cayetano, Sitio Gual-
pa, Cardona 2133 (VEN); Depto. Piar, La Gamilera, 40
km W de El Manteco, Delascio s.n. (VEN); La Gran
Sabana, 2 km W de Kamarata, Koyama 7256 (VEN);
20 km E de La Paragua, Liesner 5458 (MO, VEN); 30
km E de Upata, Little 15926 (VEN); Alto Rio Cuyuni,
Maguire 46987 (VEN); 15 km SE de El Callao, Little
17528 (MER); Rio Caura, arriba del Salto Para, en las
islas, 2-3 km del campamento Las Pavas, Steyermark
112938(\ EN); cerca de El Palmar, Wessels 2071 (MER);
Bajo Paragua, rebalses de San Mateo, Williams 12815
(VEN). GARABORO: cerca de San Joaquin, Fittier 7624
(VEN). Delta Amacuro: Depto. Tucupita, 13-14 km SE
de Los Castillos, Little 17671 (MER). DISTRITO FEDERAL:
bosque alrededor de Los Caracas, Aristeguieta 3088
(VEN); Los Caracas, Brito 7 (VEN); Suapo, Hda. El
Limon, Gomez 43 (VEN); La Guaira, Delgado 133 (VEN);
7 km E de Los Caracas, Steyermark 91508 (VEN).
Miranda: carretera Higuerote-Chuspa, 20 km NW de
Higuerote, 10°35'N, 66°12’W, Agostini 2661, 2662
(both VEN); W de Chirimena, 1 km E de la desembo-
cadura del rio Aricagua, Berry 968 (MO, VEN); Taca-
rigua, Curran 2, 4 (both MER); sendero a El Frio, Santa
Lucia, Soto 5 (VEN); Depto. Brion, NE de Pueblo Seco,
7 km E de Aricagua, Steyermark 106914 (VEN); car¬
retera Higuerote-Chirimena, Velasquez 70, 94 (both
VEN). MONAGAS: El Balneario, entre Las Piedras y Sabana
de Las Piedras, al lado del rio Guarapiche, Badillo 7314
(MY); Chapopotal, cerca de Quiriquire, Lasser 4096
(VEN); entre Caripito y Quiriquire, Tamayo 3512 (VEN);
entre La Pica y Cano Colorado, E de Maturin, Wurdack
39530 (VEN). SUCRE: Cumana, Funck 6 (K); sabanas en
cuencas del rio Cariaco, Castillo 56 (VEN); Cerro Palo
de Agua, SE de El Rincon, 7 km N de El Pilar, Morillo
3767 (VEN); alrededores de Cumanacoa, quebrada Jua-
juas, Fittier 14670 (VEN); Depto. Sucre, Quebrada Zu-
rita, Paso Hondo, S de Limonal y Santa Fe, Steyermark
107813 (VEN); Serrania de La Paloma, entre Ajies y
Guariquen, Steyermark 121357 (V EN); Depto. Benitez,
E de Los Pozotes, entre Guaraunos y Ajies, Steyermark
121271 (VEN).
2. Hrownea eoccinea Jacq. subsp. coccinea.
Brownea aroensis Pittier, Contr. U.S. Natl. Herb. 18:
152. 1916. TYPE: Venezuela. Lara: Palmasola,
Aroa River, Fittier 6367 (holotype, US not seen;
photo, VEN).
Brownea guaraba Pittier, Contr. U.S. Natl. Herb. 18:
152. 1916. TYPE: Venezuela. Lara: Palmasola,
Aroa River, Pittier 6366 (holotype, US not seen;
photo, VEN).
Brownea guaramaco Pittier, Cat. Flora Ven. 1: 381.
1945, nom. nud.
Brownea longiflora Pittier, Cat. Flora Ven. 1: 381. 1945,
nom. nud.
Brownea superba Pittier, Cat. Flora Ven. 1: 381. 1945,
nom. nud.
Distribution. Venezuela, in the western part of
the Cordillera de la Costa, Cordillera de los Andes
and Zulia state, from 60 to 1,300 m.
Specimens examined. Venezuela. FALC6n: Depto. Sil¬
va, cerro Chichiriviche, laderas Sur, entre La Luisa y La
Soledad, Steyermark 110908 (VEN); Sierra de San Luis,
entre La Negrita y La Chapa, Steyermark 99159 (MO,
VEN); lado sur del rio Tocuyo, Steyermark 94483 (VEN);
Depto. Acosta, Municipio Jacura, Cerro de La Mina, Ruiz
652 (MER); Depto. Colina, El Paramito, Ruiz 964 (VEN);
Depto. Zamora, Cerro Santa Isabel, Ruiz 1825 (VEN);
Meachiche, Lasser 188 (VEN); Zamora, Cerro Mampos-
tal, Gonzalez 997 (VEN); 20 km de Puerto Cumarebo,
Ewel 1 (MY); N de Sanare, Ferrari 922 (MY); Paraguana,
1.5 km de Pueblo Nuevo, via a Santa Ana, Breteler 4393
(VEN); Depto. Democracia, entre Cerro Pozo Azul y Cerro
Maporal, Agostini 1028 (VEN); Depto. Democracia, al¬
rededor de Burnica, quebrada Araguato, Agostini 1023
(VEN). LARA: Carretera Barquisimeto- Aroa, 15 km NW
de Duaca, quebrada de Oro, Agostini 2549 (VEN); Si-
carigua, Burkart 16627 (VEN); entre Yaritagua y Duaca,
Volume 2, Number 2
1992
Velazquez
Brownea coccinea subsp. capitella
175
Saer 339 (VEN). m£rida: carretera Panamericana, cano
Zancudo, Bernardi 2178 (MER); El Vigia, Little 15227
(VEN); Depto. Miranda, Municipio Palmira, Arapuey,
Lopez-P. 3391 (MER); carretera de penetracion Los Gi-
ros Panamericana, Quintero 1569 (MER); Rio Perdido,
a 7 km de la carretera Panamericana, T. Ruiz 571
(MER); debajo Mesa Bolivar, Steyermark 56690 (VEN).
tachira: Urena, Aguas Calientes, Tamayo 3692 (VEN).
Trujillo: entre Agua Viva y Carora, 20 km NW de Agua
Viva, Bunting 2820 (MO, MY); cerca de El Cacao,
saliendo de Cuicas, Trujillo 1150 (MY); Sabana de Men¬
doza, caserio San Alejo, Quintero 1494 (MER); entre
Motatan y Agua Viva, Tamayo 1917 (VEN). yaracuy:
selvas de Yurnare, Bernardi 6938 (MER); 7 km W de
Guayabo, cerca de La Carbonera, Steyermark 94463
(VEN); San Felipe, Chiguara, Jahn 648 (VEN); camino
al Fundo El Chorro, Aristeguieta 1236 (VEN). ZULIA:
Cano Maria, 15 km N de El Vigia, Breteler 4421 (MER);
cerca de Mene Grande, Pittier 10614 (VEN); selvas del
Rio Lora, Pittier 10920 (VEN).
Acknowledgments. This work was made possible in
part thanks to the support of the Decanto de Investigacion
y Desarrollo at the Universidad Simon Bolivar, Caracas,
Venezuela. I thank Paul Berry for his assistance and
revisions to the original manuscript.
Literature Cited
Velasquez, D. & G. Agostini. 1981. Dos nuevas especies
de Brownea (Leguminosae-Caesalpinioideae). Ern-
stia 5: 1-13.
Klitgaard B. B. 1991. Ecuadorian Brownea and Brow-
neopsis (Leguminosae-Caesalpinioideae): taxonomy,
palynology, and morphology. Nordic J. Bot. 11: 433-
449.
A Further Note on Myrsine perpauciflora Pipoly (Myrsinaceae)
John J. Pipoly III
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT. Myrsine perpauciflora Pipoly is vali¬
dated by the designation of a holotype.
through an inadvertent error, a holotype was not
designated for Myrsine perpauciflora Pipoly, No-
von 1: 207. 1991. According to ICBN Art. 45.1,
“When the various conditions for valid publication
are not simultaneously fulfilled, the date [of valid
publication] is that on which the last is fulfilled. . . .
A name published on or after 1 Jan. 1973 for which
the various conditions for valid publication are not
simultaneously fulfilled is not validly published unless
a full and direct reference (Art. 33.2) is given to
the places where these requirements were previously
fulfilled.” Therefore, to fulfill the requirements ol
valid publication for this name, as fully and directly
referenced above, I designate the holotype as fol¬
lows:
Myrsine perpauciflora Pipoly, sp. nov. I YPE:
Venezuela. Territorio Federal Amazonas: Cerro
de la Neblina, Camp VII, 5 km NE of Pico
Phelps, 00°50'40"N, 65°58'10"W, 1,850 m,
1 Feb. 1985 (fr), M. Nee 30697 (holotype,
VEN; isotypes, NY, US). Figure 3.
Acknowledgments. I thank John Pruski of NY
for kindly pointing out this error, and Dan Nicolson
of US for discussing the correction with me.
Novon 2: 176. 1992.
A Further Note on Justicia parguazensis Wasshausen (Acanthaceae)
Dieter C. Wasshausen
Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.
ABSTRACT. Justicia parguazensis Wasshausen is
the correct name in Justicia for Chaetochlamys
wurdackii Leonard.
Paul Berry (MO) has pointed out that 1 inadver¬
tently added an extra “a” to the new name I pub¬
lished for Chaetochlamys wurdackii Leonard in
Novon 2: 64, 1992.
The epithet I chose is based on the name of the
river known as Rio Parguaza, spelled incorrectly
in the protologue as “Paraguaza," and the name of
the species should be Justicia parguazensis.
1 thank Paul Berry for bringing this to my at¬
tention.
Novon 2: 177. 1992.
Volume 2, Number 1, pp. 1-80 of NOVON was published on 18 March 1992.
Volume 2
Number 3
1992
NOVON Ny
A New Species of Solarium Section Micracantha (Solanaceae) from
Northeastern Brazil
Maria dc Fatima Agra
Laboratorio de Tecnologia Farmaceutica, Universidade Federal da Paraiba,
Caixa Postal 5009, Cidade Universitaria, Joao Pessoa, PB, Brazil
Abstract. Solarium paraibanum is a new species
from the wet forest of northeastern Brazil; it is found
only in Joao Pessoa, State of Paraiha. The new taxon
is described and illustrated, and its closest relation¬
ships are discussed.
A distinct species of the scandent and prickly
section Mi cracantha Dunal of Solarium L. was not¬
ed from recent collections made by the author in
Joao Pessoa. State of Paraiba, and is described here.
It is apparently rare and has been collected only
three times. Its closest relationships are with a small
group of species of this section restricted to the
northeast of Brazil.
Solanum paraibanum Agra, sp. nov. TYPE: Bra¬
zil. Paraiba: municipio de Joao Pessoa, Cidade
Universitaria, 6 km Sudeste do centro, 7°06'S,
34°53'W, 30 45 m, 10 nov. 1989, Agra 1101
(holotype, JPB; isotypes, PEUFR, MO). Fig¬
ure 1 ,
Frutex scandens. Folia lanceolata sinuato-angulata,
breviter petiolata, scabra supra strigosa, subtus pilis stel-
latis pulverulento-punctata. Inflorescentia scorpiodeo-cy-
inosa, simplex, interaxillaris; pedicello 1.0-1.2 cm longo;
calycis laciniis 0.8-1.0 cm longis, lineari-lanceolatis re-
volutis; corolla violacea, profunde lobata, extus stellato-
tomentosa.
Scandent shrub 4-10 in high, with a fuscous
pubescence, armed with short, yellow, recurved
prickles of various sizes up to 0.4 cm. Stem terete,
dark brown hispid-pubescent with mostly sessile pau-
ciradiate stellae bearing greatly prolonged 3-4-celled
midpoints, sparsely armed with small, broad-based.
ferruginous prickles to 0.2-0.3 cm that are re¬
curved at the apex. Leaves petiolate; petiole terete,
to 1-3.5 cm long, stellate-scabrous, the hairs and
prickles similar to those of the stems, but more
compact; leaf blades chartaceous, ovate-lanceolate,
the larger 8-15 cm long, 3.5-8.0 cm broad, acute
at apex, broadly cuneate at base, usually sinuate-
lobed, the lobes 3-4 pairs, acute, above stellate-
hispid, the hairs sessile with greatly prolonged mid¬
points, sparsely prickly on the principal veins, be¬
neath stellate-pubescent, the hairs sessile, porrect-
stellate, 6 8-celled; sparsely armed on the midrib
or minor veins, the prickles yellowish, short and
recurved at apex. Inflorescences lateral, extra-ax-
illary monochasial, unbranched, remote from the
leaves, some of the distal flowers with reduced gy-
noecia; pedicels 0.6-1.0 cm long, becoming longer
and stouter in fruit, stellate-pubescent and prickly
with hairs similar to those of the branches and leaves
and small, yellow prickles, ca. 0.1-0.15 cm at an-
thesis, enlarged in fruit, ca. 2-2.5 cm long, 0.3-
0.5 cm wide, with small prickles and pubescence
similar to those of peduncle and branches. Flowers
heterostylous, the basal (two or more) with elongate
and functional styles ca. 1.2-1.5 cm long, exceeding
the anthers, broad and glabrous to a clavate stigma;
the distal flowers with short thick and abortive style
and stigma. Calyx densely stellate-pubescent, armed
with short, acute, yellow prickles, deeply lobed about
44 the length, the tube ca. 0.1 cm, the lobes linear-
lanceolate and revolute. Corolla blue or violet, deep¬
ly parted, the lobes gradually recurved, narrowly
lanceolate, 1.0-1.5 cm long, 0.3-0.4 cm wide above
the tube, the tube short, 0.1 -0.2 cm long, densely
Novon 2: 179-181. 1992.
180
Novon
E
Figure 1. Solarium paraibanum Agra (drawing from M. F. Agra 1101). —A. Habit. —B. Bud. —C. Staminate
flower. —D. Fruit. —E. Leaf, stellate hair, upper surface. —F. Leaf, stellate hair, lower surface.
stellate-pubescent, the hairs sessile with fully de¬
veloped midpoints, sometimes with lateral rays want¬
ing and the hairs apparently simple and stiff. Fil¬
aments 0.2-0.3 cm long, 0.1 0.2 cm wide, glabrous.
Anthers equal, oblong-linear, 1.0-1.2 cm long, 0.2-
0.3 cm wide at base, cordate at base, attenuate to
apex, the pores minute. Ovary densely stellate-pu¬
bescent. Berry red-orange, ca. 2.5 3 cm diam.,
Volume 2, Number 3
1992
Agra
Solanum paraibanum
181
densely hirsute until maturity; the hairs consisting
of prolonged uniseriate midpoints; seeds numerous,
black, lenticular, 0.40-0.45 cm across.
Paratypes. Brazil, paraiba: Joao Pessoa, Campus 1
da Universidade Federal da Paraiba, Mourn 10 (JPB),
Agra 448 (JPB, BB).
Solanum paraibanum is known only from the
type locality, in the Atlantic Coastal Forest in Pa¬
raiba, in wet forest, at elevations near sea level.
This vegetation includes floristic elements related to
the Amazon forest and consists mainly of tall trees,
many of large diameter. The rainfall of this area is
strongly seasonal, with most rains falling between
May and October; it is only during these months
that S. paraibanum flowers.
Solanum paraibanum is related to S. rupincola
Sendtner and S. rupincola var. piauhyensis Dunal,
both members of section Micracantha sensu D’Arcy
(1973), characterized by glabrous and pubescent
berries. It is distinguished from those species by the
smaller size of the calyx, corolla, and fruit, the shape
of the linear-lanceolate and revolute calyx lobes,
and the more scabrous pubescence. These and other
unnamed species are restricted to the northeast of
Brazil and constitute a small, closely related group
of Micracantha that is characterized by large ber¬
ries that are densely hirsute until maturity, bearing
sessile stellate trichomes with prolonged midpoints.
Acknowledgments. I am grateful to Enrique For-
ero for facilities at the Missouri Botanical Garden
and for reading the manuscript; Geraldo Mariz for
encouragement and help with the Latin diagnosis;
William G. D'Arcy for corrections to the manuscript
and for many helpful suggestions; and Delby F.
Medeiros, Director, Laboratorio de Tecnologia Far-
maceutica and Jnanabrata Bhattacharyya for en¬
thusiasm and encouragement in the study of So¬
larium. Madson H. Sousa provided the illustration
of the type. Research at MO was funded in part by
the l inker Foundation.
Literature Cited
D’Arcy, W. G. 1973. Solanaceae./a. Flora of Panama.
Ann. Missouri Bot. Card. 60: 573-780.
A New Lowland Species of Tropaeolum (Tropaeolaceae) from the
Venezuelan Guayana
Paul E. Perry
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, IJ.S.A.
ABSTRACT. Tropaeolum orinocense from southern
Venezuela, the first species in the genus recorded
from the upper Orinoco basin, is described.
Tropaeolum is a genus of southern temperate
origin, with about 90 species native to South and
Central America. The tropical and subtropical spe¬
cies are restricted to cool, montane areas, especially
m the Andes, except for a few species that enter
into coastal lowland Ecuador and Peru. In the most
recent monograph of the family (Sparre & Anders-
son, 1991), no species was reported to occur in
lowland Amazonia or Guayana.
In 1951, Leon Croizat participated in the Franco-
Venezuelan expedition to the headwaters of the Ori¬
noco River and made over 1,000 plant collections.
One of these, Croizat 790, was a Tropaeolum grow¬
ing on the banks of the upper Orinoco at about 300
m elevation. It was reported by Holst & Todzia
(1990) as T. fintelmannii Wagener ex Schldl., a
species otherwise known from the Coastal Cordillera
of Venezuela and the Andes of Colombia, Ecuador,
and Peru. In 1989, as part of a Venezuelan group
retracing the previous expedition to the Orinoco
headwaters, I found the same Tropaeolum at a single
locality along the Orinoco above the mouth of the
Hio Ugueto, probably at the same site where Croizat
collected it previously. Study of pickled flowers and
additional dried specimens showed significant dif¬
ferences from T. fintelmannii and other related spe¬
cies of Tropaeolum. Consequently, the Orinoco col¬
lections are described as a new species and the only
one known from Amazonian South America.
Tropaeolum orinocense P. Berry, sp. nov. TYPE:
Venezuela. Territorio Federal Amazonas: Rio
Orinoco below Raudal and Salto El Tobogan,
above the mouth of Rio llgueto, 2°14'N,
63°45'W, elevation 370 m, 21 Nov. 1989,
Berry 4783 (holotype, MO; isotypes, MYF,
TFAV, VEN). Figure I.
Species haec Tropaeolo repando Heilborn affinis, sed
floribus minoribus, calcari 24-27 mm longo, petiolo a
basi folii 3-5 mm inserto, foliis manifeste quinquelobatis
differt.
Slender climbing herb to 5 m long, stem glabrous
to lightly strigillose. Leaves peltate, insertion index
(ratio of the length of the longitudinal axis of the
blade distal to the point of petiole insertion vs. the
proximal part) 9:1, stipules lanceolate, ca. 1 mm
long; petiole 5—7(—11) cm long, glabrous to strigil¬
lose, inserted 3-5 mm from base of leaf; blade thin-
membranous, glabrous, light green on upper surface,
glaucous on lower surface, ± depressed ovate, 3.2-
5.5 cm long, 4.5-9.0 cm wide, the length:width
ratio usually 2: 3 or less often 1 : 2, truncate at the
base, shallowly 5-lobed, the central 3 lobes shortly
mucronate, principal ascending veins 3 from point
of petiole insertion, the lateral ones dichotomously
branched close to the base. Flowers orange-red, with
slender, pendent pedicels 45-75 mm long. Calyx
lobes elliptic, the uppermost lobe 5 x 4 mm, the
lowermost one 9-10 x 5-6 mm, with a narrowly
conical spur either straight or slightly downcurved
in distal Vi to l A, the spur 24-27 mm long and 5-
6 mm diam. at point of pedicel insertion. Petals
ciliate-dentate at the apex, the cilia 2-2.5 mm long,
the upper two petals (opposite the pedicel) cuneate,
5-6 x 3-4 mm (excluding the cilia), the lower three
spathulate-unguiculate, the blade 2.5-3.5 x 3-4
mm, somewhat concave, the claw smooth and 5-6
mm long. Stamens 8, slightly uneven, filaments 4-
6 mm long, anthers 2 x 1 mm. Ovary 3-angled;
style 2-3 mm long, trifid at apex, one lobe longer
than the other two. Mericarps 1 (—3?), ca. 5x3
mm, brown.
Paratypes. VENEZUELA. AMAZONAS: Ugueto, upper Rio
Orinoco, Croizat 790 (VEN).
Morphologically, this species strongly resembles
Tropaeolum repandum Heilborn, a lowland species
from coastal Ecuador and northern Peru. The two
are very similar in the apical position of the petal
cilia and in the shape of the upper and lower petals;
the depressed ovate to reniform leaf shape is also
common to both species. Tropaeolum orinocense
differs, however, in its smaller flowers and the leaves
with a higher petiole insertion ratio and a higher
length to width ratio. Details of petal coloration, a
useful character at the species and sectional level,
require further observation of fresh flowers.
Novon 2: 182-184. 1992.
Volume 2, Number 3
1992
Berry
Tropaeolum
183
Figure 1. Tropaeolum orinocense P. Berry, drawn from Berry 4783. —A. Habit. B. Flower, with sepals and
petals partially removed to reveal the staminal arrangement. —C. Upper petal. —D. Lower petal. E. Gynoecium
at anthesis. —F. Single mericarp.
Tropaeolum orinocense is also similar in leaf shape
and apically serrate-ciliate petals to T. lasseri Sparre,
from northeastern Venezuela, and T. emarginatum
Turcz., from northeastern Colombia. The flowers of
T. lasseri are much smaller, however, and T. emar¬
ginatum has leaves with the petiole inserted much
farther from the margin and petals with yellow and
purple coloration.
Specimens of Tropaeolum orinocense were pre¬
viously determined as T. fintelmannii, a species from
northern Venezuela and Andean Colombia to Peru,
but that species has a shorter calyx tube, the upper
petals with fewer apical cilia, and the lower petals
larger, more elliptic, and serrate-ciliate along the
entire blade (not just the apex).
Sparre & Andersson (1991) established ten sec¬
tions in Tropaeolum in their monograph of the fam¬
ily. They placed T. repandum in section Serrato-
ciliata, a group of mostly red-flowered species
centered in the northwestern Andes, whereas T.
emarginatum, T. lasseri, and T. fintelmannii were
placed in section Tropaeolum, a wide-ranging group
from southern Mexico to Peru. There is no clear
delimitation between these two sections, however,
which leaves the sectional placement of T. orino¬
cense in doubt. Pollen morphology favors its place¬
ment in section Tropaeolum, since it has type A
grains (equilateral triangle in polar view, with three
identical pores), following the classification of pollen
types in the family by Ricardi et al. (1957) and later
modified by Huynh (1968). Type A grains occur in
section Tropaeolum, but not in section Serratoci-
liata, which has type B grains (isosceles triangle in
polar view, with one smaller pore), as in 7. repan¬
dum, or else type D grains (biaperturate and elon¬
gate).
Acknowledgments. I gratefully acknowledge the
support of the U.S. National Science Foundation
Grant BSR-8717303 for study on the Flora of the
Venezuelan Guayana and Radio Caracas Televi-
sion/CORAVEN in Caracas, Venezuela, for inviting
184
Novon
me to participate in their 1989 expedition to the
upper Orinoco. Bruno Manara made the illustration,
kindly reviewed the manuscript, and corrected the
Latin diagnosis. I also thank Lennart Andersson,
one of the reviewers, for helpful comments.
Literature Cited
Holst, B. K. & C. A. Todzia. 1990. Leon Croizat's
plant collections from the Franco-Venezuelan expe¬
dition to the headwaters of the Rio Orinoco. Ann.
Missouri Bot. Card. 77: 485-516.
Huynh, K. 1968. Morphologie du pollen des tropaeo-
lacees et des balsaminacees. 1. Grana Palynol. 8: 88-
184.
Ricardi, M., C. Marticorena & F. Torres. 1957. Nota
preliminar sobre la morfologia de los polenes de Tro-
paeolaceae chilenas. Bol. Soc. Biol. Concepcion 32:
17-19.
Sparre, B. & L. Andersson. 1991. A taxonomic revision
of the Tropaeolaceae. Opera Bot. 108: 1-140.
Nomenclatural Changes of Some Californian
Castilleja (Scrophulariaceae)
7. I. Chuang
Department of Biological Sciences, Illinois State University, Normal, Illinois 61761, U.S.A.
L. R. Heckardf
Jepson Herbarium, University of California, Berkeley, California 94720, U.S.A.
ABSTRACT. In anticipation of the upcoming revi¬
sion of the Jepson Manual of the Flowering Plants
of California, the following 10 new nomenclatural
combinations of Californian Castilleja subg. Cas¬
tilleja at the subspecific level are proposed: C. affinis
subsp. litoralis (Pennell) Chuang & Heckard, C.
affinis subsp. neglecta (Zeile) Chuang & Heckard,
C. applegatei subsp. disticha (Eastwood) Chuang
& Heckard, C. applegatei subsp. martinii (Abrams)
Chuang & Heckard, C. applegatei subsp. pallida
(Eastwood) Chuang & Heckard, C. applegatei subsp.
pinetorurn (Fernald) Chuang & Heckard, C. his-
pida subsp. brevilobata (Piper) Chuang & Heckard,
C. lanata subsp. hololeuca (Greene) Chuang &
Heckard, C. minor subsp. spiralis (Jepson) Chuang
& Heckard, and C. subinclusa subsp. franciscana
(Pennell) Chuang & Heckard. The justification for
these nomenclatural changes is briefly summarized.
The genus Castilleja, especially subgenus Cas¬
tilleja, is notorious for its taxonomic difficulty, which
is due at least partly to natural hybridization and
associated polyploidy. We recently proposed to sub¬
divide the expanded genus Castilleja into three sub¬
genera (Chuang & Heckard, 1991): suhgenus Co-
lac us, with three sections, Oncorhynchus, Pilosae,
and Pallescentes, to include bee-pollinated plants
with bracts and calyx colored other than red and a
relatively well-developed lower corolla lip, usually
with some pouch development; subgenus Gentrya,
a monotypic taxon from the Sierra Surutato of north¬
ern Sinaloa, Mexico, which exhibits a distinctive
curved corolla and is probably also bee-pollinated;
and subgenus Castilleja, comprising the majority of
the species, which have a flower basically modified
for hummingbird pollination, with bract and calyx
tips predominantly red, a well-developed corolla tube,
an upper lip much longer than the lower, and a
lower lip reduced to three small teeth and lacking
any pouch.
The purpose of this paper is to make appropriate
nomenclatural changes in Californian Castilleja subg.
Castilleja in anticipation of the upcoming revision
of the Jepson Manual of the Flowering Plants of
California (Jepson, 1925). The following 10 new
combinations are proposed at the subspecific level.
This study is based on field observations and col¬
lections of Castilleja in California during the past
two decades and abundant herbarium specimens de¬
posited at CAS, JEPS, and UC.
1. Castilleja affinis Hooker & Arnott subsp. li¬
toralis (Pennell) Chuang & Heckard, comb,
nov. Castilleja litoralis Pennell, Proc. Acad.
Nat. Sci. Philadelphia 99: 183. 1947. Castil¬
leja wightii Elmer subsp. litoralis (Pennell)
Munz, Aliso 44: 98. 1958. TYPE: U.S.A. Or¬
egon: Coos County, Bandon. 19 July 1931,
Pennell 15651 (holotype, PH).
2. Castilleja affinis Hooker & Arnott subsp. neg¬
lecta (Zeile) Chuang & Heckard, comb, et stat.
nov. Castilleja neglecta Zeile in Jepson, Man.
FI. PI. Calif. 936. 1925. TYPE: U.S.A. Cali¬
fornia: Marin County, Tiburon, 7 July 1907,
Hrandegee s.n. (holotype, JEPS).
Castilleja affinis is a highly intractable polyploid
complex, which has six levels of polyploidy, ranging
from n = 12 to 72 (Heckard, 1968; Chuang &
Heckard, unpublished data). Pennell (1951) rec¬
ognized six species in the C. affinis complex. These
species (C. affinis, C. californiea Abrams, C. doug-
lasii Bentham, C. inflata Pennell, C. litoralis, and
C. neglecta) were placed in four different sections
of the genus on the basis of such morphological
features as shape of leaf and calyx lobes, and the
pubescence of the upper corolla lip (beak). There is
too much variability and overlap in these features
within the group to delimit satisfactorily distinct
species. Some populations might warrant subspecific
recognition but do not necessarily coincide with poly¬
ploidy levels, and various polyploidy levels inter¬
grade to such an extent that it is impossible to make
a useful taxonomic separation. Furthermore, Cas-
Novon 2: 185-189. 1992.
186
Novon
tilleja affinis itself is not sharply delimited from
neighboring species, either because of its probable
partial origin from them in the past or because of
more recent hybridizations with them. Therefore,
taxonomic recognition of any entities other than the
three subspecies, subspecies affinis , subspecies li-
toralis, and subspecies neglecta, seems impractical.
The taxonomic difficulties of the C. affinis complex
are comparable to those in the Californian compo¬
nent of such other complex species as Dactylis
glomerata Linn. (Stebbins & Zohary, 1959), Am¬
brosia dumosa (A. Cray ex Torr.) Payne (Haven et
al., 1968), Eriophyllum lanatum (Pursh) Forbes
(Mooring, 1975), and Epilobium [ Zauschneria ]
canum (Greene) Haven (Haven, 1977).
Castilleja affinis subsp. affinis (n = 12, 24, 36,
48) is widespread over much of cismontane Cali¬
fornia at lower elevations in sandy and rocky soil
in chaparral and coastal scrub. It is generally char¬
acterized by bristly puberulent herbage, 3- to 5-
lobed leaves, acute bract and calyx lobes, and larger
(25-40 mm), scarlet to orange-red flowers. A coast¬
al-bluff form (Pt. Heyes Peninsula south to northern
Santa Cruz) with rather fleshy leaves, distally inflated
calyx, and less protruding corolla beak has been
named C. inflata (= C. ivightii subsp. infiata (Pen¬
nell) Munz). Another coastal sand-dune form (San
Luis Obispo and Santa Barbara counties) has vari¬
able branched hairs in the herbage; this form has
been named C. affinis var. contentiosa (J. F. Mac-
bride) Bacigalupi, which perhaps represents a past
introgressant between affinis and C. mollis. Cas¬
tilleja affinis subsp. litoralis {n = 48, 60, 72) is
distributed on coastal bluffs from northern California
to northern Oregon. This subspecies is distinguish¬
able by its usually glabrous herbage, entire leaves,
obtuse to rounded bract and calyx lobes, and larger
(25-40 mm), scarlet to orange-red flowers. Castil¬
leja affinis subsp. neglecta (n — 36) is a rare ser¬
pentine endemic found only on Tiburon Peninsula
and south of Nicasio Reservoir in Marin County and
American Canyon in Solano County, and is char¬
acterized by its bristly puberulent herbage, 3- to 5-
lobed leaves, acute bract and calyx lobes, and small¬
er (18-22 mm), yellow flowers.
3. Castilleja applegatei Fernald subsp. disticha
(Eastwood) Chuang & Heckard, comb, et stat.
nov. Castilleja disticha Eastwood, Proc. Calif.
Acad. Sci. III. 2: 289. 1902. TYPE: U.S.A.
California: Fresno County, Converse Basin,
South Fork of King’s River, 1-3 July 1899,
Eastwood s.n. (holotype, CAS).
Castilleja quibellii Beane, Contr. Dudley Herb. 4: 37.
1950. TYPE: LI.S.A. California: Fresno County,
Rancheria Camp Ground, 11 July 1949, Beane
1536 (holotype, DS; isotype, JEPS).
4. Castilleja applegatei Fernald subsp. martinii
(Abrams) Chuang & Heckard, comb, et stat.
nov. Castilleja martinii Abrams, Bull. S. Calif.
Acad. Sci. 1: 69. 1902. TYPE: U.S.A. Cali¬
fornia: Los Angeles County, Wilsons Peak, San
Gabriel Mountains, 10 July 1901, Abrams IBBI
(holotype, DS; isotype, DS).
Castilleja clokeyi Pennell, Proc. Acad. Nat. Sci. Phila¬
delphia 89: 420. 1937. Castilleja martinii var.
clokeyi (Pennell) N. Holmgren, Mem. New York
Bot. Card. 21: 55. 1971. TYPE: U.S.A. Nevada:
Clark County, Charleston Mountains, 8 July 1936,
Clokey & Clokey 7322 (holotype, PH; isotypes,
BRY, CAS, DS, MO, NY, ORE, OSC, RSA, SD,
UC, US, UTC, WTU).
Castilleja roseana Eastwood, Leaf!. W. Bot. 2: 104.
1938. TYPE: U.S.A. California: Monterey County,
between San Lucas and Priest V alley, 11 May 1936,
Eastwood & Howell 2460 { holotype, CAS; isotypes,
CAS, GH, RSA).
Castilleja hoffmannii Eastwood, Leaf!. W. Bot. 3: 116.
1942. TYPE: U.S.A. California: Ventura County,
between Chula Vista Camp and summit of Mount
Pinos, 26 May 1928, Howell 3840 (holotype, CAS).
Castilleja gyroloba Pennell, Proc. Acad. Nat. Sci. Phil¬
adelphia 99: 186. 1947. TYPE: U.S.A. California:
Los Angeles County, near Oak Flat Camp, 18 Apr.
1940, Pennell 25140 (holotype, PH; isotypes, DS,
GH, NY, PH).
Castilleja montigena Heckard, Syst. Bot. 5: 83. 1980.
TYPE: U.S.A. California: San Bernardino County,
N of Baldwin Lake, San Bernardino Mountains, 12
June 1976, Heckard & Morris 4240 (holotype,
JEPS; isotypes, NY, RSA, WTU).
5. Castilleja applegatei Fernald subsp. pallida
(Eastwood) Chuang & Heckard, comb. nov.
Basionym: Castilleja breweri var. pallida
Eastwood, Leafl. W. Bot. 2: 284. 1940. Cas¬
tilleja glandulifera subsp. pallida (Eastwood)
Pennell, in Abrams, Illust. FI. Pac. States 3:
832. 1951. Castilleja applegatei var. pallida
(Eastwood) N. Holmgren, Mem. New York Bot.
Card. 21: 37. 1971. TYPE: U.S.A. California:
Alpine County, Carson Pass, 17 June 1940,
Eastwood & Howell B449 (holotype, CAS; iso¬
type, CAS).
Castilleja breweri Fernald, Erythea 6: 49. 1898. Cas¬
tilleja applegatei var. breweri (Fernald) N. Holm¬
gren in Cronquist et ah, Intermountain FI. 4: 486.
1984. TYPE: U.S.A. California: Tuolumne County,
Mount Dana, 29 June 1863, Brewer 1744 (holo¬
type, GH; isotypes, UC, US).
Castilleja adenophora Eastwood, Leafl. W. Bot. 3: 87.
1941. TYPE: U.S.A. California: Inyo County, Mono
Pass Trail, 22 July 1941, Howell 16400 (holotype,
CAS; isotypes, CAS, GH, PH, US).
Volume 2, Number 3
1992
Chuang & Heckard
Californian Castilleja
187
6. Castilleja applegatei Fernald subsp. pine-
torum (Fernald) Chuang & Heckard, comb,
et stat. nov. Castilleja pinetorum Fernald, Er-
ythea 6: 50. 1898. Castilleja applegatei var.
pinetorum (Fernald) N. Holmgren in Cronquist
et al., Intermountain FI. 4: 486. 1984. TYPE:
U.S.A. Oregon: Klamath County, Swan Lake
Valley, 30 June 1896, Applegate 415 (holo-
type, GH; isotypes, I)S, PH).
Castilleja brooksii Eastwood, Proc. Calif. Acad. Sci. III.
2: 288. 1902. TYPE: U.S.A. California: Fresno
County, Bubba Creek of King’s River, 1-13 July
1899, Eastwood s.n. (holotype, CAS).
Castilleja trisecta Greene, Leaf!. Bot. Observ. Crit. 1:
78. 1904. TYPE: U.S.A. California: Tulare County,
Hockett’s Meadow, 18 July 1904, Baker 4431 (ho¬
lotype, NDG; isotypes, CAS, GH, NY, RSA).
Castilleja pinetorum var. fragilis Zeile in Jepson, Man.
FI. PI. Calif. 938. 1925. Castilleja fragilis East-
wood ex C. F. Baker, W. Ainer. Pis. 3: 4. 1904,
nomen nudum. Castilleja applegatei var. fragilis
(Zeile) N. Holmgren, Mem. New York Bot. Card.
21: 38. 1971. TYPE: U.S.A. California: Siskiyou
County, Sisson, 14 Aug. 1903, Copeland 3883
(holotype, CAS; isotypes, DS, GH, JEPS, MO, NY,
RSA; distributed as Castilleja fragilis).
Castilleja excelsa Eastwood, Lead. W. Bot. 2: 241. 1940.
TYPE: U.S.A. California: Siskiyou County, near Spirit
Lake, Marble Mountains, 4 Aug. 1939, Howell
15058 (holotype, CAS).
Castilleja dolichostylis Eastwood, Lead. W. Bot. 3: 88.
1941. TYPE: U.S.A. California: Tehama County,
near Government Flat, 9 July 1941, Eastwood &
Howell 9837 (holotype, CAS; isotypes, CAS, GH,
PH, US).
Castilleja wherryana Pennell, Proc. Acad. Nat. Sci. Phil¬
adelphia 99: 180. 1947. TYPE: LJ.S.A. Oregon:
Baker County, Dooley Mountain, 4 July 1931, Pen¬
nell 15454 (holotype, PH; isotypes, MO, NY, US).
Castilleja latifoliata Pennell ex Edwin, Lead. W. Bot.
9: 46. 1959. TYPE: U.S.A. Nevada: Washoe Coun¬
ty, S of Mount Ross, 24 July 1940, Pennell 26267
(holotype, PH; isotypes, CAS, UT, UTC).
Castilleja applegatei is one of the most polytypic
species of Californian Castilleja, but it can be dis¬
tinguished from all others by its glandular-viscid
herbage and usually wavy-margined leaves. Pennell
(1951) recognized six species (C. applegatei, C.
breiveri, C. disticha, C. ewanii Eastwood, C. gy-
roloba, and C. martinii) in two sections of this
complex, while Munz & Keck (1959) listed five (C.
applegatei, C. breweri, C. disticha, C. martinii,
and C. roseana). In his taxonomic revision ol the
C. viscidula group, Holmgren (1971) treated Cal¬
ifornian members of the C. applegatei complex as
comprising C. applegatei (with two varieties, var.
fragilis and var. pallida ), C. disticha, and C. mar¬
tinii (with three varieties, var. clokeyi, var. ewanii,
and var. martinii). Later, Heckard et al. (1980)
added C. montigena from the San Bernardino
Mountains of southern California, an allopolyploid
(n = 24, 36) derived from diploid races (n = 12)
of C. chromosa A. Nelson (= C. angustifolia (Nutt.)
G. Don) and C. martinii var. martinii. The binomial
C. ewanii has been misapplied to polyploid C. mon¬
tigena, because the type specimen of C. ewanii (n
= 12) falls within the diploid introgressants, and that
name was consequently placed in synonymy under
C. chromosa (Heckard et ah, 1980).
A sufficient number of chromosome counts (Heck¬
ard, 1968; Chuang & Heckard, unpublished data)
have been made in this group to show that consid¬
erable polyploidy (n = 12, 24, 36, 48) is present.
The C. applegatei complex is extremely difficult
taxonomically, and the complex pattern of variation
has resulted in diverse treatments by earlier workers,
as indicated above. The key characters used by
Pennell (1951), Munz & Keck (1959), and Holm¬
gren (1971) to identify members of this group are
such differences as height of plant, degree of glan¬
dular puberulence in the herbage, leaf shape, length
of corolla, and lengths of upper and lower corolla
lips. In most instances, their keys allow for over¬
lapping variation, which suggests taxonomic diffi¬
culty. It thus appears preferable to consider C. ap¬
plegatei to be a single polytypic species. We propose
the recognition of subspecies disticha, subspecies
martinii, subspecies pallida, and subspecies pine¬
torum for the Californian members of this complex.
These four subspecies can be separated imperfectly
by the following key:
Key to the subspecies of Castilleja applegatei
la. Leaves mostly 3-lobed; calyx 13 15 mm, di¬
vided ca. Vi on the sides; subalpine in high Sierra
Nevada .subsp. pallida
lb. Leaves mostly entire; calyx 12-22 mm, gen¬
erally divided less than V b on the sides (except
some subsp. pinetorum)', often below subalpine.
2a. Calyx lobes usually obtuse to rounded; cen¬
tral and southern California . . subsp. martinii
2b. Calyx lobes usually acute.
3a. Calyx 12-18 mm; central and south¬
ern Sierra Nevada .subsp. disticha
3b. Calyx 16-22 mm; widespread in
northern California . . . subsp. pinetorum
7. Castilleja hispida Bentham subsp. brevilo-
bata (Piper) Chuang & Heckard, comb, et stat.
nov. Castilleja brevilobata Piper, Proc. Biol.
Sci. Wash. 33: 104. 1920. TYPE: U.S.A. Or¬
egon: Josephine County, 8 mi. S of Waldo, 14
June 1904, Piper 6118 (holotype, US).
According to Ownbey (1959), Castilleja hispida
is a common, complex, and variable species most
closely related to C. angustifolia (= C. chromosa).
The diploid coastal plants of northern Oregon and
188
Novon
the tetraploid plants of the Rocky Mountains and
the Cascades of Washington are strikingly similar
(Heckard, 1968). This species apparently inter¬
grades with C. angustifolia and C. miniala where
their ranges juxtapose. For example, the polyploid
C. peckiana Pennell (n = 36, 48, 60; Heckard,
1968) is in fact a morphological intermediate be¬
tween C. hispida and C. miniata and was reduced
to synonymy under C. miniata (Ownbey, 1959).
When describing C. brevilobata. Piper (1920)
allied it with C. angustifolia, hut noted that all its
parts were smaller, its leaves were shorter-lobed,
and it possessed somewhat hispidulous pubescence.
Holmgren (1971) placed it in the C. viscidula al¬
liance because of its glandular pubescence, its some¬
what crisped-margined leaves, and its geographical
location. Some specimens of C. brevilobata are
strikingly similar in habit, leaf and bract shape, and
lobing pattern to C. hispida of coastal central and
northern Oregon and W ashington, except that the
former exhibits glandular puberulent herbage and
has a more restricted range confined to the corner
of northwestern California and southwestern Ore¬
gon. Because C. hispida is a highly polymorphic
species and none of the criteria used to separate the
two is absolute, and the variation in morphological
features within each group overlaps, we prefer to
treat C. brevilobata as a subspecies of C. hispida.
8. Castilleja lanata A. Cray subsp. hololeuca
(Greene) Chuang & Heckard, comb, et stat.
nov. Castilleja hololeuca Greene, Pittonia 1:
39. 1887. TYPE: U.S.A. California: Santa Bar¬
bara County, Island of San Miguel, Sep. 1886,
Greene s.n. (holotype, NDC; isotype, CAS).
Castilleja lanata and C. hololeuca, along with
C. foliolosa Hooker & Arnott and C. grisea Dunkle,
have been included in section Lanatae because of
their grayish or white tomentose herbage, branched
hairs, and calyx lobes, which are rounded or wholly
united laterally (Pennell, 1951). Castilleja lanata
is widespread from northern Mexico and western
Texas to southern Arizona, while C. hololeuca is
restricted to the northern Channel Islands of Cali¬
fornia. These two taxa can be distinguished by the
fact that Castilleja hololeuca has a smaller calyx
(15-18 mm) and corolla (20-25 mm). We propose
to reduce C. hololeuca to subspecific status under
C. lanata. Castilleja lanata differs from the closely
related C. Joliolosa and C. grisea by its white-woolly
felt of long, interwoven, slightly branched hairs, and
entire leaves.
9. Castilleja minor (A. Gray) A. Gray subsp.
spiralis (Jepson) Chuang & Heckard, comb.
nov. Basionym: Castilleja spiralis Jepson, FI.
W. Mid. Calif. 412. 1901. Castilleja sten-
antha subsp. spiralis (Jepson) Munz, Aliso 4:
98. 1958. TYPE: U.S.A. California: Napa
County, Butt’s Canyon, 13 July 1897, Jepson
21113 (holotype, JEPS).
Castilleja stenantha A. Gray, Syn. FI. N. Amer. 2: 295.
1878. TYPE: U.S.A. California: Hartweg 1897
(holotype, GH).
Castilleja minor, one of a few annual species in
subgenus Castilleja, is widely distributed in the west¬
ern United States and adjacent Mexico, growing in
wet places (usually alkaline), such as marshes,
streambanks, and valley hot springs. Customarily,
three or four species (C. minor , C. exilis A. Nelson,
C. spiralis, and C. stenantha) have been recognized
in this complex. They are recognizable only by minor
or very inconstant morphological features such as
differences in herbage indument, length of corolla,
shape of bracts, and color of lower corolla lip. Most
of these characters show overlapping variation (Pen¬
nell, 1951), suggesting taxonomic difficulty. It seems
preferable to treat this complex as a single variable
species with two subspecies: subspecies minor and
subspecies spiralis. Castilleja minor subsp. minor
(including C. exilis) is widespread in the western
United States east of the Sierra-Cascade crest and
in adjacent Mexico, and is distinguished by a shorter
corolla (15—20(—30) mm); in contrast, subspecies
spiralis (including C. stenantha) is found only in
cismontane California and has a longer corolla (25-
35 cm).
10. Castilleja subinclusa Greene subsp. fran-
ciscana (Pennell) Chuang & Heckard, comb,
et stat. nov. Castilleja franciscana Pennell,
Proc. Acad. Nat. Sci. Philadelphia 99: 188.
1947. TYPE: U.S.A. California: San Mateo
County, Crystal Springs Lake, 15 May 1940,
Pennell & Keck 25420 (holotype, PH).
Castilleja subinclusa is most closely related to
C. linariifolia Bentham. Both species share such
remarkable features as an unevenly cleft calyx and
a corolla generally curved forward through a calyx
sinus. The calyx divides more deeply in front (34)
than in back {V b ~ x A), with the lobes generally curved
upward. Pennell (1951) treated these two species
plus C. franciscana Pennell in section Linariaefol-
iae. Later, Bacigalupi & Heckard (1966) described
C. jepsonii, which is distributed in the inner South
Coast Ranges from southeastern San Benito County
and southward to the Sierra San Pedro Martir in
northern Baja California. They concluded that “The
new species is somewhat intermediate in taxonomic
Volume 2, Number 3
1992
Chuang & Heckard
Californian Castilleja
189
position as well as its geographic distribution between
C. affinis and C. linariifolia. A hybrid origin is a
possibility with the resulting hybrid derivative form¬
ing a stable self-perpetuating entity over a consid¬
erable geographic area.’’ In that treatment, Baci-
galupi & Heckard (1966) proposed to include C.
franciscana in C. subinclusa. The key characters
used to separate C. subinclusa and C. jepsonii com¬
prised such variable features as color and shape of
leaves, thickness of upper corolla lip, and color and
shape of bract and calyx. We prefer to regard C.
subinclusa as a highly variable species consisting of
two subspecies: subspecies subinclusa (including C.
jepsonii) and subspecies franciscana. The two sub¬
species can be somewhat arbitrarily distinguished by
the following key.
Key to the subspecies of Castilleja subinclusa
la. Corolla yellow-orange; calyx divided 2-4 mm
on the sides, lobes clearly curved upward; Cen¬
tral and North Coast Ranges from Santa Cruz
County north to Mendocino County .
.subsp. franciscana
lb. Corolla reddish; calyx divided 4-7 mm on the
sides, lobes barely curved upward; widespread
from South Coast Ranges south to northern Baja
California and central and southern Sierra Ne¬
vada foothills .subsp. subinclusa
Acknowledgments. We thank Lincoln Constance, James
C. Hickman, and David M. Thompson for critical reading
of the manuscript.
Literature Cited
Bacigalupi, R. & L. R. Heckard. 1966. A new Castil¬
leja from south-central and Baja California. Leaf!.
W. Bot. 10: 281-286.
Chuang, T. I. & L. R. Heckard. 1991. Generic realign¬
ment and synopsis of subtribe Castillejinae (Scroph-
ulariaceae — tribe Pediculareae). Syst. Bot. 16: 644-
666 .
Heckard, L. R. 1968. Chromosome numbers and poly¬
ploidy in Castilleja (Scrophulariaceae). Brittonia 20:
212-226.
-, M. I. Morris & T. I. Chuang. 1980. Origin
and taxonomy of Castilleja montigena. Syst. Bot.
5: 71-85.
Holmgren, N. H. 1971. A taxonomic revision of the
Castilleja viscidula group. Mem. New York Bot.
Card. 21: 1-63.
Jepson, W. L. 1925. A Manual of Flowering Plants of
California. Berkeley, California.
Mooring, J. S. 1975. A. cytogeographic study of Er-
iophyllum lanatum (Compositae, Helenieae). Amer.
J. Bot. 62: 1027-1037.
Munz, P. M. & D. D. Keck. 1959. A California Flora.
Univ. California Press, Berkeley, California.
Ownbey, M. 1959. Castilleja. Pp. 295-326 in C. L.
Hitchcock, A. Cronquist, M. Ownbey & J. W.
Thompson (editors), Vascular Plants of the Pacific
Northwest, part 4. University of Washington Pub¬
lications in Biology, vol. 17. Univ. Washington Press,
Seattle, Washington.
Pennell, F. W. 1951. Scrophulariaceae. Pp. 686-859
in L. Abrams (editor), Illustrated Flora of the Pacific
States, vol. 3. Stanford Univ. Press, Stanford, Cal¬
ifornia.
Piper, C. V. 1920. Some new plants from the Pacific
Northwest. Proc. Biol. Soc. Wash. 33: 103-106.
Raven, P. H. 1977. Generic and sectional delimitation
in Onagraceae, tribe Epilobieae. Ann. Missouri Bot.
Card. 63: 326-340.
-, D. W. Kyhos, D. E. Breedlove & W. W. Payne.
1968. Polyploidy in Ambrosia dumosa (Composi¬
tae: Ambrosieae). Brittonia 20: 205-211.
Stebbins, G. L. & D. Zohary. 1959. Cytogenetic and
evolutionary studies in the genus Dactylis. I: Mor¬
phology, distribution, and interrelationships of the
diploid subspecies. Univ. Calif. Publ. Bot. 31: 1-40.
Jaltomata grandijlora (Solanaceae): A Rare Mexican Species
William G. D'Arcy
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Thomas Mione
University of Connecticut, Department of Ecology and Evolutionary Biology,
Storrs, Connecticut 06269, U.S.A.
Tilton Davis II
Wyoming State Crime Laboratory, 316 22nd Street, Cheyenne, Wyoming 82002, U.S.A.
Abstract. Saracha grandijlora is transferred to
Jaltomata as J. grandiflora. The species appears
to be rare: only three collections are known, all
lrom the same locality. A closely related species of
Jaltomata, J. procumbens, is undergoing progres¬
sive domestication in Mexico, and this species ex¬
pands the representation of the genus in the region.
The generic concepts of Saracha Ruiz & Pavon
and Jaltomata Schldl. have long been confused, and
although recent workers (Gentry, 1973; Davis, 1980)
have clarified the separation of the two groups, some
species that were described as Saracha remain to
be transferred to Jaltomata. This paper makes one
such transfer.
Our interest in this species stems from seed col¬
lected ( Davis 1114) from Mexico that has been
grown and propagated over several years. The ear¬
liest name we found for the species is now in Saracha
and not in Jaltomata, where it should be placed.
Because the species is hitherto unknown except lor
the brief original description, we provide an illus¬
tration and an amplified description made from our
cultivated plants and the holotype.
Jaltomata grandiflora (Robinson & Greenmann)
D'Arcy, Mione & Davis, comb. nov. Basionym:
Saracha grandijlora Robinson & Greenmann,
Amer. J. Sci., ser. 3. 161. 1895. TYPE: Mex¬
ico. Michoacan: hills near Patzcuaro, 22 July
1892, Pringle 5273 (GH). Figure 1.
Sprawling perennial herb to 1 m long, the ultimate
branchlets ascending, growth often plagiotropic and
the stems appearing somewhat zigzagged; rootstock
large (14 cm long X 3 cm wide or larger) and
starchy; pubescence white, erect, simple, multicel¬
lular hairs, these sometimes exuding minute clear
droplets (glandular). Stems subterete or somewhat
Novon 2: 190-192. 1992.
pentagonal, drying angled, sometimes with reddish
longitudinal stripes, weak, becoming stout, the pith
large, becoming hollow with age, villous-velutinous
with persistent, erect, weak, whitish, sometimes
glandular hairs ca. 1 mm long and occasional hairs
to 2 mm long. Leaves alternate along the stems or
subequal-paired in the flowering regions, the nodes
mostly 1-5 cm apart; lamina ovate, mostly 4-8(-
12) cm long, 3-5(-8) cm wide, apically broadly
acute or obtuse, basally short-cuneate and winging
the distal portion ol the stems, the margins entire
or 2-3 sinuate-lobed on each side, softly membra¬
nous, major veins ca. 5 on each side, slightly arcuate,
bifurcating near the margins into one trace leading
into a lobe, and another looping and anastomosing
to form a weak, undulating submarginal vein, the
venation pellucid, above fine and slightly impressed,
beneath thick, elevated, and conspicuously white-
strigulose, villous overall with weak erect hairs, more
densely so proximally and above, ciliate, slightly
discolorous, slightly shiny above, drying paler be¬
neath; petioles 1 —2.5(—5) cm long, longer and some¬
times reddish beneath in age, flattened above, evenly
pubescent. Inflorescences racemose, arising in the
axils of a pair of leaves near the branch tips; (1 -)2(-
3)-flowered; peduncles terete, green, evenly pubes¬
cent, ca. 8 mm long; pedicels resembling the pe¬
duncles but slightly thinner, mostly 12-15 mm long,
expanded slightly upward, hardly accrescent. Flow¬
ers all perfect, buds turbinoid, becoming flat-topped,
prominently 5-angled; calyx green, 10 15 mm
across, membranaceous, lobed about halfway, the
lobes deltoid-obtuse, soon porrect, keeled, minutely
puberulent inside, villous outside; corolla rotate, yel¬
lowish white with green maculae forming a star
occupying the central l A- l A of the limb, pentagonal,
25 mm across, pubescent on the veins outside and
near the center within, ciliate, the costae evident,
fine, elevated outside, the lobes deltoid-acute; sta-
Volume 2, Number 3
1992
D’Arcy et al.
Jaltomata grandiflora
191
Figure 1. Jaltomata grandiflora (Robinson & Greenmann) D’Arcy, Mione & Davis. —A. Habit. —B. Flower and
flower bud. —C. Young fruit. —D. Mature fruit. —E. Androecium and corolla pattern. —F. Seed. (All after D’Arcy
17749, MO.)
192
Novon
mens subequal, anthers yellow, ovoid, 3-3.5 mm
long before dehiscence (2 mm after) and lobed ca.
1 mm below the connective, ventrifixed near the
base of the connective, deeply creased on the dorsal
side and along the stomia, not apiculate, filaments
white, erect, subulate, apically glabrous, basally tuft¬
ed, ca. 7 mm long; pollen yellow; disk pale yellow
or orange, surrounding the basal half of the ovary
and immersed beneath the corolla tube and filament
bases; ovary green, hemispherical, style ca. 7 mm
long, glabrous, stigma dark green, hemispherical, 1
mm across, situated among the anthers; ovules ca.
200. Berry black, juicy, depressed globose, 9.2 12
x 14-18 mm across; fruiting calyx porrect, be¬
coming brownish; seeds ca. 90. Chromosomes (mi¬
totic) 2n = 24.
Additional material examined. MEXICO. MICHOACAN:
near Km 35 on route 120 to Santa Clara from Patzcuaro
in old pine-oak forest, 2,004 in, 24-25 July 1977 (in
bud), Davis 776 (MO); Patzcuaro area ca. 1 mi. past
microwave tower, 1.8 mi. past Los Tanques pueblo, 3
July 1982 (st), Davis 1114 (MO); progeny of seed from
Davis 1114 grown at Missouri Botanical Garden, D'Arcy
17749 (MO); grown at University of Connecticut, Mione
454 (COLO, CONN, MEXU).
Jaltomata grandiflora is known only from three
collections made in the Patzcuaro region of central
Michoacan, which is about halfway between Morelia
and Uruapan, 19°30'N, 101°35'W. The region is
of Quaternary and Tertiary extrusive exposures (At¬
las Nacional del Medio Fisico, 1981: 145).
As more material is grown, duplicates will be
distributed to other institutions.
In cultivation, the species appears to tolerate a
limited temperature range, 18°-23°C. At lower and
higher temperatures, growth declines and plants de¬
crease in size, flower buds fail to appear, and buds
already present fail to develop into flowers. This
narrow temperature tolerance may partly account
for the limited distribution of the species in nature,
although many other species have similar temper¬
ature requirements. For plants in our North Amer¬
ican greenhouses, this means blooming is restricted
to a few weeks in spring and autumn. Flowering
plants were found in nature in July and August.
Jaltomata grandijlora is easily distinguished from
other members of the genus by its velutinous, soft
leaves and stems, and by its large, pentagonal, spot¬
ted flowers. Although minute droplets can be seen
at the tips of many of the hairs of the indumentum,
the plants are not viscid. The markings on the co¬
rolla, green against a yellowish white background,
are a series of spots arranged in a star, resembling
the markings in Leucophysalis grandijlora (Hook.)
Rydberg, L. viscosa (Schrader) Hunz, Physalis
franchetii Masters, P. greenmannii Waterfall, and
P. stapelioides (Regel) Bitter. These species are all
members of the physaloid group (subtribe Physali-
dinae Miers) of Solanaceae, which includes about a
dozen genera centered in northern Mexico ( Cha-
maesaracha, Jaltomata, Marga ran thus, Physalis,
Quincula) but is also represented in temperate Asia
( Physaliastrum, Physalis) and in tropical America
(Deprea ). Corollas of most other members of the
group are either unmarked or the maculae are in
solid blotches. The spotted corolla of Jaltomata
grandi flora is shown in Figure IE, and that of
Leucophysalis grandijlora is illustrated by D’Arcy
et al. (1990). Another unusual feature is the root-
stock, which one collector (Davis) recorded as “Wow!
what a large rootstock!” Such enlarged roots may
be otherwise unknown in the physaloid group of
genera.
Jaltomata grandijlora, based on our analysis of
morphological similarity (including fruit color) and
chloroplast DNA characters, is clearly a member of
the herbaceous, black/purple fruited clade (one of
two major clades of Jaltomata). This species, based
on cpDNA characters, is basal; it is the sister taxon
to all other species of the black/purple fruited clade.
Acknowledgments. Supported in part by a Na¬
tional Science Foundation dissertation grant to T.
M ione and G. Anderson.
Literature Cited
Atlas Nacional del Medio Fisico. 1981. Secretaria de
programacion y presupuesto. Mexico, D.F.
D’Arcy, W. G., K. Pickett & R. C. Keating. 1990.
Investigation into Leucophysalis grandijlora. Wild-
flower, Fall/Winter 1990: 21-26.
Davis, T., IV. 1980. The generic relationship of Sar-
acha and Jaltomata (Solanaceae: Solaneae). Rho-
dora 82: 345-352.
Gentry, J. L. 1973. Restoration of Jaltomata (Sola¬
naceae). Phytologia 27: 286-288.
Two New Species of Paspalum (Poaceae: Paniceae) from
Venezuela and Colombia
Gerrit Davidse
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Fernando Zuloaga
Instituto de Botanica Darwinion, Casilla de Correo 22, San Isidro (1642), Argentina
Abstract. Morphological characters support the
description of two new species of Paspalum: P.
atabapense, from Depto. Atabapo, Venezuela, in
group Alterniflora; and P. tillettii, from Comisaria
del Guainia, Colombia, and Territorio Federal Ama¬
zonas, Venezuela, in an unspecified taxonomic group
but clearly related to P. schultesii Swallen.
Preparatory to completing an account of the ge¬
nus Paspalum for the Flora of the Venezuelan
Guayana, two new species are described and illus¬
trated.
Paspalum L., with more than 350 species, is one
of the largest genera of grasses. It is confined pri¬
marily to the New World, although a few species
occur in Africa, Asia, and Oceania. Several species
are pantropical or nearly so, and two weedy species
have now become widely naturalized as weeds in the
tropics. A few have been widely distributed as forage
grasses and have subsequently become naturalized
in new areas. The most useful taxonomic treatment
of this genus remains the monograph of the North
American species (including Mesoamerica) by Chase
(1929).
PaspaSum atabapense Davidse & Zuloaga, sp.
nov. TYPE: Venezuela. Amazonas: Depto. Ata¬
bapo, SE bank of the middle part of Cano Yagua
at Cucurital de Yagua, 3°36'N, 66°34'W, 120
m, 8 May 1979, G. Davidse, O. Huber & S.
5. Tillett 17414 (holotype, MO; isotypes, IAN,
SI, US, VEN). Figure 1.
Gramen perenne caespitosum; culmi 45-75 cm alti.
Folia basalia; ligulae 0.5-1.2 mm longae, glabrae; laminae
6-18.5 cm x 0.3-0.4 mm, filiformes. Racemus 1, 3-
9.5 cm longus; rhachis 0.4-0.5 mm lata, exalata vel
interdum anguste alata basin versus. Spiculae 2.6 3.6 x
1.0-1.3 mm, ellipticae anguste, singulares, pubescentes;
gluma inferna absens; gluma supera 0.3-0.7 mm spicula
brevior. Flosculus superus 2.6-3.5 x 1.0-1.3 mm, stra-
mineus. Antherae 1.5-1.9 mm longae.
Caespitose, perennial herbs. Culms 45-75 cm
long, erect, unbranched above the base, with 2 3
nodes above the base; internodes cylindric, glabrous;
nodes dark, glabrous or pilose. Leaves basal. Sheaths
striate, rounded, densely pilose toward the base with
appressed, whitish hairs, to sparsely pilose toward
the apex, the margins glabrous. Ligules 0.5-1.2 mm
long, membranous, glabrous, erose. Blades 6-18.5
cm long, 0.3-0.4 mm wide, filiform, involute, the
adaxial surface scabrous, prominently ridged, the
abaxial surface glabrous, the apex permanently in¬
volute, somewhat pungent. Inflorescences terminal,
long-exserted; peduncle 4-17 cm long, cylindric,
glabrous; raceme 1, 3.0-9.5 cm long, slightly arch¬
ing, densely pilose at its base; rachis 0.4 0.5 mm
wide, puberulent to glabrous, concavo-convex, un¬
winged or sometimes with partially developed wings
0.1-0.2 mm wide near a portion of the base, ter¬
minating in a spikelet; pedicels 0.1 0.3 mm long,
densely short-pilose. Spikelets 2.6 3.6 mm long,
1.0-1.3 mm wide, solitary, plano-convex, narrowly
elliptic, pilose. Lower glume always absent. Upper
glume 2.4-3.0 mm long, 0.3-0.7 mm shorter than
the upper floret, acute, 3-nerved. Lower floret ster¬
ile. Lower lemma as long as the spikelet, 3-nerved,
glabrous to puberulent in the middle portion, pilose
toward the margins. Lower palea usually absent,
rarely developed and then to % as long as the lower
lemma. Upper floret 2.6-3.5 mm long, 1.0 1.3
mm wide, as long as the spikelet, bisexual, narrowly
elliptic, stramineous, chartaceous, smooth, papillose,
the papillae regularly distributed; lemma 3-nerved,
short-pilose, with prickle hairs at the apex. Stamens
3; anthers 1.5—1.9 mm long, purple. Styles 2, sep¬
arate; stigmas about as long as the styles, plumose.
Caryopsis (1.2—) 1.4-1.5 mm long, 0.9-1.0 mm
wide, obovate in outline; hilum punctiform; embryo
ca. % as long as the caryopsis.
Paratypes. VENEZUELA. AMAZONAS: Depto. Atabapo,
area between the W base of Cerro Yapacana and the
headwaters of Cano Cotua, 3°38'N, 66°52'W, 6 May
1979, Davidse, Huber & Tillett 17251 (K, MO, SI, US,
VEN); Depto. Atabapo, sabanas y bosques riberenos en
los alrededores de Canaripo, en la margen izquierda (Sur)
del bajo Rio Ventuari, a unos 20 km al E de la confluencia
Novon 2: 193-197. 1992.
194
Novon
Figure 1. Panicum atabapense Davidse & Zuloaga (Davidse el nl. 17414, MO). —A. Habit. —B. Leaf, ligular
region. —C. Spikelet, upper glume view. —D. Spikelet,
Upper floret, palea view. —G. Caryopsis, embryo side.
con el Rio Orinoco, 4°03'N, 66°49'W, 98 m, 30 May
1978, Huber 190b (MO, NY, VEN); Depto. Atabapo.
tercera sabana al pie W del Cerro Yapacana, 3°38'N,
66°52'W, 100 m, 3 June 1978, Huber 2023 (MO, NY,
VEN).
This species is known so far only from a ca. 70 x
40-km area in the Depto. Atabapo, Territorio Fed¬
eral Amazonas, Venezuela, where it grows in open.
lower lemma view. —E. Upper floret, lemma view. —F.
H. Caryopsis, hilum side.
white-sand savannas, at approximately 1 00 m ele¬
vation.
Paspalum atabapense appears to fit best in the
informal West Indian group Alterniflora of Chase
(1929) because it is perennial, caespitose, and has
filiform basal leaves and solitary inflorescences with
solitary spikelets. Although this group appears to be
a heterogenous assemblage, P. atabapense is almost
Volume 2, Number 3
1992
Davidse & Zuloaga
Paspalum
195
certainly closely related to the Cuban endemic P.
rottboellioides C. Wright. Paspalum rottboel-
lioides differs from P. atabapense in its winged
rachis up to 1.5 mm wide (vs. essentially wingless
rachis 0.4-0.5 mm wide), spikelets with the upper
glume as long as the upper floret and covering it
(vs. 0.3-0.7 mm shorter than the upper floret and
exposing its apex), lower lemma pilose at the base
(vs. glabrous to puberulent in the middle and pilose
along the margins), and blades densely papillose-
pilose (vs. scabrous adaxially and glabrous abaxially).
The specific epithet refers to the Depto. Atabapo,
where all four known collections originated.
Paspalum tillettii Davidse & Zuloaga, sp. nov.
TYPE: Venezuela. Amazonas: Depto. Atabapo,
Cucurital de Caname, S bank of the middle part
of Cano Caname, 3°40'N, 67°22'W, 100 m,
30 Apr. 1 May 1979, G. Davidse, 0. Huber
& S. S. Tillett 16920 (holotype, MO; isotypes,
SI, US, VEN). Figure 2.
Gramen perenne caespitosum; culmi 23-62 cm alti.
Folia basalia; ligulae 0.2-0.5 mm longae, ciliolatae; lam¬
inae 4-17 cm x 0.3-0.6 mm (1.0-1.5 mm latae ubi
coinplanatae). Racemi 2, 1.5-6 cm longi, conjugati; rhachis
0.2-0.5 mm lata, exalata alis. Spiculae 1.4-1.8(-2.2) x
0.7-0.8(-1.0) mm, ellipticae, singulares, pubescentes;
gluma inferna absens vel raro Vi-Vz- plo spicula longior;
gluma supera plerumque 0.3-0 mm spicula brevior, raro
ad 0.3 mm spicula longior. Flosculus superus 1.4-1.8
(-2.0) x 0.7-0.8(-l .0) mm, ellipticus, atrobrunneus ni-
tidus. Antherae 0.8-1.0 mm longae.
Tufted, c.aespitose, perennial herbs. Culms 23-
62 cm tall, erect, unbranched above the base; in¬
ternodes glabrous or sparsely pilose; nodes dark,
sparsely pilose at the basal nodes, sparsely glabrous
or pilose on the flowering culms. Leaves mostly basal,
erect. Sheaths striate, rounded, the margins mem¬
branous, the basal ones strongly pilose toward the
base, the upper ones usually glabrous to sparsely
pilose, sometimes the entire sheath prominently pi¬
lose. Ligules 0.2-0.5 mm long, membranous, cili-
olate, often with hairs to 2.3 mm long in the hack
at the base of the blade. Blades 4-17 cm long, 0.3-
0.6 mm wide, filiform, usually involute, sometimes
flattened in the middle and then 1.0-1.5 mm wide,
usually glabrous except for a dense row of whitish
hairs to 3.2 mm long behind the ligule and at the
sheath apex and sometimes with the lowest portion
of the margins ciliate, or sometimes the lower half
or the entire abaxial surface pilose, the adaxial sur¬
face prominently ridged; upper blades reduced. In¬
florescences terminal, exserted; peduncle 8-22 cm
long, cylindric, glabrous; racemes 2, 1.5-6 cm long,
conjugate, usually with a dense tuft of white hairs
2-4 mm long at the base; rachis 0.2 0.5 mm wide,
flexuous, glabrous, triquetrous, without wings, ter¬
minating in a spikelet; pedicels 0.2-0.5 mm long,
glabrous to sparsely pilose. Spikelets 1.4—1,8(—2.2)
mm long, 0.7-0.8(-1.0) mm wide, solitary, plano¬
convex, elliptic, dark, densely to sparsely pilose.
Lower glume usually absent, rarely present ( Davidse
et al. 16920), lanceolate, Vz-Vs as long as the spike-
let, pilose. Upper glume usually as long as or to 0.3
mm shorter than the upper floret, rarely to 0.3 mm
longer, obtuse, covered with capitellate hairs toward
the margins, nearly glabrous in the center to uni¬
formly pilose, 3-nerved with obscure nerves. Lower
floret sterile. Lower lemma 2-3-nerved, the nerves
obscure, short-pilose toward the upper margin, oth¬
erwise glabrous or sparsely pilose. Lower palea ab¬
sent. Upper floret 1.4—1.8(—2.0) mm long, 0.7-
0.8(—1.0) mm wide, bisexual, elliptic, dark brown,
smooth and shining, indurate, minutely papillose with
microhairs at the lower margins of the lemma, oth¬
erwise glabrous. Stamens 3; anthers 0.8 1.0 mm
long. Styles 2, separate; stigmas about as long as
the styles, plumose, purple. Caryopsis 0.8-1.0 mm
long, 0.6 0.8 mm wide, obovate in outline; hilum
punctiform; embryo ca. % as long as the caryopsis.
Paratypes. VENEZUELA. AMAZONAS: Depto. Atabapo,
Cano Caname (afluente derecho del medio Rio Atabapo),
sabanas de Cucurital, aprox. 20 km al E de la boca,
3°40'N, 67°22'W, 100 m, Huber, Tillett & Davidse
3702 (MO, VEN); Depto. Rio Negro, extensa sabana en
la margen derecha del bajo Rio Pasimoni, 1°35'N,
66°33'W, 125 m, 8 Feb. 1981, Huber & Medina 5845
(MO, VEN); pequena sabana ubicada en la margen de¬
recha (E) del bajo Rio Pasimoni, a unos 3 km al E del
Rio, 1°38'N, 66°32'W, 125 in, 9 Feb. 1981, Huber &
Medina 5889 (VEN); Depto. Atabapo, white-sand savan¬
na on the N bank of the lower part of Cano Caname, ca.
3 km W of Macaval, 3°41’N, 67°23'W, 95 m, 2 May
1979, Davidse, Huber & Tillett 17078 (K, MO, PRE,
VEN); Depto. Atabapo, white-sand savannas on the N
bank of Cano Caname, nearly opposite Cucurital de Can¬
ame, 3°40'N, 67°22'W, 95 m, 2 May 1979, Davidse,
Huber & Tillett 17050 (MO, VEN); Depto. Atabapo,
area between the W base of Cerro Yapacana and the
headwaters of Cano Cotua, 100 m, 3°38'N, 66°52'W,
100 m, 6 May 1979, Davidse, Huber & Tillett 17216
(INA, MEXU, MO, US, VEN). Colombia, cuainia: near
Coitara, ca. 7 km S of San Fernando de Atabapo, white-
sand area adjoining a laja on the W bank of the Rio
Atabapo, 3°55'N, 67°43'W, 95 m, 28 Apr. 1979, Dav¬
idse 16824 (MO).
This species is known from the Comisaria del
Cuainia, Colombia, and the Territorio Federal Ama¬
zonas, Venezuela, where it grows in open white-sand
savannas at an elevation of 95-220 m.
Paspalum tillettii is very closely related to P.
schultesii Swallen, which is only known from the
nearby Comisaria del Vaupes, Colombia. Both spe-
196
Novon
Figure 2. Paspalum tillettii Davidse & Zuloaga (Davidse et al. 16920, MO). — A. Habit. — B. Portion of a
raceme with spikelets and pedicels. —C. Spikelet, upper glume view. —D. Spikelet, lower lemma view. —E. Upper
floret, lemma view. —F. Upper floret, palea view. —G. Caryopsis, embryo side. —H. Caryopsis, hilum side.
cies have filiform, involute leaf blades, conjugate
racemes, and a dark brown, shiny upper floret. This
combination of characters is not known in any of
the informal groups recognized by Chase (1929),
and this combination also caused Swallen (1967) to
exclude P. schultesii from any known group when
describing this species. The dark brown, shiny upper
floret is characteristic of all species of the Plicatula
group, but conjugate racemes are not known in that
group. Conjugate racemes (P. multicaule Poiret, P.
clavuliferum C. Wright) and capitellate hairs (P.
parviflora Rhode, P. multicaule, P. clavuliferum)
occur in the Parviflora group, but none of the species
in this group have a shiny, dark brown upper floret.
Paspalum schultesii differs from P. tillettii in
having leaves about as long as (vs. V\-Vi as long as)
Volume 2, Number 3
1992
Davidse & Zuloaga
Paspalum
197
the flowering culms, basal sheaths with rufous (vs.
white) hairs, spikelets lanceolate (vs. elliptic), 2.4-
3x 1.1-1.2 mm (vs. 1.4-1.8(-2.2) x 0.7-0.8(-l .0)
mm), and an acute upper glume 0.2 0.5 mm longer
than the upper floret (vs. an obtuse upper glume as
long as or 0.1-0.3 mm shorter). Although most
spikelets are glabrous in P. schultesii, a few of the
spikelets of the type collection have capitellate hairs.
In both species these hairs are only inconspicuously
enlarged at their tips in comparison to those of the
Parviflora group. Nevertheless, all these capitellate
hairs may he fundamentally of the same kind, and
because such specialized hairs are relatively uncom¬
mon in the genus, they may prove to be good in¬
dicators of relationships. Paspalum tillettii, as here
circumscribed, is variable in leaf pubescence and
spikelet shape and pubescence.
The two collections (Huber & Medina 5815,
5889) from Depto. Rio Negro have leaves that are
conspicuously pilose nearly throughout, in contrast
to the more nearly glabrous leaves of the northern
collection from Colombia and Depto. Atabapo.
Certain collections (i.e., Davidse et al. 17216,
17078) appear to consist of two kinds of plants,
one with spikelets 1.5-1.8 mm long and another
with spikelets 1.8-2.2 mm long hut about as wide
as the first. The latter kind has obtuse upper glumes
as long as or to 0.2-0.3 mm longer than the upper
floret, and it varies in the direction of P. schultesii.
This pattern of variation suggests the possibility of
hybridization. It would be worthwhile in future work
to look for additional intermediate populations to
determine whether such plants represent hybrid
swarms between two hybridizing species. The white-
sand savannas in the eastern Colombian llanos are
interspersed with other kinds of soils, and this creates
a mosaic of vegetation types, certainly a setting that
might be favorable for the maintenance of hybrid
Another collection (Venezuela. Amazonas: Depto.
Atures, transecto desde matorral ribereno hasta bos-
que bajo ralo, en la margen izquierda del Cano
“Cabeza de Manteco,” afluente del Rio Autana, en
el ‘*Raudal Manteco,” 4°52'N, 67°27'W, 100-120
m, 10 Nov. 1984, Gudnchez & Melgueiro 3123
(MO)) is related to this species pair but is at the
moment excluded from both P. schultesii and P.
tillettii. In its overall leaf, inflorescence, and spikelet
morphology it resembles both species. Its leaves are
as long as the flowering culms and the hairs on the
basal sheaths are rufous; and its spikelets are 1.8-
2.3 x 1-1.1 mm. In the first two characters it
resembles P. schultesii, and in the latter, P. tillettii.
However, it differs from both in its unusually broad
pedicels.
This species is dedicated to a valued field com¬
panion, Stephen S. Tillett, Universidad Central de
Venezuela, Caracas, Venezuela, in recognition of his
outstanding contributions to Venezuelan botany
through teaching, collecting, curation, and research.
Acknowledgments. Fieldwork by Davidse was
conducted with the support of NSF grant 1NT grant
76-14750 (United States) and a CONIC1T grant to
Otto Huber (Venezuela). Davidse thanks Otto Huber
and Stephen S. 'Tillett for the opportunity to join
their stimulating field expeditions. We thank Vla-
dimiro Dudas for the illustrations, and Tarciso S.
Filgueiras for critical comments on the manuscript
and help with the Latin descriptions.
Literature Cited
Chase, A. 1929. The North American species of Pas¬
palum. Contr. U.S. Natl. Herb. 28: 1-310, i-xvii.
Swallen, J. R. 1967. New species of Paspalum. Phy-
tologia 14: 358-389.
swarms.
Pa ssijlora brachyantha (Passifloraceae), a New Species from the
Andes of Southern Ecuador
Linda k. Escobar
Department of Biology, University of Puerto Rico,
Rio Piedras, Puerto Rico 00931
ABSTRACT. A new species of Passijlora subg. Tac-
sonia is described from the Andes of southern Ec¬
uador. Passijlora brachyantha is placed in section
Bracteogama, where it most closely resembles an¬
other short-flowered species, P. glaberrima, which
is endemic to Peru.
Members of Passijlora subg. Tacsonia (A. L.
Juss.) Triana & Planchon, are tend riled lianas con¬
fined to high Andean habitats, distributed from the
Cordillera de Merida in northwestern Venezuela to
Boli via. One cultivated species, Passijlora pinna-
tistipula Cav., is found in mesic sites in the Andes
of Chile. The plants are characterized by long-tubed,
highly colored, hummingbird-visited flowers, and the
geographical distribution of the plants coincides with
that of Ensifera ensifera, the sword-billed hum¬
mingbird (Escobar, 1980, 1989b; Schauensee,
1982). Hypanthium length in the subgenus varies,
however, and falls mainly into two groups: (1) long-
tubed species probably pollinated by Ensifera en¬
sifera, as is the most widespread species, Passijlora
mixta L. f. (Snow & Snow, 1980), and (2) shorter-
tubed species presumably pollinated by shorter-billed
hummingbirds. There are few species of intermediate
hypanthium length (Escobar, 1980). Since all of the
subgenera of Passijlora that are confined to the
high Andes produce flowers adapted for humming¬
bird pollination (Escobar, 1989b, and unpublished
data), it is likely that the short-tubed taxa, which
are most common in southern Ecuador and northern
Peru, have speciated in response to the selective
pressures exerted by their pollinators.
Section Bracteogama (DC.) L. Escobar com¬
prises 14 of the 47 species placed by Escobar in
subgenus Tacsonia (Escobar, 1980, 1987, 1988,
1989a) and has its center of diversity in the moist
montane forests of southern Ecuador and the eastern
Andean slopes of northern Peru around the Huaca-
bamba depression. The members of the section are
recognized by the presence of large, coriaceous,
mostly connate [tracts that enclose and presumably
protect the bud during development (Escobar, 1980;
Harms, 1925). Both long-flowered and short-flow-
Novon 2: 198 200. 1992.
ered species occur in this section. The new species
described here is another short-flowered member of
section Bracteogama.
Passiflora brachyantha L. Escobar, sp. nov.
TYPE: Ecuador, f „oja: 8 km W of Loja on road
to Catacocha, 7,400 ft., 30 Jan. 1979 (fl, fr),
R. M. king & E. Almeda 7887 (holotype,
US). Figure 1.
Differt a Passijlora glaberrima (A. L. Juss.) Poiret
caulibus foliis bracteisque pubescentibus, foliis majoribus
apicibus acuminatis, et floribus roseis.
Lianas with stout tendrils, pubescent on stems,
petioles, peduncles, bracts, and abaxial surface of
lea ves and stipules, with straight to wavy, trans¬
parent trichomes to 0.5 mm long. Stems angulate,
striate. Leaf blades 3-lobed, 3.5-7.3 cm long, 7.8-
11.0 cm wide, parted ca. 3 A their length into 3
elliptic segments, acuminate at apex of lobes, shal¬
lowly cordate at base, glandular-serrate at margins,
coriaceous; lateral segments 2.2-5.5 cm long, 1.6-
3.4 cm wide, divergent from mid-segments ca. 90°;
mid-segments 3.5-7.3 cm long, 1.6-3.3 cm wide;
petioles 1.2-2.7 cm long, with 3-5 subspherical
nectaries 0.7-1.2 mm long, 0.8-1.0 mm wide, scat¬
tered on upper half of adaxial surface; stipules ren-
iform, concave, 1.2-1.5 cm long, ca. 6 mm wide,
attenuate at apex, oblique at base, shallowly glan¬
dular-serrulate at margins, coriaceous, glabrous on
adaxial surface. Peduncles slender, 2.7-3.4 cm long,
bearing pendent flowers; bracts oblong, connate ca.
3 A their length from base, forming an ampliate tube
at base of hypanthium, each bract 3.1-3.5 cm long,
1.0-1.1 cm wide, acuminate at apex. Flowers 6.4-
6.8 cm long, ca. 3.5 cm wide, bright pink; hypan¬
thium 3.9-4.0 cm long, 0.4-0.6 cm wide (pressed),
dilated at base, pubescent on outer surface with
straight, transparent trichomes ca. 0.3 mm long;
sepals oblong, ca. 2 cm long, ca. 0.6 cm wide,
pubescent with subterminal aristae ca. 1 mm long
on abaxial surface; petals subequal to sepals; corona
tuberculate, ca. 0.3 mm long; ovary ellipsoidal, pu¬
bescent with fine, straight trichomes ca. 0.1 mm
Volume 2, Number 3
1992
Escobar
Passiflora brachyantha
199
Passiflora brachyantha L. Escobar
dct. Linda K. Escobar (UPRRP) 199]
Passiflora mauhavsii (Mast IKilIip
Subg Tacsooia
Del L B Holm-Nielsen 6.
P M lamcosen (AAU) 19S6
PLANTS OF ECUADOR
(ytologit.il jnd/or Hiothcmit.il Vouthcrs
ROHI RT MFRRII.I KINC. 30 January , 9 79
AM) FRANK AI MI IM. NO 7887
UNITED STATES
2850561
Loja: 8 kin W of Loja on the road to Catacocha.
Disturbed pastures. Elevation 7400 ft.
twining vine about 4 m long. Perianth intense pink;
anthers yellow.
NATIONAL HERBARIUK
Spram-n . »!U
Imittn/toi/
titiJ for the 1'ntn J Statei National Herbarium Smith* omiaii
jiiJ ll<, ll. flum. „( tin Citifnrnu Audim) "! Stir*in.
Figure 1. Holotype of Passiflora brachyantha L. Escobar, deposited at US.
200
Novon
long. Fruit ellipsoidal, ca. 5.5 cm long, ca. 2.2 cm
wide (pressed), strongly coriaceous, green; seeds ob-
ovate, ca. 5.5 mm long, ca. 3.8 mm wide, with
reticulate pitting on testa.
Passiflora brachyantha most closely resembles
the Peruvian species P. glaberrima (A. L. Juss.)
Poiret by virtue of the general shape of the leaves,
the slender peduncles, and small flowers. It differs
from P. glaberrima in being pubescent, possessing
much larger leaves with acuminate apices, and in
having pink rather than blue-violet flowers. It is
known only from the type collection.
Acknowledgments. 1 gratefully acknowledge the
University of Antioquia, Medellin, Colombia, and
the Fulbright Commission, Ecuador, for financial aid
tor the original studies of subgenus Tacsonia, and
that of COLCIENCIAS Grant 10019-1-39-82 for
subsequent work. The Plant Resources Center, Uni¬
versity of Texas, Austin, has provided space and
facilities for the present study. I am indebted to Cuy
Nesom for the Latin diagnosis and to B. L. Turner
for a critical reading of the manuscript. Finally, I
thank the staff of US for loan of the specimen here
designated as the holotype of Passiflora brachyan¬
tha.
Literature Cited
Escobar, L. K. 1980. Interrelationships of the edible
species of Passiflora centering around Passiflora
mollissima( H.B.K.) Bailey, subgenus Tacsonia. Ph.D.
Dissertation, University of Texas, Austin.
-. 1987. Novedades en Passiflora (Passiflora-
ceae) de Colombia. Mutisia 54: 1-8.
-. 1988. Passiflora subgeneros: Tacsonia, Ka-
thea, Manicata y Distephana. Monografia 10. In:
Polidoro Pinto & Gustavo Lozano (editors), Flora de
Colombia. Imprenta Universidad Nacional de Colom¬
bia, Bogota, D.E.
-. 1989a. A new subgenus and five new species
in Passiflora (Passifloraceae) from South America.
Ann. Missouri Bot. Gard. 76: 877-885.
-. 1989b. Speciation in the Andes: The genus
Passiflora. Amer. J. Bot. Suppl. 76: 212-213. [Ab¬
stract.]
Harms, H. 1925. Passifloraceae In: Engler & Prantl,
Die Natiirlichen Pflanzenfamilien, ed. 2: 470-507.
Schauensee, R. M. de. 1982. A Guide to the Birds of
South America. Pan American section of the Inter¬
national Council for Bird Preservation. [Reprint.]
Snow, D. W. & B. K. Snow. 1980. Relationships be¬
tween hummingbirds and flowers in the Andes of
Colombia. Bull. Brit. Mus. (Zool.) 38: 105-139.
Gurania sessiliflora (Cucurbitaceae), a New Species from Panama
Rachel Jane Hampshire
The Natural History Museum, Cromwell Road, London SW7 5BD, England
ABSTRACT. A new Panamanian species, Gurania
sessiliflora, is described. The species appears most
similar to Gurania coccinea Cogn. Description of
G. sessiliflora brings the number of Gurania species
in Mesoamerica to seven. A key to the Mesoamer-
ican species of Gurania is given, followed by a list
with synonyms and distributions in Mesoamerica.
During my study of Central American specimens
of Gurania and Psiguria for Flora Mesoamericana,
the new species Gurania sessiliflora was found.
Gurania sessiliflora R. J. Hampshire, sp. now
TYPE: Panama. Darien: Cerro Pirre, along
river by Rancho Frio, 7°58'N, 77°42'W, 600
m, 9 Aug. 1986 (fl), McDonagh, Lewis, Gum-
pel & Plumptre 620 (holotype, BM). Figure 1.
Species G. coccineae Cogn. similis, sed floribus sessi-
libus, planta omnino glabrata (nec pilosa nec villosa), foliis
integris vel raro lobatis (nunquam foliolatis) differt.
Monoecious vines; tendrils simple; stems slender,
striate, glabrate, green or pale brown, the epidermis
sometimes flaking. Leaf blades 1217.5 x 7-13
cm, cordiform to broadly ovate, simple, unlobed, or
rarely 2-3-lobed, the apex long-acuminate, gla¬
brate, the base cordate or truncate, the margin
remotely dentate; petioles 3.5-7 cm, the epidermis
of the basal half flaking, glabrate. Staminate inflo¬
rescences axillary, subcapitate; peduncles 9-12.5
cm, ± glabrous; flowers sessile, the apex of the
peduncle clearly scarred where earlier flowers have
been, the scars congested; calyx tube 4-6 mm,
urceolate, glabrous, orange, the lobes 5, ca. 2 mm,
conical, glabrous; corolla lobes 5, 2-3 mm, lanceo¬
late, papillose, yellow; anthers 2, ca. 3 mm, oblong,
initially straight, spiraling slightly after anthesis, the
appendage ca. 0.5 mm, papillose. Female flowers
not seen. Fruit not seen.
Paratype. Panama. DARIEN: Cerro Pirre, cloud forest
and/or mossy forest, ca. 2,500-4,500 ft., 9-10 Aug.
1967, Duke & Elias 13686E (MO).
Gurania sessiliflora is clearly distinct from other
species of Gurania, but is closest to G. coccinea.
However, G. coccinea has distinctly pedicellate flow¬
ers (with pedicels 4-15 mm long) and pedicel scars
that are evenly spaced at the apex of the peduncle,
while G. sessiliflora has sessile flowers, with the
scars from fallen flowers very congested at the tip
of the peduncle. Gurania coccinea is usually pilose,
or sometimes villous or glabrate, while the new spe¬
cies is glabrate. The leaves of G. coccinea are un¬
lobed, 2-3-lobed or (in Panama) 3-foliolate, while
those of the new species are unlobed or rarely 2-
3-lobed, but never foliolate. It is assumed that, like
other species of Gurania and Psiguria, G. sessili¬
flora will prove to be monoecious (Condon & Gilbert,
1990). A dissected flower of the paratype had three
smaller anthers, but is assumed to be an aberrant
specimen.
The new species is found in cloud or elfin forest
at elevations of 600-1,500 m in the Cerro Pirre
area of Darien, Panama. The holotype was collected
by four undergraduates from the University of Bris¬
tol, England, who made a collection of interesting
material while on an expedition to Panama.
Many authors have said the genera Psiguria and
Gurania need revision. Wunderlin (1978) was un¬
certain whether the two genera were distinct, while
Condon & Gilbert (1990) and Jeffrey (1978) ac¬
knowledged the need for revision of both genera.
For Flora Mesoamericana I have treated the species
in the broadest sense, reducing to synonymy names
which, following a more detailed study, may prove
distinct. Following this policy, G. sessiliflora brings
the total number of Gurania species in Mesoamerica
(from Tabasco, Yucatan, and Chiapas to the Pan-
ama-Colombia border) to seven. A key to the Me-
soamerican species of Gurania is given below, fol¬
lowed by a list with synonyms and distributions in
Mesoamerica.
la. Older steins corky; calyx tube of male flowers longer than 1 cm; male flowers on pedicels 1-3 cm .
. 7. G. tubulosa
lb. Stems never corky; calyx tube of male flowers usually less than 1 cm; male flowers on pedicels usually less
than 1 cm.
2a. Calyx lobes of staminate flowers ca. 2 mm (Darien Province, Panama).
Novon 2: 201-203. 1992.
202
Novon
5 cm
Figure 1. Gurania sessiliflorci R. J. Hampshire. —A. Map of Panama, showing the position of Cerro Pirre, the
type locality. —B. Habit. —C. Dissected male flower showing position of the anthers.
Volume 2, Number 3
1992
Hampshire
Gurania sessiliflora
203
3a. Male inflorescences on peduncles longer than 35 cm; leaves hispidulous beneath . . 1. G. brevisepala
3b. Male inflorescences on peduncles shorter than 13 cm; leaves glabrate .5. G. sessiliflora
2b. Calyx lobes of staminate flowers longer than 2 mm (Chiapas-Panama).
4a. Fruit villous; stems villous with hairs 4-6 mm long; male flowers subsessile . 3. G. eriantha
4b. Fruit glabrous; stems sometimes villous, the hairs usually less than 4 mm long, but if longer, the
male flowers on pedicels at least 6 mm.
5a. At least some leaves foliolate .2. G. coccinea
5b. Leaves not foliolate.
6a. Male inflorescences on peduncles longer than 29 cm . 6. G. spinulosa
6b. Male inflorescences on peduncles shorter than 24 cm.
7a. Lowest male flowers on pedicels 1-3 mm .4. G. makoyana
7b. Lowest male flowers on pedicels at least 4 mm. 1. G. coccinea
1. Gurania brevisepala Cuatr.
Panama.
This species, known in Mesoamerica from two
collections (Uhitefoord & Eddy 470, BM, and Gar¬
wood 747, F), represents a new record for the flora
of Panama. One collection is from Mamey, the other
from near Jacque, both in Darien Province.
2. Gurania coccinea Cogn.
Synonyms: G. costaricensis Cogn., G. racemifera
Standley.
Nicaragua, Costa Rica, and Panama.
3. Gurania eriantha (Poeppig & Endl.) Cogn.
Synonym: G. hirsuta Cogn.
Costa Rica and Panama.
4. Gurania makoyana (Lem.) Cogn.
Synonyms: G. donnell-smithii Cogn. ex J. D.
Smith, G. levyana Cogn., G. seemaniana Cogn.,
G. tonduziana J. D. Smith.
Chiapas, Belize, Cuatemala, Honduras, Nicara¬
gua, Costa Rica, and Panama.
5. Gurania sessiliflora R. J. Hampshire
Panama.
6. Gurania spinulosa (Poeppig & Endl.) Cogn.
Nicaragua and Panama.
This species represents a new record for Nica¬
ragua (Jeffrey, in press) and Panama.
7. Gurania tubulosa Cogn.
Synonyms: G. megistantha J. D. Smith, G. sub-
erosa Standley.
Costa Rica and Panama.
Acknowledgments. I thank Charles Jeffrey (K)
for advice and encouragement, Margaret Tehbs (BM)
for advice on preparation of the illustration, and
Norman Robson (BM) for help with the Latin di¬
agnosis.
Literature Cited
Condon, M. A. & L. E. Gilbert. 1990. Reproductive
biology and natural history of the neotropical vines
Gurania and Psiguria. Pp. 150-166 in D. M. Bates,
R. W. Robinson & C. Jeffrey (editors), Biology and
Utilization of the Cucurbitaceae. Comstock Publish¬
ing Assocs., Cornell Univ. Press, Ithaca & London.
Jeffrey, C. 1978. Further notes on Cucurbitaceae: IV.
Some New-World taxa. Kew Bull. 33: 347-380.
Wunderlin, R. P. 1978. Cucurbitaceae. In: Flora of
Panama. Ann. Missouri Bot. Card. 65: 285 366.
New Western North American Taxa of Arceuthobium (Viscaceae)
Frank G. Hawksworth
USDA Forest Service, Rocky Mountain Forest and Range Experiment Station,
Fort Collins, Colorado 80526, U.S.A.
Delbert Wiens
Department of Riology, University of Utah, Salt Lake City, Utah 85112, U.S.A.
Daniel L. Nickrent
Department of Plant Biology, Southern Illinois University, Carbondale, Illinois 62901, U.S.A
ABSTRACT. Several new names are proposed ior a
taxonomic revision of the dwarf mistletoes {Arceu¬
thobium). Two new species from northwestern Cal¬
ifornia and southwestern Oregon are described: A.
siskiyouense, a parasite of knobcone pine, and A.
monticola, a parasite of western white pine. Also
described as new is A. littorum, a parasite of Mon¬
terey and bishop pines in coastal California. Arceu¬
thobium tsugense is segregated into two subspecies:
subspecies tsugense, primarily parasitic on western
hemlock, and subspecies mertensianae, primarily
parasitic on mountain hemlock.
In preparing a revision of our monograph on
dwarf mistletoes ( Arceuthobium, Viscaceae) (Hawk¬
sworth & Wiens, 1972) and for our treatment of
the Viscaceae for the new Jepson Manual oj the
Flowering Plants of California, we have continued
to investigate the systematics of this economically
important genus of conifer parasites (Hawksworth
& Wiens, 1976, 1977, 1980, 1984, 1989; Nick-
rent, 1986, 1987; Nickrent & Butler, 1989, 1990,
1991; Nickrent & Stell, 1990; Nickrent et al.,
1984). Here we describe three new species that
parasitize pines in Oregon and California, and we
also segregate the hemlock dwarf mistletoe into two
subspecies. Two of the new species are endemic to
the Siskiyou-Klamath Mountains floristic province
of northwestern California and southwestern Ore¬
gon, an area well known for its high degree of plant
endemism (Smith & Sawyer, 1988).
Only brief descriptions of the new taxa, their
relationships to closely related taxa, and selected
collections representing the host and geographic dis¬
tribution of the new taxa are cited here. Additional
descriptions, illustrations, complete specimen cita¬
tions, and distribution maps will be presented in our
revised monograph of the genus, which is nearing
completion. The methods used for our morphological
analyses are as described by Hawksworth & Wiens
(1972).
Arceulhobiuni siskiyouense Hawksworth, W iens
& Nickrent, sp. nov. TYPE: U.S.A. Oregon:
Josephine County, Oregon Mountain Road, 6
mi. SW of O’Brien, parasitic on Pinas atten¬
uate T. 41 S„ R. 9 W„ Sect. 9, 42°01'N,
123°46'W, elevation 650 m, 20 July 1987,
I). Wiens 6756 (holotype, US; isotypes, FPF,
MO, ORE, OSC, UC).
Plantae 6-10 (8) cm altae; surculi brunnei, parce fla-
bellate ramosi; surculi principals basi 2-2.5 mm diam.,
internodiis tertiis 6-15 (9) mm longis, 2 mm latis; flores
staminati 3-4-ineri; fructus maturi 4 mm longi, 2.5 mm
lati; anthesis mense Septembri; fructus maturitas mense
Septembri-Octobri. In Pino attenuata parasiticae.
Plants 6-10 (mean 8) cm tall, brownish, flabel-
lately branched, predominant shoots 2 2.5 mm diam.
at base, third internode from base 6 15 (mean 9)
mm long and 2 mm wide; staminate flowers 3- or
4-partite; mature fruit 4 mm long and 2.5 mm wide;
anthesis in September; fruits mature in September
and October; parasitic principally on Pinus atten¬
uata.
We previously included this dwarf mistletoe under
A. campylopodum (Hawksworth & Wiens, 1972,
1984), but noted an anomalous situation near Gas-
quet, Del Norte County, California, where a dwarf
mistletoe (then presumed to be A. campylopodum)
was common on Pinus attenuata but rare on as¬
sociated P. ponderosa (Hawksworth & Wiens,
1972). This situation is now readily explained, be¬
cause the taxon on P. attenuata is A. siskiyouense,
not A. campylopodum.
Arceuthobium siskiyouense is a local endemic
restricted to the Siskiyou-Klamath Mountains of
southwestern Oregon (Curry and Josephine counties)
Novon 2: 204-211. 1992.
Volume 2, Number 3
1992
Hawksworth et al.
Arceuthobium
205
Table 1. Comparison of some characters of Arceuthobi
Klamath Mountains. Means are in parentheses.
urn siskiyouense and
1. campylopodum in the Siskiyou-
Character
A. siskiyouense
(19 collections)
A. campylopodum
(17 collections)
Shoot height
6-10 (8) cm
10-14 (12) cm
Basal shoot diameter
2-2.5 mm
3-6 mm
Third internode
6-15 (9) x 2 mm
11-17 (14) x 3-3.5 mm
Mature fruit
4 x 2.5 mm
6 x 3.5 mm
Witches’ brooms
None
Well developed
Peak anthesis
September
August
Peak seed dispersal period
September-October
September-October
Elevational range
400-1,200 m
50-2,400 m
Parasitism of Pinus ponderosa and P. jeffreyi
Rare
Common
Parasitism of P. attenuata
Common
Rare
and adjacent northwestern California (Del Norte,
Humboldt, and Siskiyou counties). Its known ele-
vational range is from 400 to 1,200 m. In marked
contrast, A. campylopodum has a wide distribution
from northern Idaho and northern Washington to
Baja California. The two species are sympatric in
several areas but retain their distinctive morpholog¬
ical shoot and fruit characteristics. Their flowering
periods partially overlap, but there is no evidence
of hybridization between them, thus further sup¬
porting their specific status.
Pinus attenuata is the most common host of A.
siskiyouense. The mistletoe will rarely parasitize P.
ponderosa, P. jeffreyi, and P. contorta subsp. con¬
torta, but only in areas where these species grow
in close association with infected P. attenuata. Ar¬
ceuthobium siskiyouense retains its morphological
integrity in such ‘‘host crossovers,” providing fur¬
ther evidence supporting its specific segregation.
Furthermore, inoculation tests on Pinus attenuata
with A. siskiyouense (from the Casquet area) in the
central Sierra Nevada foothills at Camino (elevation
900 m) resulted in the exceptionally high infection
rate of 86 percent. Inoculations on the same host
with A. campylopodum seed from P. jeffreyi on
the Plumas National Forest resulted in only 11 per¬
cent infection (H. F. Scharpf, pers. comm.).
Arceuthobium siskiyouense differs from A. cam¬
pylopodum primarily in its shorter, more slender
shoots, smaller fruits, later flowering period, absence
of witches’ broom formation, and varying host re¬
lationships (Table 1). Electrophoretic evidence also
confirms the distinctness of this taxon (Nickrent &
Butler, 1991).
Paratypes: on Pinus attenuata, except as noted. U.S. A.
CALIFORNIA: Del Norte Co., Gasquet, Tracy 16464 (CAS,
JEPS, UC, VS, WTU); Gordon Mtn., Newcomb 165 (UC);
Bear Wallows, 2 mi. N of Sanger Peak, Kildare 8785
(CAS); 6 mi. NE of Gasquet on old toll road, on P.
ponderosa, Hawksworth & Wiens 867 (FPF) and on P.
contorta subsp. contorta, Hawksworth & Wiens 866
(FPF); 18 Mile Creek Canyon, Elk Camp Ridge, on P.
contorta subsp. contorta. Parks & Parks 24063 (UC);
2.5 mi. NW of Black Butte, Tinnin <$c Kirkpatrick 1A
(FPF); 2 mi. S of Hazelview Summit on Rte. 199, Hawk¬
sworth 2272 (FPF). Humboldt Co., NW of Scotia, Miller
FP98033 (PFRS). Siskiyou Co., Russian Creek, Butler
273 (UC); 3 mi. N of Branch Guard Sta., Kuijt 1277
(UC); 16 mi. N of Happy Camp on O’Brien rd., Hawk¬
sworth & Hinds 1001 (FPF). OREGON: Curry Co., 12 mi.
NNW of Agness, Graham s.n. (FPF); W side of Vulcan
Peak, Denton 3689 (WTU); SW slope of Iron Mtn.,
Baker 3100 (ID). Josephine Co., O'Brien, on Pinus con¬
torta subsp. contorta, Weir 3187 (ILL); and on Pinus
jeffreyi, Lewis, 1917 (FPF, ILL); 5 mi. W of Grants
Pass, on P. jeffreyi, Graham, 1964 (FPF); Bain Station
on Forest Service road 112, on P. contorta subsp. con¬
torta. Mathiasen 8605 (FPF); 2 mi. E of Cave Jet.,
Theisen, 1965 (FPF); near Lookout Gap, 20 air mi. S of
Galice, on P. jeffreyi, Graham, 1965 (FPF); 0.5 mi. S
of Galice-Agness road on Hobson Horn road, Hawksworth
2333 (FPF); Wonder, on P. jeffreyi, Mitchell FP68212
(FPF); Mt. Peavine, near Galice, Childs FP91615 (OSC).
Arceuthobium monticola Hawksworth, Wiens
& Nickrent, sp. nov. TYPE: U.S.A. Oregon:
Josephine County, 7 mi. S of O’Brien on Old
Gasquet Toll Hoad (Oregon Mountain Road);
parasite on Pinus monticola, T. 41 S., R. 9
W., Sect. 9, 42°01'N, 123°46'W, elevation
650 m, 20 July 1987, D. Wiens 6757 (ho-
lotype, US; isotypes, FPF, MO, ORE, OSC,
UC).
Plantae 5-10 (7) cm altae; surculi brunnei, parce fla-
bellate ramosi; surculi principales basi 2-4 (3) mm diam.,
internodiis tertiis 8-15 (12) mm longis, 1.5-2 mm latis;
ffores staminati 3-meri; fructus maturi 4-4.5 mm longi,
2-2.5 mm lati; anthesis mense Julio-Augusto; fructus
maturitas mense Octobri-Novembri. In Pino monticola
parasiticae.
Plants 5-10 (mean 7) cm tall, brownish, flabel-
lately branched; dominant shoots 2-4 (mean 3) mm
206
Novon
Table 2. Comparison of some
parentheses.
characters of Arceuthobium monticola and
A. californicum. Means are in
A. monticola
A. californicum
Character
(26 collections)
(8 collections)
Shoot color
Brownish to reddish
Yellow to greenish
Shoot height
5-10 (7) cm
6-14 (8) cm
Basal shoot diameter
2-4 (3) mm
1.5-4 (2) mm
Third internode
8-15 (12) x 2 mm
5-16 (10) x 1.5 mm
Mature fruit
4x2 mm
4 x 2.5 mm
Fruit color
Bluish
Greenish
Peak anthesis
July-August
July
Peak seed dispersal
Parasitism of:
October-November
September
Pinus monticola
Common
Rare
Pinus lambertiana
Rare
Common
diam. at the base; third internode from the base 8-
15 (mean 12) mm long and 1.5-2 mm wide; sta-
minate flowers 3-partite; mature lruit 4-4.5 mm
long and 2-2.5 mm wide; anthesis July and August;
fruits mature in October and November; parasite
principally on Pinus monticola.
Previously we included this taxon under Arceu-
thobium californicum (Hawksworth & Wiens,
1972). However, subsequent field and laboratory
studies have demonstrated that it is a distinct species
endemic to the Siskiyou and Klamath mountains ot
southwestern Oregon (Coos, Curry, and Josephine
counties) and adjacent northwestern California (Del
Norte County). Its known elevational range is from
700 to 1,600 m. To our knowledge, it is not sym-
patric with A. californicum, which is apparently
confined to California.
Arceuthobium monticola differs from A. cali¬
fornicum in its much darker shoot color, later flow¬
ering and seed dispersal periods, and occurrence
primarily on Pinus monticola (Table 2). Electro¬
phoretic evidence also confirms the distinctness of
these two taxa (Niekrent & Butler, 1991).
Paratypes: on Pinus monticola except as noted. U.S.A.
CALIFORNIA: Del Norte Co., 0.5 mi. E of Sourdough Jet.,
ca. 10 air mi. N of Gasquet, Miller, 1969 (FPF); Black
Butte, Theisen <£• Bynum, 1968 (FPF), and on Picea
breweriana, Mathiasen #609 (FPF); 2.4 mi. W of Smith
River near Lee Brown Crossing, Mathiasen 8606 (FPF);
0.6 mi. N of Ship Mt. Lookout on Little Jones Creek
road, Mathiasen 8502 (FPF); 0.5 mi. N of Black Butte,
Mathiasen 8607 (FPF); 3.0 mi. E of Smith River bridge,
10 mi. N of Gasquet, Niekrent & Wiens 2705 (FPF,
ILL); Lower Coon Mtn., 1 mi. SW of Camp 6 Lookout,
Mastrogiuseppe & Mastrogiuseppe 153 (WS). OREGON:
Coos Co., 13 air mi. S of Powers, Mathiasen 8610 (FPF).
Curry Co., Saddle Mtn., ca. 15 air mi. ESE of Gold Beach,
Bynum, 1967 (FPF) and on Picea breweriana, Bynum,
1967 (FPF); 0.2 mi. N of Snow Camp Fire Lookout,
Mathiasen 8601 (FPF), and on P. jeffreyi, Mathiasen
8602 (FPF); 14 mi. ESE of Gold Beach, Wiens 6 793
(FPF); Windy Creek, N of Loeb State Park, Wiens 6794
(FPF). Josephine Co., Oregon Mt., 14 mi. SW of Cave
Jet., on P. lambertiana, Weir 3191 (FPF, ILL).
Arceuthobium littorum Hawksworth, Wiens &
Niekrent, sp. nov. TYPE: U.S.A. California:
Mendocino County, 3.0 mi. E of Noyo on State
Highway 20, on Pinus muricata, elevation 30
m, 12 Dec. 1989, R. L. Mathiasen 8940
(holotype, LIS; isotypes, CAS, FPF, HSC, MO,
UC).
Plantae 10 20 (12) cm altae; surculi brunneo-virides,
parce flabellate ramosi; surculi basi 2-5 (3.5) mm diam.,
internodiis tertiis 10-20 (15) mm longis, 2-5 (3.5) mm
latis, flores staminati 4-meri; fructus maturi 3-5 mm longi;
anthesis mense Augusto-Septembri; fructus maturitas
mense Septembri-Octobri. In Pino radiata et Pino mur¬
icata parasiticae.
Plants 10-12 (mean 1 2) cm tall; brown to green,
flabellately branched; dominant shoots 2 5 (mean
3.5) mm diam. at base; third internode 10 20 (mean
15) mm long and 2-5 (mean 3.5) mm wide; sta-
minate flowers 4-partite, mature fruit 3-5 mm long;
anthesis August and September; fruits mature in
September and October; parasite principally on Pi¬
nus radiata and P. muricata.
Previously this taxon was included in Arceu¬
thobium occidentals Engelm. (Hawksworth &
Wiens, 1972, 1984). We now separate A. littorum
as a distinct species parasitic on Pinus radiata I).
Don, P. muricata D. Don, and rarely P. contorta
Dougl. ex Loud, subsp. bolanderi (Pari.) Critch. It
occurs from Fort Bragg (Mendocino County) to
Cambria (San Luis Obispo County) and ranges in
elevation from near sea level to about 250 m.
Arceuthobium littorum differs from A. occiden-
Volume 2, Number 3
1992
Hawksworth et al.
Arceuthobium
207
Table 3. Comparison of some
characters of Arceuthoblium littorum and A.
occidentale. Means are in parentheses.
A. littorum
A. occidentale
Character
(8 collections)
(9 collections)
Shoot height
10-20 (12) cm
6-12 (8) cm
Mean shoot diameter
2-5 (3.5) mm
1.5-4 (2.3) mm
Third internode
15 x 3.5 mm
12x2 mm
Shoot color
Dark brown to olive green
Light green to straw
Mature fruit length
4-5 mm
3.5 4.5 mm
Staminate flowers
4-partite
3- or 4-partite
Shoot habit
In dense globose clusters
Open, scattered shoots
Witches’ brooms
Large, nonsystemic
Usually none
Peak anthesis
August-September
October-November
(24 observations)
(176 observations)
Peak seed dispersal
September-October
October December
(31 observations)
(187 observations)
Mean fruit maturation
14 months
13 months
Elevational range
Near sea level to 250 in
100-1,200 m
tale in several features. Arceuthobium occidentale
is primarily a parasite of Pinas sabiniana in the
foothills surrounding the Central Valley of California.
It is not sympatric with A. littorum, which is con¬
fined to a narrow band within about 5 km of the
sea. Verticillate branching of A. littorum is some¬
what more common than for A. occidentale (Mark
& Hawksworth, 1981). Electrophoretic studies con¬
firm that A. littorum is the most distinct member
of the A. campylopodum-A. occidentale complex
(Nickrent & Butler, 1990). Additional differences
between A. littorum and A. occidentale are given
in Table 3.
Pleistocene fossils of A. littorum (originally re¬
ported as A. campylopodum) were found associated
with P. muricata and P. radiata in Marin County
(Tomales Bay) and Santa Barbara County (Carpin-
teria and Santa Cruz Island) (Chaney & Mason,
1933). These 30,000-40,000-year-old populations
are characterized by mostly 4-partite staminate flow¬
ers, a characteristic of modern A. littorum. This
dwarf mistletoe is still extant near Tomales Bay, but
is extinct at the other two localities.
Paratypes. U.S.A. CALIFORNIA: Alameda Co., Berke¬
ley, on planted P. radiata, Offord & Scharpf 1964
(FPF). Marin Co., Inverness Ridge, on P. muricata, How¬
ell 19686 (CAS, UC). Mendocino Co., Fort Bragg, on P.
muricata. Mason 5639 (UC); mouth of Gualala River,
on P. muricata, Bacigalupi 1808 (CAS); Van Damme
State Park, on P. muricata, Kuijt 1214 (UC); 2.9 mi.
E of Point Arena, on P. muricata, Nickrent 2719 (FPF,
ILL); 2.7 mi. SE of Fort Bragg, on P. contorta subsp.
bolanderi, Nickrent 2716 (FPF, ILL); Van Damme State
Park, on P. contorta subsp. bolanderi, Kuijt 1215 (UC);
just E of airport near Albion, on P. contorta subsp.
bolanderi, Peterson 65-116 (FPF); White Sands near
Mendocino City, on P. muricata, Eastivood 18836 (CAS);
2 mi. S of Anchor Cove, on P. muricata. Wolf 1342
(CAS). Monterey Co., Pebble Beach, Carmel, on P. ra¬
diata, Boyce 33 (FPF); Carmel, on P. radiata, von
Schrerik, 1920 (MO); Carmel Highlands above Yankee
Point, on P. radiata. Balls 23608 (WTU); Point Lobos
State Park, on P. radiata, Lee & Mason 9153 (UC);
Gibson Creek, on P. radiata, Wheeler 4452 (POM);
Monterey, on P. radiata. Parry, 1850 (MO); Pacific
Grove, on P. radiata, Davy 7064 (UC); Cypress Point,
on P. radiata, Abrams 7660 (RM, US); Huckleberry
Hill, Carmel, on P. muricata. Mason 5515 (UC). San
Luis Obispo Co., Cambria, on P. radiata, Gill FP68079
(FPF); between Cambria and Cambria Pines on Roberts
road, on P. radiata, Hawksworth 2264 (FPF). Sonoma
Co., Fort Ross, on P. muricata. Mason 4285 (UC); Kruse
Rhododendron State Park on Rte. 1, on P. muricata,
Nickrent 2722 (FPF, ILL).
Arceuthobium tsugense (Rosendahl) G. N. Jones,
Univ. Wash. Publ. Biol. 5: 139, 1936.
In our monograph (Hawksworth & Wiens, 1 972),
we commented on the unusually broad host range
of A. tsugense, which not only encompassed both
western species of Tsuga, but also several species
of Abies, Picea, and Pinus. The occurrence of a
form or “pathotype” (Hunt & Smith, 1978) on
shore pine in British Columbia and the San Juan
Islands of Washington had long been known (Hawk¬
sworth & W iens, 1972; Hunt & Smith, 1978; Smith,
1971, 1974; Smith & Wass, 1976, 1979; Wass,
1976; Kuijt, 1956), although its taxonomic status
has not been previously established. Field and lab¬
oratory studies of A. tsugense now show that this
species is comprised of at least three segregates
(Hawksworth, 1987; Mathiasen & Hawksworth,
1988).
The populations on mountain hemlock (subsp.
rnertensianae) and western hemlock (subsp. tsu¬
gense) are described here as subspecies because of
208
Novon
Table 4. Comparison of the shoot height of the
two subspecies
of Arceuthobium
tsugense.
Shoot height
Number of
Mean and
Subspecies
collections
Range
standard error
subsp. tsugense (western hemlock race)
86
3-13 cm
6.9 ± 0.2 cm
subsp. mertensianae
58
3-9 cm
5.3 ± 0.2 cm
their distinct host ranges, shoot sizes, and marked
isozyme differences (Nickrent, 1988; Nickrent &
Stell, 1990). The populations of subspecies tsugense
on western hemlock and shore pine are considered
to he host races because they are morphologically
and phenologically similar. Also, the isozyme pat¬
terns of the two races are similar, and they are
sympatric in many stands (Nickrent & Stell, 1990).
Arceuthobium tsugense (Rosendahl) G. N. Jones
subsp. tsugense, Univ. Wash. Publ. Biol. 5:
139, 1936.
As presently interpreted, this subspecies consists
of two host races, one primarily on Tsuga hetero-
phylla and one on Pinus contorla subsp. contorta.
An intensive series of field and inoculation studies
in British Columbia (Smith, 1971, 1974; Smith &
Wass, 1976, 1979; Wass, 1976) showed that the
two races are similar morphologically hut differ in
their host relationships. In inoculation tests, the 7.
heterophylla race produced very low infection levels
on P. contorta subsp. contorta, but the few infec¬
tions produced developed abundant aerial shoots. In
contrast, the P. contorta subsp. contorta race pro¬
duced a moderate infection level on /. heterophylla,
but only a few of these produced aerial shoots (Smith
& Wass, 1979). Maximum shoot height of the 7.
heterophylla race was about 30 percent taller than
shoots of the P. contorta subsp. contorta race (Smith,
1971). Results from inoculation trials in two areas
in British Columbia confirm the morphological sim¬
ilarity of the two races (E. Wass, pers. comm.). The
only statistically significant difference between the
populations in the two areas was that the fruits of
the P. contorta subsp. contorta race were slightly
longer and wider than those of the T. heterophylla
race. Also, the peaks in anthesis and seed dispersal
occurred about 10 days earlier in the P. contorta
subsp. contorta race than in the T. heterophylla
race in the same areas.
See Hawksworth & Wiens (1972) for information
on the distribution of the two races. In general:
1. The western hemlock race is distributed from
southeast Alaska (from near Haines, at nearly 60°N)
throughout the coastal western hemlock forests of
British Columbia, Washington, and Oregon to Hum¬
boldt and Del Norte counties in northwestern Cali¬
fornia (Hawksworth, 1987; Hawksworth & Wiens,
1972; Wood, 1986). Its principal host is Tsuga
heterophylla. Abies amabilis and A. procera are
frequently parasitized, but usually only where these
trees are associated with infected /’. heterophylla.
This race is rare on associated Tsuga mertensiana
and Pinus monticola. Inoculation tests have shown
that several other trees from outside the natural
range of western hemlock race are susceptible (Smith,
1965, 1970a, b, 1972, 1974; Smith & Craig, 1968;
Smith & Wass, 1979; Weir, 1918). The western
hemlock race is distributed from elevations near sea
level in Alaska, British Columbia, Washington, Or¬
egon, and California to about 1,200 in in southern
Oregon.
2. The shore pine race is local in southern British
Columbia (on the east shore of Vancouver Island
and along the adjacent mainland coast) and on Orcas
Island, Washington (Hawksworth, 1987; Smith &
Wass, 1976; Wass, 1976). The northern limits of
the race are poorly known. Two additional collec¬
tions of a dwarf mistletoe on shore pine more than
450 km north of the main range of the shore pine
race have been reported: at Port Clements on the
Queen Charlotte Islands and at Terrace on the main¬
land (Wass, 1976). Studies of these populations are
needed to determine whether they are indeed the
shore pine race or merely rare crossovers ol the
western hemlock race. The principal host of the
shore pine race is Pinus contorta subsp. contorta.
It is usually rare on associated Tsuga heterophylla,
but in some areas on Vancouver Island both trees
are infected in mixed stands. Possibly these cases
represent mixed populations of the two races. The
shore pine race occasionally parasitizes associated
Pinus monticola (Kuijt, 1956). Inoculation tests
have shown that several other tree species from
outside the natural range of the shore pine race are
susceptible (Smith, 1970a, b, 1974; Smith & Craig,
1968; Smith & Wass, 1979). The shore pine race
is distributed from about 100 to 800 m.
Volume 2, Number 3
1992
Hawksworth et al.
Arceuthobium
209
Table 5. Differential parasitism of selected hosts of the three segregates of Arceuthobium tsugense.
Taxon
Tsuga
heterophylla
Tsuga
mertensiana
Pinus contorta
subsp. contorta
Pinus rnonticola
subsp. tsugense
western hemlock race
Common
Very rare
Rare
Rare
shore pine race
Rare
*
Common
Rare
subsp. rnertensianae
Immune
Common
*
Common
* Tree does not occur naturally within the range of the dwarf mistletoe.
Arceuthobium tsugense (Rosendahl) G. N. Jones
subsp. inertensianae Hawksworth & Nick-
rent, subsp. nov. TYPE: U.S.A. Oregon: Doug¬
las County, 16 mi. N of Union Creek on State
Highway 230, on Tsuga mertensiana, eleva¬
tion ca. 1,500 m, 2 Jan. 1990, H. L. Mathi-
asen 9002 (holotype, US; isotypes, FPF, MO,
ORE, OSC, UC, UWT).
Plantae 3-9 (5) cm altae; anthesis mense Augusto-
Septembri; fructus maturitas mensis Septembri-Octobri.
In Tsugo mertensiana parasiticae.
Plants similar to subspecies tsugense but are 3-
9 (mean 5) cm tall; anthesis in August and Septem¬
ber; fruits mature in September and October; par¬
asite principally on Tsuga mertensiana.
The two subspecies are generally similar mor¬
phologically, but in subspecies rnertensianae the
shoots average about 30 percent shorter than in the
western hemlock race of subspecies tsugense (Table
4). The differences are statistically highly significant
(t = 47.08, P = .0001). We do not have sufficient
material to make a comparable morphological anal¬
ysis of the shore pine race of A. tsugense.
The phenology of flowering of the two subspecies
is similar except that anthesis averages about 1 week
earlier in the western hemlock race of subspecies
tsugense (peak anthesis in August, with extremes
from late July to late September, 56 observations)
than for subspecies rnertensianae (peak anthesis
from mid-August to mid-September, 33 observa¬
tions). The phenology data are based on observations
throughout the range of the two taxa. In contrast
to flowering, the seed dispersal period for subspecies
tsugense (late September to early November, 57
observations) averages about 2 weeks later than for
subspecies rnertensianae (early September to late
October, 31 observations), thus the fruit maturation
period averages about 13 months in subspecies tsu¬
gense and 14 months for subspecies rnertensianae.
Perhaps the marked differences in susceptibility
of the two hemlocks to the two subspecies of the
parasite is explained by the wide taxonomic differ¬
ences of the two trees. For example. Page (1988)
indicated that mountain hemlock is such a distinct
entity that it should be segregated from Tsuga as a
monotypic genus, Hesperopeuce Lemmon. If this
classification is accepted, the scientific name for
mountain hemlock becomes Hesperopeuce merten¬
siana (Bong.) Rydb. The two genera are said to
differ in needle anatomy and morphology, shoot and
cone morphology, bark features, and pollen mor¬
phology (Page, 1988). In another study, Farjon
(1988) named the southern populations of Tsuga
mertensiana as subspecies grandicona (the cones
being 4-8 cm long, compared to 2-5 cm long for
cones of the typical subspecies, mertensiana). All
populations of Tsuga mertensiana from California
and Nevada are referable to subspecies grandicona.
The northern limits of this subspecies are not clear,
but it ostensibly occurs as far north as the Siskiyou
Mountains near the Califomia-Oregon border. Thus,
both subspecies are parasitized by A. tsugense subsp.
rnertensianae, but the mistletoe is much more com¬
mon within the range of subspecies grandicona. In
any case, the strong taxonomic differences between
the two species of hemlock may help explain the
unique host relations of the two subspecies of A.
tsugense.
The three segregates of A. tsugense are most
easily distinguished by their differential parasitism
of Tsuga heterophylla, Tsuga mertensiana , Pinus
con tort a subsp. contort a, and Pinus rnonticola (Ta¬
ble 5). Further information on the host relationships
of these mistletoes will be given in our revised mono¬
graph.
Arceuthobium tsugense subsp. rnertensianae
ranges from extreme southern British Columbia to
the central Sierra Nevada in California, but it is
most common from central Oregon southward. Its
elevational range is from about 800 m in British
Columbia to 2,500 m in central California. The
northern limits of subspecies rnertensianae have not
been determined, but it occurs at least as far north
as the Cypress Bowl area just north of Vancouver,
British Columbia (Van Sickle & Fiddick, 1982). It
is common at the Mt. Baker Ski Area in northern
Washington. There are a number of collections, of
210
Novon
what is presumably this dwarf mistletoe, on the east
side of the Cascade Mountains in Chelan County,
Washington, hut the distribution of the taxon there
is poorly documented.
Arceuthobium tsugen.se subsp. tsugense rarely
parasitizes Tsuga mertensiana in areas of Alaska
where this tree is associated with infected Tsuga
heterophylla (Shaw, 1982) and perhaps elsewhere.
The principal host of A. tsugense subsp. merten-
sianae is Tsuga mertensiana, and we have not found
it parasitizing Tsuga heterophylla, even where this
tree is closely associated with infected 7. merten¬
siana. The mistletoe also parasitizes Firms monti-
cola, P. albicaulis, Abies procera, A. amabilis,
and A. lasioearpa, but usually only in areas where
these trees are closely associated with infected Tsuga
mertensiana.
Additional specimens examined: on Tsuga merten¬
siana, except as noted. U.S.A. CALIFORNIA: Alpine Co.,
Mosquito Lake, 11 mi. NE of Alpine, Hawksworth &
Scharpf 665 (FPF), and on Pinus monticola, Hawk¬
sworth & Scharpf 666 (FPF). Placer Co., Emigrant Gap,
Jones, 1881 (POM). Plumas Co., Mt. Elwell, Lieberg
5363 (ILL, US); 0.4 mi. NE of Hawlsey Lake, Mathiasen
8513 (FPF). Shasta Co., Lassen National Park, 2.5 mi.
N of S entrance, Wiens 3235 (COLO, FPF); 0.7 mi. E
of Kings Creek Ground on Rte. 89, Mathiasen 8511
(FPF). Sierra Co., Hawlsey Lake, 3 mi. W of Gold Lake,
on Pinus monticola, Mathiasen 8514 (FPF). Siskiyou
Co., Marble Mtns., Kidder Creek, Gill <£' Sargent
FP68l80 (FPF); Chimney Rock Lake, Hemphill, 1968
(FPF, UT) and on Picea breweriana, Hemphill, 1968
(FPF, LIT); near Chimney Rock, on Pinus monticola,
Hemphill, 1968 (FPF, UT); Little Marble Valley, Ma¬
thiasen 8623 (FPF); Siskiyou Mtns., Applegate Creek
Divide, Meinecke FP97938 (PFRS) and on Picea brew¬
eriana, Meinecke FP9794I (PFRS); N Slope of W hite
Mtn., Mathiasen 8629 (FPF), and on P. monticola,
Mathiasen 8628 (FPF); Trinity Alps, Snowslide Lake,
Mathiasen 9016 (FPF). Tehama Co., Mineral, Lory
FP97940 (PFRS). Trinity Co., 1-2 mi. S of Helen Lake,
Scharpf FP38027 (PFRS). OREGON: Clackamas Co., near
Frog Lake, Wapinita Pass, Mathiasen 8643 (FPF). Coos
Co., Iron Mtn., Baker, 1946 (ID, OSC); 10 air mi. NNE
of Agness, Graham, 1965 (FPF). Curry Co., Snow Camp
Mt n., on Pinus monticola, Nickrent & Wiens 2701 (FPF,
ILL). Deschutes Co., 11 mi. W of Sisters on McKenzie
Pass road, Hawksworth & Hinds 991 (FPF), and on
Abies lasioearpa, Hawksworth & Hinds 992 (FPF); 1
mi. W r of Elk Lake near trail to Horse Lake, Mathiasen
8645 (FPF). Douglas Co., just N of Crater Lake National
Park on Diamond Lake road, Graham, 1964 (FPF); F
side of Diamond Lake, 1 mi. N of jet. ol Rtes. 230 &
138 on 138, Mathiasen 8616 (FPF); 4 mi. SW of Win-
digo Pass on Lemolo Lake road, Hawksworth 2339 (FPF),
and on Pinus monticola, Hawksworth 2340 (FPF). Jack-
son Co., Union Creek, Applegate 6038 (CAS); Huckle¬
berry Mtn., 10 mi. NE of Prospect, Graham, 1965 (FPF).
Klamath Co., Crater Lake National Park, Annie Springs,
Gill FP68184 (FPF); Wineglass, on Pinus albicaulis.
Gill FP68182 (FPF); 5 mi. E of main road on N rim of
Crater Lake, Hawksworth & Wiens 624 (FPF), and on
Pinus albicaulis, Hawksworth & Wiens 625 (FPF); 2
mi. N of Windigo Pass, Hawksworth A W iens 621 (FPF)
and on Pinus monticola, Hawksworth & Wiens 622
(FPF); 8 air mi. N of Lake of the Woods on Cold Springs
road, Hawksworth 2375 (FPF). Lane Co., McKenzie
Pass, on Pinus albicaulis. Gill FP68188 (FPF). Linn
Co., Minto Mtn., Mielke FP40694 (OSC); Marion Lake,
Coville & Applegate 1156 (US); Wildcat Mtn., Hawk¬
sworth 2358 (FPF), on Abies amabilis, Hawksworth
2355 (FPF), and on Abies procera, Hawksworth 2356
(FPF); H. J. Andrews Experimental Forest, near Frissell
Point, Hawksworth 2359 (FPF), on Abies amabilis,
Hawksworth 2360 (FPF), and on Abies procera, Hawk¬
sworth 2361 (FPF). WASHINGTON: Whatcom Co., Mt.
Baker Ski Area, Mathiasen 9005 (FPF), and on Abies
amabilis, Mathiasen 9006 (FPF); Twin Sister Range,
Meunschler 10431 (CAS). Chelan Co., Lyman Glacier
W of Lucerne, Weir 2456 (ILL). Pierce Co., Upper
Nisqually Valley, Allen 303 (CAS, MO, UT). Canada.
BRITISH COLUMBIA: Cypress Bowl Park, Hollyburn rd.,
Wood, 1982 (DAVFP); Lost Lake, Brothers Creek, ad¬
jacent to Cypress Bowl Park, Alexander, 1972 (DAVFP).
Acknowledgments. We thank Robert L. Mathi¬
asen for making the type collections for two of the
new taxa and Richard B. Smith, Robert F. Scharpf,
Robert (). Tinnin, Robert L. Mathiasen, Donald M.
Knutson, and Ed Wass for information and manu¬
script reviews. We also thank Job Kuijt for criticisms
of an earlier draft of the manuscript and John Hough
for assistance with the Latin descriptions.
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- & -. 1976. Arceuthobium oxycedri
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clatural changes, and range extensions in Mexican
Arceuthobium (Viscaceae). Phytologia 66: 5-11.
Hunt, R. S. & R. B. Smith. 1978. Natural infection of
two new hosts by hemlock dwarf mistletoe in British
Columbia. Canada Plant Dis. Surv. 58(2): 31-32.
Volume 2, Number 3
1992
Hawksworth et al.
Arceuthobium
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Kuijt, J. 1956. A new record of dwarf mistletoe on
lodgepole pine and western white pine. Madrono 13:
170-172.
Mark, W. R. & F. G. Hawksworth. 1981. Taxonomic
implications of branching patterns in the dwarf mis¬
tletoes (Arceuthobium). Phytologia 48: 441-444.
Mathiasen, R. L. & F. G. Hawksworth. 1988. Dwarf
mistletoes on western white pine and whitebark pine
in northern California and southern Oregon. Forest
Sci. 34: 429-440.
Nickrent, D. L. 1986. Genetic polymorphism in the
morphologically reduced dwarf mistletoes ( Arceu¬
thobium , Viscaceae). Amer. J. Bot. 73: 1492-1502.
-. 1987. Systematics and population biology of
two sibling species of Arceuthobium (dwarf mistle¬
toes, Viscaceae). Pp. 597-611 in H. Chr. Weber
& W. Forstreuter (editors), Proc. IV International
Symposium Parasitic Flowering Plants, Marburg,
West Germany.
-. 1988. Electrophoretic characterization of host
races in hemlock dwarf mistletoe ( Arceuthobium tsu-
gense). Amer. J. Bot. 75(6), Part 2: 196. [Abstract.]
- & T. L. Butler. 1989. Electrophoretic evi¬
dence supports the recognition of a new species of
Arceuthobium (Viscaceae) from coastal California.
Amer. J. Bot. 76(6): 261-262 (supplement). [Ab¬
stract.]
- & -. 1990. Allozymic relationships of
Arceuthobium campylopodum and allies in Califor¬
nia. Biochem. Systematics and Ecol. 18: 253-265.
- & -. 1991. Genetic relationships in
Arceuthobium monticola and A. siskiyouense (Vis¬
caceae): new dwarf mistletoes from California and
Oregon. Biochem. Systematics and Ecol. 19: 305-
313.
- & A. Stell. 1990. Electrophoretic evidence for
genetic differentiation in two host races of hemlock
dwarf mistletoe (Arceuthobium tsugense, V iscaceae).
Biochem. Systematics and Ecol. 18: 267-280.
-, S. I. Guttman & W. H. Eshbaugh. 1984.
Biosystematics and evolutionary relationships among
selected taxa of Arceuthobium. Pp. 20-35 in Gen.
Techn. Rep. RM-111, Biology of Dwarf Mistletoes,
Proc. of the Symposium. USDA Forest Serv., Fort
Collins, Colorado.
Page, C. N. 1988. New and maintained genera in the
conifer families Podocarpaceae and Pinaceae. Notes
Roy. Bot. Card. Edinburgh 45: 377-395.
Shaw, C. G., III. 1982. Mountain hemlock is occasional
host for hemlock dwarf mistletoe in Alaska. Plant
Dis. 66: 852 854.
Smith, J. P., Jr. & J. O. Sawyer, Jr. 1988. Endemic
plants of northwestern California and southwestern
Oregon. Madrono 35: 54-69.
Smith, R. B. 1965. Some results of artificial inoculation
with western hemlock dwarf mistletoe seed. Canad.
Dept. Forest., Bi-Monthly Progr. Rep. 21(6): 3-4.
-. 1970a. Infection of western larch by hemlock
dwarf mistletoe. Canad. Dept. Fish. Forest., Bi-
Monthly Res. Notes 26: 16-17.
-. 1970b. Infection of Engelmann spruce by
hemlock dwarf mistletoe. Canad. Dept. Fish. Forest.,
Bi-Monthly Res. Notes 26: 14.
-. 1971. Development of dwarf mistletoe (Ar¬
ceuthobium) infections on western hemlock, shore
pine and western larch. Canad. J. Forest. Res. 1:
35-42.
-. 1972. Relative susceptibility of coastal and
interior western hemlock to hemlock dwarf mistletoe
(Arceuthobium tsugense). Canad. Dept. Environ.,
Forest. Serv., Bi-Monthly Res. Notes 28: 17-18.
-. 1974. Infection and development of dwarf
mistletoe on plantation-grown trees in British Colum¬
bia. Canad. Forest. Serv., Pacific Forest Res. Centre
Rep. BC-X-97.
- & H. M. Craig. 1968. Infection of Scots,
Monterey and ponderosa pines by western hemlock
dwarf mistletoe. Canad. Dept. Forest. Rural Devel¬
opment, Bi-Monthly Res. Notes 24: 10-11.
- & E. F. Wass. 1976. Field evaluation of
ecological differentiation of dwarf mistletoe on shore
pine and western hemlock. Canad. J. Forest Res. 6:
225-228.
- & -. 1979. Infection trials with three
dwarf mistletoe species within and beyond their known
ranges in British Columbia. Canad. J. PI. Pathol. 1:
47-57.
Van Sickle, G. A. & R. L. Fiddick. 1982. Pacific
Region. Pp. 71-79 in Canad. Forest. Serv., Forest
Insect and Dis. Surv. Annual Rep. 1979.
Wass, E. F. 1976. Ecological study of shore pine stands
infested with dwarf mistletoe on southeastern Van¬
couver Island. Canad. Forest. Serv., Pacific Forest
Res. Centre Rep. BC-X-142.
Weir, J. R. 1918. Experimental investigations on the
genus Razoumofskya. Bot. Gaz. (Crawfordsville) 56:
1-31.
Wood, C. 1986. Distribution maps of common tree
diseases in British Columbia. Canad. Forest. Serv.,
Pacific Forest Centre Inform. Rep. BC-X-281.
New Combinations in Stigmatodactylus (Orchidaceae)
Paul Kores
Ecology, Evolution, and Organismal Biology Department, Tulane University,
New Orleans, Louisiana 70118, U.S.A.
ABSTRACT. Recent anatomical and morphological
studies of the Acianthinae indicate that the four
species of Acian thus from Papuasia are better placed
within Stigmatodactylus. This transfer requires three
new combinations.
Stigmatodactylus Maxim, ex Makino is a small
genus of ground orchids that was established in
Makino’s (1891) Illustrations of the Flora oj Ja¬
pan. As currently delimited, it contains approxi¬
mately seven species and is found in Japan, northern
India, Java, Borneo, the Celebes, the Moluccas, New
Guinea, and the Solomon Islands (Bougainville and
Guadalcanal). The genus is closely allied to Acian-
thus R. Br. and townsonia Cheesm., but is readily
distinguished from these genera by its massive, sub¬
entire or 3-lohed basal callus on the labellum, ven-
trally keeled or appendaged column, and prominent
dactyliform appendage on the posterior margin of
the stigma. In addition, Stigmatodactylus has ax¬
illary subterranean tubers (as opposed to terminal
ones in Acianthus); the plants are unifoliate (as
opposed to multifoliate in Townsonia); and the cells
that line the anther have type IV endothecial thick¬
enings (Freudenstein, 1991) (as opposed to type I
or type life thickenings in Acianthus or type I
thickenings in Townsonia).
After examining these genera in detail, I have
concluded that the three species I previously de¬
scribed from Papuasia (Kores, 1991) are incorrectly
placed within Acianthus and should be transferred
to Stigmatodactylus. New combinations are here
proposed for the three species not previously trans¬
ferred to Stigmatodactylus. It should also be noted
that with the exclusion of the Papuasian taxa from
Acianthus, the geographic distribution of the genus
is restricted to New Caledonia, New Zealand, and
Australia.
Stigmatodactylus gibbsae (Kores) Kores, comb,
nov. Basionym: Acianthus gibbsae Kores in
Lindleyana, 6: 166, f. 2. 1991.
Stigmatodactylus croftianus (Kores) Kores,
comb. nov. Basionym: Acianthus croftianus
Kores in Lindleyana, 6: 168, f. 3. 1991.
Stigmatodactylus variegatus (Kores) Kores,
comb. nov. Basionym: Acianthus variegatus
Kores in Lindleyana, 6: 171, f. 4. 1991.
Literature Cited
Freudenstein, J. V. 1991. A systematic study of en¬
dothecial thickenings in the Orchidaceae. Amer. J.
Bot. 78: 766-781.
Kores, P. 1991. A revision of the genus Acianthus
(Orchidaceae) in Papuasia. Lindleyana 6: 162-173.
Makino, T. 1891. Illustrations of the Flora of Japan,
vol. 1, fasc. 7: 70, 81-82, pi. 43. Keigyosha, Tokyo.
Novon 2: 212. 1992.
New Combinations in Californian Lessingia (Compositae: Astereae)
Meredith A. Lane
R. L. McGregor Herbarium and Department of Botany, University of Kansas,
Lawrence, Kansas 66047, U.S.A.
Abstract. As treated for the forthcoming revision
of the Jepson Manual of the Flowering Plants of
California, there are 14 species of Lessingia Cham,
in California. Benitoa Keck and Corethrogyne DC.
are considered to be congeneric with Lessingia.
A forthcoming publication will provide more sup¬
porting data for the transfer of Benitoa occidentalis
(H. M. Hall) Keck, Corethrogyne filaginifolia
(Hook. & Arn.) Nutt., and C. californica DC. into
Lessingia Cham., and discuss the treatment of Les¬
singia by Spence (1963). However, new combina¬
tions to accommodate these three taxa in Lessingia
are needed for the revision of the Jepson Manual
of the Flowering Plan ts of California. A brief jus¬
tification for these transfers follows.
Benitoa Keck has an annual habit, turbinate in¬
volucres, and disk corollas that are funnelform, yel¬
low tinged with deep purple-red, and deeply lobed,
and style-branch appendages that are linear; it shares
these characters with several undisputed species of
Lessingia (Howell, 1929; Spence, 1963). Benitoa
is monotypic, and is found on serpentine-derived
soils in southern San Benito, southwestern Fresno,
and southeastern Monterey counties (Keck, 1956).
Though Benitoa can be a much larger plant than
most lessingias, in branching pattern, leaf mor¬
phology, pubescence, and glandularity it is very
similar to them.
Corethrogyne DC. taxa have a perennial her¬
baceous habit, turbinate to hemispheric involucres,
disk corollas that are linear, yellow, and shallowly
lobed, and style-branch appendages that are tufted;
they share these characters with several other un¬
disputed species of Lessingia (Spence, 1963). Co¬
rethrogyne has been treated as having various num¬
bers of taxa (Canby, 1927; Howell, 1929; Keck.
1 959), because it is extremely variable both within
and between populations. Here it is treated as having
a single species with two highly polymorphic vari¬
eties, based on an unpublished revision of Corethro¬
gyne by J. P. Saroyan, D. R. Parnell, and J. L.
Strother. The range of variation in vestiture of the
foliage, the capitulescence, and the shape ol the
involucres of Corethrogyne taxa is all encompassed
by that of Lessingia.
The three genera have been maintained in the
past because taxa of Lessingia sensu stricto are all
eradiate and the others have ray florets, and because
the rays of Benitoa are yellow while those of Coreth¬
rogyne are purple. The presence or absence of ray
florets is probably only a single-gene character (Jack-
son & Dimas, 1981; Gottlieb. 1984). Given that
species of Lessingia have yellow, white, pink, or
purple disk corollas, it does not seem logical to use
corolla color as a generic marker. Also, the members
of all three genera have phyllaries with resin-canals
and (usually) gland-tipped apices, obconic, trinerved,
mottled achenes with a pappus of several (late-)de-
ciduous bristles, and chromosome numbers of n =
5. In addition, they share at least 20 unique re¬
striction site mutations (and differ among themselves
in none) found among 63 taxa of Astereae in a
chloroplast DNA study using 17 enzymes (M. Lane,
unpublished data).
Lessingia filaginifolia (Hook. & Arn.) M. A.
Lane, comb. nov. Basionym: Aster ? filagini-
folius Hook. & Arn., Bot. Beech. Voy. 146.
1833. Corethrogyne californica DC. [var. \ fi¬
laginifolia (Hook. & Arn.) Kuntze, Rev. Gen.
PI. 1: 330. 1891, illegit. TYPE: U.S.A. [Cal¬
ifornia]: presumably Lay and/or Collie s.n.
(Beechey voyage) (holotype, E fide Saroyan et
ah).
Kuntze’s combination is illegitimate because its
basionym. Asterfilaginifolius, is older than Coreth¬
rogyne californica.
Lessingia filaginifolia (Hook. & .Arn.) M. A.
Lane var. californica (DC.) M. A. Lane, comb,
et stat. nov. Basionym: Corethrogyne califor¬
nica DC., Prodr. 5: 215, 1836. TYPE: “Nova
California,” Douglas, 1833 (holotype, C-DC
fide Saroyan et ah; isotypes, BM, K fide Sa¬
royan et ah; microfiche, KANU).
Lessingia occidentalis (H. M. Hall) M. A. Lane,
comb. nov. Basionym: Haplopappus occiden¬
talis H. M. Hall, Carnegie Inst. Wash. Publ.
389: 214. 1928. Benitoa occidentalis (II. M.
Hall) Keck, Leaf!. W. Bot. 8: 25-40. 1956.
Novon 2: 213-214. 1992.
214
Novon
TYPE: IJ.S.A. California: Fresno Co., E side
of summit of Parkfield Crade, Hall 11769 (ho-
lotype, UC).
Acknowledgments. Supported in part by NSF
grant BSH 89-08963. I thank John Strother, Hon
Hartman, John Semple, Creg Brown, J. P. Saroyan,
and I). R. Parnell for assistance.
Literature Cited
Canby, M. L. 1927. The genus Corethrogyne in south¬
ern California. Bull. S. Calif. Acad. Sci. 26: 8 16.
Gottlieb, L. D. 1984. Genetics and morphological evo¬
lution in plants. Amer. Naturalist 123: 681-709.
Howell, J. T. 1929. A systematic study of the genus
Lessingia Cham. Univ. Calif. Publ. Bot. 16: 1-44.
Jackson, R. C. & C. T. Dimas. 1981. Experimental
evidence for systematic placement of the Haplo-
pappus phyllocephalus complex (Compositae). Syst.
Bot. 6: 8 14.
Keck, D. D. 1956. Benitoa, a new genus of Compositae
from California. Leaf!. W. Bot. 8: 25 40.
-. 1959. Corethrogyne. Pp. 1204-1207 in P.
A. Munz, A California Flora. Univ. California Press,
Berkeley & Los Angeles.
Spence, W. L. 1963. A biosystematic study of the genus
Lessingia Cham. (Compositae). Unpublished Ph.D.
dissertation, University of California, Berkeley. (Uni¬
versity Microfilms order no. 64-2136.)
New Combinations in Californian Grindelia (Compositae: Astereae)
Meredith A. Lane
R. L. McGregor Herbarium and Department of Botany, University of Kansas,
Lawrence, Kansas 66047, U.S.A.
ABSTRACT. As treated for the forthcoming revision
of the Jepson Manual of the Flowering Plants of
California, there are six species of Grindelia in
California. Hybridization among them is common,
and is apparently the reason for proliferation of
epithets in the group. Thorough study of over 6,000
specimens (including the types for all epithets in¬
volved) from 34 herbaria has necessitated the rec¬
ognition of several new combinations, and synony-
mization of many others.
Grindelia Willd. (Compositae: Astereae) is even
more highly variable in California than elsewhere in
its range in the western United States, Mexico, and
South America. Several of the taxa apparently hy¬
bridize freely when they are in contact; this is the
primary source of the taxonomic confusion that has
led to the erection of more than 60 epithets for taxa
of California alone. Though a more thorough dis¬
cussion of the systematics of this group will be pub¬
lished elsewhere, it is necessary now to provide new
combinations that may be used in a treatment of
Grindelia for the forthcoming revision of the Jepson
Manual of the Flowering Plants oj California.
Based on examination of the types for all the bas-
ionyms involved and over 6,000 specimens from 34
herbaria, I have come to the conclusion that it is
most reasonable to recognize six species for the state.
One of these, Grindelia squarrosa (Pursh) Dunal,
represented in California by G. squarrosa var. ser-
rulata (Rydberg) Steyermark, is probably adventive,
and occurs in only a few localities. It is not a major
player in the hybridization reticula, and is easily
distinguished from other California specimens by its
bluntly dentate-serrulate leaf margins.
Another species, Grindelia fraxino-pratensis Re¬
veal & Beatley, is known only from the Ash Mead¬
ows area of Nye Co., Nevada, and Inyo Co., Cali¬
fornia, and is not involved in hybridizations with
other taxa. The involucres of G. fraxino-pratensis
are broadly turbinate; all other grindelias in Cali¬
fornia have broadly campanulate to hemispheric
heads.
Grindelia nana Nutt, is found in northeastern
California, and though it is variable and probably
participates in hybridizations with other taxa, the
nomenclatural situation surrounding it (in California,
at least) is straightforward. Steyermark (1934: 541-
546) treated it as having three varieties in California;
I agree with Keck (1959: 1 166) that there are
insufficient characters for such subdivision of the
taxon. Plants of this species may be recognized by
the tightly coiled phyllaries, coppery-red stems, and
sharply serrate leaves.
The remaining three species have been treated
differently by each worker who has encountered
them, and for that reason synonymies for each new
and recognized combination are included here. 1
have adopted a broad species-concept, accepting a
wide range of variability for each species; geograph¬
ical components of the variability have been rec¬
ognized at the varietal level. The characters that
distinguish the taxa are few enough and intergrade
sufficiently that the recognition of more taxa than
those listed below is unwarranted. Combinations that
have been applied to California specimens of Grin¬
delia but that do not appear in the lists of synonyms
below represent hybrids, and cannot legitimately be
assigned as synonyms to either of the parent taxa.
Grindelia stricta DC. and its varieties are coastal
in distribution and are found on beaches and at
the edges of marshes and sloughs. They have very
robust pappus awns (> 0.3 mm wide at base) that
are generally V-shaped in cross section, and the
leaves are usually more or less succulent. The va¬
rieties are distinguished by habit of the plant and
shape of the leaves. The other two species have
slender pappus members (< 0.3 mm wide) and are
mostly found inland. Grindelia camporum E. Greene
and its variety have stiff phyllaries usually with
reflexed apices, mostly white to buff-colored stems
and light green, very resinous leaves, and are found
in the San Francisco Bay area, the Great Valley,
and the lower foothills of the Sierra Nevada. Grin¬
delia hirsutula Hook. & Aril, and its varieties
usually have somewhat flexible phyllaries with as¬
cendant to reflexed apices, coppery-buff to deep red-
brown stems, and leaves that are variously invested
hut not highly resinous; they are found in the San
Francisco Bay area and the Coast Ranges.
Novon 2: 215-217. 1992.
216
Novon
Grindelia camporum E. Greene, Man. Bot. S.
F. Bay Beg. 171. 1894. TYPE: U.S.A. Cali¬
fornia: Antioch, Greene, 1881 (lectotype, MO,
selected here).
Grindelin robusta Nutt. var. (?) rigida A. Gray, Geol.
Surv. Cal. Bot. 1: 304. 1876. TYPE: U.S.A. Cal¬
ifornia: (holotype, GH).
Grindelin procera E. Greene, Man. Bot. S. F. Bay Reg.
171. 1894. TYPE: U.S.A. California: San Joaquin
Co., Banta, llioletti, 1892 (lectotype, selected here,
TJC). [Contrary to Howell (1931), I agree with Jepson
(1901) that this is a synonym of G. camporum.]
Grindelin camporum E. Greene var. parviflora Steyer-
mark, Ann. Missouri Bot. Gard. 21: 534. 1934.
TYPE: U.S.A. California: Contra Costa Co., Liberty
School on Marsh Creek road, Howell 549b (holo¬
type, CAS; isotypes, GH, 1ND not seen, MO).
Grindelin camporum E. Greene var. abbreviata Stey-
ermark, Ann. Missouri Bot. Gard. 21: 228. 1934.
TYPE: U.S.A. California: Los Angeles Co., Lancas¬
ter, Hoffmann, 1927 (holotype, SBBG; photograph,
MO).
Grindelin camporum E. Greene var. australis Steyer-
mark. Anti. Missouri Bot. Gard. 21: 228. 1934.
TYPE: U.S.A. California: Los Angeles Co., 3 mi. E
of Elizabeth Lake, Munz & Johnston 11157 (ho¬
lotype, POM; isotype, LL; photograph, MO).
Grindelia camporum E. Greene var. bracteosa
(J. Howell) M . A. Lane, comb. nov. Basionym:
Grindelin bracteosa J. Howell, Madrono 2: 22.
1931. G. rubricaulis DC. var. bracteosa (J.
Howell) Steyermark, Ann. Missouri Bot. Gard.
21: 227. 1934. G. robusta Nutt. var. brac¬
teosa (J. Howell) Keck, Aliso 4: 102. 1958.
TYPE: U.S.A. California: Orange Co., N side
of Santa Ana Canyon, J. Ilowell 2786 (holo¬
type, CAS; isotypes, CAS, MO, BSA; photo¬
graphs, F, GH, US).
Grindelin rubricaulis DC. var. elata Steyermark, Ann.
Missouri Bot. Gard. 21: 227. 1934. Syn. nov. TYPE:
U.S.A. California: Ventura Co., 7 mi. SW of Simi,
J. Howell 6574 (holotype, CAS; photograph, MO).
Grindelia hirsutula Hook. & Arn., Bot. Beech.
Voy. 147. 1833. TYPE: U.S.A. California: in
vicinity either of Monterey or of San Francisco
Bay (A. Gray in Geol. Surv. Gal. Bot. 1: 304.
1876), Lay and/or Collie s.n. (holotype, E;
isotype, K; photographs, GH, UC).
Grindelia humilis Hook. & Arn., Bot. Beech. Voy. 147.
1833. Syn. nov. TYPE: U.S.A. California: in vicinity
either of Monterey or of San Francisco Bay (A. Gray
in Geol. Surv. Cal. Bot. 1: 304. 1876), Lay and/
or Collie s.n. (holotype, E not seen; isotype, K;
fragments, CAS, GH; photographs, GH, LJS). [In the
past, this name consistently has been misapplied,
based on a misinterpretation of the type; however,
the type is clearly nonspecific with G. hirsutula. The
plant that has been called G. humilis is G. stricta
var. angustifolia.]
Grindelia rubricaulis DC., Prod. 5: 316. 1836. Syn.
nov. G. hirsutula Hook. & Arn. subsp. rubricaulis
(DC.) Keck, Aliso 4: 102. 1958. TYPE: U.S.A.
California: Douglas 55 (holotype, G-DC; isotypes,
BM, K; fragment, GH; photographs, GH, MO, UC).
Grindelia pacifica M. E. Jones, Bull. Torrey Bot. Club
9: 31. 1882. TYPE: U.S.A. California: Santa Cruz
Co., Santa Cruz, M. E. Jones 2750 (holotype, POM;
isotypes, GH, NY).
Grindelia hirsutula Hook. & Arn. var. subintegra Stey¬
ermark, Ann. Missouri Bot. Gard. 21: 229. 1934.
TYPE: U.S.A. California: Ventura Co., 2 mi. E of
Ojai, J. Howell 11414 (holotype, MO; isotypes,
CAS—2, F, GH, K, LL, NY).
Grindelia hirsutula Hook. & Arn. var. brevisquama
Steyermark, Ann. Missouri Bot. Gard. 21: 572.
1934. TYPE: U.S.A. California: Sonoma Co., Black
Point Road, /. Howell 5338 (holotype, CAS; isotype,
UC; photograph, MO).
Grindelia hirsutula Hook. & Arn. var. calva Steyer¬
mark, Ann. Missouri Bot. Gard. 21: 575. 1934.
TYPE: U.S.A. California: San Luis Obispo Co., San
Luis Obispo, Roadhouse 65 (holotype, UC; photo¬
graph, MO).
Grindelia hirsutula Hook. & Arn. var. davyi
(Jepson) M. A. Lane, comb. nov. Basionym: G.
robusta Nutt. var. davyi Jepson, FI. W. Mid.
Cal. 554. 1901. G. camporum E. Greene var.
davyi (Jepson) Steyermark, Ann. Missouri Bot.
Gard. 21: 534. 1934. TYPE: U.S.A. Califor¬
nia: Alameda Co., Berkeley, Peralta Park, Davy,
1896 (holotype, JEPS; isotypes, UC—2; pho¬
tographs, F—3, GH—2, MO).
Grindelia rubricaulis DC. var. interioris Jepson, Man.
FI. PI. Cal. 1021. 1925. G. camporum E. Greene
var. interioris (Jepson) Steyermark, Ann. Missouri
Bot. Card. 21: 534. 1934. TYPE: U.S.A. California:
Calaveras Co., Copperopolis, Davy 1383 (holotype,
UC; photograph, F).
Grindelia hirsutula Hook. & Arn. var. hallii
(Steyermark) M. A. Lane, comb. nov. Basio¬
nym: G. hallii Steyermark, Ann. Missouri Bot.
Gard. 21: 229. 1934. TYPE: U.S.A. Califor¬
nia: San Diego Co., Cuyamaca Lake, Abrams
3957 (holotype, NY; isotypes, CAS, E, F, GH,
K, MO, POM).
Grindelia hirsutula Hook. & Arn. var. mariti-
ma (E. Greene) M. A. Lane, comb. nov. Bas¬
ionym: G. rubricaulis DC. var. maritima E.
Greene. Pittonia 2: 289. 1892. G. robusta
Nutt. var. maritima (E. Greene) Jepson, FI.
W. Mid. Cal. 554. 1901. G. maritima (E.
Greene) Steyermark, Ann. Missouri Bot. Gard.
21: 576. 1934. TYPE. U.S.A. California: San
Volume 2, Number 3
1992
Lane
Grindelia
217
Francisco Co., Point I.obos, E. Greene , 1892
(holotype, ND-G; isotvpe, UC).
Grindelia stricta DC., Prod. 7: 28. 1838. TYPE:
“in America boreali-occid., ad portum Mul-
grave,” Haenke s.n. (holotype, G-DC not seen;
photographs, CAS, GH, UC).
Grindelia blakei Steyermark, Ann. Missouri Bot. Card.
21: 567. 1934. G. stricta DC. subsp. blakei (Stey¬
ermark) Keck, Aliso 4: 102. 1958. TYPE: U.S.A.
California: Humboldt Co., Eureka, Moore & Stey¬
ermark 3686 (holotype, MO; isotypes, MO, UC).
Grindelia stricta DC. var. angustifolia (A. Gray)
M. A. Lane, comb. nov. Basionym: G. robusta
Nutt. var. angustifolia A. Gray, Geol. Surv.
Cal. Bot. 1: 304. 1876, as nom. nov. for G.
cuneifolia Nutt., Trans. Amer. Philos. Soc. n.s.
7: 315. 1841. TYPE: U.S.A. California: Santa
Barbara Co., Santa Barbara, Nuttall s.n. (ho¬
lotype, PH not seen; isotype, BM; fragment,
GH). [The type of Grindelia robusta Nutt, is
a hybrid between G. hirsutula var. hirsutula
and another variety of G. stricta; therefore,
the autonym does not take precedence.]
Grindelia stricta DC. var. platyphylla (E.
Greene) M. A. Lane, comb. nov. Basionym: G.
robusta var. platyphylla E. Greene, Pittonia
2: 289. 1892. G. rubricaulis DC. var. pla¬
typhylla (E. Greene) Steyermark, Ann. Mis¬
souri Bot. Gard. 21: 227. 1934. G. latifolia
Kellogg subsp. platyphylla (E. Greene) Keck,
Aliso 4: 102. 1958. TYPE: U.S.A. California:
Monterey Co., Monterey, Howe , 1892 (holo¬
type, UC; isotype, UC; photograph, MO).
Grindelia arenicola Steyermark, Ann. Missouri Bot. Gard.
21: 229. 1934. TYPE: U.S.A. California: Mendo¬
cino Co., 4.5 mi. N of Fort Bragg, Howell 5473
(holotype, CAS; isotypes, GH, MO, POM, UC).
Grindelia arenicola Steyermark var. pachyphylla Stey¬
ermark, Ann. Missouri Bot. Gard. 21: 596. 1934.
TYPE: U.S.A. California: Mendocino Co., Point Are¬
na, Hoivell 8106 (holotype, CAS; isotypes, GH, MO).
Grindelia stricta DC. var. procumbens Steyermark, Ann.
Missouri Bot. Gard. 21: 559. 1934. TYPE: U.S.A.
California: Humboldt Co., 10 mi. N of Trinidad,
Moore & Steyermark 3687 (holotype, MO; isotype,
MO).
Grindelia venulosa Jepson, Man. FI. PL Calif. 1021.
1925. G. stricta DC. subsp. venulosa (Jepson) Keck,
Aliso 4: 102. 1958. TYPE: U.S.A. California: Hum¬
boldt Co., Big Flat, Bolander 6493 (holotype, JEPS;
isotypes, K, US; photograph, F).
Acknowledgments. Supported by NSF grant BSR
89-08963 and a Grant-in-Aid of Research from the
University of Colorado Council on Research and
Creative Work. The curators of ABIZ, BM, CAN,
CAS, CM, E, F, GH, ID, IDS, INI), JEPS, K, LL,
MO, MONT, MONTH, ND-G, NY, ORE, OSC, PH,
POM, RM, RSA, SBBG, SD, TEX, UBC, UC, US,
UTC, V, WS, and WTU graciously loaned speci¬
mens and types. I thank Shannon Curry, Barbara
Ertter, Elva Lane, Jianwei Li, Carol She, John
Strother, Philip Thompson, Robyn Tierney, and Die¬
ter Wilken for assistance.
Literature Cited
Howell, J. T. 1931. Plantae Occidentales. —11. Ma¬
drono 2: 18 23.
Jepson, W. L. 1901. A Flora of Western Middle Cal¬
ifornia. Encina Publishing, Berkeley.
Keck, D. D. 1959. Grindelia. Pp. 1164-1166 in P.
A. Munz, A California Flora. Univ. California Press,
Berkeley & Los Angeles.
Steyermark, J. A. 1934. Studies in Grindelia. II. Ann.
Missouri. Bot. Gard. 21: 434-608.
Antiostelma (Asclepiadaceae), a New Genus from China
Li Ping-tao
South China Agricultural University, Guangzhou 510642, Guangdong,
People’s Republic of China
ABSTRACT. Antiostelma (Asclepiadaceae), previ¬
ously recognized as a section of Hoy a, is raised to
generic rank. The new combinations Antiostelma
lantsangense (Tsiang & P. T. Li) P. T. Li and A.
manipurense (D. B. Deb) P. T. Li are proposed.
As a result of recent study of Chinese material
of Asclepiadaceae and Apocynaceae from the Mis¬
souri Botanical Garden, the Smithsonian Institution,
Harvard University, and the California Academy of
Sciences, and during preparation of the accounts of
these families for the Flora of China, it became
evident that the delimitation of certain genera need¬
ed critical study, and several nomenclatural adjust¬
ments needed to be made. This publication deals
with a new genus of Asclepiadaceae, Antiostelma.
It was originally described as a monotypic section
of Hoya R. Br. The new combinations are proposed
in order to make the names available for the Flora
of China and other floristic works in progress.
Antiostelma (Tsiang & P. T. Li) P. T. Li, comb,
et stat. nov. Basionym: Hoya sect. Antiostelma
Tsiang & P. T. Li, Acta Pbytotax. Sin. 12:
126. 1974. TYPE: Antiostelma lantsangensis
(Tsiang & P. T. Li) I’. T. Li.
Lacticiferous epiphytic subshrubs. Leaves oppo¬
site, obdeltoid or obovate. Cymes extra-axillary, um¬
bellate, subsessile or sessile. Calyx eglandular inside;
corolla cylindric, lobes erect, aestivation twisted left¬
ward; corona lobes 5, fleshy, erect, quadrate, adnate
adaxially to gynostegium, bilateral margin mani¬
festly recurved at back; stamens 5, anthers erect,
with acuminate apical membranes; pollinia subquad¬
rate, basal margin translucent; ovaries free; style
apex beaked, exceeding apical membrane of anthers.
Follicles linear-lanceolate. Seeds comose.
Antiostelma includes two species, A. lantsan¬
gensis and A. manipurense, that are endemic to
China and India, respectively. The genus was orig¬
inally recognized as a section of Hoya R. Br. (Tsiang
& Li, 1974, 1977), but critical examination of sev¬
eral collections reveal that it is best treated as an
independent genus readily distinguished from Hoya
and Dischidia by the several characters summarized
in I able 1.
The two species of Antiostelma are easily distin¬
guished by the following key.
la. Branches and leaves glabrous; leaf apex retuse;
China. A. lantsangense
lb. Branches and leaves densely pubescent; leaf
apex truncate; India . A. manipurense
Antiostelma lantsangense (Tsiang & P. T. Li)
P. T. Li, comb. nov. Basionym: Hoya lant¬
sangensis Tsiang & P. T. Li, Acta Pbytotax.
Sin. 12: 126. 1974. TYPE: China. Yunnan:
Table 1.
Comparison of Antiostelma, Hoya,
and Dischidia.
Antiostelma
Hoya
Dischidia
Corolla
cylindric
rotate
urceolate
Aestivation
twisted leftward
valvate
valvate
Corolla lobes
erect, much shorter than
tube
reflexed, as long as or
longer than tube
patent, shorter than tube
Corona lobes
erect, quadrate
stellate-patent, ovate or
elliptic
erect, anchor-shaped
Caudicle
not dilated
not dilated
apex dilated
Style apex
exserted from apical anther
membranes
included in apical anther
membranes
included in apical anther
membranes
Inflorescence
sessile or subsessile
long-pedunded
short-peduncled
Novon 2: 218-219. 1992.
Volume 2, Number 3
1992
Li Ping-tao
Antiostelma
219
Mo-Jiang, 1,000 m, A. Henry 13689 (holo-
type, NY; isotype, IBSC).
Antiostelma manipurense (D. B. Deb) P. T. Li,
comb. nov. Basionym: Hoyn manipurensis D.
B. Deb, J. Indian Bot. Soc. 34: 50. 1955.
TYPE: India. Manipur, Litan, 1 Sep. 1953,
D. H. Deb 1081 (holotype, CAL).
Acknowledgments. I am grateful to Peter H.
Haven, William Tai, W. Douglas Stevens, Laurence
Skog, Dan Nicholson, and David Boufford for their
support of my visit to the United States. I am also
grateful to Ihsan /U-Shehbaz for help with the manu¬
script.
Literature Cited
Tsiang, Y. & P. T. Li. 1974. Praecursers Flora As-
clepiadacearum Sinensium. Acta Phytotax. Sin. 12:
79-149.
- & -. 1977. Asclepiadaceae. Flora Rei-
publicae Popularis Sinicae 63: 475-559.
Note Added in Proof
M. Gilbert (MO) recently brought to our attention the
fact that Maxwell & van Donkelaar (Nat. Hist. Bull. Siam
Soc. 39: 78. 1991) have treated, with scant discussion,
Hoya lantsangensis as a synonym of Dischidia obcor-
data and thus, by implication, Hoya sect. Antiostelma,
which was termed “an erroneous and quite unnecessary
taxonomic complication,” as a synonym of Dischidia .—
Editor
Cynanchum wangii (Asclepiadaceae), a New Species from China
Li Ping-tao
South China Agricultural University, Guangzhou 510642, Guangdong,
People’s Republic ol China
Walter kittredge
Harvard University Herbaria, 22 Divinity Avenue, Cambridge,
Massachusetts 02138, U.S.A.
ABSTRACT. Cynanchum wangii, a new species from
Yunnan, China, is described, and a photograph of
the type specimen is given.
Cynanchum is a genus of about 200 species, of
which 53 species and 12 varieties were recognized
in China by Tsiang & Li (1077). With the addition
of five other species by P. T. Li (1983, 1990) and
C. wangii, the total number of species of Cynan¬
chum in China is now 59. The genus is difficult
taxonomically, and 1). Z. Li (1990) and Qiu (1989)
have treated it as several segregate genera. How¬
ever, most Chinese taxonomists (e.g., Tsiang & Li,
1977; P. T. Li, 1991) agree that the larger concept
represents a natural genus.
Cynanchum wangii P. T. Li & kittredge, sp.
nov. TYPE: China. Yunnan: Meng-Nun, Jenn-
Yeh Hsien, 750 m, riverbank, Nov. 1936, C.
If . Wang 80897 (holotype, A; isotypes, 1BSC,
KUN, PE). Figure 1.
Herba perennis erecta 10-20 cm alta; caule ramoso
puberulo. Folia linearia glabra, 5-20 x 1-2 mm. Cymae
3 7-florae, extra-axillares vel terminales; pedunculis gra-
cilibus 3-8 cm longis, puberulis. Flores calyx intus as
bases sinuum 5-glandulifero, lobis ovato-triangularibus,
glabris, ca. 0.5 x 0.5 mm; corolla viridi-flava, rotata,
extus glabra, intus dense pubescente; corona applanata
ad dorsum gynostegii adnata gynostemio breviore 5-par¬
tita, lobis semiorbiculatis, ca. 0.5 x 0.5 mm; staminibus
stigma aequantibus, antheris oblongis membranaceis, con-
nectivis orbiculatis, polliniis longe ovoideis, pendulis, re-
tinaculo oblongo, caudicula brevissima; ovariis glabris,
stigmate basi discoideo-pentagono; ovulis numerosis.
Erect perennial herbs 10 20 cm tall; stem many-
branched, ca. 1 mm diam.; uniformly puberulous,
internodes 0.8-3.5 cm long; branchlets delicate,
puberulous. Leaves opposite, linear, 5-20 x 1-2
mm, glabrous, base cuneate, apex rounded or obtuse;
midrib Hat on both surfaces, lateral veins obscure.
Cymes umbelliform or corymbose, 3- to 7-flowered,
extra-axillary or terminal, solitary; peduncles deli¬
cate, 3 8 cm long, puberulous; pedicels 4-7 mm
long. Calyx with 5 basal glands inside, calyx lobes
ovate-triangular, 0.5 x 0.5 mm, glabrous; corolla
greenish yellow, rotate, membranous, tube ca. 1
mm, glabrous, lobes oblong-lanceolate, ca. 4 mm
long, 1 mm wide, glabrous outside, densely pubes¬
cent inside; outer corona absent, inner one inserted
at base of gynostegium, 5-partite, lobes semiorbi-
cular, 0.5 x 0.5 mm, erect, somewhat fleshy, apex
rounded, nearly half as long as anther membrane;
anthers oblong, apical membrane rounded, as high
as stigma; pollinia oblong-ovoid, pendulous, retinac¬
ulum oblong, caudicle very short; ovaries long-ovoid,
glabrous; stigma pentagonal-disklike; apex slightly
elevated; ovules numerous in each carpel.
Flowering May through August, fruiting October
through November; grows along riverbanks at al¬
titudes of ca. 700-900 m.
Paratypes. China. YUNNAN: Mong-Hain, Gan-Lan-Ba,
Ching-Hung Hsien, Wang 79920(A)-, Lann-Tsang Hsien,
Wang 76710 (A).
Cynanchum wangii, which is so far only known
from three collections, is most closely related to C.
stenophyllum Hemsley, a species distributed in Hu¬
bei and Guizhou provinces, China. It is readily dis¬
tinguished from the latter by its small, slender leaves
1-2 mm wide, umbellate or corymbose inflores¬
cences longer than leaves, greenish yellow flowers,
oblong-lanceolate corolla lobes, and 5-partite corona
nearly half as long as the anther membrane.
Acknowledgments. We are grateful to David E.
Boufford, Bryan Dutton, and Ihsan Al-Shehbaz for
help with the manuscript. Thanks also to Hoy Gereau
for checking the Latin.
Literaure Cited
Li, D. Z. 1990. The chemotaxonomy of Cynanchum
and its allied genera. Acta Phytotax. Sin. 28: 461-
470.
Novon 2: 220-222. 1992.
Volume 2, Number 3
1992
Li Ping-tao & Kittredge
Cynanchum wangii
221
Collected In cooperation between the Arnold Arboretum of Harvard
University ami the Fan Memorial Institute of Biology.
Figure 1. Cynanchum wangii P. T. Li & Kittredge, C. If . Wang 80897 (holotype, A).
el
V.t. i: & <a. e.jrvw** ,
y..r, —- - /
No.
p. T Li vT
SOLTH CHINA AGRICLLTLR
,ANT8 OF YUNNAN PROVINCE, CHINA
e.ATr.^.* v [W1
SOLTH CHINA AGRICULTl RAl. UNIVERSITY (CANTip^. , .
?6tl7
.'Memorial Institute
'* of Biology
FLORA OF YUNRAN
PI.* NO..* < * W . __ l>.to HOVlja- «*»-
Locality «ggL.tt« (•••ST???.* *«*** ‘""I
. Altitude_Sfl... m-
H.bit.«_.„ ilcor t»nk..tin» rortca
Habit____i- 1 --
Height_
Flown __
Ftni*_- . -— --
F unity—A twin-
Col.ocunJE#« O. W.W-,
222
Novon
Li, P. T. 1983. Three new species of Asclepiadaceae
from China. Bull. Bot. Res. 3: 103-109.
-. 1990. New materials of Asclepiadaceae from
Chinese Hengduanshan Region. Acta Bot. Yunnan.
12: 19-21.
-. 1991. A new genus of Euphorbiaceae and
some new nomenclatural combinations of the Ascle¬
piadaceae plants. J. South China Agr. Univ. 12(3):
38-42.
Qiu, S. X. 1989. Chemotaxonomy of Cynanchum and
its allied genera, with notes on the generic charac¬
teristics of Vincetoxicum. Acta Bot. Yunnan. 11:
41-50.
Tsiang, Y. & P. T. Li. 1977. Cynanchum. Flora Rei-
publicae Popularis Sinicae 63: 309-384.
A New Species of Sarcostemma (Asclepiadaceae) from Malawi
Sigrid Liede and Ulrich Meve
Institut filr Botanik, Schlossgarten 3, W-4400 Munster, Germany; Liede’s present address:
Ahteilung Spezielle Botanik (Biologie V), Albert-Einstein-Allee II, W-7900 Ulm, Germany
Abstract. A new species of Old World Sarcos¬
temma , Sarcostemma mulanjense from Malawi, is
described.
Sarcostemma virninale (L.) R. Br. and its allies
are widespread throughout the arid areas of the Old
World. Because dried specimens rarely show such
key characters as habit, morphology of the stem
base, and floral color, the group is taxonomically
difficult and still lacks a clear-cut concept applicable
over its entire range. However, field-based studies
in more limited areas have resulted in the distinction
of several closely related species and subspecies
(Adams & Holland, 1978a, b, for East Africa; For¬
ster, 1988, for Australia). Recent fieldwork in Ma¬
lawi has resulted in the discovery of a new, clearly
delimited species endemic to the southern Malawi
mountains.
Specimens have been studied from BM, MO, and
NU. Fieldwork in Malawi (by S. Liede) has yielded
living material, cultivated at the Institut fiir Botanik,
Munster (Germany), Kenyatta University, Nairobi
(Kenya), and the National Botanic Gardens, Kir-
stenbosch (South Africa). Corona terminology, fully
explained in the description, is based on a recent
study of corona types in Asclepiadaceae and Peri-
plocaceae (Liede & Kunze, unpublished). To ac¬
count for the present unsatisfactory taxonomic sit¬
uation of the Old World species of Sarcostemma ,
the term “5. virninale and its allies” is used to
circumscribe all Old World members of the genus,
except for S. pearsonii N. E. Br., while the term
“S. virninale ” excludes the segregated species.
Sarcostemma mulanjense Liede & Meve, sp.
nov. TYPE: Malawi. Mt. Mulanje, outer slopes
at Chitakale stream, W branch, 15°57'S,
35°36'E, 1,250 m, 27 Nov. 1985, Chapman
& Chapman 6892 (holotype, MO). Figures
1 , 2 .
Sarcostemma virninale R. Br. affinis, sed differt sur-
culis omnino glabris, vivide viridis et altitudine gynostegii
quam diametro majore.
Plants succulent, without rhizome, not forming
an aboveground trunk, much branched basally, ex¬
uding white latex. Shoots to 1.5 m long, 3 5 mm
diam., bright green, not glaucous, totally glabrous,
erect when young, later bending over, but never
twining; internodes 15-40(-60) mm long; nodes
slightly flattened, without corky sheath; normally
not rooting at the nodes, heaves reduced to scales,
1 mm long, 1 mm wide, ovate, apiculate, green
when young, soon turning brown apically, deciduous.
Inflorescences terminal, umbelliform, 5-7-flowered,
sessile; pedicels 4-7 mm long, sparsely indumented
with trichomes 150-170 /am long. Buds conical,
3.5-4 mm long, 2.5-3 mm diam., petals imbricate
in bud, dextrorse. Flowers sweetly scented, scent
Philadelphus- like. Calyx rotate; sepals 0.9 mm long,
0.8 mm wide, ovate, sparsely indumented with tri¬
chomes 150-170 /am long. Corolla rotate; corolla
lobes fused only at the base, 4-6 mm long, 1.4-
1.6 mm wide, cream-colored with reddish tinged
base and median stripe, glabrous, patent to almost
horizontal at anthesis, oblong, apically blunt with a
tip, flat, slightly undulating, with slightly revolute
margins. Gynostegial corona white, consisting of
an outer ring of fused connate staminal (Gs) and
interstaminal (Ci) parts, (C (l ,,), plus five inner staminal
parts (Cs), connate to C |1S) . C (is) annular, Cs 1.4 mm
high, slightly cucullate, Ci 1 mm high, erect. Inner
Cs 2.8-3 mm high, shorter than the gynostegium,
cochleariform, connate to the back of the stamens
to about 2 A of anther height. Gynostegium sessile,
3.0-3.5 mm high, 1.4-2 mm diam., height clearly
exceeding the diameter. Stamens trapezoidal, abax-
ially rounded, filament differentiated, 1.8 mm high;
anther wings 0.6 mm long, clearly differentiated,
paralleling the anther, parallel to each other, basally
forming a distinct “moutb.” consisting of outer rail
alone; connective appendages 0.5 mm long, 0.5 mm
wide, debate, equaling the stamen in width, slightly
indexed. Pollinarium: corpusculum 200 220 /am
long; caudicles 60-80 yarn long, cylindrical, s-shaped,
convex-concave, slightly thickened at the insertion
of the pollinium; pollinia apically inserted, 280-380
/am long, 80-100 /im wide, elliptical in cross section,
pyriform. Style apex cream, 0.9-1 mm diam, 0.7-
0.8 mm high, upper part 0.4-0.5 mm high, higher
than the lower part, conical to depressed-conical.
Novon 2: 223-226. 1992.
224
Novon
Figure 1. Sarcostemma mulanjense Liede & Meve. —A. Inflorescence. —B. Corona and gynostegium. C.
Pollinarium (Liede 2900). Ci: interstaminal part of corona; Cs: staminal part of corona; C,„,: fused staminal and
interstaminal parts of corona.
Volume 2, Number 3
1992
Liede & Meve
Sarcostemma mulanjense
225
Figure 2. Sarcostemma mulanjense in habitat on the
Lichenya Plateau, Mt. Mulanje (Liede 2899). Note the
cushionlike growth form and the dead branches of the
last season.
Follicles and seeds unknown. Chromosome number:
2 n = 22 (Voucher: Liede 2900).
Distribution. Malawi, 15 17°S, 35-36°E; 900-
2,100 m; on granitic outcrops between sedges, Aloe,
and Velloziaceae (Figs. 2, 3).
Flowering time. November-April (according to
dates on flowering specimens).
Paratypes. MALAWI. MULANJE: Mt. Mulanje, approach
to Cilemba from the Likhubula valley, 15°57'S, 35°36'E,
2,100 m, 13 Mar. 1986, Chapman <£■ Chapman 7285
(MO); ascent from Likhubula via Chapaluka path, 19 Jan.
1967, Hilliard & Ilurtt 4504 (NU); Lichenya Plateau,
near the Crater, 1,900 m, 16 Apr. 1991, Liede 2899
(Inst. Bot. Minister); Lichenya Plateau, between hut and
the Crater, 1,900 m, 16 Apr. 1991, Liede 2900 (Inst.
Bot. Munster); foot near Likhubula Forest station, 900
m, 17 Apr. 1991, Liede 2902 (Inst. Bot. Munster).
ZOMBA: Nkhoronje Hill, 15 0 25'S, 35°20'E, 1,120 in, 12
Apr. 1983, La Croix 2425 (BM); Zomba Mt., 15°17'S,
35°17'E, 20 Nov. 1979, Banda & Salubani 1617, 1,600
m, 10 Mar. 1985, La Croix 2731, 27 Jan. 1978, Seyani
765 (all MO).
Sarcostemma mulanjense possesses the corona
structure C (l ,, + Cs (an outer ring consisting of fused
staminal and interstaminal parts plus five inner sta-
minal lobes) characteristic of Sarcostemma (Fig.
IB). Cynanchum, in contrast, always lacks inner
staminal lobes; its corona can be described as C w
(a single corona consisting of fused staminal and
interstaminal parts). From all other African Sar¬
costemma species, S. mulanjense is readily distin¬
guished morphologically by the height of the gy-
nostegium exceeding its diameter. In addition, the
cochleariform inner Cs, the well-differentiated parts
of Cs and Ci in the outer C(is) and the flat, slightly
undulating, cream-colored corolla lobes set it apart
from ,S. viminale. Ecologically, it is different in its
preference for high altitudes and undisturbed veg¬
etation (Fig. 2).
Species rank seems appropriate for this taxon,
because the changes in gynostegium proportions in¬
dicate a considerable time of independent evolution,
due to its isolation in the southern Malawi mountains.
Mechanical difficulties in inserting the slightly larger
pollinia of S. viminale into the guide rails of S.
mulanjense support at least some degree of repro¬
ductive isolation. No difficulties have been encoun¬
tered when introducing ,S. mulanjense pollinia into
guide rails of S. viminale; however, no fruit set has
resulted from all crosses tried. In nature, interbreed¬
ing of the two taxa is highly unlikely, because pop¬
ulations of S. viminale are known only from localities
considerably further south (Zimbabwe) or further
north (Rumpi District).
Specimens from high altitudes of Mt. Mulanje
226
Novon
exhibit a dense cushion-shaped habit. Longer shoots
die back during the winter, probably due to Irost,
and only the lowermost 4-6 internodes survive.
Resprouting occurs from the nodes underneath the
black remnants of the shoots of the previous season
(Fig. 2). These resprouts remain comparatively short
and erect. In cultivation, however, these specimens
show the same arching growth form as those irom
the base of the mountain and the Zomba plateau.
Acknowledgments. We thank F. Alhers, Mini¬
ster, for the use of facilities. Fieldwork was sponsored
hy the Deutsche Forschungsgemeinschaft (Liede,
grant No. 477/25/91). E. v. Jaarsveld, Kirsten-
bosch, and L. E. Newton, Nairobi, have enthusi¬
astically participated in fieldwork. Loan of material
from the herbaria mentioned is gratefully acknowl¬
edged.
Literature Cited
Adams, B. R. & R. W. K. Holland. 1978a. The genus
Sarcostemma in East Africa. Cact. Succ. J. (U.S.)
50: 107-111.
- Si -. 1978b. The genus Sarcostemma
in East Africa. Cact. Succ. J. (U.S.) 50: 166-169.
Forster, P. I. 1988. The genus Sarcostemma in Aus¬
tralia. Anacampseros: 73-79.
Two New Combinations in Sechium (Cucurbitaceae) from
Centra] America, and a New Species from Oaxaca, Mexico
Rafael Lira and Fernando Chiang
Herbario Nacional de Mexico, Apdo. Postal 70-367, 04510 Mexico, D.F., Mexico
ABSTRACT. Based on studies of herbarium collec¬
tions, fieldwork in Mexico and Central America, and
palynological evidence, two species of Frantzia (F
panamensis and F. venosa) are transferred to Se¬
chium, and a species in this genus (S. chinantlense)
from the lowlands of Oaxaca is described as new.
A key to the 10 species of Sechium is presented.
According to the most recent classification of the
Cucurbitaceae (Jeffrey, 1990), the genus Sechium
P. Browne belongs to tribe Sicyeae, suhtrihe Sicyi-
nae. Ever since Jeffrey’s (1978) work, in which a
broader generic circumscription of Sechium was
presented (encompassing Ahzolia Standley & Stey-
erm., Frantzia Pittier, and Polakowskia Pittier),
there has been a growing interest in the genus. Gruz-
Leon (1985-1986) and Cruz-Leon & Querol-Lip-
covich (1985) reported wild populations of 5. edule
(Jacq.) Sw. found in 1982—1983 in Veracruz, Mex¬
ico. Newstrom (1985, 1986, 1989, 1990, 1991)
presented the results of her research on the origin
and evolution of this species, reporting new localities
for wild populations of the species in Oaxaca, Mex¬
ico, and proposing a different generic circumscrip¬
tion.
Studies of collections (including type material) of
all species of Sechium and related genera, fieldwork
in Mexico and Central America (Lira, 1990, 1991;
Lira & Soto, 1991), and palynological studies of
species of most of the genera of suhtrihe Sicyinae
(Lira & Alvarado, 1991) have revealed new evi¬
dence supporting Jeffrey’s (1978) generic circum¬
scription. Taking into account all the above-men¬
tioned works, we propose two new combinations and
a new species of Sechium.
Sechium panamense (Wunderlin) Lira & Chiang,
comb. nov. Basionym: Frantzia panamensis
Wunderlin, Bull. Torrey Bot. Club 104: 102.
1977. TYPE: Panama. Chiriqui: Boquete Dis¬
trict, Bajo Chorro, 6,000 ft., Davidson 416
(holotype, F, photograph).
Geographical distribution. Apparently endemic
to Panama, known only from the type locality, al
elevations between 1,800 and 3,000 m.
Additional specimens examined. PANAMA. CHIRIQUI:
top of peak between Baru and Respinga, ca. 3,000 m,
27 Nov. 1975, D'Arcy 10135 (MO); Cerro Pata de
Macho, ca. 5 mi. NE of Boquete, trail to continental
divide leading to Finca Serrano (Francisco Serrano, Pacific
slope), 22 Nov. 1979, Antonio 2687 (MO); Bugaba,
Cerro Punta, along ridge to watershed to Bocas del Toro,
8°52’N, 82°33'W, 2,200 m, 26 Jan. 1985, van der
Werff & Herrera 6 458 (MO).
Sechium venosum (L. D. Gomez) Lira & Chiang,
comb. nov. Basionym: Frantzia venosa L. D.
Gomez, Phytologia 53: 447. 1983. TYPE: Cos¬
ta Rica. Limon: along road from JJone Creek
to Bribri, Gomez et al. 20483 (holotype, MO;
isotypes, BM, CR, K).
Geographical distribution. Originally reported
(Gomez & Gomez, 1983) as endemic to the lowlands
of Caribbean Costa Rica, it has now been found at
higher elevations (1,300-1,650 m) and also in Pan¬
ama.
Additional specimens examined. Costa Rica, limon:
Talamanca, 7 km SW' of BriBri, 100 m, 4 May 1983,
Gomez et al. 20361 (MEXU, MO), 20459 (BH, BM, K,
MEXU, MO); Limon, 1-3 km N of BriBri, Rio Sixaola
drainage, 9°38'N, 82°50’W, 250 m, evergreen premon-
tane wet and tropical moist forest formation and cocoa
plantation, 9-10 Sep. 1978, Burger Antonio 11014
(CR, F); Costado E, Lago Dabagri, 9 Nov. 1984, Gomez
et al. 23309-B (MO); Hitori Cerere Reserve and vicinity
in Valle La Estrella S of Finca Concepcion in woods on
slope along Rio Cerere, 9°42'N, 83°02'W, 100 m, 31
July 1985, Hammel & Grayum 14297 (CR); forested
hill above BriBri, 100 m, 2 Feb. 1984, Bennington &
Poveda 11465 (CR); base of hills between Punta Codes
and Punta Uva (E of Puerto Viejo de Talamanca), 9°38'N,
82°43'W, 20 60 m, forested ravine, 6 Nov. 1984, Gra¬
yum et al. 4426 (BM, CR, MO); San Jose Cabecar, 4-
23 Mar. 1982, Ocampo s.n. (CR); Canton Talamanca,
ca. 3 km sobre el camino maderero que va a Cataratas,
9°38'N, 82°55'W, borde de selva alta perturbada con
Cecropia, Heliconia spp., Urticaceae y Malvaceae, 150
m, 10 Aug. 1990, Lira & Ocampo 1036 (MEXU);
camino al puente del Rio Uatsi (ahora destruido), viniendo
de Cruce Bribri y Uatsi, 9°37'N, 82°52'W r , selva alta
perturbada, con cultivos de platano y cacao, 100 m, 10
Aug. 1990, Lira & Ocampo 1039 (MEXU) (only fruits
were collected). PUNTARENAS: foothills of the Cordillera de
Talamanca, between Agua Caliente and the Rio Canasta,
8°57'-9°2'N, 82°56'-82°59'W, 1,300-1,650 m, mon¬
tane forest remnants scattered among pastures, 5 Sep.
Novon 2: 227-231. 1992.
228
Novon
Figure 1. Sechium chinantlense Lira & Chiang. —A. Habit. —B. Staminate flower. —C. Pistillate flower. —D.
Staminate flower of S. hintonii. —E. Staminate flower of S. pittieri. —F. Staminate flower of wild S. edule. —G.
Apical view of the fruit showing the cleft.
Volume 2, Number 3
1992
Lira & Chiang
New Combinations in Sechium
229
1984, Davidse et al. 28360 (MO). Panama, bocas dei.
TORO: vicinity of Nievecita, 0-50 m, 8-19 Aug. 1938,
Woodson Jr. et al. 1841 (MO); Nievecita, 3-20 Aug.
1940, Woodson Jr. & Schery 1026 (NY).
Sechium chinantlense Lira & Chiang, sp. nov.
TYPE: Mexico. Oaxaca: Municipio San Lucas
Ojitlan, del poblado El Zapotal a Mata de Cana,
relictos de selva alta perennifolia, 60 m, 23
Jan. 1989, Calzada 14297 (leaves, stam. fl.,
and mature fr.) (holotype, MEXU). Figure 1.
Sechio eduli (Jacq.) Sw. proximum, a quo differt ped-
icellis florum masculorum longioribus, filamentis starnin-
alibus per '/£-% longitudinis connatis, columnam forman-
tibus, hac in 5 ramis divisa, unoquoque ramo in 2 ramis
inaequalibus diviso, et fructu constanter glabro, inermi,
lateraliter compresso, laete viridi.
Stout, monoecious, perennial vine, or sometimes
perennating through the massive roots. Stems sul-
cate, smooth, almost glabrous, much branched,
thickened and woody toward the hase. Leaves her¬
baceous to papery membranous when dried, cordate-
ovate to suborbicular, 10-16(-17.5) cm long, 12-
22.5(-24) cm broad, slightly 3-angulate-lobed, the
lobes obtuse, acute to acuminate, margins remotely
denticulate, base cordate, sinus usually semiclosed,
both surfaces essentially glabrous, the adaxial one
scabrous, slightly puberulent only on the veins, mi¬
nutely white-pustulate, the abaxial one smooth; pet¬
ioles sulcate, 12 16(-22) cm long, glabrous. Ten¬
drils 4-6-parted, sulcate, glabrous. Staminate
inflorescences axillary, racemose-paniculate, pedun¬
culate, 12 27 cm long, puberulent to glabrous, the
flowers in fascicles or in short racemelike branches
distributed at intervals along the rachis, the branches
not exceeding 18 mm; pedicels slender, 4-12 mm
long, puberulent to glabrous; receptacle disk-shaped,
ca. 1 mm long, 3-5 mm wide, glabrous; sepals 5,
narrowly lanceolate to subulate, 1-3.5 mm long;
petals 5, spreading, white to greenish white, trian¬
gular-ovate, obtuse to acute, 4-8 mm long, 2.5-
4.5 mm broad, with 7-9 parallel nerves, puberulent,
glabrescent; stamens 5, 3-4 mm long; filaments
fused Vi-34 their length, forming a thin column,
divided into 5 branches somewhat broadened and
flat, spreading, these again divided into two unequal¬
sized and curved branches; antheriferous tissue at
the tip of the branches, visible mainly at the margins
and underneath; nectaries 10, porelike, surrounding
the staminal column at the base of the receptacle,
puberulent on the surface, slightly projected under¬
neath the calyx or not at all projected. Pistillate
flowers in the same axil as the staminate inflores¬
cences, 1-3 on individual pedicels, sometimes gem¬
inate on the same pedicel; pedicel slender, sulcate.
ca. 1 cm long, elongating up to 10 cm in fruit,
glabrous; ovary ovoid, constricted at the tip, gla¬
brous, smooth, 1-ovuled; perianth as in the staminate
flowers but somewhat reduced; styles fused into a
slender column, 3 mm long; stigma 2-lobed, each
lobe 3-fid; nectaries as in the staminate flowers, but
less evident. Fruits fleshy, obovoid, compressed, 6-
9 cm long and nearly as broad in the middle, delt
at the apex, glabrous, unarmed, bright green when
mature, pulp pale green to whitish, bitter; seed ovoid,
flattened, smooth, 3.5-6 cm long.
Sechium chinantlense is known only from north¬
ern Oaxaca, from the region known as “La Chin-
antla" in the municipios Valle Nacional, Tuxtepec,
Chiltepec, Santa Maria Jacatepec, and San Lucas
Ojitlan, where it thrives at altitudes between 20 and
800 m, in evergreen forest or at higher elevations,
in the transition between evergreen and montane
forest. The populations from south of Valle Nacional
start blooming in August and set fruit in October or
earlier, whereas the ones from Chiltepec and San
Lucas Ojitlan produce flowers until November and
fruits from December to January and February. An
explanation for these differences could be found in
the higher humidity and lower temperatures present
in the Valle Nacional area, compared to ihe warmer
and somewhat drier climate of the north.
Paratypes. MEXICO. OAXACA: Mpio. San Lucas Ojitlan,
Ejido 0. Teran, Presa Cerro de Oro (Santo Domingo), al
S de San Lucas Ojitlan, 18°2'N, 96°25'W, relictos de
selva mediana con Bursera creciendo sobre lomerios muy
pedregosos y rodeados de planicies con suelos muy hu-
medos, 60 m, 24 Oct. 1990, Lira & Soto 1173 (MEXU);
Mpio. Santa Maria Jacatepec, Plan Martin Chino, acahual
derivado de selva alta perennifolia, 60 m, suelo negro
rocoso con lomerios, 25 Oct. 1989, Calzada 15043
(MEXU); Mpio. Chiltepec, Cerro Faisan, 6 km al W de
Chiltepec, 17°58'N, 96°15'W, milpas en lomerios rocosos
rodeados con relictos de selva mediana con Bursera,
Gliricidia, Ceiba, Enterolobium, Araceae y Sapindaceae,
y estos a su vez rodeados por grandes extensiones de
potreros, 60 m, 24 Oct. 1990, Lira & Soto 1176 (MEXU);
Chiltepec and vicinity, in llanos, 20 m, 12 Dec. 1940,
Martinez-Calderon 369 (GH, MEXU), 14 Jan. 1941,
458 (GH); Mpio. Valle Nacional, km 61 de la carretera
Tuxtepec-Oaxaca, selva mediana, 730 m, 28 31 Oct.
1991, Castrejon & Concepcion 183, 194, 212 (MEXU);
12 km al S de Valle Nacional, carretera 175 a Oaxaca,
17°41.3'N, 96°20.5'W, ladera humeda con una caida de
agua, selva alta perennifolia perturbada con Cecropia,
720 m, 25 Oct. 1990, Lira & Soto 1185 (MEXU); 12.5
km al S de Valle Nacional, carretera 175 a Oaxaca,
17°41.2'N, 96°19.5'W, ladera humeda con una caida de
agua, selva alta perennifolia perturbada, 720 m, 25 Oct.
1990, Lira & Soto 1186 (MEXU); 13.5 km al S de
Valle Nacional, carretera 175 a Oaxaca, 17°41'N,
96°19.5'W, ladera humeda con una caida de agua, selva
alta perennifolia, 730 m, 25 Oct. 1990, Lira & Soto
1187 (MEXU); 15 km al S de Valle Nacional, casi 1 km
230
Novon
al N de Puerto Eligio, carretera 175 a Oaxaca, 17°38'N,
96°20'W, ladera humeda con una cai'da de agua, selva
alta perennifolia perturbada, 660 m, 26 Oct. 1990, Lira
& Soto 1188 (MEXU); steep mountain sides above and
generally 9 mi. S of Valle Nacional, along the highway
to Oaxaca, 800 m, 10 Oct. 1962, McVaugh 21801
(MICH); all km al S de Valle Nacional, carretera Tux-
tepec-Oaxaca, selva alta perennifolia, 850 m, Soto et al.
13191 (MEXU).
Specimens of Sechium chinantlense have been
identified and/or reported as 5. compositum (J. I).
Smith) C. Jeffrey, or as a hybrid between the latter
and S. edule or a different species (Newstrom, 1985,
1986, 1989, 1990, 1991). The characteristics
shown by the stamens and fruits permit the sepa¬
ration of S. chinantlense from other species of the
genus (see Fig. 1). Furthermore, the absence of S.
compositum in Oaxaca does not support the con¬
tention that the new species is a hybrid between S.
Key to the Sections and Species of Sechium
compositum and any other species. In this regard,
populations of wild S. edule can be found growing
in the vicinity (although at higher elevations) of
populations of ,S. chinantlense in the south of Valle
Nacional, Oaxaca. However, an examination of both
populations did not yield evidence of hybridization,
indicating that, notwithstanding their vicinity and
similar phenology, these two species are probably
reproductively isolated. On the other hand, in the
northern part of the range of ,S. chinantlensis, there
are no wild populations of 5. edule.
With these three additions, Sechium is now con¬
sidered to include 10 species forming the two sec¬
tions proposed by Jeffrey (1978). Based on our
observations, as well as some additional bibliographic
information (Wunderlin, 1976, 1977, 1978), we
propose the following key to distinguish the sections
and the species.
la. Nectaries surrounding base of reproductive structures, pore- or pouchlike, sunken into the base of receptacle
and ± protruding beneath .section Sechium
2a. Filaments completely fused; anthers free, sessile on the tip of the filament column; floral nectaries
conspicuously protruding beneath the receptacle.
da. Lower surface of the leaves pubescent; floral nectaries tomentellous on upper surface
. S. talamancense (Wunderlin) C. Jeffrey
db. Lower surface of the leaves glabrous; floral nectaries glabrous on upper surface .
. S. tacaco (Pittier) C. Jeffrey
2b. Filaments partially fused, about l A-M their length and then divided into d-5 branches; anthers d-5
(sometimes more) at the tip of the filament branches; floral nectaries not protruding beneath the
receptacle, or only slightly so.
4a. Filament branches not or inconspicuously divided; mature fruits variable in size, shape, indument,
and armature.
5a. Male inflorescence paniculate; pedicels 7-20 mm long; mature fruits 6-10 cm long, fleshy,
with longitudinal ridges and short spines on the ridges, to completely smooth and unarmed
(Chiapas and Guatemala) . 5. compositum (J. D. Smith) C. Jeffrey
5b. Male inflorescence racemose, or if more divided, the branches very short and the pedicels
less than 6 mm; mature fruits fleshy to woody-fibrous with or without ridges, angles or furrows
and spines.
6a. Male inflorescence erect; filaments fused more than 3 A their total length; mature fruits
fleshy, variable in size, shape, and armature; wild (bitter fruits; Mexico in the states of
Veracruz, Hidalgo, Puebla, and Oaxaca) and cultivated (nonbitter fruits) plants .
. S. edule (Jacq.) Sw.
6b. Male inflorescence pendulous; filaments fused X A-V\ their total length; mature fruits d-
4.5 cm long, woody-fibrous, with 5 longitudinal ridges, the ridges with thin spines covered
by retrorse barbs; wild plants (Estado de Mexico and Guerrero) .
.5. hintonii (P. G. Wilson) C. Jeffrey
4b. Filament branches divided into two unequal-sized and curved branches; mature fruits 6-9 cm
long, ovate-cordate, consistently smooth, unarmed, and bright green (Oaxaca) .
.S. chinantlense Lira & Chiang
lb. Nectaries surrounding base of reproductive structures, covered by a cushionlike compressed structure, not
sunken into the base of the receptacle .section Frantzia
7a. Male inflorescence racemose to paniculate, erect or pendulous (Nicaragua to Panama).
8a. Male inflorescences erect, racemose to racemose-paniculate; flowers conspicuously pedicellate;
fruits ovoid to fusiform.
9a. Male inflorescences densely villous; sepals linear lanceolate, 6-8 mm long; fruits fusiform
(Costa Rica) . S. villosum (Wunderlin) C. Jeffrey
9b. Male inflorescences not villous; sepals triangular, up to 1 mm long; fruits ovoid (Nicaragua
to Panama). S. pittieri (Cogn.) C. Jeffrey
8b. Male inflorescences pendulous, racemose; flowers very shortly pedicellate to almost sessile; fruits
ovoid to globose (Costa Rica and Panama) . 5. venosum (L. D. Gomez) Lira & Chiang
7b. Male inflorescences umbelloid (Panama) .S. panamense (Wunderlin) Lira & Chiang
Volume 2, Number 3
1992
Lira & Chiang
New Combinations in Sechium
231
Acknowledgments. This paper forms part of the
project “Taxonomic and Ecogeographic Studies in
Latin American Cucurbitaceae,” sponsored by the
International Board for Plant Genetic Resources,
conducted by R. Lira in the Herbario Nacional de
Mexico, Instituto de Biologia, U.N.A.M. We thank
the curators of BH, BM, CHAPA, CR, ENCB, F,
GH, K. MICH, MO, MU, NY, l!C, US, USF, and
XAL for loan of specimens. The illustration was
prepared by Albino Luna, Instituto de Biologia,
U.N.A.M.
Literature Cited
Cruz-Leon, A. 1985-1986. ^Chayote o cruzas inter-
genericas? Hallazgo y caracteristicas. Revista de
Geografia Agricola, Chapingo Nos. 9-10: 100-106.
- & D. Querol-Lipcovich. 1985. Catalogo de
Recursos Geneticos de chayote (Sechium edule Sw.)
en el Centro Regional Universitario Oriente. Univ-
ersidad Autonoma de Chapingo, Mexico.
Gomez, L. D. & J. Gomez. 1983. Plantae Mesoamer-
icanae Novae. IX. Phytologia 53: 447-448.
Jeffrey, C. 1978. Further notes on Cucurbitaceae. IV.
Some New World taxa. Kew Bull. 33: 347-380.
Jeffrey, C. 1990. An outline classification of the Cu¬
curbitaceae. Appendix. Pp. 449-463 in D. M. Bates,
R. W. Robinson & C. Jeffrey (editors). Biology and
Utilization of the Cucurbitaceae. Cornell Univ. Press,
Comstock, New 1 York.
Lira, R. 1990. Taxonomic and Ecogeographic Studies
in Latin American Cucurbitaceae. 1st Progress Re¬
port (January-August 1990). Project supported by
FAO/International Board for Plant Genetic Re¬
sources. Rome.
-. 1991. Taxonomic and Ecogeographic Studies
in Latin American Cucurbitaceae. 2nd Progress Re¬
port (August 1990-January 1991). Project sup¬
ported by FAO/International Board for Plant Genetic
Resources. Rome.
-& J. L. Alvarado. 1991. A palynological study
of Sechium (Cucurbitaceae) and its allies. 42nd AIBS
Annual Meeting, San Antonio, Texas, U.S.A. (4 8
August 1991). Amer. J. Bot. (Supplement) 78(6):
233. [Abstract.]
- & J. C. Soto. 1991. Sechium hintonii (P. G.
Wilson) C. Jeffrey (Cucurbitaceae): rediscovery and
observations. FAO/IBPGR PI. Genet. Res. Newslet¬
ter 87: 5-10.
Newstrom, L. 1985. Collection of Chayote and its wild
relatives. FAO/IBPGR PI. Genet. Res. Newsletter
64: 14-20.
-. 1986. Studies in the origin and evolution of
Sechium edule (Jacq.) Sw. Ph.D. Thesis, University
of California, Berkeley.
-. 1989. Reproductive biology and evolution of
the cultivated chayote Sechium edule: Cucurbita¬
ceae. Pp. 491-509 in G. H. Bock & Y. B. Linhart
(editors), Evolutionary Ecology of Plants. Westview
Press, Boulder, Colorado.
-. 1990. Origin and evolution of Chayote, Se¬
chium edule. Pp. 141-149 m D. M. Bates, R. W.
Robinson & C. Jeffrey (editors), Biology and Utili¬
zation of the Cucurbitaceae. Cornell Univ. Press,
Comstock, New York.
-. 1991. Evidence for the origin of chayote
Sechium edule (Cucurbitaceae). Econ. Bot. 45: 410-
428.
Wunderlin, R. P. 1976. Two new species and a new
combination in Frantzia (Cucurbitaceae). Brittonia
28: 239-244.
-. 1977. A new species of Frantzia (Cucurbi¬
taceae) from Panama. Bull. Torrey Bot. Club 104:
102-104.
-. 1978. Cucurbitaceae. In: R. E. Woodson et
al. (editors). Flora of Panama. Ann. Missouri Bot.
Card. 65: 285-366.
New Combinations and Nomenclatural Notes on the Genera Salacia
and Tontelea (Celastraceae, Sensu Lato) in the Neotropics. V.
Alberta M. W. Mennega
Department of Plant Ecology & Evolutionary Biology, Herbarium Division,
University of Utrecht, Heidelberglaan 2, P.0. Box 80.102,
3508 TC, Utrecht, The Netherlands
Abstract. The increase in collections of Hippo-
crateaceae, in particular from Andean regions, has
led to several taxonomic and nomenclatural changes
in the genera Salacia and Tontelea. Miers’s Ther¬
mophila cordata is reestablished as Salacia cor-
data; S. acreana and .S’, megistophylla are con¬
sidered to be synonyms. Salacia pittieriana is placed
in synonymy with S. multiflora. Salacia mucronata
is treated as a subspecies of .S’, multiflora. Tontelea
richardii is considered as a subspecies of 7. oval-
ifolia, and /’. ftuminensis placed in synonymy with
the latter species. Other changes in Tontelea con¬
cern the transfer of T. ulei to Salacia ulei. Krukoff
5141 , the type of Smith’s Salacia petiolata, later
transferred by him to Tontelea ulei, is treated here
as T. petiolata.
Salacia cordata (Miers) A. M. Mennega, comb,
nov. Basionym: Thermophila cordata Miers,
I'rans. Linn. Soc. London 28: 401. 1872.
TYPE: Peru. Pavon s.n. (holotype, BM; iso¬
type, G).
Salacia acreana A. C. Smith, Bull. Torrey Bot. Club 66:
243. 1939. Syn. nov. TYPE: Brazil. Acre: Krukoff
5523 (holotype, NY; isotypes, A, F, MICH, S, U,
US).
Salacia megistophylla Standi., Publ. Field Mus. Nat.
Hist. Bot. Ser. 4: 222. 1929. Syn. nov. TYPE:
Panama. Bocas del Toro: Crica Mola Valley, Cooper
503 (holotype, F).
In 1872, M iers came across a Pavon collection
in the British Museum which, despite the lack of
flowers on the inflorescence, he recognized as be¬
longing to the Hippocrateaceae. He placed it in the
genus Thermophila and named it 7. cordata Miers,
which he considered closely related to 7. macro-
phylla Miers (= Salacia miqueliana Loes.).
I studied additional material of the Pavon collec¬
tion in the herbarium at Geneva. These sheets also
lack flowers, but they have the same characteristic
large leaves with a more or less cordate base and a
rather short, several-branched inflorescence. I sup¬
port Miers' opinion that the Pavon material repre¬
sents a good species, despite the absence of flowers.
Because Thermophila is now placed in synonymy
with Salacia, this species is here named Salacia
cordata (Miers) A. M. Mennega.
Salacia multiflora (Lam.) DC. Prodr. 1: 570.
1824. Basionym: Hippocratea multiflora Lam.
Tab. Encycl. 1: 101. 1791. TYPE: French
Guiana. Richard s.n. (holotype, P).
Salacia pittieriana A. C. Smith, Brittonia 3: 440. 1940.
Syn. nov. TYPE: Venezuela. Aragua: Parque Na¬
tional, L. Williams 10332 (holotype, F; isotype,
US).
Salacia multiflora subsp. mucronata (Busby) A.
M. Mennega, comb. nov. Basionym: Salacia
mucronata Rusby, Descr. S. Amer. PI. 53.
1920. TYPE: Venezuela. Delta Amacuro: low¬
er Rio Orinoco, Rusby & Squires 130 pro
parte (holotype, NY; isotypes, A, F, G, GH,
MICH, US).
In his treatment of Salacia, Smith (1940) rec¬
ognized a number of species groups, two of which
are closely related: Multiflorae, with the single spe¬
cies S. multiflora, and Mucronatae, with two species
(S. mucronata and ,S. pittieriana). The differenti¬
ating characters between these groups are slight
variations in the form of the disk and the mode of
dehiscence of the anthers.
After studying a large number of specimens be¬
longing to this alliance I found it hard to distinguish
5. pittieriana from S. multiflora. The coriaceous
leaves of S. pittieriana were also occasionally found
in S. multiflora, and the disk characters were not
clearly distinctive. Salacia mucronata, though
closely related to .S', multiflora, can be distinguished
from the latter by a strictly dichotomously branched
inflorescence with smaller-sized flowers. I consider
it to be a subspecies of S. multiflora.
Tontelea ovalifolia (Miers) A. G. Smith, Brittonia
3: 476. 1940. Basionym: Cuervea ovalifolia
Miers, Trans. Linn. Soc. London 28: 371. 1872.
Novon 2: 232-234. 1992.
Volume 2, Number 3
1992
Mennega
Salacia and Tontelea
233
TYPE: Colombia. Meta: Villa Vicencio, Triana
s.n. (holotype, K).
Tontelea ovalifolia subsp. richardii (Peyr.) Gorts
& A. M. Mennega, comb, et stat. nov. Salacia
richardii Peyr. in C. Martius, FI. Bras. 11 (1):
148. 1878. Syn. nov. TYPE: French Guiana.
Richard s.n. (holotype, P).
Salacia jiuminensis Peyr. in C. Martius, FI. Bras. 11
(1): 149. 1878. Syn. nov. Tontelea jiuminensis
(Peyr.) A. C. Smith, Brittonia 3: 477. 1940. TYPE:
Brazil. Riedel s.n. (holotype, F; isotypes, GH, K, P,
US).
According to Smith’s (1940) classification, the
three species cited above belong in the species group
Attenuatae, together with several other species; in
his key to this group, the three are placed close
together. The main differences between the taxa
occur in the shape of the leaves and the more or
less immersed or prominulous venation. In his dis¬
cussion on T. ovalifolia, Smith pointed out the great
variation in leaf size, but he concluded that there
was sufficient agreement in leaf shape and texture.
Furthermore, he stated that specimens referred to
this species and the following (T. richardii and T.
jiuminensis) are grouped only with difficulty. With
the numerous collections received for identification
from Central America and northern South America
over a number of years, it became increasingly dif¬
ficult to distinguish among the three species. Con¬
sequently, T. richardii and T. Jiuminensis are now
placed under synonymy of T. ovalifolia. However,
because the fruits show a remarkable difference in
size and structure, two subspecies are recognized.
Fruits of T. ovalifolia subsp. ovalifolia are of mod¬
erate size, 3-5 x 5 cm, and the pericarp is thin
and leathery, whereas the fruits of 7. ovalifolia
subsp. richardii are very large and more or less
globose, 8x10 cm, and the pericarp is woody, ca.
10 mm thick.
The holotype of T. richardii from French Guiana
is a flowering specimen without fruits. Specimens
with big woody fruits were collected in Panama, in
the Canal Zone on Barro Colorado Island by several
collectors, e.g., Wetmore, Abbe & Shattuck 92,
Shattuck 658. Smith identified these collections as
7. richardii, notwithstanding the distributional gap.
Fruits of T. jiuminensis were described by Riedel
(ex Peyritsch, 1878) as “maximus, magnitudine
capitis infantis,” which agrees well with the T. ri¬
chardii material. Among the scarce fruiting speci¬
mens recently collected is M. Monsalve R 762 from
Colombia, Valle del Cauca. This material has flowers
and big, thick-walled fruits. It was first determined
as T. jiuminensis, but is now considered to be T.
ovalifolia subsp. richardii.
Salacia ulei Loes., Verh. Bot. Vereins Prov. Bran¬
denburg 48: 178. 1907. Tontelea ulei (Loes.)
A. C. Smith, Brittonia 3: 498. 1940. TYPE:
Brazil. Amazonas: Rio Jurua, Marary, l le 5050
(holotype, B destroyed; photo of type, F; lec-
totype, selected here, G; isotype, L).
Salacia cuspidata A. C. Smith, Bull. Torrey Bot. Club
66: 248. 1939. Syn. nov. TYPE: Brazil. Amazonas:
near Palmares, Sao Paulo de Oliven^a, Krukojf 8415
(holotype, NY; isotypes, BM, F, K, S, U, US; para-
type, same locality, Krukojf 8255, A, F, NY, S, U).
When Loesener (1907) described Salacia ulei,
based on Ule 5050, he noted the unusual form of
the disk. This disk consists of two parts: a low, thin
outer ring, and an inner part that is higher and
thicker. Such a disk had not yet been described in
Brazilian species of Salacia, and Loesener doubted
if Salacia was the correct genus.
In his treatment of New World Hippocrateaceae,
Smith (1940) transferred Salacia ulei to Tontelea.
He did not see the specimen, but based his decision
on a photograph of the type and on Loesener s
description. He regarded the double disk as "trau¬
matic.” If the double disk is not taken into account,
the plant might indeed be accommodated in Ton¬
telea, a genus with a cup-shaped disk. In Tontelea,
however, the stigmas are usually well developed,
whereas Ule 5050 was described by Loesener as
having a style with only minute stigmatic points. In
Smith’s (1940) treatment of Tontelea, there are five
species that lack prominent stigmas. These were
placed in the species group Nectandrifoliae. The
Ule specimen was treated as T. ulei in this rather
aberrant group. Smith also assigned several other
collections to 7. ulei: Cardenas 1763 from Bolivia,
and Krukojf4755 and 5141 from Amazonas, Brazil.
I studied the type specimen, as well as the other
specimens cited above. The flowers of Ule 5050, in
accordance with Loesener’s (1907) ample descrip¬
tion, show a disk consisting of two parts. At the time
of Loesener’s paper, such a disk was unknown in
Salacia. In later collections, however. Smith noticed
in three more species of Salacia a disk with a flat
basal part and a conical inner part, very similar to
the disk in Ule 5050; this was found each time
combined with a branched inflorescence. He placed
these species in his group Arboreae of Salacia. One
of the species in this group, which includes five
species in total, is S. cuspidata A. C. Smith. 1 he
type specimen of S. cuspidata, Krukojf 8115, re¬
sembles Ule 5050 in several respects. In addition
234
Novon
to the identical disks, both collections share the
absence of stigmas, the short-branched inflores¬
cences, and the characteristic narrowly cuspidate
leaf apex. Therefore, I consider the two collections
as conspecific and belonging in Salacia , species
group Amplectentes (Mennega, 1984).
Recent collections of Salacia ulei include: Peru.
LORETO: Iquitos, Asplund 14505 { S), Vasquez 3373 (MO,
U); Rio Ucayala, Vasquez et al. 7027 (MO, U).
As stated above. Smith also included a fruiting
specimen, Cardenas 1763 (US), in Tontelea ulei.
The leaves ol this plant, however, lack the char¬
acteristic cuspidate apex of Tontelea ulei, and they
much better resemble the leaves of 7. ovalifolia.
Therefore, I exclude this collection from 7. ulei.
Krukoff 4755 (NY), a fruiting specimen, does
not belong in 7. ulei either. It has large concolorous
leaves with an obtuse apex that in venation and
texture suggests Salacia cordata (Miers) A. M.
Mennega. The globose brownish fruits also agree
with this species. The third collection, Krukoff 5141
(A, NY), is a vine that is the type of Salacia pe-
tiolata A. C. Smith. This species was included in
the synonymy of Tontelea ulei by Smith in 1940.
From Smith’s description and from my own obser¬
vations of the only flower present on the sheet, it is
clear that this plant truly belongs in Tontelea and
in the species group Nectandrifoliae, necessitating
the new combination below.
Tontelea petiolata (A. C. Smith) A. M. Mennega,
comb. nov. Basionym: Salacia petiolata A. C.
Smith, Bull. Torrey Bot. Club 66: 242. 1939.
TYPE: Brazil. Amazonas: basin of Hio Jurua,
near mouth of Bio Embira, Krukoff 5141 (ho-
lotype, NY; isotype, A).
Acknowledgments. I thank the curators of A,
BM, F, G, MO, NY, P, S, US for loan of specimens.
I am much indebted to A. B. A. Gorts-van Bijn for
her cooperation, and to L. Y. Th. Westra for his
critical reading of the English text.
Literature Cited
Loesener, T. 1907. Hippocrateaceae. Verb. Bot. Ver-
eins Prov. Brandenburg 48: 179.
Mennega, A. M. W. 1984. Notes on New World Sal-
acioideae I. Proc. Kon. Ned. Akad. Wetensch. Ser.
C, Biol. Med. Sci. 87(1): 81-90.
Peyritsch, J. 1878. Hippocrateaceae. In: C. Martius,
Flora Brasiliensis 11(1): 125-164.
Smith, A. C. 1940. The American species of Hippo¬
crateaceae. Brittonia 3: 341-555.
Validation of Subclass and Superordinal Names in Magnoliophyta
James L. Reveal
Department of Botany, University of Maryland,
College Park, Maryland 20742-5815, U.S.A.
Abstract. Three subclass and 40 superordinal
names now in current use, but not validly published,
are validated.
In recent years, several authors have adopted
subclass and superordinal names in Magnoliophyta,
the flowering plants (Cronquist, 1981; R. Dahlgren,
1980; Dahlgren & Clifford, 1981, 1982; Dahlgren
& Bremer, 1985; Dahlgren et ah, 1985; G. Dahl¬
gren, 1989a, b; Takhtajan, 1959, 1906, 1969,
1970, 1973, 1980, 1986, 1987; Thorne, 1968,
1976, 1981, 1983, 1992). Unfortunately, except
for those proposed by Takhtajan in 1966, almost
none have been validly published in accordance with
provisions in the International Code of Botanical
Nomenclature (Greuter et ah, 1988). Although the
principle of priority is not mandatory tor names
above the rank of family (Art. 11.4), all names
regardless of rank must be effectively and validly
published (Art. 6).
All subclass and superordinal names not validly
published, but now in current use, are validated
below with the exception of two superordinal names
(based on Annonaceae and Chenopodiaceae) used
by Thorne (1992), for which the validly published
Magnolianae Takhtajan (1966) and Caryophyllanae
Takhtajan (1966) are available, and a subclass name
based on Cornaceae used by Frohne & Jensen (1985).
The latter has not been adopted by Cronquist or
Takhtajan.
All names proposed here are based on and au¬
tomatically typified by the type of the genus of an
included family name (Art. 16.1) and validated by
a direct reference to a validly published description
(Art. 32). The terminations are in accordance with
those given in Article 16.2 and Recommendation
16A. The rationale for the termination “-anae” for
superorders has been discussed elsewhere (Reveal
& Bedell, 1983) and is now uniformly adopted by
the major workers who use the rank (Takhtajan,
1987; Thorne, 1992).
All proposed new names are currently accepted
by Dablgren (in G. Dahlgren, 1989a, b), Takhtajan
(1987), or Thorne (1992). Authors adopting these
names are noted by the use of D, T, or Th respec¬
tively.
Subclass Lamiidae (Findley) Takhtajan ex Reveal,
stat. nov. Based on Lamiaceae Lindley, Nat.
Syst. Bot. ed. 2, 275. 1836. TYPE: Lamium
Linnaeus (1753). T.
Subclass Ranunculidae (A. L. de Jussieu) Takh¬
tajan ex Reveal, stat. nov. Based on Ranun-
culaceae A. L. de Jussieu, Gen. PI. 231. 1789.
TYPE: Ranunculus Linnaeus (1753). T.
Subclass Triurididae (G. Gardner) Takhtajan ex
Reveal, stat. nov. Based on Triuridaceae G.
Gardner, Trans. Linn. Soc. 19: 160. 1843.
TYPE: Triuris Miers (1841). T.
Superorder Aranae (A. L. de Jussieu) Thorne ex
Reveal, stat. nov. Based on Araceae A. L. de
Jussieu, Gen. PL 23. 1789. TYPE: Arum Lin¬
naeus (1753). D, T, Th.
Superorder Balanophoranae (L. C. Richard & A.
Richard) R. Dahlgren ex Reveal, stat. nov.
Based on Balanophoraceae L. C. Richard & A.
Richard, Mem. Mus. Hist. Nat. 8: 429. 1822.
TYPE: Balanophora J. R. & G. Forster (I 775).
D.
Superorder Bromelianae (A. L. de Jussieu) R.
Dahlgren ex Reveal, stat. nov. Based on Bro-
meliaceae A. L. de Jussieu, Gen. PI. 49. 1789.
TYPE: Bromelia Linnaeus (1753). D, T.
Superorder Hutomanae (L. C. Richard) Takhtajan
ex Reveal, stat. nov. Based on Butomaceae L.
C. Richard, Mem. Mus. Hist. Nat. 1: 366.
1815. TYPE: Butomus Linnaeus (1753). T.
Superorder Campanulanae (A. L. de Jussieu)
Takhtajan ex Reveal, stat. nov. Based on Cam-
panulaceae A. L. de Jussieu, Gen. PI. 163.
1789. TYPE: Campanula Linnaeus (1753).
T.
Superorder Cornanae (Dumortier) Thorne ex Re¬
veal, stat. nov. Based on Cornaceae Dumortier,
Anal. Fam. PI. 33, 34. 1829. TYPE: Cornus
Linnaeus (1753). D, T, Th.
Superorder Cyclanthanae (Dumortier) Thorne ex
Reveal, stat. nov. Based on Cyclanthaceae Du¬
mortier, Anal. Fam. PI. 65, 66. 1829. TYPE:
Cyclanthus P. A. Poiteau ex A. Richard (1824).
D, T, I h.
Superorder Eucommianae (Engler) Takhtajan ex
Reveal, stat. nov. Based on Eucommiaceae En-
Novon 2: 235-237. 1992.
236
Novon
gler, Syllabus ed. 6, 145. 1909. TYPK: Fu-
commia I). Oliver (1890). T.
Superorder Euphorbianae (A. L. de Jussieu)
Takhtajan ex Reveal, stat. nov. Based on Eu-
phorbiaceae A. L. de Jussieu, Gen. PI. 384.
1789. TYPE: Euphorbia Linnaeus (1753). T.
Superorder Fabanae (Lindley) R. Dahlgren ex Re¬
veal, stat. nov. Based on Fabaceae Lindley,
Nat. Syst. Bot. ed. 2, 148. 1836. TYPE: Fab a
J. Hill (1753). T.
Superorder Gentiananae (A. L. de Jussieu) Thorne
ex Reveal, stat. nov. Based on Gentianaceae
A. L. de Jussieu, Gen. PI. 141. 1789. TYPE:
Gentiana Linnaeus (1753). I), T, Th.
Superorder Geranianae (A. L. de Jussieu) Thorne
ex Reveal, stat. nov. Based on Geraniaceae A.
I., de Jussieu, Gen. PI. 268. 1789. TYPE:
Gentiana Linnaeus (1753). Th.
Superorder Ilydatellanae (Hamann) Takhtajan ex
Reveal, stat. nov. Based on Hydatellaceae Ha¬
mann, New Zealand J. Bot. 14: 195. 1976.
TYPE: Hydatella Diels (1904). T, Th.
Superorder Juglandanae (A. Richard ex Kunth)
Takhtajan ex Reveal, stat. nov. Based on Ju-
glandaceae A. Richard ex Kunth, Ann. Sci.
Nat. (Paris) 2: 343. 1824. TYPE: Juglans
Linnaeus (1753). T.
Superorder Eecythidanae (P. A. Poiteau) Takh¬
tajan ex Reveal, stat. nov. Based on Lecythi-
daceae I*. A. Poiteau, Mem. Mus. Hist. Nat.
13: 143. 1845. TYPE: Lecythis Loefling
(1758). T.
Superorder Loasanae (Dumortier) R. Dahlgren ex
Reveal, stat. nov. Based on Loasaceae Du¬
mortier, Comment. Bot. 58. 1822. TYPE:
Loasa Adanson (1763). D, T, Th.
Superorder Najadanae (A. L. de Jussieu) Takh¬
tajan ex Reveal, stat. nov. Based on Najadaceae
A. L. de Jussieu, Gen. PI. 18. 1789. TYPE:
Najas Linnaeus (1753). T.
Superorder Nelumbonanae (Dumortier) Takhta¬
jan ex Reveal, stat. nov. Based on Nelumbon-
aceae Dumortier, Anal. Fam. PI. 20, 24. 1829.
TYPE: Nelumbo Adanson (1763). T.
Superorder Nepenthanae (Dumortier) Takhtajan
ex Reveal, stat. nov. Based on Nepenthaceae
Dumortier, Anal. Fam. PI. 14, 16. 1829. TYPE:
Nepenthes Linnaeus (1753). T.
Superorder Nymphaeanae (R. A. Salisbury) Thorne
ex Reveal, stat. nov. Based on Nymphaeaceae
R. A. Salisbury, Ann. Bot. (Konig & Sims) 2:
70. 1805. TYPE: Nymphaea Linnaeus, nom.
cons. (1753). I), T, Th.
Superorder Pandananae (R. Brown) Thorne ex
Reveal, stat. nov. Based on Pandanaceae R.
Brown, Prodr. 340. 1810. TYPE: Pandanus
Linnaeus (1753). D, T, Th.
Superorder Plumbaginanae (A. L. de Jussieu)
Takhtajan ex Reveal, stat. nov. Based on Plum-
baginaceae A. L. de Jussieu, Gen. PL 92. 1789.
TYPE: Plumbago Linnaeus (1753). I), T.
Superorder Podostemonanae (L. C. Richard ex
C. A. Agardh) R. Dahlgren ex Reveal, stat.
nov. Based on Podostemonaceae L. C. R ichard
exC. A. Agardh, Aphor. Bot. 125. 1822. TYPE:
Podostemon Michaux (1803).
Superorder Polygonanae (A. L. de Jussieu) Takh¬
tajan ex Reveal, stat. nov. Based on Polygon-
aceae A. L. de Jussieu, Gen. PI. 82. 1789.
TYPE: Polygonum Linnaeus (1753). D, T.
Superorder Pontederianae (Kunth) Takhtajan ex
Reveal, stat. nov. Based on Pontederiaceae
Kunth in Humboldt, Bonpland & Kunth, Nov.
Gen. Sp. 1: ed. fol. 211; ed. qu. 265. 1816.
TYPE: Pontederia Linnaeus (1753). T.
Superorder Primulanae (Ventenat) R. Dahlgren
ex Reveal, stat. nov. Based on Primulaeeae
Ventenat, Regne Veg. 2: 285. 1799. TYPE:
Primula Linnaeus (1753). 1).
Superorder Rafflesianae (Dumortier) Thorne ex
Reveal, stat. nov. Based on RafUesiaceae Du¬
mortier, Anal. Fam. PI. 13, 14. 1829. TYPE:
Kafflesia R. Brown (1821). T, Th.
Superorder Ranunculanae (A. L. de Jussieu)
Takhtajan ex Reveal, stat. nov. Based on Ran-
unculaceae A. L. de Jussieu, Gen. PI. 231.
1789. TYPE: Ranunculus Linnaeus (1753).
D, T.
Superorder Rhamnanae (A. L. de Jussieu) Takh¬
tajan ex Reveal, stat. nov. Based on Rham-
naceae A. L. de Jussieu, Gen. PI. 376. 1789.
TYPE: Rhamnus Linnaeus (1753). T.
Superorder Santalanae (R. Brown) Thorne ex Re¬
veal, stat. nov. Based on Santalaceae R. Brown,
Prodr. 350. 1810. TYPE: Santalum Linnaeus
(1753). D, Th.
Superorder Sarracenianae (Dumortier) Thorne ex
Reveal, stat. nov. Based on Sarraceniaceae Du¬
mortier, Anal. Fam. PI. 53. 1829. TYPE: Sar-
racenia Linnaeus (1753). T.
Superorder Solananae (A. L. de Jussieu) R. Dahl¬
gren ex Reveal, stat. nov. Based on Solanaceae
A. L. de Jussieu, Gen. PI. 124. 1789. TYPE:
Solarium Linnaeus (1753). I), T, Th.
Superorder Theanae (D. Don) Thorne ex Reveal,
stat. nov. Based on Theaceae D. Don, Prodr.
FI. Nepal. 224. 1825. TYPE: Then Linnaeus
(1753). I), T, Th.
Su, )erorder Triuridanae (G. Gardner) Thorne ex
Reveal, stat. nov. Based on Triuridaceae G.
Volume 2, Number 3
1992
Reveal
Names in Magnoliophyta
237
Gardner, Trans. Linn. Soc. 19: 160. 1843.
TYPE: Triuns Miers (1841). D, T, Th.
Superorder Troehodendranae (Prantl) Takhtajan
ex Reveal, stat. nov. Based on Trochodendra-
ceae Prantl in Engler & Prantl, Nat. Pflanzen-
fam. Ill, 2: 21. 1888. TYPE: Trochodendron
Zuccarini (1839). T.
Superorder Typhanae (A. I., de Jussieu) Thorne
ex Reveal, stat. nov. Based on Typhaceae A.
L. de Jussieu, Gen. PI. 25. 1 789. TYPE: Typha
Linnaeus (1753). T.
Superorder Urticanae (A. L. de Jussieu) Takhtajan
ex Reveal, stat. nov. Based on Urticaceae A.
L. de Jussieu, Gen. PI. 400. 1789. TYPE:
Urtica Linnaeus (1753). T.
Superorder Violanae (Batsch) R. Dahlgren ex Re¬
veal, stat. nov. Based on Violaceae Batsch, Tab.
Alhn. Regni Veg. 57. 1802. TYPE: Viola Lin¬
naeus (1753). I), T, Th.
Superorder Vitanae (A. L. de Jussieu) Takhtajan
ex Reveal, stat. nov. Based on Vitaceae A. L.
de Jussieu, Gen. PL 267. 1789. TYPE: I it is
Linnaeus (1753). D, T.
Superorder Zingiberanae (Lindley) Takhtajan ex
Reveal, stat. nov. Based on Zingiberaceae Lind¬
ley, Key Bot. 69. 1835. TYPE: Zingiber G.
R. Boehmer, nom. cons. (1760). 1), T.
Acknowledgments. I thank R. D. Hoogland, A.
Cronquist, A. L. Takhtajan, and R. E. Thorne for
comments and help. This work is supported, in part,
by National Science Foundation Grant BSR-8812816
to the Missouri Botanical Garden. This is Scientific
Article A-6273, Contribution No. 8442, of the
Maryland Agricultural Experiment Station and Co¬
operative Extension Service.
Literature Cited
Cronquist, A. 1981. An Integrated System of Classifi¬
cation of Flowering Plants. Columbia Univ. Press,
New York.
Dahlgren, G. 1989a. The last Dahlgrenogram: System
of classification of the dicotyledons. Pp. 249-260 in
K. Tan, R. R. Mill & T. S. Elias (editors), Plant
Taxonomy, Phytogeography and Related Subjects.
Edinburgh Univ. Press, Edinburgh.
-. 1989b. An updated angiosperm classification.
J. Linn. Soc., Bot. 100: 197-203.
Dahlgren, R. 1980. A revised system of classification
of the angiosperms. J. Linn. Soc., Bot. 80: 91 124.
- & K. Bremer. 1985. Major clades of the
angiosperms. Cladistics 1: 349-368.
- & H. T. Clifford. 1981. Some conclusions
from a comparative study of the monocotyledons and
related dicotyledonous orders. Ber. Deutsch. Bot.
Ges. 94: 203-227.
- & -. 1982. The Monocotyledons: A
Comparative Study. Academic Press, London.
-, - & P. F. Yeo. 1985. The Families
of the Monocotyledons: Structure, Evolution, and
Taxonomy. Springer-Verlag, New York.
Frohne, D. & U. Jensen. 1985. Systematik des Pflan-
zenreichs. G. Fischer Verlag, Stuttgart.
Greuter, W., FI. M. Burdet, W. G. Chaloner, V. Demoulin,
R. Grolle, D. L. Hawksworth, D. H. Nicolson, P. C.
Silva, F. S. Stafleu, E. G. Voss & J. McNeill (editors).
1 988. International code of botanical nomenclature
adopted by the Fourteenth International Botanical
Congress, Berlin, July-August 1987. Regnurn Veg.
1 18.
Reveal, J. L. & H. G. Bedell. 1983. Add the rank
superorder denoted by -anae. Taxon 32: 661-662.
Takhtajan, A. L. 1959. Die Evolution der Angiosper-
men. G. Fischer Verlag, Jena.
-. 1966. Systema et Phylogenia Magnoliophy-
torum. Nauka, Lenningrad.
-. 1969. Flowering Plants: Origin and Dispersal.
Oliver & Boyd, Edinburgh.
-. 1970. The Origin and Dispersal of Flowering
Plants. Nauka, Leningrad. [In Russian.]
-. 1973. Evolution und Ausbreitung der Blu-
tenpflanzen. G. Fischer Verlag, Stuttgart.
-. 1980. Outline of the classification of the flow¬
ering plants (Magnoliophyta). Bot. Rev. (Lancaster)
46: 225-359.
-. 1986. Floristic Regions of the World. Univ.
California Press, Berkeley.
-. 1987. Systema Magnoliophytorum. Nauka,
Lenningrad.
Thorne, R. F. 1968. Synopsis of a putatively phylo¬
genetic classification of the flowering plants. Aliso
6(4): 57-66.
-. 1976. A phylogenetic classification of the
angiospermae. Evol. Biol. 9: 35-106.
-. 1981. Phytochemistry and angiosperm phy¬
togeny: A summary statement. Pp. 233-295 in D.
A. Young & D. S. Seigler (editors), Phytochemistry
and Angiosperm Phylogeny. Praeger Scientific, New
York.
-. 1983. Proposed new realignments in the an¬
giosperms. Nordic J. Bot. 3: 85-117.
-. 1992. An updated phylogenetic classification
of the flowering plants. Aliso 13: 365-389.
Validation of Ordinal Names of Extant Vascular Plants
James L. Reveal
Department of Botany, University of Maryland,
College Park, Maryland 20742-5815, U.S.A.
ABSTRACT. Thirty-two ordinal names in current
use are validly published.
Of the more than 400 ordinal names based on a
validly published generic name currently available
for extant vascular plants, more than 60 are not
validly published in accordance with provisions in
the International Code of Botanical Nomenclature
(Greuter et al., 1988). Although the principle of
priority is not mandatory for names above the rank
of family (Art. 1 1.4), all names regardless of rank
must be effectively and validly published (Art. 6).
Of the some 60 invalid ordinal names, slightly more
than half are in current use (Gronquist, 1981, I 988;
G. Dahlgren, 1989a, b; Takhtajan, 1985, 1987;
Thorne, 1992). These names are validated below
by direct reference to a valid description (Art. 32).
A catalog of the validly published, automatically
typified ordinal names is in preparation and will be
presented in the near future.
All names proposed here are based on and au¬
tomatically typified by the type of the genus of an
included family name (Art. 16.1). All proposed new
names are currently accepted by Gronquist (1981,
1988), Dahlgren (in G. Dahlgren, 1989a, b), Takh¬
tajan (1986, 1987), or Thorne (1992). Authors
adopting these names are noted by the use of C, D,
T, or Th respectively.
Actinidiales (J. Hutchinson) Takhtajan ex Reveal,
stat. nov. Based on Actinidiaceae J. Hutchison,
Fam. FI. PI. 1: 177. 1926, nom. cons. TYPE:
Actinidia Findley (1836). T.
Ancistrocladales (J. E. Planchon ex Walpers)
Takhtajan ex Reveal, stat. nov. Based on An-
cistrocladaceae J. E. Planchon ex Walpers, Ann.
Bot. Syst. 2: 175. 1851, nom. cons. TYPE:
Ancistrocladus Wallich (1829), nom. cons. T.
Aralidiales (Philipson & B. G. Stone) Takhtajan
ex Reveal, stat. nov. Based on Aralidiaceae
Philipson & B. G. Stone, Taxon 29: 402. 1980.
TYPE: Aralidium Miquil (1856). T.
Austrobaileyales (Groizat) Takhtajan ex Reveal,
stat. nov. Based on Dilleniaceae subfam. Aus-
trobaileyoideae Groizat, J. Arnold Arbor. 21:
404. 1940. TYPE: Austrobaileya C. T. W hite
(1933). T.
Buxales (I)umortier) Takhtajan ex Reveal, stat.
nov. Based on Buxaceae Dumortier, Comment.
Bot. 54. 1822, nom. cons. TYPE: Buxus Lin¬
naeus (1753). D, T, TH.
Cercidiphyllales (Engler) H-H. Hu ex Reveal,
stat. nov. Based on Cercidiphyllaceae Engler,
Syllabus ed. 6, 132. 1909, nom. cons. TYPE:
Cercidiphyllum Siebold & Zuccarini (1846).
D, T, TH.
Chlorantbales (Blume) Conzatti & L. C. Smith ex
Reveal, stat. nov. Based on Chloranthaceae
Blume, Enum. PI. Javae 1: 78. 1827, nom.
cons. TYPE: Chloranthus Swartz (1789). D, T.
Crossosomatales (Engler) Takhtajan ex Reveal,
stat. nov. Based on Crossosomataceae Engler
in Engler & Prantl, Nat. Pflanzenfam. Nachtr.
[1:J 185. 1897, nom. cons. TYPE: Crosso-
soma Nuttall (1848). T.
Dioncophvllales (Gilg) Takhtajan ex Reveal, stat.
nov. Based on Flacourtiaceae subtrib. Dion-
eophyllinae Gilg in Engler & Prantl, Nat. Pflan¬
zenfam. ed. 2, 21: 390, 420. 1925. TYPE:
Dioncophyllum Baillon (1890). T.
Euponiatiales (Endlicher) Takhtajan ex Reveal,
stat. nov. Based on Eupomatiaceae Endlicher,
Ench. Bot. 425. 1841, nom. cons. TYPE: Eu-
pomatia R. Brown (1814). T.
Eupteleales (K. Wilhelm) H-H. Hu ex Reveal, stat.
nov. Based on Eupteleaceae K. Wilhelm, Sa-
menpfl. 17. 1910, nom. cons. TYPE: Euptelea
Zuccarini (1840-1841). C. T.
Fouquieriales (A. P. de Candolle) Takhtajan ex
Reveal, stat. nov. Based on Fouquieriaceae A.
I’, de Candolle, Prodr. 3: 349. 1828, nom.
cons. TYPE: Fouquieria Kunth (1823). 1), T,
TH.
Geissolomatales (Endlicher) Takhtajan ex Reveal,
stat. nov. Based on Geissolomataceae Endlich¬
er, Ench. Bot. 214. 1841, nom. cons. TYPE:
Geissoloma Lindley ex Kunth (1830). I), T.
Glaucidiales (Tamura) Takhtajan ex Reveal, stat.
nov. Based on Glaucidiaceae Tamura, Bot. Mag.
(Tokyo) 85: 40. 1972. TYPE: Glaucidium Sie¬
bold & Zuccarini (1846). T.
Novon 2: 238-240. 1992.
Volume 2, Number 3
1992
Reveal
Ordinal Names of Extant Vascular Plants
239
Gunnerales (Meisner) Takhtajan ex Reveal, stat.
nov. Based on Gunneraceae Meisner, PI. Vase.
Gen.: Tab. Diagn. 345, 346, Comm. 257. 1841,
nom. cons. TYPE: Gunnera Linnaeus (1767).
D, T.
Hanguanales (Airy Shaw) R. Dahlgren ex Reveal,
stat. nov. Based on Hanguanaceae Airy Shaw,
Kew Bull. 18: 260. 1965. TYPE: Hanguana
Blume (1827). D.
Hydnorales (C. A. Agardh) Takhtajan ex Reveal,
stat. nov. Based on Hydnoraceae C. A. Agardh,
Aphor. Bot. 88. 1821, nom. cons. TYPE: Hyd-
nora Thunberg (1775). T.
Hydrostachyales (Engler) Diels ex Reveal, stat.
nov. Based on Hydrostachyaceae Engler, Syl¬
labus ed. 2, 125. 1898, nom. cons. TYPE:
Hydrostachys l)u Petit-Thouars (1806). 1), T.
Lactoridales (Engler) Takhtajan ex Reveal, stat.
nov. Based on Lactoridaceae Engler in Engler
& Prantl, Nat. Pflanzenfam. Ill, 2: 19. 1888,
nom. cons. TYPE: Lactoris R. A. Philippi
(1865). D. T.
Melanthiales (Batsch) R. Dahlgren ex Reveal, stat.
nov. Based on Melanthiaceae Batsch, Tab. Af-
fin. Regni Veg. 133. 1802, nom. cons. TYPE:
Melanthium Linnaeus (1753). D.
Ochnales (A. P. de Candolle) ,}. Hutchinson ex
Reveal, stat. nov. Based on Ochnaceae A. 1’.
de Candolle, Ann. Mus. Natl. Hist. Nat. 17:
410. 1811, nom. cons. TYPE: Ochna Linnaeus
(1753). T.
Paracryphiales (Airy Shaw) Takhtajan ex Reveal,
stat. nov. Based on Paracryphiaceae Airy Shaw,
Kew Bull. 18: 265. 1965. TYPE: Paracry-
phia E. G. Baker (1921). T.
Podocarpales (Endlicher) Pulle ex Reveal, stat.
nov. Based on Podocarpaceae Endlicher, Syn.
Conif. 203. 1847, nom. cons. TYPE: Podo-
carpus L'Heritier ex Persoon (1807), nom.
cons. T.
Khoipteleales (Handel-Mazzetti) Novak ex Re¬
veal, stat. nov. Based on Rhoipteleaceae Han¬
del-Mazzetti, Repert. Spec. Nov. Regni Veg.
30: 75. 1932, nom. cons. TYPE: Rhoiptelea
Diels & Handel-Mazzetti (1932). T.
Salvadorales (Lindley) R. Dahlgren ex Reveal,
stat. nov. Based on Salvadoraceae Lindley, Nat.
Syst. Bot. ed. 2, 269. 1836, nom. cons. TYPE:
Salvadora Linnaeus (1753). I).
Simmondsiales (Muller Argoviensis) Reveal, stat.
nov. Based on Buxaceae trib. Simmondsieae
Miiller Argoviensis in A. P. de Candolle, Prodr.
16(1): 9, 22. 1869. TYPE: Simmondsia Nut-
tall (1844). T.
Stylidiales (R. Brown) Takhtajan ex Reveal, stat.
nov. Based on Stylidiaceae R. Brown, Prodr.
565. 1810, nom. cons. TYPE: Stylidium
Swartz ex Willdenow (1805), nom. cons. D, T.
Toricelliales (Wangerin) Takhtajan ex Reveal, stat.
nov. Based on Cornaceae trib. Toricellieae
Wangerin in Engler, Pflanzenr. IV, 229(Heft
41): 18, 31. 1910. TYPE: Toricellia A. P. de
Candolle (1830). T.
Tropaeolales (A. L. de Jussieu ex A. P. de Can¬
dolle) Takhtajan ex Reveal, stat. nov. Based
on Tropaeolaceae A. L. de Jussieu ex A. P. de
Candolle, Prodr. 1: 683. 1824, nom. cons.
TYPE: Tropaeolum Linnaeus (1753). 1), T.
Velloziales (Endlicher) R. Dahlgren ex Reveal,
stat. nov. Based on Velloziaceae Endlicher,
Ench. Bot. 101. 1841, nom. cons. TYPE: Vel¬
io zia Vandelli (1788). D, T.
Welwitschiales (Markgraf) Pulle ex Reveal, stat.
nov. Based on Welwitschiaceae Markgraf in
Engler & Prantl, Nat. Pflanzenfam. ed. 2, 13:
419. 1926, nom. cons. TYPE: Welwitschia J.
D. Hooker (1862), nom. cons. T.
Winterales (R. Brown ex Lindley) A. C. Smith ex
Reveal, stat. nov. Based on Winteraceae R.
Brown ex Lindley, Intr. Nat. Syst. Bot. 26:
1830, nom. cons. TYPE: IT interin J. A. Mur¬
ray (1784), nom. illeg. = Drimys J. R. & G.
Forster (1775). D, T.
Acknowledgments. I thank R. D. Hooglaiul, A.
Cronquist, A. L. Takhtajan, and R. F. Thorne for
comments and help. This work is supported, in part,
by National Science Foundation Grant BSR-8812816
to the Missouri Botanical Garden. This is Scientific
Article A-6284, Contribution No. 8453, of the
Maryland Agricultural Experiment Station and Co¬
operative Extension Service.
Literature Cited
Cronquist, A. 1981. An Integrated System of Classifi¬
cation of Flowering Plants. Columbia Univ. Press,
New York.
-. 1988. The Evolution and Classification of
Flowering Plants, 2nd ed. New York Botanical Gar¬
den, Bronx.
Dahlgren, G. 1989a. The last Dahlgrenogram: System
of classification of the dicotyledons. Pp. 249-260 in
K. Tan, R. R. Mill & T. S. Elias (editors), Plant
Taxonomy, Phytogeography and Related Subjects.
Edinburgh Univ. Press, Edinburgh.
-. 1989b. An updated angiosperm classification.
J. Linn. Soc., Bot. 100: 197-203.
Greuter, W., H. M. Burdet, W. G. Chaloner, V. Demoulin,
R. Grolle, D. L. Hawksworth, D. H. Nicolson, P. C.
Silva, F. S. Stafleu, E. G. Voss & J. McNeill (editors).
1988. International code of botanical nomenclature
adopted by the Fourteenth International Botanical
240
Novon
Congress, Berlin, July “August 1987. Regnum Veg.
118.
Takhtajan, A. L. 1985. Floristic Regions of the World.
Univ. California Press, Berkeley.
-. 1987. Systems Magnoliophytorum. Nauka,
Lenningrad.
Thorne, R. F. 1992. An updated phylogenetic classi¬
fication of the flowering plants. Aliso 13: 365-389.
New Natural Hybrids and Nomenclatural Novelties in Catasetum
(Orchidaceae) from the Guianas, Ecuador, and Peru
Gustavo A. Romero
Oakes Ames Orchid Herbaria, Harvard University Herbarium,
22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A.
Rudolf Jenny
Moosweg 9, 3112 Allmendingen, Switzerland
ABSTRACT. One new Catasetum natural hybrid from
Guyana is described and illustrated, Catasetum
X guianense, and two taxa previously described as
species are shown to be hybrids, Catasetum x so-
diroi Schltr. and C. x violascens Reichb. f. & Warzc.
The latter two are neotypified and lectotypified, re¬
spectively. In addition, a lectotype is designated for
Catasetum incurvum Klotzsch.
An understanding of the pollination biology of
Catasetum species assemblages and careful atten¬
tion to reproductive isolation mechanisms have helped
to distinguish species from natural hybrids (Dodson,
1962; Romero & Carnevali, 1992, and references
therein). Here we present three natural hybrids from
distinct geographical areas where species from Ca¬
tasetum subg. Catasetum and Catasetum subg.
Pseudocatasetum are found within euglossine bee
flight range. The hybrids discussed encompass two
of three possible combinations between these two
groups: one between Catasetum longifolium Lindley
(subg. Pseudocatasetum) and C. macrocarpum
(subg. Catasetum , sect. Catasetum) from the Guian¬
as; one between Catasetum macroglossurn and C.
expansum (both in sect. Catasetum) from Ecuador;
and one between Catasetum discolor Lindley (subg.
Pseudocatasetum ) and C. incurvum Klotzsch (subg.
Catasetum , sect. Catasetum) from Peru.
Catasetum x guianense G. Romero & Jenny, no-
thosp. nov. TYPE: Guyana. Essequibo District,
Dawa, Lake Tapakuma, 30 m, 18 Oct. 1974,
C. H. Dodson, ex hort. Marie Selby Botanical
Gardens sub Dodson 4476 (holotype, SEL; ho-
lotype fragment, AMES). Figure 1.
Planta intermedia ad Catasetum macrocarpum Rich,
ex Kunth vergens, sed ab eo floribus minoribus, columnae
antennis parallelis brevioribus et clinandrio apiculo brev-
iore recedit.
Plant epiphytic, vegetatively indistinguishable from
other members of the genus except for the pendent
habit and the generally longer, narrower, linear leaves
(to 75 cm long and 4 cm wide). Staminate inflores¬
cences pendent, racemose, to 10-flowered, to 30
cm long, developing from fully developed pseudo¬
bulbs at the base of the first internode. Staminate
flower resupinate. Sepals and petals yellow, wine
red, or greenish yellow spotted wine red. Dorsal sepal
slightly concave, ovate, acute, 23-25 mm long, 11-
13 mm wide; lateral sepals slightly concave, oblong-
obovate, acute, slightly oblique, 25-27 mm long,
11-13 mm wide; petals broadly elliptic, acute to
shortly apiculate, 18-19 mm wide, 28 29 mm long.
Labellum greenish yellow to yellow, internally yellow
with wine red spots toward the base or entirely wine
red, saccate, the sac 15-17 mm deep, slightly con¬
stricted and internally thickened at the opening, the
opening subcircular, 12-17 mm diam.; labellum
shallowly trilobate apically, with a ridge inside near
the apex uniting the lateral lobes behind the mid¬
lobe, the ridge separated from the mid-lobe by a
shallow depression; lateral lobes erect patent, up to
10 mm wide, margins dentate, fimbriate toward the
base; middle lobe 2-5 mm long, linguiform, the apex
acuminate, acute, to truncate. Column light greenish
white to gold, semiterete, straight, apically thick¬
ened, shortly rostrate, 16 mm long, 6 mm high, and
10 mm wide; clinandrium shortly apiculate; anten¬
nae tapering to a fine point, bilaterally symmetrical,
6-7 mm long, 1.25 mm diam. at the base. Anther
orange to yellowish green, 8 mm long, 5 mm wide;
viscidium semicircular, 6-7 mm diam.; stipe, before
folding, cymbiform, 11-12 mm long and 5-6 mm
wide; pollinia two, cleft, waxy, yellow, compressed,
4 mm long and 3 mm wide. Pistillate inflorescence
and flower unknown.
Paratypes. Guyana. Essequibo District, Dawa, Lake
Tapakuma, 30 m, 1 Dec. 1975, C. H. Dodson, ex hort.
Marie Selby Botanical Gardens sub Dodson s.n. (SEL).
French Guiana. Without locality, 1 Nov. 1990, Lafon-
taine s.n. (AMES). SURINAME. Without locality, ex hort.
G. Schiller, W (Herb. Reichenbach No. 24634).
Novon 2: 241-248. 1992.
242
Novon
Figure 1. Catasetum x guianense G. Romero & Jenny. — A, B. Front and side view of staininate flower. —C.
Anther. — D. Pollinium. —E. Pollinarium (pollinia removed). —F. Sepals and petals. (A F drawn from Lafontaine
s.n.)
Volume 2, Number 3
1992
Romero & Jenny
Hybrids and Novelties in Catasetum
243
Etymology. Named after the Guianas, a region
in northern South America encompassing Guyana,
Suriname, and French Guiana, the known geograph¬
ical range of this natural hybrid.
Dodson (1978) first reported this natural hybrid
between Catasetum macrocarpum Rich, ex Kunth
and C. longifolium Lindley without providing a for¬
mal name for the taxon. Photographs of the same
hybrid swarm had previously been published by Tan
(1971, fig. 8). Lafontaine (1990) recently identified
the hybrid as Catasetum blepharochilum Schltr.
However, this binomial (Schlechter, 1920, 1929) is
a synonym of C. maculatum Kunth and is not the
hvbrid described here. According to Dodson (1978),
the hybrid origin of C. x guianense “. . . was con¬
firmed by later hybridization experiments.”
Catasetum x guianense is morphologically in¬
termediate between the two parents. It differs from
C. longifolium in the larger flower (dorsal sepal to
25 mm long in the hybrid vs. 15 mm long in C.
longifolium) and longer antennae (to 7 mm in the
hvbrid vs. 3 mm in C. longifolium). It differs from
C. macrocarpum in the orientation of the inflores¬
cence (pendent in the hybrid vs. erect in C. ma¬
crocarpum) and the antennae (bilaterally symmet¬
rical in the hybrid vs. bilaterally asymmetrical in C.
macrocarpum). The hybrid is probably pollinated
by the same euglossine bees that pollinate the par¬
ents: Eulaema bombiformis, E. cingulata , and E.
meriana. Pollen flow between the parents of this
hybrid, as in other Catasetum hybrids (e.g., C.
x dunstervillei; Romero & Carnevali, 1989a), is
unidirectional: only the pollinia of Catasetum lon¬
gifolium can pollinate female flowers of C. macro¬
carpum (the pollinia of C. macrocarpum are too
big to fit in the stigmatic cleft of female flowers of
C. longifolium). This hybrid has been found on
Mauritia palms (Dodson, 1978).
Catasetum incurvum Klotzsch, Allg. Gartenzei-
tung 22: 178. 1854. Catasetum saccatum
Lindley var. incurvum (Klotzsch) Mansf., Re-
pert. Spec. Nov. Regni Veg. 30: 272. 1932.
TYPE: Peru. Without precise locality, J. War-
scewicz ex hort. Mathieu (holotype, B de¬
stroyed; lectotype, here designated, W (Herb.
Reichenbach No. 24568, pro parte)). Figures
2, 3B.
Catasetum stupendum Cogn., J. Orchidees 6: 13. 1895.
TYPE: Peru. Without precise locality, ex hort.
L’Horticulture Internationale (type not located).
Catasetum trautmanii Senghas, Orchidee 41: 216. 1990.
Syn. nov. TYPE: Peru. Without precise locality,
Dec. 1986, G. Trautmann ex Hort. Botanischer
Garten Heidelberg 643 (holotype, HEID not seen).
Other specimens examined. Peru. AMAZONAS: La Peca,
1,000 m, 21 Sep. 1963, F. Woytkowski 2 (AMES, F,
K, MO, US), 20 Aug. 1964, Woytkowski 4 (AMES), 9
June 1966, Woytkowski 8 (AMES); canyon of the Rio
Maranon, below Milagro on the Mesones Muro highway,
Quebrada Aramango, 24 June 1966, P. C. Hutchinson
1547 (BR); canyon of the Rio Maranon, between km 247
and 250 of the Mesones-Muro highway, 1959, Hutch¬
inson 1559 (AMES); Rio Utcubamba, 6 km E of Due-
brada Honda, km 276 E of Olmos, 26 Aug. 1966, Hutch¬
inson & J. K. Wright 6776 (AMES); Rio Utcubamba
Canyon, 1,800 m, 6 Apr. 1965, F. Antich s.n. (AMES);
vicinity of Campamento Ingenio 1-3 km up road to Po-
macocha (and Rioja), 1,300 1,400 m, 27 Jan. 1964,
Hutchinson & Wright 3836 (AMES).
Catasetum incurvum has been treated as a va¬
riety of C. saccatum. However, we believe this taxon
should be recognized at the rank of species based
on its larger size and the shape of the labellum
(Lindley, 1855). Furthermore, C. incurvum is re¬
stricted to mid- to high-altitude rainforests (600-
l, 800 m) in the Peruvian Amazon, whereas C. sac¬
catum is widespread in lowland forests (below 500
m) in the Guianas, Brazil, Ecuador, Peru, and Bo¬
livia. The holotype of C. incurvum was presumably
destroyed during the bombing of Berlin. Fortunately,
Reichenbach made a drawing of one of the flowers
described by Klotzsch during one of his visits to this
important herbarium. This drawing is selected as
lectotype (Fig. 3B).
Catasetum macroglossum Reichb. f. Card. Chron.
n.s. 8: 552, 1877. TYPE: Ecuador. May 1877,
ex hort. Strickland (lectotype, here designated,
W (Herb. Reichenbach No. 38701)).
Other specimens examined. Ecuador. LOS RIOS: km
56 Quevedo-Santo Domingo, Rio Palenque Biological Sta¬
tion, 150-220 m, 6 Mar. 1974, Dodson 5491; Quevedo,
700 m, 5 June 1978, ex hort. Marie Selby Botanical
Gardens, Dodson 3236 (SEL). GUAYAS: near Bucay, 150
m, 28 Aug. 1978, ex hort. Marie Selby Botanical Gardens,
Dodson 3235 (SEL); El Progreso, Guayas, 14 Aug. 1940,
ex hort. Puerto Rico Experimental Station, A. G. Ker-
orkian 6199 (AMES).
Reichenbach included several “varieties” of C.
macroglossum in his protologue (“I have had them
ochre coloured, with green sepals and petals, quite
light green, dark green, light yellow, dark yellow,
and brownish purple. I have seen it rather frequent¬
ly, first from Messrs. Veitch, at different times from
Mr. Day, and then from Sir C. W. Strickland, and
various other correspondents since last November”).
Among these varieties were several representatives
of the natural hybrid between C. macroglossum and
C. expansum , as well as what we currently know
as C. macroglossum (Dodson, 1962: 52, fig. 10;
Dodson & Dodson, 1980; Arosemena et al., 1988:
244
Novon
Figure 2. Catasetum incurvum Klotzsch. —A, B. Side and front view of staminate flower. —C. Pollinarium (pollinia
removed). —D. Sepals and petals. —E. Pollinium. (A-E drawn from Hutchinson <£■ Wright 3836.)
Volume 2, Number 3
1992
Romero & Jenny
Hybrids and Novelties in Catasetum
245
Figure 3. —A. Lectotype of Catasetum macroglossum (from Flerb. Reichenbach 38701.) —B. Lectotype of
Catasetum incurvum Klotzsch (from Herb. Reichenbach 24568).
37, fig. 38). Reichenbach annotated at least two
specimens as C. macroglossum in his herbarium
without designating a type. We have chosen a lec¬
totype (Fig. 3A) that maintains the currently ac¬
cepted concept of C. macroglossum. There are six
paintings of “ Catasetum macroglossum ” in John
Day’s scrapbooks (K). Of these, only one corre¬
sponds to our current concept of C. macroglossum
(Book 20: 51, December 1876, as Catasetum ma¬
croglossum var. viridi-pallens). The other five de¬
pict different forms of C. x sodiroi (Book 20: 79,
1 Feb. 1877; Book 20: 81, 3 Feb. 1877; Book 21:
61, 18 Nov. 1879; Book 23: 13, 26 Dec. 1876;
Book 23: 59, 22 Apr. 1878).
Catasetum x sodiroi Schltr. (pro sp.)
Catasetum sodiroi Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 8: 91. 1921. TYPE: Ecuador. Chimborazo:
in silvis tropicis, Puente de Chimbo (holotype, B
destroyed). Ecuador. Road Guayaquil-Quevedo, ca.
km 78, 30 m, 5 Apr. 1960, Dodson 38 (neotype,
here designated, SEL; isoneotypes, MO, SEL).
Catasetum triloba!um Senghas, Orchidee 41: 218. 1990.
Syn. nov. TYPE: Ecuador. Cotopaxi: near Pucay-
acu, ca. 600 m, W. Rauh ex hort. Botanischer
Garten Heidelberg 34459 (holotype, HE1D).
Other specimen examined. Ecuador, guayas: Pales-
tina, 70 m, 6 July 1978, ex Hort. Marie Selby Botanical
Gardens, Dodson 3234 (SEL).
Dodson (1962) proposed Catasetum macroglos¬
sum Reichb. f. and C. expansum Reichb. f. as the
parents of this hybrid swarm, providing solid evi¬
dence to support his hypothesis. We selected Ca¬
tasetum sodiroi as the earliest name because plants
collected in the type locality of C. sodiroi Schltr.
(currently known as Bucay) bear flowers that re¬
semble the natural hybrid (C. H. Dodson, pers.
comm.). A neotype was chosen because a search of
those herbaria with known Sodiro’s collections (BP,
BR, K, MO, NY, Q, QPLS, US) failed to produce
authentic material of Catasetum sodiroi.
As in the case of Catasetum X tapiriceps Reichb.
f. (Romero & Carnevali, 1989b, 1990), C. x sodiroi
exhibits a wide range of variation in flower mor¬
phology, resupination, and color. It should be em¬
phasized, however, that “. . . a nothotaxon is cir¬
cumscribed so as to include all individuals (as far as
246
Novon
they can he recognized) derived from the crossing
of representatives of the stated parent taxa (i.e., not
only the F,, but subsequent filial generations, and
also back-crosses and combinations of these)” (Greu-
ter, 1988: 83, article H.4.1). Thus, the name Ca-
tasetum x sodiroi Schltr. applies to the entire hybrid
complex, regardless of its morphological variability.
The hybrid differs from C. expansum in the labellum
that is generally trilobate (vs. entire in C. expansum)
or hooded (vs. open, cup-shaped in C. expansum).
It differs from C. macroglossum in the resupinate
flowers (vs. nonresupinate in C. macroglossum ), the
presence of a conspicuous, triangular callus in the
labellum (vs. a semicircular callus in C. macroglos¬
sum), and the yellow spotted with red to orange
color of the flowers (vs. green to greenish yellow in
C. macroglossum). Both the hybrids and the parents
are pollinated by Eulaema cingulata, E. meriana,
and E. speciosa (Dodson, 1962). Color photographs
of the hybrids may be seen in Arosemena et al.
(1988, cover and pp. 27-36).
Catasetum x violascens Reichb. f. & Warsc. (pro
sp.)
Catasetum violascens Reichb. f. & Warsc., Bonplandia
2: 97. 1854. TYPE: Peru. Sources of the Maranon,
May 1853, J. von Warscewicz 14, (lectotype, des¬
ignated here, K). Figure 4.
Other specimen examined. Peru. Moyobamba, G.
Monnier 1030 (AMES).
In 1854, Josef von Warscewicz sent several ship¬
ments of plants collected in Peru to horticultural
houses in Europe. As in the case of Catasetum
macroglossum, some of the collections included a
Catasetum natural hybrid and one of its parents.
The parent was first described as Catasetum incur-
vum by Klotzsch lrom a plant cultivated in Germany
(see C. incurvum above). Bindley used the name
Catasetum secundum in his herbarium and in one
of Warscewicz’s catalogs, but the binomial was never
validly published. Reichenbach and Warscewicz lat¬
er described this collection as C. violascens. Ca¬
tasetum x violascens is similar to C. x dunstervillei
G. Romero & Carnevali, a natural hybrid between
C. pileatum and C. discolor (Romero & Carnevali,
1989a, 1991). This similarity reveals a well-estab¬
lished pattern in the inheritance of certain char¬
acters in Catasetum hybrids: crosses between spe¬
cies in subgenus Pseudocatasetum (with minute
antennae, usually under 3 mm long) and species in
section Catasetum (with bilaterally asymmetric, long
antennae, usually over 15 mm long) produce prog¬
eny with short, stout, bilaterally symmetric anten¬
nae. We propose C. incurvum (Fig. 2) and C. dis¬
color as putative parents based on the following
evidence: (1) the morphology of the hybrid (inter¬
mediate between the proposed parent species) in¬
cluding the shape of the antenna mentioned above;
(2) they are the only possible species in the known
range of the hybrid capable of producing such an
intersubgeneric hybrid.
Catasetum x violascens differs from C. discolor
in the conspicuous antennae (vs. inconspicuous in
C. discolor) and the resupinate staminate flowers
(vs. nonresupinate in C. discolor). It differs from C.
incurvum in the parallel, stout antennae (vs. long,
slender, bilaterally asymmetrical antennae in C. in¬
curvum) and the much smaller flowers (compare
Figs. 2 and 4). Plants of Catasetum x violascens
were found on the side of a road, growing with
grasses in clay soils (G. Monnier, pers. comm.). This
growth habit is probably inherited from C. discolor,
also a terrestrial species: Catasetum incurvum is
always epiphytic. Little is known about the polli-
nator(s) of the hybrid or the putative parents except
that in Venezuela and the Guianas C. discolor is
pollinated by Eulaema bombiformis, E. cingulata,
and E. meriana (Dodson, 1978; G. A. Romero,
unpublished data).
A single flower of Warscewicz’s original material
found at K is here designated as lectotype for Ca¬
tasetum violascens; a tracing of this flower was
found in Vienna (Herb. Reichenbach No. 24609,
upper right corner). What appears to be the original
of Warscewicz’s drawing 14 is at W (Herb. Rei¬
chenbach No. 24628) and a tracing of it at K.
Acknowledgments. We thank the curators of BP,
BR, F, K. MO, NY, SEL, US, and W for providing
access to their collections; G. Monnier and F. Paget
for providing specimens from their live collections;
G. Carnevali and E. Christenson for useful comments
on an earlier version of this manuscript; and K.
Brown-Wing and B. Burkhardt for the line drawings.
Literature Cited
Arosemena, A., C. de Jurado, R. Estrada G. & M. Konanz
M. 1988. Orchids from the coast of Ecuador. Aso-
ciacion Ecuatoriana de Orquideologia. Guayaquil, Ec¬
uador.
Dodson, C. H. 1962. Pollination and variation in the
subtribe Catasetinae (Orchidaceae). Ann. Missouri
Bot. Gard. 49: 35-56.
-. 1978. The Catasetums (Orchidaceae) of Ta-
pakuma, Guyana. Selbyana 2: 159-168.
- & P. M. Dodson. 1980. Catasetum macrog¬
lossum Reichb. f. Icon. PI. Trop., ser. 1, 1: t. 18.
Greuter, W. 1988. International Code of Botanical
Nomenclature. Koeltz Scientific Books, Konigstein,
Germany.
Lafontaine, A. 1990. Quelques representants du genre
Volume 2, Number 3
1992
Romero & Jenny
Hybrids and Novelties in Catasetum
247
5 mm +■
Figure 4. Catasetum ~x violascens Reichb. f. & Warsc. —A, B. Side and front view of staminate flower. —C.
Sepals and petals. —D. Pollinium (A-D drawn from G. Monnier 1030.)
248
Novon
Catasetum L. C. Richard. L'Orchidophile No. 90:
21-25.
Lindley, J. 1855. Catasetum incurvum Klotzsch. Card.
Chron. 1855: 4.
Romero, G. A. & G. Carnevali. 1989a. Novelties in
the orchid flora of southern Venezuela. Ann. Missouri
Bot. Card. 76: 454-461.
- & -. 1989b. A new combination for
the orchid flora of southern Venezuela: Catasetum
x tapiriceps Reichb. f. Lindleyana 4: 127 134.
-&-. 1990. Catasetum natural hybrids
from southern Venezuela. I: Catasetum x tapiriceps
Reichb. f. Amer. Orchid Soc. Bull. 59: 1214 1220.
- & -. 1991. Catasetum natural hybrids
from southern Venezuela. II: Catasetum xdurister-
villei G. Romero & Carnevali. Amer. Orchid Soc.
Bull. 60: 115-120.
-&-. 1992. Catasetum natural hybrids
from southern Venezuela IV: Biology and nomencla¬
ture. Amer. Orchid Soc. Bull. 61: 355-360.
Schlechter, R. 1920. Die Orchideenfloren der Siida-
merikanischen Kordillerenstaaten. II. Colombia. Re-
pert. Spec. Nov. Regni Veg. Beih. 7: 158.
-. 1929. Figuren-Atlas zu den Orchideenfloren
der Siidamerikanischen Kordillerenstaaten. Repert.
Spec. Nov. Regni Veg. Beih. 57: t. 56, fig. 216.
Tan, K. W. 1971. Orchids of Dawa, Guyana. Amer.
Orchid Soc. Bull. 40: 585-591.
Taxonomic Notes on Mesoamerican Annona Section
Atta (Annonaceae), Including Annona pruinosa sp. nov.
George E. Sehatz
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Abstract. A new species of Annona sect. Atta
from the upper altitudinal limit of the Pacific dry
forest of Nicaragua and Costa Rica is proposed based
upon a distinctive abaxial leaf epidermis. In addition,
the analysis of flowers discovered for Annona lon-
gipes necessitates its transfer to section Atta.
During preparation of treatments of the Anno¬
naceae for the Flora de Nicaragua and the Manual
to the Plants of Costa Rica , a new species of
Annona L. sect. Atta C. Martius has been identified
based on material from Nicaragua and Costa Rica.
In addition, recent flowering collections of Annona
longipes Saff. from Veracruz, Mexico, reveal its
incorrect placement by Safford (1913) in section
Pilannona Saff., and indicate its proper placement
in section Atta.
Annona (section Atta C. Martius) pruinosa Sehatz,
sp. nov. TYPE: Nicaragua. Carazo: Finca de
Minco Cabrales, 700 m al sur del reloj, 18
June 1982 (fl, fr), L. Reyes V. 64 (holotype,
MO; isotype, HNMN). Figure 1.
A speciebus aliis Annonae sectionis Attae laminis subtus
pruinosis differt.
Tree to 10 m tall; young branches very sparsely
white-tomentose, at length glabrescent. Petiole 0.8-
1.3 cm long, slender, deeply canaliculate, very
sparsely tomentose, drying black; lamina charta-
ceous, elliptic to obovate-elliptic, 6.4-15 cm long,
3.2- 8.3 cm broad, the base cuneate to obtuse, the
apex acute to rounded, the upper surface glabrous,
the lower surface granular pruinose and initially
sparsely white-puberulous, at length glabrescent, the
venation eucamptodromous with 9-14 secondary
veins per side, the primary vein slightly impressed
adaxially, prominently elevated and sparsely golden
puberulous, as are the slender secondary veins,
abaxially. Flowers solitary, terminal but appearing
leaf-opposed or supra-axillary by displacement dur¬
ing growth of the renewal shoot; pedicel slender,
1.3- 2.4 cm long, to 3.1 cm long, 0.2 cm diam.
and becoming rigid in fruit, white-tomentose, bearing
a minute bract 0.8 0.9 cm from the base; sepals
triangular, 0.2-0.3 cm long, 0.3 cm broad, the apex
acute, densely white-tomentose; petals 3 (the outer
petal whorl only, the inner whorl absent or at most
represented by tiny vestigial petals), fleshy, trique¬
trous, narrowly oblong-elliptic, 1.8-2.4 cm long,
0.5-0.8 cm broad, the apex acute to obtuse, the
base concave inside, the outer surface white-seri¬
ceous toward the base, becoming sparsely puberu¬
lous toward the apex, the inner surface densely
tomentose; stamens numerous, 1 mm long, the con¬
nective expanded truncate discoid, minutely papil¬
lose, yellow, contrasting with the white thecae; car¬
pels 25-36. Fruit syncarpous, subglobose to broadly
ovoid, to 4.5 cm long, to 4.5 cm diam., the apex
rounded, the surface initially areolate, but then
smooth with no evidence of the individual carpels
at maturity, green, or light and dark green mottled,
initially puberulous, at length glabrescent; seeds flat¬
tened ellipsoid-obovoid, to 1.6 cm long, 0.9 cm
broad, 0.6 cm thick, the seed coat very thin, smooth-
reticulate, light brown.
Paratypes. COSTA RlCA. GUANACASTE: Parque Rincon
de la Vieja, Hacienda Santa Maria, sendero a las pailas,
10°48'N, 85°10'W, 700-800 m, 13 Aug. 1987 (fr), G.
Herrera 693 (CR, MO); El Mirador, Rio Negro,
10°47'40"N, 85°18'35"W, 1 Oct. 1990 (fr), G. Rivera
679 (C.R, MO). PUNTARENAS: San Luis, Monteverde, Rio
Guacimal, 10° 16 ; N, 84°49'W, 700 m, 24 June 1988
(fr), Bello rt al. 24 (CR, MO, U), Canton de Puntarenas,
Monteverde, cliff edge on Pacific slope, Rajo Tigre trail,
10°18'N, 84°48'W, 900 m, 20 Oct. 1990 (fr), Haber
& Zuchowski 10110 (CR, MO), 27 Dec. 1991 (fr), Sehatz
et al. 3230 (MO). Nicaragua, chontai.es: ca. 2.8 km
above (N of) Cuapa, ca. 12°17'N, 85°23'W, 400-500
m, 4 Sep. 1977 (fr), Stevens 3634 (HNMN, MO, WIS);
2 3 km NE of Cuapa, 12°17'N, 85°22'W, 400 m, 24
Sep. 1983 (fr), Nee & Sebastian 28486 (HNMN, MO,
WIS). BOACO: Las Pitas, carretera a Camoapa (No. 19),
1 2°28'N, 85°35'W, ca. 400 m, 29 Aug. 1981 (fr), Mo¬
reno 10651 (HNMN, MO).
Distribution and habitat. Annona pruinosa is
known from southern Nicaragua and the Pacific
slope of the Tilaran range in northwestern Costa
Rica, in moist forest between 400 and 800 m, i.e.,
the upper altitudinal limit of the Pacific dry forest.
The affinities of Annona pruinosa within section
Atta lie probably with several Caribbean species (A.
urbaniana R. E. Fries, from Haiti; A. praetermissa
Fawcett & Rendle, from Jamaica; and A. cubensis
Novon 2; 249-251. 1992.
250
Novon
Figure 1. Annona pruinosa Schatz, flowering and fruiting branch (from L. Reyes V. 64). Inset: magnification of
abaxial leaf surface, x200.
K. E. F ries, from Cuba), all of which possess glaucous
leaf undersides. In Costa Rica, collections of A.
pruinosa have been referred to A. lutescens Saff.,
described from southern Mexico and Guatemala,
which, however, possesses a fruit twice as large as
A. pruinosa and lacks the pruinose leaf underside.
Annona lutescens is probably better placed in syn¬
onymy as merely a broad-leaved variant of A. re¬
ticulata L.
Annona (section Atta C. Martius) longipes Safi’.,
Contr. U.S. Natl. Herb. 16(10): 269, pi. 89.
1913. TYPE: M exico. Veracruz: Canton de los
Tuxtlas, near the outlet of Lake Catemaco, 28
Apr. 1894 (lr), Nelson 430 (holotype, US;
isotype, NY).
Additional specimens examined. MEXICO. VERACRUZ:
Laguna Encantada, 2 Nov. 1971 (st), Beaman 5242
(XAL); 5 km SW of Santiago Tuxtla and Hwy. 180 at
bridge over the Rio Tepango (Rio Grande), 18°27'N,
95°19'W, 160 m, 14 Dec. 1985 (fl). Nee 32131 (NY,
WIS, XAL), 5 Apr. 1983 (fl, fr), Nee & Taylor 26490
(F, WIS, XAL), 30 May 1986 (fl, fr), Schatz & Alverson
1161 (MEXU, U, WIS, XAL).
Although flowers were lacking, Safford (1913)
nonetheless confidently assigned Annona longipes
to his newly circumscribed section Pilannona, pre¬
sumably based on the indument, which, as he ac¬
knowledged, is erect in contrast to appressed (se¬
riceous) in Annona sericea Dunal and its allies.
Nevertheless, he remarked that A. longipes lacked
a muricate fruit surface, an essential characteristic
of other members of section Pilannona. Fries (1931:
Volume 2, Number 3
1992
Schatz
Annona section Atta
251
Figure 2. Annona longipes, flowering and fruiting
branches (Schatz & Alverson 1161).
245-246) followed Safford in retaining A. longipes
in section Pilannona, stating that "Da indessen die
Bliiten nicht bekannt sind, kann die Stellung der Art
nicht mit voller Sieherheit angegeben werden” [in¬
sofar as the flowers are unknown, the position of
the species cannot be determined with complete
certainty]-
Flowers remained unknown until 1983, when Nee
& Taylor (26490) re-collected A. longipes near
Santiago Tuxtla. In contrast to the broadly ovate
outer petals of species in section Pilannona, A.
longipes possesses outer petals typical of species in
section Atta, i.e., distinctly keeled on the inner sur¬
face or triquetrous (Safford, 1914) (Fig. 2). Within
section Atta, A. longipes is most closely related to
several species with similar indument, including A.
lorigiflora S. Watson from western Mexico, and the
cultivated A. cherimola Miller, probably native to
Ecuador. As such, A. longipes holds promise for
possible hybridization with A. cherimola in the de¬
velopment of a second lowland “custard apple” cul-
tivar; the “Atemoya,” a hybrid between A. cheri¬
mola and A. squamosa L., has already proven
successful in southern Florida. Plants originating
from seed of Schatz & Alverson 1161 are now in
cultivation in the Climatron at the Missouri Botanical
Garden. Known only from the above-cited collec¬
tions, A. longipes is apparently rare in the wild. Its
range appears to fall within the rain shadow of the
coastal Los Tuxtlas volcanic peaks, and therefore,
climatically within a somewdiat drier phase of “Selva
Alta Perennifolia,” which has now largely been con¬
verted to pasture and agriculture.
Acknowledgments. I thank M. Nee for directing
me to Annona longipes, W. Haber lor showing me
Annona pruinosa, and J. Myers for assistance with
the illustration.
Literature Cited
Fries, R. E. 1931. Revision der Arten einiger Anona-
ceen-Gattungen. II. Acta Horti Berg. 10(2): 129-
341, pi. 1-27.
Safford, W. E. 1913. Annona sericea and its allies.
Contr. U.S. Natl. Herb. 16(10): 263-276, pi. 85-
99.
-. 1914. Classification of the genus Annona with
descriptions of new and imperfectly known species.
Contr. U.S. Natl. Herb. 18: 1 XII, 1-68, pi. 1-41.
Croton ascendens (Euphorbiaceae), a New Liana from
Eastern Amazonia
Ricardo de S. Secco and Nelson A. Rosa
Museu Paraense Emilio Goeldi, Depto. de Botanica,
Cx. Postal 399, 66.040, Belem, Para, Brazil
ABSTRACT. A new species, Croton ascendens, is
described as the second liana in its genus from South
America. The new species is distinguished from Cro¬
ton pullei, the other liana in the genus described
from Surinam, by its palmately lobed leaves, bisexual
racemes in inflorescences, and slightly cucullate lin-
ear-spatulate petals of staminate flowers. Croton as¬
cendens is placed in section Croton, subsection Cro¬
ton .
Croton L. is one of the largest genera in number
of species (ca. 1,000 according to Webster & Burch,
1967) of the Euphorbiaceae, and is in urgent need
of careful revision. It was last studied comprehen¬
sively by Mueller (1866, 1873). Although a natural
group, the genus displays considerable polymor¬
phism. The majority of species occur in the West
Indies and South America, and their identification
is very difficult. Recent papers on South American
Croton (Lanjouw, 1931; Croizat, 1940, 1941, 1944;
Jablonski, 1965) have not included the identification
of the large number of herbarium specimens cur¬
rently available in Amazonian herbaria (IAN, INPA,
MG). Mueller’s treatments (1866, 1873) give some
help, hut unfortunately they are out of date.
As part of ongoing research on the Euphorbiaceae
of Amazonia, the authors are currently concentrat¬
ing on the genus Croton. Fieldwork in Paragominas,
Para, Brazil has yielded a new species of liana, which
is herein described.
Croton ascendens R. Secco & N. A. Rosa, sp.
nov. TYPE: Brazil. Para: Paragominas, Fazen¬
da Vitoria, 10 Apr. 1989, Rosa & Uhl 5212
(holotype, MG; isotypes, K, MO, U). Figure 1.
Liana monoica, pilis stellatis vestita. Folia 3-lobata ad
basin cordata. Inflorescentia bisexualis ex racemo glom-
eruloruin constans. Flores masculini numerosi; calyce 5-
lobato pilis stellatis obtecto; receptaculo piloso; petalis 5
liberis, lineari-spathulatis ad apicem cucullatis; staminibus
1 1, eorurn 10 basi connatis, uno libero centralique. Flores
feminei 1-3 in partibus basalibus medianisque racemi cum
floribus inasculinis nonnullis commixti; calyce 5-lobato;
ovario subgloboso, stellato-pubescenti, disco undulato sty-
loque 6-ramoso instructo.
Liana climbing on shrubs or trees, monoecious.
Leaves alternate, chartaceous, trilobate, the lobes
acute or acuminate, 15-20 cm long, 10 19 cm
wide, with stellate hairs on both surfaces, glands
irregularly distributed on the ahaxial side, and to¬
ward the apex on the adaxial side, base cordate,
with 5 pairs of lateral veins, veins prominulous on
the ahaxial side, petiole 5-11 cm long, cylindric,
with two flat glands at the apex, stellate hairs cov¬
ering the whole surface. Inflorescence a raceme of
glomerules, terminal, solitary or geminate, bisexual,
pistillate flowers 1-3 basal and medial, mixed with
some staminate flowers, staminate flowers in glom¬
erules along most of the rachis. Staminate flowers
numerous, pedicels slender, 2 mm long, stellate-
lepidote, calyx 5-lohate, lobes sagittate, stellate-lep-
idote without, glabrous within; receptacle pilose; pet¬
als 5, free, ca. 5 mm long, linear-spatulate, slightly
cucullate at the apex, pilose; stamens 11, 10 connate
basally, one free central, 5-5.5 mm long, with abun¬
dant long, silky hairs, disc 5-glandular. Pistillate
flowers fewer, pedicels subulate, 2.5 mm long, stel-
late-lepidote, calyx 5-lobate, slightly connate at base,
lobes acuminate, stellate-lepidote hairs outside, long
silky hairs inside at the apex of the lobes; ovary
subglobose, tricarpellate, ca. 2 mm long, 2.5 mm
wide, with stellate hairs, disc undulate, with long
and silky hairs, the styles 6 times bifid, 3-5 mm
long, with long stellate hairs, apex glabrous. Fruit
seen only in an early stage.
Distribution. Known only from mata de terra
firme (Amazon forest) and capoeira of Paragominas,
in Para, Brazil.
Vernacular name: Algodao bravo (Rosa <& Uhl
5212).
Paratype. Brazil, para: Paragominas, Fazenda Vi¬
toria. 8 Oct. 1990, Secco et al. 781 (MG).
Lianas are uncommon within Croton. Lanjouw
(1931) described Croton pullei from Surinam, in¬
dicating it as the first species of the genus with this
habit. He later (1939a) described variety glabrior,
indicating that, according to the collector Rombouts,
Novon 2: 252-254. 1992.
3mm
Volume 2, Number 3
1992
Secco & Rosa
Croton ascendens
253
Figure 1. Croton ascendens (Rosa & Uhl 5212). —A.
flowers. —B. Staminate flower, without petals and central
E. Pistillate sepals, abaxial view on left, adaxial on right.
this was a tree. Likewise, he suggested that C. pullei
must be a Iree and that the annotation made by its
original collector, Stahel, must have been a mistake,
as no species of Croton were known to be lianas.
Habit with inflorescence showing staminate and pistillate
stamen. —C. Central stamen. —D. Staminate petal. —
— F. Ovary.
Lanjouw (1939b) published a correction for the habit
of C. pullei as lollows: “page 34, line 15, instead
of ‘Scandent shrub’ read: ‘tree or shrub (scan-
dent?)’.” However, recent studies on Amazonian
254
Novon
Tablk 1 . Comparison of morphological characters of
Croton ascendens and C. pullei.
C. ascendens
C. pullei
Palmately lobed leaves
Unlobed pinnately veined
leaves
Inflorescence raceme of
glomerules bisexual;
pistillate flowers basal
and medial; staminate
flowers along most of
the rachis
Inflorescence unisexual,
racemose, the pistillate
ones a simple raceme,
the staminate ones a
raceme of glomerules
Petals of staminate flow¬
ers linear-spatulate,
slightly cucullate at
the apex
Petals of staminate flow¬
ers obovate
Croton made by the senior author confirm that C.
pullei is really a liana.
The affinities of Croton ascendens are not entirely
clear, since no species closely resembling it has been
found among the large collection of Amazonian Cro¬
ton at the herbaria of MC/, IAN, and IN PA, and
South American Croton (including Brazil) at the
herbaria of RB, R, and SP. Although a liana, C.
ascendens is quite different from C. pullei, es¬
pecially in the characters cited in Table 1.
Because the female flower of C. pullei from the
analyzed collections (Stahel 76, holotype, IJ) is in
an early stage, it is difficult to compare its morpho¬
logical characteristics with those of C. ascendens.
Croton ascendens is placed in section Croton
subsect. Croton because of the bisexual lower cy-
mules, the pentamerous, equal-lobed calyx in both
the staminate and pistillate flowers, and the pilose
receptacle of the staminate flowers.
Acknowledgments. The senior author expresses
appreciation to the Conselho Nacional de Desen-
volvimento Cientifico e Tecnologico (CNPq) for a
research grant (no. 301.252/86-ZO-FV) to study
the Euphorbiaceae; to Enrique Forero and David
Brunner, Missouri Botanical Carden (MO), for the
translation of the text from Portuguese to English;
to William Rodrigues (1NPA, Manaus) for the Latin
diagnosis; to Christopher Uhl, (EMBRAPA, Belem)
for logistical support; to Jose Maria Albuquerque,
Ana Rita Alves, and William Overal, for suggestions
on the manuscript; to the curator of the Utrecht
Herbarium (U) for loan of the type material of
Croton pullei Lanj.; to Raphael Alvarez for his il¬
lustration of the habit of the species; to the reviewers
for important comments on the manuscript; and to
Celso Moraes for assistance in preparing the manu¬
script on diskette.
Literature Cited
Croizat, L. 1940. Thirty-five new species of American
Croton. J. Arnold Arbor. 21: 76-107.
-. 1941. Preliminaries for the study of Argentine
and Uruguayan species of Croton. Darwiniana 5:
417-462.
-. 1944. Additions to the genus Croton L. in
South America. Darwiniana 6: 442-468.
Jablonski, E. 1965. Euphorbiaceae. In: B. Maguire &
Collaborators, Botany of the Guayana Highland, Part.
VI Mem. New York Bot. Gard. 12(3): 150-178.
Lanjouw, J. 1931. The Euphorbiaceae of Surinam. Am¬
sterdam.
-. 1939a. New or noteworthy Euphorbiaceae
from Suriname. Rec. Trav. Bot. Neerl. 36(2): 697-
704.
-. 1939b. Euphorbiaceae. In: A. Pulle (editor),
Flora of Suriname. Additions and corrections to vol.
2(1): 457-470.
Mueller, J. 1866. Euphorbiaceae. In: A. De Candolle
(editor), Prodromus Systematica Universalis Regni
Vegetabilis 15(2): 512-700.
-. 1873. Euphorbiaceae. In: C. F. P. Martius
& A. G. Eichler (editors), Flora Brasiliensis 11(2):
1-752.
Webster, G. L. & D. Burch. 1967. Euphorbiaceae. In:
R. E. Woodson, Jr. and Collaborators, Flora of Pan¬
ama, part 6. Ann. Missouri Bot. Gard. 54: 211 —
350.
Ozobryum ogalalense (Pottiaceae), a New Moss Genus and
Species from the American Great Plains
G. L. Smith Merrill
Division of Biology, Kansas State University, Manhattan, Kansas 66506-4901, U.S.A.
ABSTRACT. Among the bryophytes collected in
northwestern Kansas by Vernon L. Wranosky of
Colby, Kansas, during the summer of 1990 is a new
genus and species of Pottiaceae, Ozobryum ogal¬
alense. Subsequent collecting in the vicinity has
resulted in two additional localities for this moss,
one in Kansas and one in Nebraska.
Ozobryum ogalalense Merrill, gen. et sp. nov.
TYPE: U.S.A. Kansas: Decatur County, on
Ogallala Formation mortarbeds above south fork
of Sappa Creek, 25 km SW of Oberlin, 39°40'N,
100°43'W, ca. 800 m, 11 Aug. 1990, If ra-
nosky 62 (holotype, KSC; isotypes, DUKE,
MICH, MO). Figures 1, 2.
Planta insignis foliis obtusatis margine incrassatis quasi
succulentis, cellulis utroque folii paginae grosse mamil-
losis, costa debili obscura utrinque cellulis mamillosis oc¬
culta bene distincta.
Plants minute, growing in soft, compact cushions,
to 2 cm high, dark green to yellowish green at
surface of tufts, without a central strand. Leaves
erect when dry, spreading at 45° or less when moist,
concave, oblong-elliptic, rounded at the apex, 0.35
0.55 mm long, margins plane or erect. Costa ending
well below the apex, thin and obscure, with 2 guide
cells but no stereids, covered on both surfaces by
short, bulging-mamillose cells similar to those of the
lamina. Upper leaf cells bistratose (or 3-stratose) at
the margins and in patches internal to the margins,
rounded-hexagonal to subquadrate, ca. 7-9 gm,
green and obscure, bulging on both surfaces, crowned
with a broad, circular, knoblike thickening of the
outer wall, centered over the lumen. Basal cells
hyaline and subquadrate in a short area. Archegonial
buds scattered along the stem, sometimes easily de¬
tached, with a tuft of rhizoids at the base. Peri-
chaetial leaves smooth, ovate, acuminate, entire to
unevenly serrate, the costa ending in the apex. Per-
igonia not seen.
Paratypes. U.S.A. Kansas: Decatur County, type lo¬
cality, 30 Apr. 1991, Merrill 12888 ; on N-facing mor-
tarbed cliffs, “Elephant Rock,” 2.5 km S of Traer, 30
Apr. 1991, Merrill 12903 ; Rawlins County, on mor¬
tarbeds above N fork of Sappa Creek, 22 km SE of
Atwood, 11 Aug. 1990, Wranosky 69, 30 Apr. 1991,
Merrill 12901. Nebraska: Hitchcock County, on mor¬
tarbeds exposed in arroyo, 17.6 km NW of Herndon
(Kansas), 30 Apr. 1991, Merrill 12919. Voucher spec¬
imens deposited in the Reed Bryophyte Herbarium, Kan¬
sas State University Herbarium (KSC).
The genus name Ozobryum is derived from
“bryum” (moss) and from the fictional Land of Oz,
the creation of American author L. Frank Baum,
now popularly associated with the Kansas plains.
The specific epithet “ogalalense” refers to the Ogal¬
lala Formation, named for a locality in southwestern
Nebraska. The Ogallala is the principal aquifer
throughout most of the High Plains from southern
South Dakota to the Texas Panhandle.
The leaves of Ozobryum are highly distinctive
and are characterized by their oblong-elliptic shape,
the leaf cells bistratose and mamillose on both sur¬
faces, and the reduced costa obscured on both sur¬
faces by short mamillose cells. In its soft, dense
cushions and habitat beneath ledges and in crevices
on moist vertical cliff faces, Ozobryum resembles a
miniature Gymnostomum. Under the hand lens the
living plants have a peculiar frosted and succulent
appearance due to the mamillose cells and thickened
leaf margins (Fig. 2). The perichaetial leaves are
strikingly different in shape from the vegetative
leaves, the cells rhombic, unistratose, and smooth
or only slightly roughened at the extreme apex. The
female buds, which are sometimes easily detached
and have a tuft of rhizoids at the base, could function
as vegetative propagules, as improbable as this may
seem. No brood bodies have been observed so far.
At the suggestion of Richard Zander, I examined
a series of 154 specimens (BUF) from Mexico and
elsewhere representing his concept of Molendoa
sendtneriana (B. S. G.) Limpr. (see Zander, 1977).
The tropical American specimens mostly belong to
the lax, glaucous-green, linear-leaved plant that cor¬
responds to Anoectangium obtusifolium (Broth. &
Par. ex Card.) Grout, illustrated by Crum & An¬
derson (1981). Among the Mexican specimens, how¬
ever, are several that resemble Ozobryum in their
small size, short blunt leaves, and thickened leaf
margins.
Novon 2: 255-258. 1992.
256
Novon
Figure 1. Ozobryum ogalalense Merrill. —a. Habit. —b. Leaves. —c. Archegonial bud. —d. Perichaetial leaves
and archegonium. —e. Upper leaf cells at margin. —f. Marginal cells near base of leaf. —g. Cross section of leaf
in upper third. —h. Cross section of thickened margin of leaf. —i. Cross section of margin near base of leaf. Scales
j = 0.2 mm (a-d); k = 0.05 mm (e-i). (All drawings from Merrill 12888.)
Volume 2, Number 3
1992
Smith Merrill
Ozobryum ogalalense
257
Figure 2. Living plants of Ozobryum ogalalense Merrill, from the type locality (Merrill 12888); x90.
The leaf cells of Ozobryum are mamillose, and
the broad knoblike thickening of the outer cell wall
is centered over the lumen. The leaf cells of these
small Mexican plants, however, are distinctly cutic-
ular-papillose. There are always several small pa¬
pillae per cell, and they often extend over the cell
wall and onto adjoining cells. The cells at the back
of the costa are rather more distinct and elongate,
and the costa is not obscured above by bulging cells
like those of the lamina. The costa is not strongly
developed in these Mexican plants, but all the leaf
cross sections examined had a few dorsal stereids
in the costa.
The lateral female inflorescences of Ozobryum
indicate a placement in the subfamily Pleurowei-
sioideae, thus narrowing the field of potential rela¬
tionships considerably. The other familiar pleuro-
carpous genera, Anoectangium and Molendoa, have
distinctly papillose (or rarely almost smooth) leaf
cells. Brotherus’s drawing (Engler & Prantl, ed. 2)
of Pleuroweisia schliephackei Limpr. is somewhat
suggestive of Ozobryum in leaf shape and in habit.
However, examination of isotypes (H) of this rare
European species reveals that the leaves are also
papillose. Moreover, they are unistratose, and the
margins are narrowly revolute for most of their
length. The fertile branches with their convolute-
tubular perichaetial leaves are also distinctive. Zan¬
der (1977) distinguished Molendoa from Anoec¬
tangium by the tendency to develop a ventral as
well as a dorsal stereid band in the costa. Stereids
and a central strand appear to be absent in Pleu¬
roweisia , as they are in Ozobryum .
The original locality for Ozobryum in Decatur
County, Kansas, is 36 km south of the Kansas-
Nebraska state line and 114 km east of the Kansas-
Colorado line. The Rawlins County collection was
made about 8 km to the west. “Elephant Rock’ 7 is
25 km north, and the Hitchcock County, Nebraska,
station is 43 km north of the type locality.
Mortarbeds are arkosic sand and gravel deposits
that are partly or completely cemented by calcium
carbonate, and are exposed locally as eroded bench¬
es or scarps (Walters, 1 956). The name is derived
from their resemblance to mortar, as well as to their
use by early settlers in the area as natural mortar
material.
The vegetation of the area is northern grama-
buffalo grass prairie (BoutelouaBuchloe), and mixed
bluestem-grama prairie (Andropogon- Bouteloua)
258
Novon
(Kiichler, 1974). The landscape may be described
as a gently rolling, treeless plain with eroded rock
outcrops (mortarbeds) exposed in low bluffs border¬
ing the principal stream valleys, such as along Sappa
and Beaver creeks. Floodplain forest (Populus Snlix )
is confined to the valley bottoms.
Ozobryum is the second endemic species, and
the first endemic genus of mosses to be recorded
for the Great Plains. This is of considerable interest
since the region is not known for endemism of either
bryophytes or vascular plants. Aschisma kansanum
Andr. is presently known only from south-central
Kansas, growing beneath translucent, white quartz
pebbles. Ozobryum may be more common than these
few collections indicate, but its distributional range
may be limited to those areas where mortarbeds
occur at the surface. The Ogallala Formation is
composed primarily of unconsolidated sands and
gravels, and mortarbeds are discontinuous and lo¬
calized (Walters, 1956; Hodson, 1969). Although
extending over a huge area, the Ogallala is mostly
buried beneath a thick deposit of Pleistocene allu¬
vium and loess, forming a flat, almost featureless
surface with poorly developed drainage. In other
areas, the Ogallala was entirely removed by erosion
prior to the deposit of the overlying loess.
Acknowledgments. Contribution no. 91-553-J
from the Kansas Agricultural Experiment Station.
Support was provided by Konza Prairie Research
Natural Area and by NSF Grant BSR-8514327 for
Long-Term Ecological Research to Kansas State
University. I am indebted to Lewis Anderson, How¬
ard Crum, and Richard Zander for comments and
suggestions as to the identity of this moss and its
possible relationships, and to the curators of BUF
and H for the loan of specimens.
Literature Cited
Crum, H. A. & L. E. Anderson. 1981. Moss Flora of
Eastern North America. 2 vols. Columbia Univ. Press,
New York.
Hodson, W. G. 1969. Geology and ground-water re¬
sources of Decatur County, Kansas. State Geological
Survey of Kansas, Bulletin 196: 1-41.
Kiichler, A. W. 1974. The potential natural vegetation
of Kansas. Supplement to Ecology, vol. 55, no. 3.
Walters, K. L. 1956. Geology and ground-water re¬
sources of Rawlins County, Kansas. State Geological
Survey of Kansas, Bulletin 117: l-100.
Zander, R. H. 1977. The tribe Pleuroweisieae (Potti-
aceae, Musci) in Middle America. The Bryologist 80:
233-269.
Notes on Psychotria Subgenus Heteropsychotria
(Rubiaceae: Psychotrieae) in Mexico
and Northern Central America
Charlotte M. Taylor
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
David H. Lorence
National Tropical Botanical Garden, P.O. Box 340, Lawai, Kauai, Hawaii 96765, U.S.A.
ABSTRACT. Expanded descriptions, comments on
geographic ranges and infraspecific variation, and
complete synonymy are presented for Psychotria
minarum, P. galeottiana, and P. phanaerandra
(Standley & Steyerm.) Lorence, a new combination
made here that is based on Palicourea phanaeran¬
dra Standley & Steyerm. the new species Psycho¬
tria juarezana C. M. Taylor & Lorence from south¬
ern Mexico is also described.
Psychotria L. is a pantropical genus of about
1,500 species of herbs, shrubs, and small trees. It
is represented by three subgenera in the Neotropics.
All subtropical species are assignable to subgenus
Psychotria or subgenus Heteropsychotria Stey¬
erm., except P. microdon (DC.) Urban. This species
was included in subgenus Psychotria by Steyermark
(1974) and other authors, but is better placed in
subgenus Tetramerae (Hiern) E. Petit (Hamilton,
1989). Subgenus Tetramerae comprises species
found in the Palaeotropics, plus this one. It is char¬
acterized by leaves that dry pale green, pyrenes with
the abaxial (“dorsal”) surface smooth except for a
median keel, persisting marcescent stipules, and lat¬
eral branches in opposite pairs (Hamilton, 1989;
Verdcourt, 1976). In contrast, subgenus Heteropsy¬
chotria is characterized by leaves that dry dark
green to brown or gray, pyrenes with three or more
ribs on the abaxial surface, stipules that may be
persistent or deciduous but are not marcescent, and
lateral branches that are usually not opposite. Psy¬
chotria microdon is found in the Antilles and ad¬
jacent continental lowlands, and probably represents
an example of long-distance dispersal. Members of
subgenus Tetramerae characteristically have bac¬
terial nodules in their leaves (Verdcourt, 1976), a
feature not found in the neotropical members of
Psychotria (including P. microdon).
The two other subgenera found in the Neotropics
encompass at least 500 species. These have been
studied in the framework of regional floras but have
not been treated comprehensively. The species of
subgenus Psychotria have recently been treated in-
depth for Mexico and Central America (Hamilton,
1989). This is the area of highest diversity for this
subgenus, with about 60 species. Subgenus Heter¬
opsychotria includes approximately twice as many
species in this same area, and many more in South
America.
Cephaelis Sw. was studied by Steyermark (1972),
who concluded that it comprised an assemblage of
species that in many cases were more closely related
to various members of Psychotria than to other
members of Cephaelis. He therefore combined Ce¬
phaelis with Psychotria subg. Heteropsychotria,
making the appropriate combinations for the floras
of the Guayana Highlands (Steyermark, 1972) and
Venezuela (Steyermark, 1974). Combinations re¬
main to be made for many species that are not found
in Venezuela (e.g., Taylor et al., 1991).
A modern infrageneric classification of Psycho¬
tria subg. Heteropsychotria (including Cephaelis )
was also begun by Steyermark in bis South American
studies (1972), but was not extended to the Meso-
american species. Unfortunately, Steyermark's sys¬
tem did not include explicit diagnoses of his groups,
but was intended instead “to show apparent rela¬
tionships between the taxa presented.” The flora of
northern Mesoamerica includes many of the species
Steyermark classified, as well as endemic relatives
of these species, so his classification can be extended
in general to the region. However, the flora ol north¬
ern Mesoamerica is not simply a northward exten¬
sion of the South American flora, but does have its
own identity as a phytogeographic province (Takh-
tajan, 1986); the existing infrageneric classification
for subgenus Heteropsychotria is inadequate to al¬
low us to classify the species treated here.
We present here expanded descriptions, com¬
ments on geographic range and infraspecific vari-
Novon 2: 259-266. 1992.
260
Novon
ation, and complete synonymy for several poorly
known species of Psychotria, and describe and il¬
lustrate one new species, P. juarezana C. M. Taylor
& Lorence. Lists of additional specimens of P. pha-
nerandra and P. galeottiana examined are avail¬
able from the authors.
Notes on Previously Described Species
Psychotria minarum Standley & Steyerm., Publ.
Field Mus. Nat. Hist., Hot. Ser. 23: 253. 1947.
TYPE: Guatemala. El Progreso: between Cal-
era and summit of Volcan Siglo, 2,000-3,300
m, 2 Jan. 1942, ,/. A. Steyerm ark 13106
(holotype, F; isotype, US).
Psychotria lilacina Standley & Steyerm., Publ. Field
Mus. Nat. Hist., Bot. Ser. 23: 252. 1947. Syn. nov.
TYPE: Guatemala. Huehuetenango: Cerro Huitz,
between Mimanhuitz and Yulhuitz, S of Los Cuchu-
matanes, 1,500-2,600 m, 14 July 1942, J. A.
Steyermark 48564 (holotype, F).
Glabrous to puberulent shrubs to 1.5 m tall, stems
becoming terete. Lea/blades elliptic, 2-6.5 x 0.7-
2.7 cm, acute to acuminate at apex with tip to 2
cm long, acute to obtuse at base, chartaceous, pale
abaxially; secondary veins 5-6 pairs, arching, with¬
out domatia, sometimes with 1 intersecondary vein
perpendicular to costa, midrib and secondary veins
smooth adaxially, prominulous ahaxially; margins
Hat; petioles 2-6 mm long; stipules persistent, with
truncate circumferential sheath to 0.5 mm long,
with 2 lobes on each side, these narrowly triangular,
ca. 0.5 mm long, acute. Inflorescences terminal,
panicle rounded, 3-6-flowered, 1-2 cm long and
wide including corollas, erect to deflexed; peduncles
ca. 1 cm long; primary branches 1-2 pairs, axis
and primary branches simple; pedicels 0-3 mm long;
bracts narrowly triangular, acute, 0.5-3 mm long,
those subtending primary branches 1-3 mm long,
those subtending flowers 0.5-1 mm long; flowers
with hypanthium obconic, 0.8-1.2 mm long; calyx
limb green, glabrous, 1-3.5 mm long, divided par¬
tially to completely, lobes 5, triangular to ligulate,
acute; corolla funnelform, white tinged with lilac or
purple, glabrous, tube 6-8 mm long, ca. 1 mm wide
at base, ca. 2 mm wide at apex, lobes 5, triangular,
2-3 mm long, 0.5-1 mm wide, acute; stamens 5,
attached in upper part of corolla, anthers 1-1.5
mm long, included or partially exserted; style sur¬
rounded at base by annular disk ca. 0.5 mm high,
glabrous, ca. 8 mm long, with stigma ca. 0.5 mm
long. Infructescences similar to inflorescences, be¬
coming purple; fruit ellipsoid, 4.5-5 mm long, ca.
4.5 mm wide, becoming black; pyrenes 2, ca. 4 mm
long, planoconvex with ca. 5 rounded ridges on back.
Distribution and ecology. Southern Mexico to
Guatemala and adjacent El Salvador, in wet highland
forest and cloud forest at 2,000-3,300 m, fre¬
quently on windswept ridges.
Phenology. Collected in flower April-May, in
fruit August-November.
These plants are low shrubs or subshrubs that are
infrequently collected. Very few mature flowers and
fruits have been seen; whether this species is dis-
tylous cannot be determined from the material avail¬
able. The type specimens of these epithets clearly
represent the same taxon. The type collection of
Psychotria minarum has mature infructescences
and fruits, while that of P. lilacina has only im¬
mature flowers on an unexpanded inflorescence. The
name P. minarum was accompanied by a more
complete description and exemplary type specimen,
and is selected here.
Additional specimens examined. El Salvador, santa
ana: Cerro Montecristo, IT. B. Heed , 4-6 Feb. 1954
(F); Montecristo, 2,300 m, 23 May 1963, A. Molina R.
& A. R . de Molina 12673 (F). Guatemala, chiquimula:
middle slopes of Montana Norte to El Jutal, on Cerro
Brujo, SE of Concepcion de las Minas, 1,700-2,000 m,
2 Nov. 1939, J. A. Steyermark 31007 (F). EL PROGRESO:
hills N of Finca Piramonte, between Finca Piramonte and
summit of Volcan Santa Luisa, 2,400-3,333 m, 5 Feb.
1942, J. A. Steyermark 43492 (F). HUEHUETENANGO:
Cerro Canana, between Nucapuxla and Canana, Sierra
de los Cuchumatanes, 2,500-2,800 m, 18 July 1942,
J. A. Steyermark 49107 (F). zacapa: summit of Sierra
de las Minas, vicinity of Finca Planados, 2,500 m, 14
Oct. 1939, J. A. Steyermark 29961 (F). MEXICO. CHIA¬
PAS: municipio Tenejapa, barrio Banabil, paraje of Matsab,
8,800 ft., 26 Aug. 1966, D. E. Breedlove 15332 (F).
OAXACA: municipio San Miguel Chimalapa, cima del cerro
Salomon, al NO de Benito Juarez, ca. 44 km al N de San
Pedro Tapanatepec, 16°46'15"N, 94°11'45"W, 1,770
m, 11 Apr. 1986, M. Ishiki 1510 (MO).
Psychotria phanaerandra (Standley & Stey¬
erm.) Lorence, comb. nov. Basionym: Palicou-
rea phanaerandra Standley & Steyerm., Publ.
Field Mus. Nat. Hist., Bot. Ser. 23: 252. 1947.
TYPE: Guatemala. Izabal: Cerro San Gil, 1,200
m, 26-27 Dec. 1 941, J. A. Steyermark 41952
(holotype, F; isotype, US). Lorence & Dwyer
(1987: fig. 3a, b).
Psychotria luteotuba Lorence, Bol. Soc. Bot. Mexico 47:
55. 1 987. Syn. nov. TYPE: Mexico. Veracruz: mun-
icipio Hidalgotitlan, zona de Uxpanapa, Rio Solu-
suchil a 5-6 km ESE de Hermanos Cedillo, 100-
200 m, 24 Mar. 1982, D. Lorence et al. 3910
(holotype, MEXU; isotypes, BM, BR, CHAPA,
ENCB, F, MO, UC, XAL).
Glabrous or minutely hirtellous shrubs or small
trees to 4 m tall, stems becoming terete, /.cq/blades
elliptic to ovate-elliptic, 4-10(-12.5) x (1.8—)2.4—
Volume 2, Number 3
1992
Taylor & Lorence
Psychotria subgenus Heteropsychotria
261
4.7 cm, acuminate at apex with tip to 2 cm long,
acute to cuneate or attenuate or (rarely) obtuse at
base, chartaceous; secondary veins 5-10 pairs,
arching, without domatia, with l(-2) intersecondary
veins perpendicular to costa, midrib and secondary
veins prominulous on both surfaces; margins slightly
revolute; petioles 1.5-10 mm long; stipules persis¬
tent, with truncate circumferential sheath 0.4-0.8
mm long, with 2 lobes on each side, these narrowly
triangular to acicular, 0.3-0.6 mm long, acute,
deciduous. Inflorescences terminal, panicle rounded,
15-22-flowered. 25-60 mm long including corollas,
15-35 mm wide, green, erect; peduncle 15 35 mm
long; primary branches 2-4 pairs, axis and primary
branches simple or branched once; pedicels 0-1.5
mm long; bracts triangular to narrowly so, 0.5-3
mm long, those subtending primary branches 1-3
mm long, those subtending flowers 0.5-1 mm long;
flowers distylous, with hvpanthium obconic, ca. 0.5
mm long; calyx limb green, glabrous, 0.5-0.8 mm
long, divided partially to completely, lobes 5, tri¬
angular, 0.4-0.6 mm long, ca. 0.2-0.5 mm wide,
subequal, acute to obtuse; corollas funnelform, yel¬
low, glabrous or minutely hirtellous externally, in¬
ternally glabrous except for a ring 1 -2 mm wide of
sparse pilosulous pubescence just below filament at¬
tachment, tube 8-13 mm long, ca. 1 mm wide at
base, ca. 2-2.5 mm wide at throat, lobes (4 )5,
triangular to somewhat ovate or ligulate, 3-4 mm
long, acute to rounded; stamens (4—)5, in long-styled
form attached in upper third of corolla tube with
filaments 0.5-1 mm long and anthers 2-3 mm long
and included, in short-styled form attached at apex
of tube with filaments 13 mm long and anthers 2-
3 mm long and exserted; style surrounded at base
by annular disk 0.5-0.8 mm high, glabrous, in long-
styled form extending 2-3 mm beyond apex of co¬
rolla tube with stigmas ca. 0.5 mm long, in short-
styled form 6.5-8 mm long with stigmas ca. I mm
long. Infructescences similar to inflorescences, be¬
coming purple; fruit ovoid, laterally compressed,
somewhat didymous, 5-7 mm long, 5-6.5 mm wide,
becoming black; pyrenes 2, 4-5 mm long, plano¬
convex with 3-5 rounded ridges on back.
Distribution and. ecology. Southern Mexico to
Panama, though markedly less frequently collected
in the southern part of its range, in wet lowland
evergreen forests and montane wet or cloud forests
at 20-2,100 m.
Phenology. Collected in flower March-June and
in December, in fruit July December and in Feb¬
ruary.
The general appearance and biology of this spe¬
cies are described by Lorence & Dwyer (198 .). 1 be
type collection of Palicourea phanaerandra has
only immature flowers, but clearly represents the
same species. The identity of this name was unknown
for a long time; Standley & Williams did not include
it in the Flora of Guatemala (1975).
Selected specimens examined. COSTA RlCA. CARTAGO:
Valle Escondido, 750 m, 1 Apr. 1966, C. Schnell 807
(F). HEREDIA: Parque Nacional Braulio Carrillo, 600 m,
24 Mar. 1982, /. .4. Chacon 55 (CR); Estacion Carrillo
de la fila al canon del Rio Sucio, 450-700 m, 12 Nov.
1983, I. A. Chacon & G. Herrera 1670 (CR). LIMON:
Canton de Limon, El Progreso 0.5 km al E de Cerro
Muchilla, fila Matama, Valle de La Estrella, 9°47'50"N,
83°05'30"W, 850 m, 9 Apr. 1989, G. Herrera 2573
(CR, MO), 5 Apr. 1989, R. Robles & A. Chacon 2658
(CR, MO). Honduras, comayagua: ca. 10 km W of
Siguatepeque, 1,800 m, 18 May 1972, D. Burch 6106
(MO). CORTES: along Lake Yojoa ca. 5 km N of Rancho
Agua Azul, 630 m, 11 Apr. 1951, L. O. Williams & A.
Molina R. 17762 (F). MEXICO. CHIAPAS: municipio Ray¬
on, near Puerto del Viento, 9 mi. N of Pueblo Nuevo
Solistahuacan, along road to Tapiula, 6,100 ft., 30 May
1965, D. E. Breedlove 10173 (MO), guerrf.ro: Montes
de Oca, Pilas-Pasion, 2,100 m, 10-4-1937, G. Hinton
10757 (F). OAXACA: por la vereda a arroyo Plata, I6°52'N,
94°37'W, 400 m, 3 Sep. 1985, //. Hernandez G. 1439
(CHAPA, PTBG). TABASCO: municipio Huimanguillo, km
10, Ejido T. Nueva, 20 m, 4 Apr. 1972, Puig 777
(MEXU). VERACRUZ: municipio Catemaco, entre de Bas-
tonal y Arroyo Claro, 14 km al E de Lago Catemaco,
900 m, 10 June 1972, J. H. Beaman 6124 (F, MEXU,
MO). Panama, cocle: on ridge W of sawmill above El
Cope, 903-1,161 m, 21 June 1978, B. Hammel 3558
(MO).
Psychotria galeottiana (M. Martens) C. M. Tay¬
lor & Lorence, Faxon 34: 669. 1985. Pali¬
courea galeottiana M. Martens, Bull. Acad.
Roy. Soc. Sci. Bruxelles 11: 136. 1844. I r-
agoga galeottiana (M. Martens) Kuntze, Re-
vis. Gen. PL 2: 960. 1891. TYPE: Mexico.
Oaxaca: without location, Galeotti 2606 (lec-
totype, designated by Taylor & Lorence, 1985,
BR; isolectotypes, K, P). Figure 1.
Palicourea seleri Loes., Verh. Bot. Vereins Brandenburg
65: 1 14. 1923. Syn. nov. SYNTYPES: Guatemala.
Huehuetenango: zwischen Trinidad und Rosario, 13
Aug. 1896, C. Seler & E. Seler 3049 (syntype, B
destroyed, photos, F, GH, MO, NY, US, fragment,
F; isosyntype, GH); Yalambohoch, C. Seler & E.
Seler 2676 (syntype, B destroyed).
Psychotria skutchii Standley, Publ. Field Mus. Nat. Hist.,
Bot. Ser. 17: 283. 1937. Syn. nov. TYPE: Gua¬
temala. Huehuetenango: San Juan Ixcoy, 8,000 ft.
[2,581 m], 22 Aug. 1934, A. Skutch 1074 (holo-
type, F; isotype, A).
Psychotria pachecoana Standley & Steyerm., Publ. Field
Mus. Nat. Hist., Bot. Ser. 22: 205. 1940. Syn. nov.
TYPE: Guatemala. Baja Verapaz: N of divide N of
Santa Rosa, 1,650 m, 30 Mar. 1939, P. C. Standley
6 9922 (holotype, F).
Psychotria persearum Standley, Ceiba 1: 48. 1950. Syn.
262
Novon
nov. TYPE: Honduras. Francisco Morazan: Cerro
de llyuca, 1,800 m, 2 Mar. 1947, P. C. Standley
4865 (holotype, F; isotype, EAP).
Psychotria uyucana Standley, Ceiba 1: 48. 1950. Syn.
nov. TYPE: Honduras. Francisco Morazan: drainage
of Rio Yeguare, Mt. Uyuca, 14°N, 87°W, 2,000
in, 29 May 1948, A. Molina R. 919 (holotype, F;
isotype, EAP).
Psychotria orogenes L. 0. Williams, Phytologia 28: 231.
1974. Syn. nov. TYPE: Guatemala. Baja Verapaz:
Sierra de las Minas, ca. 5 km S of Purulha, 1,600
m, 2 Jan. 1973, L. O. Williams et al. 41924
(holotype, F).
Palicourea rnacrantha auct., non Loes. (Taylor, 1989).
Glabrous or puberulent to densely pilosulous shrubs
or small trees to 5 m tall with trichomes to 0.3 mm
long and sometimes yellow or yellow-brown, stems
becoming terete, Leaf blades elliptic to narrowly so,
4-15 x 1.2-4 cm, acute to acuminate at apex with
tip 1-2 cm long, acute or cuneate to rounded at
base, chartaceous, frequently pale abaxially, without
domatia; secondary veins 7 18 pairs, arching, with
1- 3 intersecondary veins perpendicular to costa,
midrib and secondary veins smooth adaxially, prom-
inulous below; margins flat; petioles 2-17 mm long;
stipules persistent, with truncate circumferential
sheath 1 -2(-4) mm long, with 2 lobes on each side,
these narrowly triangular, 0.5-4 mm long, acute.
Inflorescences terminal, panicles pyramidal to
rounded, 10-30-flowered, 1.5 9.5 cm long includ¬
ing corollas, 2-1 1 cm wide, green, erect to deflexed;
peduncles 1-4 cm long; primary branches 1-3 pairs,
axis and primary branches simple or branched once
or twice; pedicels 0-5 mm long; bracts narrowly
triangular to linear, 0.5-14 mm long, those sub¬
tending primary branches 1-14 mm long, those
subtending pedicels 0.5-3 mm long; flowers disty-
lous, with hypanthium obconic, ca. 0.8-1 mm long;
calyx limb green, puberulent to densely pilosulous
with trichomes to 0.3 mm long, 0.3-1.5 mm long,
divided nearly completely, lobes 5, triangular, often
strongly unequal, acute to obtuse; corolla funnel-
form, white sometimes flushed with pink or lavender,
externally puberulent to pilosulous with trichomes
to 0.2 mm long, internally glabrous except for a
ring 2-4.5 mm wide of sparse pubescence near
middle of tube, tube 5-10 mm long, ca. 1 mm wide
at base, 3-4 mm wide at apex, lobes 5, triangular,
2- 3 mm long, 1-2 mm wide, acute; stamens 5, in
long-styled form attached in upper third of corolla
tube with anthers subsessile, 1.8-2 mm long, and
partially exserted, in short-styled form attached at
apex of tube with filaments 1 -2 mm long and anthers
2-2.5 mm long and partially to wholly exserted;
style surrounded at base by annular disk ca. 0.5
mm high, glabrous, in long-styled form extending
12 mm beyond apex of the corolla tube with stig¬
mas 0.3-1 mm long, in short-styled form 4-7 mm
long with stigmas 1.5-2 mm long. Infructescences
similar to inflorescences, becoming purple; fruit el¬
lipsoid, somewhat compressed laterally, 3-4 mm
long and wide, becoming black; pyrenes 2, 3 3.5
mm long, with ca. 5 rounded ridges on back.
Distribution and ecology. Southern Mexico to
Nicaragua, common in Mexico, Guatemala, and
Honduras but markedly less frequently collected in
the southern portion of its range, in wet forest and
cloud forest at 1,400-3,050 m.
Phenology. Gollected in flower and fruit through¬
out the year, but more frequently collected in flower
April-June and in fruit August-December.
As circumscribed here, this species varies widely
in pubescence. Plants range from completely gla¬
brous to densely pilosulous on all parts, with the
pubescence usually golden and spreading. All inter¬
mediate pubescence conditions are represented by
specimens, including types. The most densely pu¬
bescent forms are found from Chiapas to Honduras,
but glabrous individuals occur in the same range.
Both glabrous and densely pubescent individuals are
represented from the same sites, and even in the
same collection (Croat & Hannon 64040, MO). No
other characteristics show correlation with the vari¬
ation in pubescence, and pubescence is here rejected
as a species-level characteristic. The epithets “ gal -
eottiana “ persearum ,” and “ uyucana ” were
based on glabrous specimens, “ skutchii ” and “se-
leri ” on moderately pubescent specimens with the
inflorescences and leaves smaller than average, and
“ orogenes ” and “ pachecoana ” on densely pubes¬
cent specimens with leaves larger than average.
The reproductive features, notably the inflores¬
cence and corollas, are variable as well, but they
likewise show no correlation with variation in other
characters. The inflorescences of Psychotria gal-
eottiana vary continuously from relatively open with
about 15 flowers to relatively contracted with about
50 flowers, throughout the range of the species. The
types of several of the epithets listed above, notably
“uyucana ,” show these relatively more contracted
inflorescences, but cannot be segregated on this ba¬
sis. The corolla is usually pentamerous, but some
individual flowers may occasionally be tetramerous
in the same inflorescence (Fig. 1).
Selected specimens examined. GUATEMALA. ALTA
VERAPAZ: 2 3 mi. S of Purulha on Hwy. 14 to Coban,
1,500-1,600 m, 21 July 1977, T. B. Croat 41746 (MO).
BAJA verapaz: Chilasco, 4 Aug. 1971, E. Contreras 10967
(F, MO). CHIMALTENANGO: Santa Elena, 9,000-10,000
Volume 2, Number 3
1992
Taylor & Lorence
Psychotria subgenus Heteropsychotria
263
Figure 1. Psychotria galeottiana (M. Martens) C. M. Taylor & Lorence. —A. Habit. —B. Detail of inflorescence,
showing tetra- and pentamerous corollas of short-styled flowers. Photos taken in the Sierra de Juarez of Oaxaca,
Mexico ( Lorence 4196).
ft., 18 July 1933, A. Skutch 443 (F, US), el quiche:
ca. 2 km E of Nebaj, E. Contreras 4921 (MO). SAN
MARCOS: between Todos Santos and Finca El Porvenir,
lower to middle slopes of Volcan Tajumulco, 1,300-3,000
m, 1 Mar. 1940, J. A. Steyermark 37021 (F). zacapa:
ravine bordering Quebrada Alejandrina, summit of Sierra
de las Minas, vicinity of Finca Alejandrina, 2,500 m, 13
Oct. 1939, J. A. Steyermark 29821 (F). HONDURAS. EL
PARAISO: Cerro Monserrat, cerca de Yuscaron, 2,000 m,
6-9 Oct. 1977, C. Nelson & E. Romero 4326 (MO).
Francisco morazan: Cerro de Uyuca, 6,300 ft., 10 June
1948, S. F. Classman 1532 (EAP, F, NY), 16 June
1948, S. E. Classman 1628 (EAP, F, NY), 17 July
1948, S. E. Classman 1945 (EAP, F, NY), J. W. Miller,
1950 (EAI*). OCOTEPEQUE: Mt. Merendon, 10 Sep. 1973,
D. Hazlett 806 (MO). Mexico, chiapas: municipio Jitotol,
ca. 12 km N of Jitotol along a side road to an oil well,
2,000 m, 28 Sep. 1971, D. E. Breedlove 19954 (MO),
28 Oct. 1971, D. E. Breedlove & R. E. Thorne 21495
(F, MO, NY). GUERRERO: distrito Montes de Oca, Pilas-
Pasion, 2,100 m, 10 Apr. 1937, G. Hinton et al. 10757
(MO). HIDALGO: municipio Tenango de Doria, arroyo a 5
km por camino al E de Tenango de Doria a El Cirio,
1,700 m, 9 Nov. 1985, D. //. Lorence & R. Hernandez
264
Novon
Figure 2. Psychotria juarezana C. M. Taylor & Lorence. —A. Habit. —B. Detail of inflorescence showing tetra-
and pentamerous corollas of long-styled flowers. Photos taken in the Sierra de Juarez of Oaxaca, Mexico (Lorence
4266).
M. 4921 (MEXU, MO). OAXACA: distrito Ixtlan de Juarez,
Sierra de Juarez, ruta 175 Tuxtepec a Oaxaca, 14 km
por camino al NE de Cerro Humo Chico y Cerro Pelon,
2,200 m, 28 May 1990, D. //. Lorence & H. Cedilla
T. 4175 (MO). PUEBLA: Cascada Oligui entre Teziutlan y
Tlapacovan, 1,550 in, 2 June 1968, 11. Garcia S. 73
(MO). VERACRUZ: municipio Banderilla, Rancho La Mar-
tinica, 5 km al N del pueblo de Banderilla, 19°35'N,
96°57'W, 1,500 m, 6 May 1978, J. 1. Calzada 4387
(F). Nicaragua. JINOTECA: Cerro Kilambe, falda E del
Pico Piedras Pelona, 13°34'N, 85°50'W, 1,300-1,400
m, 28 Mar. 1981, I’. P. Moreno 7775 (MO), matagalpa:
W slope and summit of Cerro El Picacho, ca. 13°00'N,
85°55'W, 1,350-1,590 m, 3 June 1983, W. 1). Stevens
& P. P. Moreno 22179 (MO). NUEVA SEGOVIA: faldas al
5 de Cerro Mogoton, 1,600 m, 12 June 1975, J. Atwood
6 I). Neill 21 (MO). ZELAYA: Cerro Saslaya, 20 km W
of Siuna, 1,100-1,400 m, 5 May 1977, 11. Neill 1845
(MO), 1,400-1,600 m, 4 May 1978, D. Neill 3855
(MO).
Description of a New Species
Psychotria juarezana C. M. Taylor & Lorence,
sp. nov. TYPE: Mexico. Oaxaca: distrito Ixtlan
Volume 2, Number 3
1992
Taylor & Lorence
Psychotria subgenus Heteropsychotria
265
de Juarez, Sierra de Juarez, ruta 175, Tuxtepec
a Oaxaca ca. 4 km al SO de La Esperanza,
1,700 m, 4 Apr. 1983, D. 11. Lorence 1266
(holotype, MO #3750634; isotype, MEXU).
Figure 2.
Species Psychotriae galeottianae (M. Martens) C. M.
Taylor & Lorence affinis, sed infiorescentia minus deflexa,
limbo calycino 1-2.5 mm longo, bracteis majoribus 2-
10 mm longis differt.
Glabrous shrubs or small trees to 2 m tall, stems
terete. LeaJ blades elliptic to slightly oblanceolate,
4-11.5 cm long, 1.4-3.8 cm wide, acuminate at
apex with tip 1-1.5 mm long, acute at base, char-
taceous, pale abaxially; secondary veins 5-10 pairs,
arching, without domatia, with l(-2) intersecondary
veins perpendicular to costa, midrib and secondary
veins prominulous on both surfaces; margins Hat;
petioles 2-22 mm long; stipules persistent, with
truncate circumferential sheath 1-1.5 mm long,
with two lobes on each side, these narrowly trian¬
gular, 1.8-4 mm long, acute. Inflorescences ter¬
minal, panicles pyramidal, 5-15-flowered, 8 25 mm
long including corollas, 8-20 mm wide, green, nod¬
ding; peduncle 15-27 mm long; primary branches
1-2 pairs, axis and primary branches simple or
rarely branched once, each terminating in a cymule
of 2-5 flowers; pedicels 0-22 mm long; bracts tri¬
angular, acute, 2-10 mm long, those subtending
primary branches 7-10 mm long and those sub¬
tending flowers 2-7 mm long; flowers distylous;
hypanthium obconic, ca. 0.8-1 mm long; calyx limb
green, glabrous, 1-2.5 mm long, divided partially
to completely to base, lobes 5, ligulate, 0.5-1 mm
wide, frequently strongly unequal, acute; corolla
funnelform, white or sometimes flushed with purple
near base, glabrous except for a ring 2-3 mm wide
of pilosulous pubescence internally at middle of tube,
tube 7-10 mm long, ca. 1 mm wide at base, ca. 4
mm wide at apex, lobes 5, triangular, 3—3.5 mm
long, acute; stamens 5, in long-styled form attached
in upper third of corolla tube with anthers sessile,
1.8-2 mm long, and partially exserted, in short-
styled form attached at apex of tube with filaments
ca. 2 mm long and anthers ca. 2 mm long and
partially to wholly exserted; style surrounded at base
by annular disk ca. 0.8 mm high, glabrous, in long-
styled form extending ca. 3 mm beyond apex of
corolla tube with stigmas ca. 0.3 mm long, in short-
styled form ca. 4-5 mm long with stigmas ca. 1
mm long. Infructescences similar to inflorescences,
becoming purple; fruit ellipsoid, ca. 5 mm long, ca.
4.5 mm wide, becoming black; pyrenes 2, ca. 4 mm
long, planoconvex with ca. 5 ridges on back.
Distribution and ecology. Southern Mexico, in
wet montane cloud forests with Pin us, Engelhard-
tia. Magnolia, Ardisia, Podocarpus, Quercus, and
ILeinmannia at 1,550-3,000 m elevation.
Phenology. Collected in flower in April, June,
and August, in fruit September October.
This new species is similar to Psychotria gal-
eottiana and is probably closely related to it. The
latter species can be separated by its erect or slightly
deflexed pyramidal inflorescences 1.5-9.5 cm long
with pedicels 0-5 mm long and linear floral bracts
0.5-3 mm long, calyx lobes 1.5 mm long or shorter
though often strongly unequal, and less strongly
exserted style and stamens. The specific epithet re¬
fers to the type locality.
Paratypes. Mexico, chiapas: municipio Jitotol, ca. 12
km N of Jitotol along a side road to an oil well, 2,000
m, 28 Sep. 1971, D. E. Breedlove 19954 (MO), 28 Oct.
1971, D. E. Breedlove & R. F. Thorne 21495 (MO).
OAXACA: distrito Ixtlan de Juarez, La Esperanza, Km 75
de la carretera Valle Nacional a Ixtlan, 3 Aug. 1981, R.
Cedilla T. et al. 923 (MEXU, MO); 4 km al SW de la
Esperanza, 1,800 m, 4 June 1983, R. Cedillo 7. R.
Torres C. 2412 (MEXU, MO); municipio de Comaltepec,
carretera Valle Nacional-Oaxaca, al NE de Cerro Pelon,
3,000 m, Mar. 1983, L. Cortes 138 (MEXU, MO); Sierra
de Juarez, along Hwy. 175 between Valle Nacional and
Oaxaca, 26 mi. above (W of) Valle Nacional, 1,900 m,
30 June 1977, T. B. Croat 39846 (MO), 39850 (MO);
distrito Ixtlan de Juarez, Sierra de Juarez, ruta 175 Tux¬
tepec a Oaxaca, 3 km al SO de la Esperanza, 1,550 m,
17 Apr. 1982, D. //. Lorence et al. 4051 (MEXU, MO);
distrito Ixtlan de Juarez, Sierra de Juarez, Ruta 175
Tuxtepec a Oaxaca, ca. 4 km al SW de La Esperanza,
1,700 m, 4 Apr. 1983, D. H. Lorence 4267 (MEXU,
MO).
Acknowledgments. We thank the curators of
EAP, F, MEXU, and NY for making specimens
available. We also thank J. D. Dwyer and W. C.
Burger for helpful discussions and review of this
manuscript, and the Dee Scholarship Fund of the
Field Museum of Natural History for support for
travel.
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Lorence, D. H. & J. D. Dwyer. 1987. New taxa in
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Standley, P. C. & L. O. Williams. 1975. Flora of
Guatemala — Part IX. (Rubiaceae). Fieldiana, Bot.
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Steyerrnark, J. A. 1972. Psychotria. In: B. M. Maguire
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(editor). Flora de Venezuela 9(3): 1111-1683. In¬
stitute Botanico, Direccion de Recursos Naturales
Renovables, Ministerio de Agricultura y Cria. Ca¬
racas, Venezuela.
Takhtajan, A. 1986. Floristic Regions of the World.
Univ. California Press, Berkeley.
Taylor, C. M. 1989. Revision of Palicoureu (Rubiaceae)
in Mexico and Central America. Syst. Bot. Monogr.
26: 1-102.
Taylor, C. M., B. E. Hammel & W. C. Burger. 1991.
New species, combinations, and records in Rubiaceae
from the La Selva Biological Station, Costa Rica.
Selbyana 12: 131-140.
Verdcourt, B. 1976. Rubiaceae (Part 1). In: R. M.
Polhill (editor), Flora of Tropical East Africa. Crown
Agents for Overseas Governments and Administra¬
tions, London.
Hiemobotrychium , a New Section of Botrychium Subgenus
Sceptridium from the Southeastern United States
It . //. Wagn er, Jr.
Department of Biology and Herbarium, University of Michigan,
Ann Arbor, Michigan 48109, U.S.A.
ABSTRACT. Ecological, morphological, and ana¬
tomical characters support the recognition of a sec¬
tion, designated here as Botrychium sect. Hiemo-
botrychium, distinct in the characters described from
section Sceptridium. It is monotypic and based on
B. lunarioides (Michaux) Sw.
The evergreen grapeferns, Botrychium subg.
Sceptridium , are represented in the New World hy
14 species, half of them found north of Mexico. The
most peculiar member of the subgenus is the winter
or prostrate grapefern, B. lunarioides, a very rare
and local species of the southeastern United States
(Wagner, 1961). Distinct in many respects from its
relatives, it is placed in a monotypic section.
Botrychium sect. Hiemobotrychium W. II.
Wagner, sect. nov. TYPE: Botrypus luna¬
rioides Michaux, Botrychium lunarioides (Mi¬
chaux) Sw.
Folium in serum autumnum et hiemem usque ad ma-
turitatem crescens. Radices 20-30 per plantain, luteo-
brunneae. Stipes communis cellulis giganteis internis sim-
ilibus tracheidarum instructus. Trophophora brevistipitata
prostrataque, plerumque 2, deltata, 2-3-pinnata, herba-
cea. Venatio pinnularum flabellata, dichotoma, sine costa.
Primordia foliorum pilis sparsis munita. Axes sporopho-
rorum late complanati, succulenti, usque ad 1.4 mm lati.
Leaf growing and maturing during late fall and
winter. Roots 20-30, yellow-brown. Common stalk
with tracheoidal idioblasts. Trophophores short -
stalked and prostrate, usually 2 per plant, debate,
2-3-pinnate, herbaceous. Pinnule venation flabel-
late, dichotomous, lacking midrib. Leaf primordia
with scattered hairs. Sporophore axes broadly flat¬
tened, fleshy, to 1.4 mm wide.
Botrychium lunarioides differs from typical
members of section Sceptridium in the following
characteristics: its curious seasonality, the plant
wholly underground and dormant for 8 to 9 months
of the year, including most of spring, summer, and
fall, rather than being essentially evergreen and
visible at all seasons as in section Sceptridium; in
the numerous, yellow-brown narrow roots rather
than few, blackish thick roots; the presence of cu¬
rious giant tracheidlike cells in the major axes (Ar-
nott, 1960, figs. 1-4) that are lacking in Sceptri¬
dium; the prostrate, short-stalked trophophores,
usually 2 per plant rather than upright, long-stalked
trophophores, 1 per plant; the flahellate-dichoto-
mous pinnule venation rather than pinnate pinnule
venation; the primordia with few, scattered hairs
rather than with numerous, densely matted hairs;
and the flattened, very fleshy axes of the sporophore
rather than nearly terete, only moderately fleshy
axes.
Recent field investigations by Thomas (1978,
1979) have not only produced more localities, but
extended the known range considerably. The species
is now known from the Coastal Plain and outer
Piedmont from the Carolinas to Arkansas and east¬
ern Texas. It is rarely collected because of its un¬
usual seasonality, growing at a time when few bot¬
anists are in the field; its flattened fronds being
commonly partially covered by adjacent vegetation;
and its occurrence in unexpected places, such as
weedy roadsides and cemeteries.
W ere it not for the somewhat intermediate spe¬
cies, Botrychium jenmanii L. Underw., section Hie¬
mobotrychium would be a candidate for recognition
as a separate subgenus of Botrychium. Technically
speaking, B. jenmanii (Alabama grapefern), an of¬
ten associated species, conforms with the major fea¬
tures of section Sceptridium, even though in a num¬
ber of respects its characters suggest an affinity to
B. lunarioides. The hypothesis has been proposed
that B. jenmanii is the ancient hybrid of B. biter-
natum (Savigny) Underw. and B. lunarioides
(Wagner, 1968: 118, 120, fig. 2). Botrychium
jenmanii commonly occurs side-by-side with B. bi-
ternatum and/or with B. lunarioides. In northward
extent, in eastern North America, B. biternatum
reaches as far as Maryland and eastern Pennsyl¬
vania, B. jenmanii as far as southwestern Virginia,
but B. lunarioides reaches only as far as South
Carolina. Both of the putative parents are diploids
with n = 45; B. jenmanii, on the other hand, with
n = 90, is the only New World polyploid in subgenus
Sceptridium.
Novon 2: 267-268. 1992.
268
Novon
Acknowledgment. I thank W. H. Anderson lor
the Latin diagnosis.
Literature Cited
Arnott, H. J. 1960. Tracheoidal idioblasts in Botry-
chium. Trans. Amer. Microscop. Soc. 79: 97-103.
Thomas, R. D. 1978. Three additions to the Ophiog-
lossaceae of Arkansas. Bull. Torrey Bot. Club 105:
234-235.
-. 1979. First records of Botrychium tuna-
rioides and Ophioglossum nudicaule var. tenerum
(Ophioglossaceae) from Texas. Southw. Naturalist 24:
395-396.
Wagner, W. H., Jr. 1961. Nomenclature and typifi-
cation of two botrychiums of the southeastern United
States. Taxon 10: 165-169.
-. 1968. Hybridization, taxonomy, and evolu¬
tion. Pp. 113-138 in V. H. Heywood (editor), Mod¬
ern Methods in Plant Taxonomy. Academic Press,
London.
Realignments in American Croton (Euphorbiaceae)
Grady L. Webster
Department of Botany, University of California, Davis, California 95616, U.S.A.
ABSTRACT. Revisional studies in Croton to achieve
a more nearly phylogenetic classification necessitate
a number of nomenclatural changes. The genera
Crotonopsis and Eremocarpus are reduced to sec¬
tions of Croton, and the new names Croton mi-
chauxii and C. willdenowii are proposed for the
two species of Croton sect. Crotonopsis. For Croton
sect. Julocroton, the 25 species accepted are enu¬
merated, including 19 new names or new combi¬
nations.
In the course of an unpublished review of the
sections of Croton L., I have found it necessary to
make a number of changes in classification. Con¬
sequently, in order to be able to cite representative
species for the 38 sections of Croton, a number of
nomenclatural changes must be validated.
The generic circumscription of Croton that has
been almost universally accepted by systematists for
more than a century is that proposed by Jean Muell¬
er (1865, 1866, 1873). In contrast to the earlier
classification of Baillon (1858), Mueller recognized
only four genera in the Croton alliance (subtribe
Crotoneae Muell. Arg.): Croton L., Julocroton Mart.,
Crotonopsis Michaux, and Eremocarpus Bentham.
These four taxa have in common (with rare excep¬
tions in individual species) the diagnostic characters
of stellate or lepidote indumentum, thyrsoid inflo¬
rescence with pistillate flowers at base, stamens in-
flexed in the bud, and carunculate seeds.
Although Bentham (1880) noted that Julocroton
was excessively close to Croton, he accepted it hes¬
itatingly. A more definite rejection of the Muellerian
circumscription was made by Macbride (1951), who
refused to recognize Julocroton as a distinct genus
and incorporated the Peruvian species into Croton.
Mueller (1866) had distinguished Julocroton from
Croton by a supposed difference in the relative po¬
sition of the sepals to the subtending bract, although
in subsequent taxonomic treatments (as by Mac-
bride), the highly asymmetrical and laciniate pistil¬
late calyx of Julocroton was used as the distinguish¬
ing character. In agreement with Bentham (1880),
who was one of the first to question the generic
distinctness of Julocroton, I find the topology of
sepal position versus bract an unusable character;
and I concur with Macbride that the irregular la¬
ciniate calyx does not adequately distinguish Julo¬
croton from various South American species of Cro¬
ton.
In a review of the Euphorbiaceae in the south¬
eastern United States (Webster, 1967), 1 reduced
Julocroton to a section of Croton, but rather diffi¬
dently accepted Crotonopsis as a distinct genus.
However, in reviewing all of the sections of Croton,
it has become apparent to me that Crotonopsis is
excessively close to Croton sect. Gynamblosis (Tor-
rey) A. Gray. In particular, the main generic char¬
acter of Crotonopsis —unicarpellate gynoecium and
indehiscent fruit—represents merely the end-point
in a reduction series from the 3-carpellate gynoe¬
cium of most Croton species through the 2-carpel-
late gynoecium of Croton monanthogynus Michaux
to the 1-carpellate gynoecium of Crotonopsis. Con¬
sequently, there appears to be a much stronger case
for treating Crotonopsis as a section of Croton than
as an independent genus.
Although I did not consider the problem of Er¬
emocarpus earlier, its situation is comparable to that
of Crotonopsis. When Hooker (1838) described
Croton setigerus Hook., he expressed doubt as to
its position in his citation of the name as Croton ?
setigerus. Following the creation of the monotypic
genus Eremocarpus to accommodate this species
(Bentham, 1844), it has been almost universally
accepted (except by Greene (1891), who is hardly
renowned as a “lumper”)- However, Eremocarpus
setigerus (Hook.) Benth. has a habit reminiscent of
various herbaceous species of North American Cro¬
ton, such as C. capitatus Michaux of section Pi-
linophyton (Klotzsch) A. Gray and C. lindheimer-
ianus Scheele of section Gynamblosis (Torrey) A.
Gray. The obsolete pistillate perianth was used by
Mueller (1866) as the main generic character for
Eremocarpus, but the staminate flowers are typical
for Croton; and the 1 -carpellate gynoecium appar¬
ently represents a reduction parallel to that discussed
above for Crotonopsis. Thus, although I earlier ac¬
cepted both Crotonopsis and Eremocarpus as dis¬
tinct from Croton (Webster, 1975), their claim to
generic separation now appears no better founded
than that for Julocroton.
For all three of these satellite genera of Croton,
a serious argument against their acceptance at the
Novon 2: 269 273. 1992.
270
Novon
generic rank is that it would lead to a blatantly
nonphylogenetic classification; or, to use cladistic
parlance, it would make Croton a thrice-paraphy-
letic genus. The only way to salvage such genera
as Julocroton would be to split Croton into a large
number ol segregate genera, as Klotzsch (1841)
attempted. Sucb an alternative, l believe, would be
highly unacceptable because of problems in defining
and recognizing the segregates, and because it would
obscure the phyletic coherence ol this great genus
of nearly 1,000 species. Consequently, the three
segregates are here treated as sections of Croton,
and the necessary nomenclatural adjustments are
proposed.
Croton L., Sp. PI. 1004. 1753. Lectotype: Croton
aromaticus L. (designated by Webster, 1967).
Section Crotonopsis
Croton sect. Crotonopsis (Michaux) Webster, stat.
nov. Basionym: Crotonopsis Michaux, FI. Bor.-
Amer. 2: 185. 1803. Leptemon Rafinesque,
Med. Repos. II. 5: 353. 1808, nom. superfl.
TYPE: Crotonopsis linearis Michaux.
Friesia Sprengel, Anleit. Halle Gewachs. 2(2): 885. 1818.
TYPE: Friesia argentea Sprengel [= Crotonopsis
linearis Michaux].
This section contains two species, sometimes com¬
bined but well discriminated by Pennell (1918), fol¬
lowed by Correll & Johnston (1970). Unfortunately,
both species have to be renamed because all available
epithets are preoccupied in Croton.
1. Croton michauxii Webster, nom. nov. Re¬
placed name: Crotonopsis linearis Michaux,
FI. Bor.-Amer. 2: 185. 1803, non Croton li¬
nearis Jacq., 1760. Crotonopsis argentea
Pursh, FI. Amer. Sept. 1: 206. 1814, nom.
superfl.; non Croton argenteus L., 1 753. TYPE:
U.S.A. South Carolina: Long Bay, Michaux
s.n. (lectotype, selected here, Michaux Her¬
barium, P; microfiche seen). The only specific
location given by Michaux was Long Bay, which
is in South Carolina according to Ewan (1974);
it seems appropriate to select it as the lectotype.
Crotonopsis spinosa Nash, Bull. Torrey Bot. Club 22:
157. 1895, non Croton spinosus Forsk., 1775.
TYPE: U.S.A. Florida: Marion Co., Dunnellon,
Swingle 1397a (holotype, NY).
2. Croton willdenowii Webster, nom. nov. Re¬
placed name: Crotonopsis elliptica Willd., Sp.
PI. 4: 380. 1805, non Croton ellipticus Gei-
seler, 1807. TYPE: U.S.A. South Carolina:
Muehlenberg s.n. (lectotype, selected here,
Willdenow Herbarium 17490, B).
Crotonopsis abnorrnis Baillon, Etude Euphorb. 381. 1858
non Croton abnorrnis Baillon, 1864. TYPE: without
locality, Leconte s.n. (P not seen).
Although Merrill (1949) listed three species of
Rafinesque as synonymous with Crotonopsis ellip¬
tica Willd., Rafinesque did not cite Willdenow, and
all three Rafinesque species—in the absence of clear
typification—could possibly be synonyms of Cro¬
tonopsis linearis Michaux. It seems preferable,
therefore, to apply a new name to the species that
can be securely based on a type specimen.
Section Eremocarpi s
Croton sect. Eremoearpus (Bentham) Webster,
stat. nov. Eremoearpus Bentham, Bot. Voy.
Sulphur 53. 1844. Piscaria Piper, Contr. U.S.
Natl. Herb. 11: 382. 1906, nom. superfl.
TYPE: Croton setigerus Hooker.
1. Croton setigerus Hooker, FI. Bor.-Amer. 2:
141. 1838. TYPE: U.S.A. Oregon: Menzies’
Island, and banks of Columbia River, Douglas
s.n. (not seen; presumably at K).
Section Jt locroton
Croton sect. Julocroton (Mart.) Webster, J. Ar¬
nold Arbor. 48: 354. 1967. Julocroton Mart.,
Flora Beibl. 1837(2): 119. 1837, nom. cons.
TYPE: Julocroton phagedaenicus Mart. [=
Croton triqueter Lam.].
Cieca Adanson, Fain. PI. 2: 355. 1763, nom. rej. TYPE:
Croton argenteus L.
Heterochlamys Turcz., Bull. Soc. Imp. Naturalistes Mos-
cou 16: 61. 1843. TYPE: Heterochlamys quin-
quinervia Turcz. [= Croton argenteus L.].
Centrandra Karsten, Linnaea 28: 440. 1857. TYPE:
Centrandra hondensis Karsten [= Croton honden-
sis (Karsten) Webster],
Julocroton subg. Eremadenia Didr., Vidensk. Meddel.
Dansk Naturhist. Foren. Kj 0 benhavn 1857: 134.
1857. TYPE: Julocroton triqueter Didr.
Julocroton subg. Oligonychia Didr., Vidensk. Meddel.
Dansk Naturhist. Foren. Kjobenhavn 1857: 132.
1857. TYPE: Croton argenteus L. (lectotype, se¬
lected here).
In the only general revision of Julocroton so far,
Croizat (1943, 1944) recognized about 25 species.
Species discrimination in the genus is difficult, and
much work remains to be done in order to evaluate
Croizat’s proposals and determine how many valid
binomials there are. Here I am making new com¬
binations only for those species being cited as rep-
Volume 2, Number 3
1992
Webster
American Croton
271
resentatives of section Julocroton in an unpublished
survey of the sections of Croton. This includes most
of the species cited by Mueller (1873), as well as
those names of Croizat that 1 have been able to
evaluate. Pending a critical revision of section Ju¬
locroton, it is not possible to evaluate all of the
binomials that have been proposed, but possible syn¬
onyms have been indicated where appropriate. In
the following enumeration the correct names under
Croton are given for those species of Julocroton
that could be verified and judged distinct. Unless
otherwise noted, microfiches and photographs cited
are at DAV.
1. Croton abutilopsis Webster, nom. nov. Re¬
placed name: Julocroton abutiloides Spencer
Moore, Trans. Linn. Soc. Bot. II. 4: 465. 1895,
non Croton abutiloides IIBK, 1817. 1YPE:
Brazil. Matto Grosso do Sul: Corumba, Moore
968 (holotype, BM not seen; photograph, B).
2. Croton ackermannianus (Muell. Arg.) Web¬
ster, comb. nov. Basionym: Julocroton ack-
ermannianu: s Muell. Arg., FI. Bras. 1 1(2): 283.
1873. TYPE: Brazil. Minas Gerais: Acker-
mann s.n. [var. 8 ovatus] (holotype, G not
seen).
3. Croton aeuminatissimus (Pittier) Webster,
comb. nov. Basionym: Julocroton acuminatis-
simus Pittier, J. Wash. Acad. Sci. 20: 11.
1930. TYPE: Venezuela. Yaracuy: between La
Piedra and Yaritagua, Pittier 11175 (holotype,
US).
4. Croton allemii Webster, nom. nov. Replaced
name: Julocroton ramboi Smith & Downs, Sel-
lowia 11: 153. 1959, non Croton ramboi Al-
lem, 1979. TYPE: Brazil. Santa Catarina: Ita-
piranga, 1951, Rambo s.n. (PACA 19824)
(holotype, US).
It is appropriate to rename this species ior
my colleague Antonio Allern, who has contrib¬
uted important publications on the systematics
of Croton in southern Brazil.
5. Croton argenteus L., Sp. PL 1004. 1753.
TYPE: America (probably in Hortus Clifforti-
anus Herbarium, BM not seen; possible isotype,
1140.8 in LINN, microfiche).
Probable synonyms, mostly indicated by Croizat
(1943), include: Julocroton camporum Chodat &
Hassler, J. elaeagnoides Spencer Moore, and J.
integer Chodat & Hassler. Although accepted by
Croizat as a distinct species, J. montevidensis
Klotzsch ex Muell. Arg. may prove to be synony¬
mous with C. argenteus; hence no new name is
provided for it here.
6. Croton calonervosus Webster, nom. nov. Re¬
placed name: Julocroton nervosus Baillon,
Adansonia I. 4: 369. 1864, non Croton ner¬
vosus Klotzsch, 1843. TYPE: Brazil. Minas
Gerais: St. Hilaire B 2 2369 (lectotype, selected
here, P not seen; photograph, DAV).
7. Croton conspurcatus Schldl., Linnaea i: 380.
1832. Julocroton conspurcatus (Schldl.)
Klotzsch, Arch. Naturgesch. 7: 193. 1841.
Julocroton triqueter var. conspurcatus (Schldl.)
Muell. Arg., DC. Prodr. 15(2): 705. 1866.
TYPE: Mexico. Veracruz (?): “in dumetis prope
Tioselo,” Schiede 39 (holotype, HAL not seen).
B. Croton cordeiroae Webster, nom. nov. Re¬
placed name: Julocroton riedelianus Muell.
Arg., FI. Bras. 1 1(2): 278. 1873, non Croton
riedelianus Muell. Arg., Linnaea 34: 104.
1865. TYPE: Brazil. Sao Paulo: Sao Carlos,
Riedel s.n. (holotype, G not seen; photograph,
DAV).
The specific epithet acknowledges the con¬
tributions of my colleague. Iris Cordeiro, to the
systematics of Euphorbiaceae (including sect.
Julocroton) in Sao Paulo and adjacent areas
of Brazil.
9. Croton didrichsenii W ebster, nom. nov. Re¬
placed name: Julocroton humilis Didr., \ i-
densk. Meddel. Dansk Naturhist. Foren. Kjob-
enhavn 1857: 132. 1857, non Croton humilis
L., 1753. TYPE: Brazil. Sao Paulo: Mugi, Lund
s.n. (holotype, C not seen; microfiche of isotype
at G).
10. Croton doratophylloides (Croizat) Webster,
comb. nov. Basionym: Julocroton doratophyl-
loides Croizat, Revista Argent. Agron. 10: 139.
1943. TYPE: Argentina. Misiones: Salto Igua-
zu, Rodriguez 438 (holotype. A).
1 1. Croton flavispicatus Rushy, Mem. New York
Bot. Card. 7: 283. 1927. Julocroton peruvi-
anus Muell. Arg. var. flavispicatus (Rusby)
Croizat, Revista Argent. Agron. 10: 136. 1943.
TYPE: Bolivia. La Paz: Ixiamas, Cardenas
2026 (holotype, NY not seen; photograph,
DAV).
Julocroton peruvianus Muell. Arg., DC. Prodr. 15(2):
704. 1866, non Croton peruvianus Briq., 1900.
TYPE: Peru. Loreto: Tarapoto, Spruce 4290 (ho¬
lotype, G, microfiche seen).
Julocroton paniculatus Pax & Hoffm., Meded. Rijks-
Herb. Leiden 40: 22. 1921, non Croton paniculatus
Lam., 1786. TYPE: Bolivia. Santa Cruz: Bio Pirai,
Herzog 1486 (holotype, B, photograph).
272
Novon
12. Croton fuscescens Sprengel, Syst. Veg. 2:
874. 1826. Julocroton fuscescens (Sprengel)
Baillon, Adansonia I. 4: 367. 1864. TYPE:
Brazil. Possibly from Rio de Janeiro, Sellow
s.n. (not seen; microfiche of possible isotype at
G).
Probable synonym: Julocroton paulensis Usteri, as point¬
ed out by Croizat (1943).
13. Croton geraesensis (Baillon) Webster, comb,
nov. Julocroton geraesensis Baillon, Adansonia
I. 4: 370. 1864. TYPE: Brazil. Minas Gerais:
St. Hilaire B' 1295 (holotype, P not seen;
photograph, DAV).
Possible synonym: Julocroton valenzuellae (Chodat &
Hassler) Croizat.
14. Croton hondensis (Karsten) Webster, comb,
nov. Basionym: Centrandra hondensis Kar¬
sten, Linnaea 28: 440. 1857. Julocroton hon¬
densis (Karsten) Muell. Arg., DC. Prodr. 15(2):
704. 1866. TYPE: Colombia. Tolima: Honda,
Karsten s.n. (not seen).
15. Croton lanceolaris Webster, nom. nov. Re¬
placed name: Julocroton lanceolatus Klotzsch
ex Muell. Arg., DC. Prodr. 15(2): 702. 1866,
non Croton lanceolatus Cav., 1800. TYPE:
Brazil. Sao Paulo: Alegres, Riedel 2806 (ho¬
lotype, B presumed destroyed; photograph,
DAV).
16. Croton microcalyx (Muell. Arg.) Webster,
comb. nov. Basionym: Julocroton microcalyx
Muell. Arg., Linnaea 34: 142. 1865. TYPE:
Brazil. Sao Paulo: Salto Ytu, Riedel 2180 (ho¬
lotype, G not seen; photograph of isotype, DAV).
17. Croton phyllanthus (Chodat & Hassler)
Webster, comb. nov. Basionym: Julocroton
phyllanthus Chodat & Hassler, Bull. Herb.
Boiss. II. 5: 501. 1905. TYPE: Paraguay. La
Cordillera (?): Rio Carimbatay, Hassler 457 6
(photograph of holotype, G).
18. Croton rupestris (Chodat & Hassler) Web¬
ster, comb. nov. Basionym: Julocroton rupes-
tris Chodat & Hassler, Bull. Herb. Boissier II.
5: 498. 1905. TYPE: Paraguay. Paraguari:
Cordillera de Altos, Hassler 8090, 3554 (syn-
types, both at G; not seen).
Julocroton velutinus (Chodat & Hassler)
Croizat, based on a variety of J. rupestris, is
doubtfully distinct.
19. Croton rutilus (Chodat & Hassler) Webster,
comb. nov. Basionym: Julocroton rutilus Cho¬
dat & Hassler, Bull. Herb. Boissier II. 5: 501.
1905. TYPE: Paraguay. ( lanindeyu: Sierra de
Maraeayu [Mbaracayu], Ipe hu [Ype Jhu],
Hassler 5226 (holotype, G not seen; isotype,
UC). Possible synonym: J. pule her Croizat.
20. Croton salzmannii (Baillon) W ebster, comb,
nov. Basionym: Julocroton salzmannii Baillon,
Adansonia I. 4: 369. 1864. Julocroton pyc-
nophyllus Schldl. ex Muell. Arg., DC. Prodr.
15(2): 705. 1866. TYPE: Brazil. Bahia: Salz-
mann s.n. (holotype, P not seen; microfiche of
possible isotype at B).
Julocroton pycnophyllus, as originally pro¬
posed by Schlechtendal (Linnaea 19: 245.
1847), was not validly published. Although
Mueller validated it in the Prodromus, Baillon
had already provided a description under the
name Julocroton salzmannii.
21. Croton solanaceus (Muell. Arg.) Webster,
comb. nov. Replaced name: Julocroton humilis
var. solanaceus Muell. Arg., DC. Prodr. 15(2):
i 01. 1866 .Julocroton solanaceus (Muell. Arg.)
Muell. Arg., El. Bras. 11(2): 279. 1873. TYPE:
Brazil. Sao Paulo: Rio Verde, Riedel 261 (ho¬
lotype, B destroyed; isotype, G not seen).
The original type collection of Sellow at B
cited by Mueller (1866) having been destroyed,
the Riedel specimen cited by Mueller (1873)
can be chosen as neotype because it has a
specific locality. There is a photograph of the
Sellow specimen at B.
Probable synonym: Julocroton typhaecephalus Croizat.
22. Croton subpannosus Muell. Arg. ex Griseb.,
PL Lorentz. 48. Dec. 1874. TYPE: Argentina.
Cordoba: Aschochinga, Lorentz 291 (bolotype,
GOET not seen; isotype, G not seen; photo¬
graph of presumed isotype at B, DAV). Julo¬
croton subpannosus Muell. Arg., J. Bot. 12:
203. July 1874. TYPE: Argentina. Cordoba:
Aschochinga, Lorentz 291 (lectotype, selected
here, G not seen).
Owing to the curious history of publication
of this species, the holotype ol Julocroton sub¬
pannosus Muell. Arg. is the same specimen as
the isotype of Croton subpannosus Muell. Arg.
ex Griseb. Croizat (1941) discussed the priority
of the names of Mueller over those of Grise-
bach.
Croton dentosus Griseb., PI. Lorentz. 49. 1874. TYPE:
Argentina. Cordoba: Cordoba, Lorentz 292 (holo¬
type, GOET not seen; photograph of isotype at B).
Julocroton serratus Muell. Arg., J. Bot. 12: 227. Aug.
Volume 2, Number 3
1992
Webster
American Croton
273
1874. TYPE: Argentina. Cordoba: Cordoba, Lor-
entz 292 (lectotype, selected here, G).
There is the same relation between the two
names and their types as in the preceding para¬
graphs. Croizat (1943) appears to be correct
in combining C. dentosus with C. subpannosus.
Julocroton brittonianum Morong, Ann. New York Acad.
Sci. 7: 222. 1893. TYPE: Paraguay. Rio Pilco-
mayo, Morong 864 (holotype, NY not seen).
Croizat (1943) reduced Morong’s species to
the synonymy of C. subpannosus.
23. Crolon stipularis (Muell. Arg.) Webster,
comb. nov. Julocroton montevidensis a [var.]
stipularis Muell. Arg., DC. Prodr. 15(2): 703.
1866. Julocroton stipularis (Muell. Arg.) Muell.
Arg., FI. Bras. 11(2): 277. 1873. TYPE: Bra¬
zil. Goyaz: Pohl 1647 (holotype, B destroyed).
24. Croton triqueter Lam., Encycl. 2: 214. 1.86.
Julocroton triqueter (Lam.) Didr., Vidensk.
Meddel. Dansk Naturhist. Foren. Kjobenhavn
1857: 134. 1857. TYPE: Brazil. Rio de Ja¬
neiro: Rio de Janeiro, Commerson s.n. (holo¬
type, Herb. Jussieu 16339, P).
Julocroton phagedaenicus Mart., Flora 20, Beibl. 2:
119. 1837. TYPE: Brazil. Martius 164 (holotype,
M not seen; microfiche of isotype at G).
25. Croton verbascoides Webster, nom. nov.
Replaced name: Julocroton verbascifolius
Muell. Arg., DC. Prodr. 15(2): 701. 1866, non
Croton verbascifolius Willd., 1803. TYPE:
Brasilia meridionalis, Sellow s.n. (lectotype, se¬
lected here, G not seen; microfiche, DAV). 1 yp-
i fie at ion was effected by the Royal Air Force
in destroying the other syntype at B.
Acknowledgments. I thank Robert Rhode for
assisting in preparation of the manuscript, and cu¬
rators of the herbaria cited for assistance in ex¬
amining specimens and references.
Literature Cited
Baillon, H. 1858. Etude generale du groupe des Eu-
phorbiacees. Victor Masson, Paris.
Bentham, G. 1844. Eremocarpus. Pp. 53-54 in Bot¬
any of the Voyage of H.M.S. Sulphur. Smith, Elder,
London.
-. 1880. Croton. In: Bentham & Hooker, Gen¬
era Plantarum 3: 293-296.
Correll, D. S. & M. Johnston. 1970. Manual of the
Vascular Plants of Texas. Texas Research Founda¬
tion, Renner.
Croizat, L. 1941. Preliminaries for the study of Argen¬
tine and Uruguayan species of Croton. Darwiniana
5: 417-462.
-. 1943. Preliminari per uno studio del genere
Julocroton Martius. Revista Argent. Agron. 10: 117—
145.
-. 1944. Note sul genere Julocroton Martius.
Revista Argent. Agron. 11: 198-102.
Ewan, J. 1974. Michaux's Flora in American Botany.
Pp. ix-xlvi in facsimile edition of A. Michaux, Flora
Boreali-Americana. Hafner Press, New York.
Greene, E. L. 1891. Flora Franciscana. Cubery Printing
House, San Francisco.
Hooker, W. J. 1838. Croton ? setigerus in Flora Bo¬
reali-Americana 2: 141. Treuttel & Wiirtz, Paris.
Klotzseh, J. F. 1841. Neue und weniger gekannte sfi-
damerikanische Euphorbiaceen-Gattungen. Arch.
Naturgesch. 7(1): 175-204.
Macbride, J. F. 1951. Euphorbiaceae. In: Flora of Peru.
Field Mus. Nat. Hist., Bot. Ser. 13, 3A(1): 3 -200.
Merrill, E. D. 1949. Index Rafinesquianus. Arnold Ar¬
boretum, Jamaica Plain.
Mueller, J. 1865. Vorlafifige Mittheilungen aus dem
fur De Candolle’s Prodromus bestimmten Manuscript
fiber diese Fainilie. [Conclusion.] Linnaea 43: 1 -224.
-. 1866. Euphorbiaceae. In: A. P. de Candolle
(editor), Prodromus Systematis Naturalis Regni Ve-
getabilis 15(2): 189-1261.
-. 1873. Euphorbiaceae, Eucrotoneae. In: C. F.
P. von Martius, Flora Brasiliensis 11(2): 81 288.
Pennell, F. 1918. Notes on plants of the southern
United States —IV. The genus Crotonopsis. Bull.
Torrey Bot. Club 45: 477-480.
Webster, G. L. 1967. The genera of Euphorbiaceae in
the southeastern United States. J. Arnold Arbor. 48:
303-430.
-. 1975. Conspectus of a new classification of
the Euphorbiaceae. Taxon 24: 593-601.
Eight New Species of Ouratea (Ochnaceae) from Mesoamerica
Caroline Whitefoord
The Natural History Museum, Cromwell Road, London SW7 5BD, England
ABSTRACT. Revision of the Ochnaceae for Flora
Mesoamericana brought to light eight new species
of Ouratea. These species are described here: O.
osaensis and 0. rinconensis from the Rincon de
Osa area of Costa Rica, and 0. jcjensis, O. knap-
piae, (). darienensis, (). stenobasis, O. sulcatiner-
via, and (). tristis from Panama.
Fhe genus Ouratea consists of about 150 species
from Central and South America, and the West
Indies; they are woody, glabrous plants character¬
ized by alternate, simple leaves, bright yellow bi¬
sexual flowers, and fruits composed of 1-seeded
glossy, black drupelets borne on an enlarged, fleshy,
purplish red receptacle or torus. Asian and African
species formerly included in Ouratea have been
transferred to Gomphia (Kanis, 1968). Riley (1924)
reviewed the Mexican and Central American species
of Ouratea, increasing the number of species in the
region from 9 to 15. Dwyer (1944) believed there
were 28 species of Ouratea from Mexico, Central
America, and the W est Indies, of which 17 occurred
in Mexico and Central America. Dwyer (1944) listed
six species of Ouratea occurring in Belize and (Gua¬
temala, whereas Standley & Williams (1961) thought
there were only three, with an additional variety in
one of these. Dwyer (1944) had believed seven
species occurred in Panama, but when he reviewed
the Panamanian Ochnaceae (Dwyer, 1967) he made
great changes, accepting eight species, with only
three names unchanged from his 1944 account. As
these earlier revisions had shown such large changes
in Central American Ouratea, a thorough generic-
revision was clearly needed for Flora Mesoameri¬
cana. Covering the area from southern Mexico (the
states of Yucatan, Tabasco, and Chiapas) to the
Panama-Colombia border, the present study re¬
vealed the following new taxa, and a total of 22
Ouratea species in Mesoamerica. Eight of these are
new and are described here so that the names can
be used in the Flora Mesoamericana treatment.
1. Ouratea darienensis Whitefoord, sp. nov.
TYPE: P anama. Darien: above Cana on trail
to top of ridge of Pirre Massif, forested slopes,
1,100-1,350 m, 3 May 1990, Mcl 5 her son
15033 (holotype, BM; isotype. MO). Figure 1.
Ab O. nitida (Sw.) Engl, foliis nervis secondariis man-
ifestiore distinctis late et irregulariter dispositis, calyce
minore laeviore, differt.
I rees 8-15 m high, with slender branchlets.
Leaves (10.5—) 15-21 x 4-7 cm, lanceolate to ob¬
long or oblanceolate, the base cuneate or rounded,
the apex acuminate; chartaceous to coriaceous, dry¬
ing red- or greenish brown, slightly glossy and ru-
gulose, the margin slightly revolnte, crenate; ve¬
nation impressed on both surfaces, with 8-13 pairs
of major secondary veins arcuate-ascending, more
clearly evident beneath; petiole 4-10 mm; stipules
ca. 6 mm, caducous. Inflorescence 8-17 cm, ax¬
illary, paniculate, the rhachis conspicuously flat¬
tened, 1-4-branched, the lower branches 3.5-7 cm
long; flowers solitary or paired, sometimes shortly
pedunculate, well-spaced; pedicel 5—6(—7) mm, slen¬
der; sepals 5, 4-5 mm long, ovate-lanceolate, thinly
coriaceous, the inner margins scarious, the sepals
and pedicel smooth and shiny; petals 5, 6 mm long,
broadly obovate, clawed; stamens 10, the anthers
4 mm; gynophore 0.5 mm; style 2.5-3 mm. Fruit
not seen.
Paratype. Panama. DARIEN: Parque Nacional del Da¬
rien, slopes of Cerro Mali, headwaters of S branch of Rio
Pucuro, ca. 22 km E of Pucuro, 1,300-1,400 in, Cuad-
ros, Hammel, de Nevers & Herrera 3966 (BM, MO).
Ouratea nitida (Sw.) Engler also has leaves with
the venation impressed on both sides, a paniculate
inflorescence, and a calyx with five sepals. Ouratea
nitida is, however, a smaller plant, rarely reaching
6 m, and grows at altitudes up to 400 m, whereas
O. darienensis is a tree of about 8 m, found at the
much higher altitudes of 1,300 1,400 m. Ouratea
nitida occurs from southern Mexico to Nicaragua,
Guyana, Cuba, and Jamaica, whereas O. darienen¬
sis is known only from the state of Darien, Panama.
2 . Ouratea jefensis Whitefoord, sp. nov. TYPE:
Panama. Panama: forested slopes of Cerro Jefe,
near large coffee fincas, 24 Jan. 1970, Wilbur,
Weaver, Foster <£• Correa 11318 (holotype,
BM; isotypes. F, MO, US). Figure 2.
Ab O. insulae Riley foliis multo brevioribus, rhachide
graciliore, floribus late dispositis pedicellis longioribus fil-
iformibus, differt.
Novon 2: 274-281. 1992.
Volume 2, Number 3
1992
Whitefoord
Our ate a
275
Figure 1. Ouratea darienensis (McPherson 15033,
BM).
Shrubs or trees 3-8.5 m high, with slender bran-
chlets. Leaves 8-11 x 2.5—4.5 cm, ovate-lanceo¬
late or elliptic, the base cuneate or rounded, the
apex acuminate; coriaceous, slightly glossy on both
surfaces, drying greenish brown, the margin plane
or slightly revolute, subentire or sometimes minutely
toothed toward the apex, the teeth well-spaced; ve¬
nation inconspicuous, the major secondary veins
mostly 5-10 mm apart, arcuate-ascending, slightly
raised above, the minor secondary and tertiary veins
nearly straight, crowded, subparallel, immersed or
scarcely raised on both surfaces; petiole 2—5 mm;
stipules to 7 mm, caducous. Inflorescence to 10 cm,
paniculate, terminal or axillary, usually broadly py¬
ramidal; rhachis slender, smooth, with 1 —3(—6) wide¬
ly spaced, spreading, slender branches; flowers wide¬
ly spaced, solitary or occasionally paired; pedicel
(5—) 10— 12 mm, filiform; sepals 5, ca. 6 mm long,
ovate-lanceolate, chartaceous, the inner margins
scarious; petals 5, 8—9 mm long, broadly obovate,
reflexed, falling before the sepals; stamens 10, the
anthers 5 mm; gynophore 0.5-1.0 mm; style 4-5
mm, often flexuous. Torus 8-12 mm, obovoid;
drupelets ca. 8 mm, subglobose to obovoid.
Figure 2. Ouratea jefensis ( Wilbur, Weaver, Foster &
Correa 11318, BM).
Paratypes. Panama, panama: beyond Goofy Lake along
road to Cerro Jefe, Correa & Dressier 473 (MO); Cerro
Jefe, 2,700 ft., D'Arcy 12183 (MO); Cerro Jefe, Duke
9405 (MO); Cerro Jefe to La Eneida, 21 2,900 ft., Duke,
Dressier & Dwyer 8191 (F, MO); 8 mi. S of Goofy Lake
toward Cerro jefe, Dwyer 7082 (MO); Cerro Jefe, to
3,100 ft., Dwyer 8482 (MO); summit of Cerro Jefe, 23
km N of Pan-American Hwy., 1,004 ft., Folsom & Kearns
2718 (MEXU, MO); Cerro Jefe, summit to 1 mi. beyond.
Gentry, Dwyer & Tyson 3517 (MO); 800 in, Gomez-
Pompa, Quero & Gonzalez 3059 (MEXU, MO); 750
m, Gomez-Pompa, Quero & Gonzalez 3550 (MEXU,
MO); 12.5 km above Pan-American Hwy. on road from
El Llano to Carti-Tupile, 350 m, Kennedy, Correa &
Dressier 2489 (MO); NE of town of Cerro Azul, 20 km
by road from Inter-American Hwy., Mori & Kallunki
3644 (MO); NE of town of Cerro Azul, 20 km by road
from Inter-American Hwy., Mori & Kallunki 3656 (MO);
slopes of Cerro Jefe between Cerro Azul and La Eneida,
ca. 15 mi. NE of Panama City, Wilbur, Almeda & Luteyn
15568 (MO); slopes of Cerro Jefe beyond Cerro Azul
between 4 and 8 mi., Wilbur & Weaver 11382 (MO).
Distribution. This species is known only from the
Cerro Jefe area in premontane forest, from 350 to
954 m.
Ouratea jefensis, like (). insulae Riley from
southern Mexico, Belize, Guatemala, and Honduras,
276
Novon
Figure 3. Ouratea knappiae, in flower ( Dressier 4332,
MO).
usually has a broadly pyramidal, paniculate inflo¬
rescence. Ouratea insulae differs in its much stouter
rhachis and crowded flowers, the pedicels to 10 mm;
O. jefensis has a slender rhachis, filiform pedicels
(5-)10 12 mm, and well-spaced flowers. Ouratea
insulae is a larger tree (to 25 m) with larger leaves
(10-31 x 6-8 cm) than O. jefensis. Some speci¬
mens of O. jefensis have been determined as O.
cocleensis Dwyer, now included in O. prominens
Dwyer, a species with a 2-sepaled calyx from Nic¬
aragua, Costa Rica, and Panama. Many herbarium
specimens of O. jefensis have been labeled O. lucens
(kuntb) Engl., a variable and widespread species.
Ouratea lucens usually has a cylindrical inflores¬
cence of crowded flowers on ascending pedicels (5-
12 mm). Stipulelike scales, lacking in O. jefensis,
tend to persist in clusters at the base of current
growth in O. lucens. In 0. lucens the leaves are
larger (up to 29 cm long) and often obovate. This
species is widely distributed, occurring from south¬
ern Mexico to Colombia, while O. jefensis is re¬
stricted to the Cerro Jefe region of Panama (after
which it is named). Ouratea lucens is a very variable
species, from which more taxa might eventually be
split.
Figure 4. Ouratea knappiae, in fruit (Knapp <6
Schmalzel 5477, MO).
3. Ouratea knappiae Whitefoord, sp. nov. TYPE;
Panama. Panama: 8 km from Pan-American
Hwy. on the El Elano-Carti road, Rio Teribe
Valley, tropical wet forest, 300-400 in, 9 June
1982, Knapp & Schmalzel 5477 (holotype,
MO; isotypes, RM, MEXU). Figures 3, 4.
Ouratea valerii Standley valde affinis, sed foliis cras-
sioribus subtilitis rugosis, toro minore, difFert.
Trees 8-18(-25) m high, with stout branchlets,
glabrous. Leaves (24-)30-40 x 6.5-10.5(-13) cm,
narrowly oblong, narrowly elliptic or oblanceolate,
the base rounded or sometimes cuneate, the apex
acute, acuminate, or obtuse; stiffly and thickly co¬
riaceous, minutely rugulose on both surfaces, drying
reddish brown with a slight sheen, the margin rev¬
olute, subentire or sometimes crenulate-serrate to¬
ward the apex; venation inconspicuous with ca. 15-
25 major and numerous minor secondary veins,
widely arcuate-ascending, slightly impressed above
and raised beneath, the tertiary veins crowded,
spreading, immersed or impressed on both surfaces;
petiole 5-10 mm, stout; stipules ca. 15 mm, some¬
times persistent. Inflorescence 14-29 cm, terminal,
broadly paniculate with ca. 4 stout branches, the
Volume 2, Number 3
1992
Whitefoord
Our ate a
277
lowest branches to ca. 18 cm long; flowers often
crowded, in stout-stalked clusters of 2-5; pedicel
10-14 mm, slender, incrassate upward; sepals 3(-5),
when 3, 2 of the sepals with a deep longitudinal
groove, 10—13 mm long, ovate-lanceolate, coria¬
ceous, persistent, the inner margins scarious; petals
5, 11-18 mm long, obovate or oblong; stamens 10,
the anthers to 12 mm; gynophore 1 mm; style 9-
10 mm. Torus 5-10 mm, depressed-globose, the
surface rough and wrinkled; drupelets to 15 mm,
ellipsoid.
Paratypes. Panama, colon: en la carretera a la zona
maderera de Santa Rita, Correa & Dressier 1206 (MO).
PANAMA: El Llano-Carti hwy., 6-10 km N of El Llano,
Dressier 4332 (MO); El Llano to Card road, 7.8 km N
of Pan-American Hwy., holsom 3559 (MO); road from
El Llano to Card, 7.4 km N of Pan-American Hwy., 300-
500 m, Folsom & Maas 5188 (MO); near border with
Comarca de San Bias, along El Llano-Carti road, ca. 340
m, McPherson 9774 (BM, MO); El Llano-Carti road, 8-
11 km from Inter-American Hwy., 300-400 m, Mori
7723 (MO); El Llano-Carti road, 10.8 km from Inter-
American Hwy., 1,100-1,200 ft., Mori & Kallunki 4138
(MO); El Llano-Carti road, 12.2 km from Inter-American
Hwy., ca. 250 m, Mori & Kallunki 6368 (MO); El Llano-
Carti road, 16-18.5 km by road N of Pan-American
Hwy. at El Llano, ca. 400-450 m, Nee & Tyson 11003
(MO); El Llano-Carti road, 8-9 mi. from Pan-American
Hwy., 250 m, Sytsma <£ Andersson 4438 (MEXU, MO).
san bi as: El Llano-Carti road, Km 19, 350 m, de Nevers,
Blake & Herrera 4235 (BM, MO); El Llano Card road,
300 m, de N'evers & Herrera 4235 (MO); El Llano-
Carti road, 26.5 km along Rio Card Chico, 200 m, de
Nevers, Herrera & Charnley 5332 (MO).
Distribution. All collections seen are from trop¬
ical wet forest in the F4 Llano-Carti and Santa Rita
Ridge area, from 200 to 500 m.
Specimens of Ouratea knappiae have been iden¬
tified as O. valerii Standley. Ouratea knappiae
differs in the venation and the finely rugose texture
of the leaves, and the smaller, depressed-globose,
rough and wrinkled torus. Ouratea valerii has thin¬
ner, smoother leaves that dry a duller, more glaucous
brown; the torus is 10-15 mm, obovoid to turbinate,
and smooth. Ouratea valerii occurs in Honduras,
Nicaragua, Costa Rica, and in Panama, west ol the
Canal, whereas O. knappiae is restricted to Panama,
east of the Canal. Ouratea knappiae is named in
honor of Sandra Knapp, one of the collectors ol the
type specimen.
4. Ouratea osaensis Whitefoord, sp. nov. TYPE:
Costa Rica. Puntarenas; high road out of Rin¬
con de Osa, Rincon to 5 km W of Rincon,
secondary forest, 0-300 m, 13 Feb. 1974, R.
Liesner 2096 (holotype, MO). Figure 5.
Ab O. pyramidalis Riley foliis majoribus latiore ellip-
ticis vel oblongis leviter nitidis, panicula breviore latiore,
pedicellis minute verrucosis, drupeolis multo minoribus,
differt.
Shrubs 2-3 m high, with slender branchlets.
Leaves 21-23 x 6.5-9 cm, broadly elliptic or ob¬
long, the base cuneate or attenuate, the apex acu¬
minate; subcoriaceous, slightly shiny above, dull be¬
neath, drying grayish green or medium-brown, the
margin plane, subentire to crenulate; venation
crowded, obscure, major secondary veins ca. 12-
18 pairs, irregularly spaced, widely arcuate-ascend¬
ing, immersed or slightly raised or silicate above,
raised beneath, the minor secondary and tertiary
venation scarcely raised on both surfaces; petiole
5-10 mm, stout; stipules not seen. Inflorescence
ca. 5.5 cm, terminal, broadly paniculate, the lower
branches to 7 cm; flowers solitary or borne in widely
spaced, stalked clusters of 2-3; pedicel 8-11 mm,
slender, the rhachis and pedicel finely striate and
minutely prickly-verruculose; flowers not seen. To¬
rus ca. 4 mm, subglobose, drupelets 3 mm, ellipsoid.
'The fruit characters are described from immature
material.
Paratype. COSTA Rica, puntarenas: Rincon de Osa,
slopes adjacent to airport, disturbed primary forest, 20-
300 m, Liesner 1862 (MO).
Ouratea osaensis is perhaps most similar to O.
pyramidalis Riley from southern Mexico, Belize,
Guatemala, Honduras, and Panama and O. insulae
Riley from southern Mexico, Belize, Guatemala, and
Honduras, both of which also have paniculate inflo¬
rescences, but neither of which occurs in Costa Rica.
Ouratea pyramidalis has smaller leaves (11-20 x
4.5-7 cm) with a dull surface and a variously toothed
margin, whereas O. osaensis has a slightly shiny
adaxial leaf surface and a subentire to crenate mar¬
gin. Ouratea pyramidalis also has a larger inflo¬
rescence (6-18 cm long), with a smooth rhachis
and laxer pedicels than O. osaensis. Ouratea in¬
sulae is generally larger, sometimes a tree to 25 m,
with larger, more robust panicles (8-27 cm long)
with stouter, more spreading branches than O.
osaensis.
5. Ouratea rinconensis Whitefoord, sp. nov.
TYPE: Costa Rica. Puntarenas: Reserva For-
estal Golfo Dulce Osa Peninsula, Trocha de La
Tarde road, 10 km SW of La Palma, S of
Rincon de Osa, primary forest along red clay
soil ridge E of the Rio Rincon valley, 150-200
m, 28 Apr. 1988, R. Hammel <£: R. Robles
16770 (holotype, BM; isotypes, CR, MO). Fig¬
ure 6.
Ab O. flexipedicellato Dwyer foliis nervis secondariis
numerosioribus regulatim dispositis acutiore arcuato-as-
278
Novon
m
Figure 5. Ouratea osaensis (Liesner 2096, MO).
cendentibus, calyce ex sepalis duobus tantum constante,
differt.
Shrubs ca. 3.5 m high, with slender branchlets,
glabrous. Leaves 19.5-23 x 5.5-7 cm, oblance-
olate, the base cuneate or narrowly rounded, long-
acuminate at the apex; subcoriaceous, the upper
surface slightly shiny, drying reddish brown, the
margin plane, crenate; major secondary veins 18-
25 pairs, penniparallel, spreading-ascending, sulcate
above, slightly raised beneath, the minor venation
slightly raised on both surfaces; petiole 3-5 mm;
stipules 10 mm, caducous. Inflorescence ca. 20 cm,
terminating branchlets, narrowly and laxly panicu¬
late, the rhachis slender, with 1 -4 longer branches
near the base, the lowest branch ca. 7 cm; flowers
well-spaced, solitary or more usually in stalked clus¬
ters of 3(—5); pedicel 8-10 mm, incrassate upward,
rhachis and pedicel smooth; sepals 2, 8 mm long,
boat-shaped, coriaceous; flowers not seen. Torus 8
mm, obovoid; drupelets 1 1 mm, ellipsoid.
Known only from the type collection.
Ouratea rinconensis resembles the Panamanian
species (). fiexipedicellata Dwyer in leaf size and
shape and its long, cylindrical inflorescences. The
two species can be distinguished easily, because the
Figure 6. Ouratea rinconensis (Hammel & Robles
16770, BM).
secondary leaf veins of 0. rinconensis are penni¬
parallel, whereas those of 0. fiexipedicellata are
irregularly spaced along the midrib and crowded
toward the margin. The flowers of 0. flexipedicel-
lata have five sepals, while those of 0. rinconensis
have two. Ouratea rinconensis is named after the
type locality.
6. Ouratea stenobasis Whitefoord, sp. nov. TYPE:
Panama. Bocas del loro: Punta Pena, vicinity
of Chiriquicito, rainforest, ca. 1,000 ft., 7 June
1967, Lewis, Escobar, MacBryde, Oliver &
Ridgway 2173 (holotype, MO; isotype, MO).
Figure 7.
Ab O. flexipedicellato Dwyer rhachidi breviore gra-
ciliore, foliis latiore ovatis nervis secondariis ordinatim
penniparallelis, differt. Species foliis lamina basi subito
contracta a congeneribus bene distincta.
Shrubs 3-4 m high, with slender flexuous bran¬
chlets, glabrous. Leaves 14.5-22 x 5.5-8.5 cm,
broadly elliptic or oblong, abruptly narrowed into a
13-mm stalk at the base above the thickened, wrin¬
kled petiole, the apex abruptly acuminate; charta-
ceous to subcoriaceous, with a dull sheen on both
surfaces, drying yellowish brown or greenish, the
Volume 2, Number 3
1992
Whitefoord
Our a tea
279
margin plane or slightly revolute, crenate, subentire
toward the base; major secondary veins ca. 15-18
pairs, arcuate-ascending, penniparallel, sometimes
impressed on the upper surface, raised beneath, the
minor tertiary venation spreading, immersed; petiole
ca. 1 cm; stipules ca. 6 mm. Inflorescence ca. 9
cm, terminal, racemose or with 1-3 short branches
near the base; flowers solitary or in well-spaced,
pedunculate clusters of 2-3; peduncles ca. 3 mm;
fruiting pedicel ca. 10 mm, markedly incrassate
upward; flowers not seen, only very small buds pres¬
ent. Torus 9 12 mm, subglobose or obovate; drupe¬
lets ca. 12 mm, ellipsoid.
Paratype. Panama, bocas DEL TORO: La Fortuna area
to Chiriqui Grande and the oil pipeline, along dirt road
10 mi. from Continental Divide, just past second large
bridge, 1 mi. N from highway, 130 m, Hammel, Mc¬
Pherson <£ Sanders 14588 (MO).
Distribution. Both collections are from the Carib¬
bean slope of Bocas del Toro, Panama.
With its narrowly cylindrical inflorescence and
oblong leaves, Ouratea stenobasis resembles O. rin-
conensis of Costa Rica and 0. flexipedicellata of
Panama, which are also low-elevation species. How¬
ever, O. stenobasis has an inflorescence ca. 9 cm
long, whereas the other two species have inflores¬
cences 20-24 cm long. Ouratea stenobasis has
penniparallel secondary leaf venation, while O. flexi¬
pedicellata has irregularly spaced secondary vena¬
tion. Ouratea stenobasis can be distinguished from
all other species of Ouratea by the stalklike base
of the leaf, which gives the appearance of a slender
petiole above a pulvinus. The specific epithet is
derived from this characteristic leaf base.
7. Ouratea sulcatinervia Whitefoord, sp. nov.
TYPE: 1 ’anama. Panama: 10 km N of Magarita
on road to Madrono then 3 km W on ridgetop
road, forest remnants, 1,800 ft., 31 Jan. 1979,
Hammel 6020 (holotype, MO). Figures 8, 9.
Ah O. tuerekheimii J. D. Smith calyce tantum ex
sepalis duobus constante, fructibus maioribus, differt.
Shrubs or trees 2-5 m high, with stout flexuous
branchlets, glabrous. Leaves 23-37 x 5.5—10( —13)
cm, narrowly or broadly oblanceolate, the base nar¬
rowly or widely subauriculate, the apex acuminate;
stiffly coriaceous, usually slightly glossy on both
surfaces, drying reddish or greenish brown, brighter
beneath, the margin revolute, subentire or serrulate;
venation conspicuous with ca. 20 pairs of major and
numerous minor secondary veins usually deeply sul-
cate above, raised beneath, the tertiary venation
very fine, scarcely impressed, immersed, or raised
above and scarcely raised beneath; petiole 2—3(—5)
mm, stout; stipules 7-8 mm, persistent. Inflores-
Figure 7. Ouratea stenobasis (Lewis, Escobar,
MacBryde, Oliver & Ridgway 2173, MO).
cence 10-29 cm, terminal or terminating short leafy
side-shoots, usually paniculate with 14 slender
rhachises unbranched or with 1-4 spreading-as¬
cending branches, or rarely racemose; paired sti¬
pulelike scales 5 7 mm occasionally persistent on
the rhachis at and 1-3 cm above the base and at
the lower branching points; flowers well-spaced, sol¬
itary or 2-3 in shortly stalked or sessile clusters;
pedicel 7-12 mm, slender, much thickened in fruit;
sepals 2, 8-11 mm long, boat-shaped, coriaceous,
the margins not scarious; petals 5, 9-10 mm long,
obovate; stamens 10, the anthers 7.5 mm; gyno-
phore 1.0-2.0 mm; style 3-6 mm. Torus 10-15
mm, narrowly or broadly obovoid; drupelets 8-10
mm, ovoid.
Paratypes. Panama. COLON: Santa Rita ridge lumber
road, Correa & Dressier 918 (MO); Santa Rita Ridge
road, 4 mi. from Transisthmian Hwy. to Agua Clara
weather station, ca. 500 m. Dressier, Berg & Gentry
8834 (MO); Santa Rita Ridge to 8 mi. E of Transisthmian
Hwy., to 800 ft., Dressier, Lallathin & Dwyer 9010
(MO); Santa Rita E ridge, Dwyer <& Correa 8398 (MO);
Santa Rita Ridge, 2 mi. from Transisthmian Hwy., Gentry
1420 (MO); Santa Rita ridge, ca. 12 km from Transisth¬
mian Hwy., Hammel 3648 (MO); Santa Rita Ridge road,
ca. 8 mi. E of Transisthmian Hwy., along trail N of road,
350-440 m, McPherson & Merello 8254 (MO); Santa
Rita Ridge road, ca. 6 km from Boyd-Roosevelt Hwy.,
280
Novon
Figure 8. Ouratea sulcatinervia, in flower (Hammel
6020, MO).
Mori <£ Kallunki 2149 (MO); Santa Rita ridge, 7 kin
by road from Transisthmian Hwy., 3 km SE of Puerto
Pilon, 200-250 m, \ee <£■ Hale 9671 (MO); Santa Rita
Ridge road, 20 km from Transisthmian Hwy., Santa Rita
hills, Smith & Smith 3431 (F); Santa Rita Ridge road,
20 km from Transisthmian Hwy., 100 1,200 ft., Sytsma
1104 (MO, MEXU); Santa Rita Ridge road, 18-20 km
from Transisthmian Hwy., 1,000-1,200 ft., Sytsma 2056
(MO). SAN BI AS: El Llano-Carti road, 20 km from Inter-
american Hwy., 100-350 m, de Nevers , Vespucio &
Herrera 3965 (MO), panama: 7 mi. N of Cerro Azul on
road to Cerro Jefe, ca. 2,600 ft., Blum, Godfrey & Tyson
1800 (MO); 2.5 mi. N of Goofy Lake on road to Cerro
Azul, Croat 11528 (F, MO); El Llano road, 3.4 mi. from
the highway, ca. 1,000 ft., Croat 49100 (MO); Cerro
Jefe, 700 m, Davidse & D'Arcy 10106 (MO); Cerro
Azul, 2,000 ft., Dwyer 1371 (MO); 10 mi. from the Pan-
American Hwy. on the El Llano-Carti road, 350 m,
Knapp. Whitten & Huft 4726 (MO); along El Llano
Carti-Tupile road, 12 mi. above Pan-American Hwy.,
200-500 m, Liesner 1233 (MO).
Distribution. All collections are from Panama,
from the Santa Rita ridge, the Goofy Lake to Cerro
Azul and Cerro Jefe area, and along the El Llano-
Carti road, tropical wet forest to cloud forest,
(30-)l 00-700 m.
Many specimens of Ouratea sulcatinervia have
been identified as O. tuerckheimii J. D. Smith, a
species with similar leaves from Mexico and Gua-
Figure 9. Ouratea sulcatinervia, in fruit (Davidse &
D'Arcy 10106, MO).
temala. However, the leaves of O. sulcatinervia are
often shorter and stiffer, with the lateral veins more
regularly spaced and more deeply sulcate (from which
the specific epithet is derived). Ouratea tuerckhei¬
mii has two sepals, while O. sulcatinervia has five.
Ouratea sulcatinervia has a torus 10-15 mm long,
while that of 0. tuerckheimii is only 5-10 mm.
8. Ouratea tristis Whitefoord, sp. nov. TYPE:
Panama. Panama: summit of Cerro Jefe and
forests along road beyond summit, 26 Aug.
1967. Hayden 1010 (holotype, MO). Figures
10 , 11 .
Ouratea podogyno J. D. Smith forma et nervatura
foliorum O. podogyno J. D. Smith similis sed foliis char-
taceis non coriaceis; ab O. podogyno rhachidi graciliore,
calyce ex sepalis tantum duobus constanti, differt.
Shrubs or trees 1 -5 m high, the hranchlets slen¬
der, glabrous. Leaves 20-38 x 3-6.5 cm, narrowly
lanceolate or narrowly ovate-lanceolate, the base
cuneate, the apex acute or acuminate; chartaceous,
minutely papillose on both surfaces, drying dull
greenish brown, the margin sometimes revolute, sub¬
entire or sometimes bluntly and widely toothed to¬
ward the apex; venation conspicuous, the major
Volume 2, Number 3
1992
Whitefoord
Ouratea
281
Figure 10. Ouratea tristis, in flower (Hayden 1010,
MO).
secondary veins 15-30 pairs, sulcate above, raised
beneath, tertiary venation scarcely raised on both
surfaces; petiole ca. 5 mm, stout; stipules not seen.
Inflorescence ca. 6.5 cm, terminal or axillary, rac¬
emose; flowers solitary, widely spaced; pedicel ca.
9 mm, pedicel and rhachis minutely prickly-ver-
ruculose; sepals 2, 7 mm long, boat-shaped, thinly
coriaceous, persistent; petals 5, ca. 7 mm long,
obovate; stamens 10, the anthers 6 mm; gynophore
1.5 mm; style 4 mm. Torus 9 mm, subglobose;
drupelets 7 mm, ohovoid.
Paratype. Panama, panama: Cerro Jefe, 1 .5 mi. down
right turnoff 6.7 mi. past Goofy Lake, disturbed cloud
forest, 700 m, Sytsma, Hahn & Antonio 2808 (MO).
Ouratea tristis resembles 0. podogyna J. D.
Smith from Guatemala in size and leaf shape; how¬
ever, 0 . podogyna differs in its coriaceous leaves
and 5-sepaled calyx. Ouratea tristis is probably
allied to O. crassinervia, which also has dull, char-
taceous leaves, minutely papillate on both surfaces,
a racemose inflorescence and a 2-sepaled calyx.
However, the leaves of O. crassinervia are crowded,
30-54 cm long, oblanceolate, serrate, with the base
cordate or subauriculate, while those of O. tristis
Figure 11. Ouratea tristis, in fruit (Sytsma, Hahn &
Antonio 2868, MO).
are 20-38 cm long, narrowly lanceolate or ovate-
lanceolate, occasionally irregularly toothed towards
the apex, with the base cuneate. Ouratea tristis is
named for its sad, drab appearance.
Acknowledgments. Thanks are due to N. k. lb
Robson for help with the Latin diagnoses, to R. .).
Hampshire for editing the manuscript, and to cu¬
rators of the herbaria mentioned in the text for
lending specimens for study.
Literature Cited
Dwyer, J. D. 1944. Taxonomy of the Mexican, Central
American and West Indian species of Ouratea (Och-
naceae). Lloydia 7: 121-145.
-. 1967. Family 119. Ochnaceae. In: R. F.
Woodson Jr., R. W. Schery & Collaborators, Flora
of Panama, part VI. Ann. Missouri Bot. Card. 54:
25-40.
Kanis, A. 1968. A revision of the Ochnaceae of the
Indo-Pacific area. Blumea 16: 51-52.
Riley, L. A. M. 1924. The Mexican and Central Amer¬
ican species of Ouratea. Kew Bull. 1924: 101-114.
Standley, P. C. & L. O. Williams. 1961. Ochnaceae.
In: P. C. Standley & L. O. Williams (editors). Flora
of Guatemala, part VII, no. 1. Fieldiana, Bot. 24(7/
1): 10-16.
A New Species of Senecio Section Dichroa
(Asteraceae-Senecioneae) from Brazil
Elsa 'Aardim
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
ABSTRACT. Senecio serranus from Parana, Brazil,
is described as new. It is related to Senecio dichrous
(Bong.) Schultz-Bip., but does not have glandular
hairs and does have leaves of the same color on
both surfaces. Both species belong to Senecio sect.
Dichroa C.abr. and unlike all other ones of the
section, have rayless heads with only one type of
(loret.
Among the collections made by Gert Hatschbach
in the state of Parana, Brazil, there is an undescribed
species of Senecio related to Senecio dichrous (Bong.)
Schultz-Bip. but without glandular hairs and with
leaves not paler on lower surface than on upper,
which is here described.
Senecio serranus Zardini, sp. nov. TYPE: Brazil.
Parana: Campina Grande do Sul, Serra Capivari
Grande, 6 Mar. 1969, Hatschbach 21212 (ho-
lotype, MBM; isotypes, MO, US not seen, dis¬
tributed as Senecio dichrous). Figure 1.
Herba erecta; caule striato profunde sulcato glabro ca.
1 m alto. Folia petiolata alterna; lamina glabra oblonga,
basi attenuata, apice acuta, margine grosse dentata, nervis
reticulatis. Capitula multa discoidea ad apices ramulorum
paniculato-corymbosa; involucro campanulato glabro ca-
lyculato ca. 9 mm alto ca. 6 mm crasso; bracteolis calyculi
4 6, lanceolatis, 9-13 mm longis; phyllariis 6 8, oblongo-
linearibus, apice acutis. Flosculi ca. 26, lutei, isomorphi,
hermaphroditi; corolla tubulosa ca. 10 mm longa, pen-
tadentata. Achaenia (immatura) costata glabra ca. 6 mm
longa; pappo albo capillari ca. 10 mm longo.
Erect herb. Stems striate, furrowed, glabrous, 1
m tall. Leaves petiolate, alternate. Blade oblong,
attenuate at base, acute at apex, coarsely dentate
at margin, with reticulate venation, glabrous or
sometimes with evanescent arachnoid tomentum be¬
low. Heads many, discoid, in terminal inflorescences
paniculate-corymbose. Involucre campanulate, gla¬
brous, 9 mm tall, 6 mm wide, with epicalyx; bracts
of the epicalyx 4-6, lanceolate, 9-13 mm long;
phyllaries 6-8, linear-oblong, acute. Florets around
26, yellow, isomorphic, hermaphrodite, with tubular
corolla 10 mm long, 5-lobed. Achenes (immature)
ribbed, glabrous, 6 mm long. Pappus white, capil¬
lary, 10 mm long.
Paratypes. Brazil. Parana: Campina Grande do Sul,
Pico Caratuva, 8 Feb. 1968, Hatschbach 18563 (MBM,
MO); Serra Capivari Grande, 18 July 1986, Silva &
Zelma s.n. (US not seen, distributed as Senecio dichrous).
Senecio serranus Zardini is similar to S. dichrous
(Bong.) Schultz-Bip. (basionym, Cacalia dichroa
Bong. Brazil. Minas Gerais: Riedel s.n., holotype,
LE not seen, photo, MO; see Fig. 2), from which it
can be differentiated by the following key.
la. Plants without glandular hairs; leaves of the
same color on both surfaces .S. serranus
lb. Plants with glandular hairs; leaves distinctly
paler on lower surface, green above, whitish
below .S. dichrous
These two species belong to Senecio sect. Dichroa,
one of the two sections of the genus characterized
by a hairy tuft at the apex of the style. The de¬
scription of this section (Cabrera, 1957) should be
amended to include rayless heads with only one type
of floret characteristic of both Senecio serranus and
S. dichrous. The latter species was treated by Ca¬
brera (1957) within section Dichroa as doubtful
because the type of its basionym, Cacalia dichroa
Bong., was not seen by him. All the other species
of the section have heads with two types of florets,
the marginal ones being ligulate.
Acknowledgments. I express special gratitude to
G. Hatschbach (MBM), to N. Imkhanitskaya of the
Komarov Botanical Institute (LE) for help with the
study of the type of Cacalia dichroa Bongard, and
to A. Cabrera (LP) for reviewing the manuscript.
Literature Cited
Bongard, H. G. 1840. Compositae Brasiliensis Novae.
Mem. Acad. Imp. Sci. Saint-Petersbourg, Ser. 6, Sci.
Math., Seconde Pt. Sci. Nat. 5: 40, pi. 7.
Cabrera, A. L. 1957. El genero Senecio (Compositae)
en Brasil, Paraguay y Uruguay. Arch. Jard. Bot. Rio
de Janeiro 15: 161-269.
Novon 2: 282-284. 1992.
Volume 2, Number 3
1992
Zardini
Senecio section Dichroa
283
Figure 1. Senecio serranus Zardini. —A. Plant. —B. Head. —C. Floret.
D. Style.
D
x 10
284
Novon
Figure 2. Photograph of the holotype of Cacalia dichroa (LF).
Volume 2, Number 1, pp. 1-80 of NOVON was published on 18 March 1992.
Volume 2, Number 2, pp. 81-178 of NOVON was published on 9 June 1992.
Volume 2
Number 4
1992
NOVON >y
Neviusia cliftonii (Rosaceae: Kerrieae), an Intriguing
New Relict Species from California
James R. Shevock
Department of Botany, California Academy of Sciences, Golden Gate Bark, San Francisco,
California 94118, U.S.A.
Barbara Ertter
University and Jepson Herbaria, University of California, Berkeley, California 94720, U.S.A.
Dean II . Taylor
Biosystems Analysis Inc., 303 Potrero Street, Suite 29-203, Santa Cruz, California 95060,
U.S.A.
ABSTRACT. A second species is added to Xet iusia
(Rosaceae: Kerrieae), previously known as a single
rare species in the southeastern United Stales. The
new species, IX. cliftonii from northern California,
differs from IX. alabamensis primarily in having
petals, fewer stamens, and more broadly ovate,
coarsely toothed leaves. The relict nature of Xev-
iusin is further supported by newly recognized fossils
from the Eocene of British Columbia, Canada.
During the course of conducting botanical inves¬
tigations in northern California in May 1992, the
third author and Glenn L. Clifton stopped at an
exposed limestone area on California Highway 299
east of Redding that had intrigued them for several
years. At the base of a shaded north-facing slope,
they collected a puzzling rosaceous shrub that they
could not recognize to genus. It was shown to the
senior author, who, after collecting additional ma¬
terial and confirming the shrub as Rosaceae, enlisted
the assistance of the second author, who was in the
midst of preparing the treatments of several rosa¬
ceous genera for The Jepson Manual: Higher Plants
of California.
After all Rosaceae in various western floras (e.g.,
Munz, 1959: Hitchcock et ah, 1961) were elimi¬
nated, the shrub was keyed to tribe Kerrieae in
Hutchinson (1964). Kerrieae has usually been cir¬
cumscribed to include three monotypic genera: ker-
ria DC. and Rhodotypos Siebold & Zucc. of eastern
Asia, and Neviusia A. Gray of the southeastern
United States. A comparison with herbarium ma¬
terial at CAS-DS and UC-JEPS indicated that the
shrub was unequivocally Neviusia , to the extent that
an initial suspicion was that the California material
represented an escape from cultivation. Subsequent
detailed morphological examination, however, sup¬
plemented by the discovery of two additional pop¬
ulations the following month, confirmed that it was
indeed a distinct new species of Neviusia, the first
addition to the tribe in 134 years. The key differ¬
ences follow:
Kky to the Species of Neywsia
la. Petals absent; leaves narrowly ovate to ovate,
finely toothed; stamens ca. 100 + , 4-7 mm
long; sepals 3.5-10 mm long, oblanceolate-ob-
ovate to elliptic, with 6+ teeth; style 5-6 mm
long; southeastern United States . . N. alabamensis
lb. Petals present, white, oblanceolate, 4-6 mm;
Novon 2: 285-289. 1992.
286
Novon
leaves ovate to cordiforin, coarsely toothed; sta¬
mens ca. 50+ , 4-5 mm long; sepals 3.5-6 mm
long, ± obovate with < 6 teeth; style ± 3 mm
long; northern California . N. cliftonii
INeviusia cliftonii .1. Shevock, B. Ertter & I).
Taylor, sp. nov. TYPE: United States. Calilor-
nia: Shasta Co., along Highway 299 7.4 road
mi. W ol Round Mountain post office, ca. 25
mi. E of Interstate Highway 5, T34N R1W
Sec. 29 NWNW, ca. 1,450 ft. elevation, un¬
derstory of mesic, north-facing, conifer-domi¬
nated forest on Hosselkus limestone, 5 May
1992, / ). If. Taylor & G. Clifton 12513 (ho-
lotype, JEPS; isotypes, CAS, CH, MO, NY).
Figure 1.
A I\. alabamense floribus petaliferis (petalis parvis
albis), foliis obovato-cordatis grosse dentatis, necnon stam-
inibus circa dimidio minus numerosis ± 50 (nec 100 + )
4-5 (vs. 4-7) mm longis differt.
Diffuse slender-hranched understory shrub, stems
erect, generally several, rarely > 1 cm diam., the
hark grayish near base, ± reddish brown above, ±
exfoliating in strips, without obvious lenticels; herb¬
age and young twigs ± strigose, the hairs ± 0.4
mm long; leaves alternate, primarily in upper Vz of
plant, generally expanded at anthesis, the stipules
linear-setaceous, tree from the 4-10(-15)-mm-long
petiole, often with small reddish glands, the leal blade
ovate to cordiforin, 2 6(9 on sterile shoots) cm
long, 1.5—5(—7) cm wide, ± bicolored, bright green
and sparsely strigose above, pallid and more densely
strigose below, the venation craspedodromus with
3-8 2° veins per side, the margin coarsely toothed
and shallowly lobed, the teeth apiculate; inflores¬
cence ± umbellate-corymbose, terminal, mostly on
short side branches, not otherwise pedunculate, the
pedicels 1-3 cm long, very slender, ca. 0.3 mm
thick (widening above); flowers (1 )3-5(-10), ap¬
pearing after or with leaves, the hypanthium ± flat,
± glabrous, 2-3 mm diam. (pressed); sepals 5 6,
± obovate, 3.5-6 mm long, 2-4.5 mm wide, veiny,
irregularly few-toothed distallv, spreading at anthe¬
sis, persisting in fruit; petals oblanceolate, 4 8 mm
long, white, quickly deciduous; stamens many, ca.
50 or more, ca. equaling sepals, the filaments 4-5
mm long, white, ± dilated, the anthers round, 0.3-
0.4 mm long, yellow; pistils 3-6, the ovary densely
white-strigose, the style ± 3 mm long, sparsely
strigose; fruit ± eccentrically ovoid achene, 3 4
mm long, brown, sparsely strigose.
Paratypes. United States. California: Shasta Co.,
type locality, 8 May 1692, Shevock 12127 (CAS, JSU,
K, KYO, MS A, UARK, UC, US), 19 May 1992, Taylor
12584 (HSC, JEPS, KUN, NA), 27 May 1992, Pap-
palardo & Molter s.n. (CHSC, Shasta-Trinity National
Forest herbarium), 13 June 1992, Shevock & Ertter
12192 (B, BM, CAS, GH, F, GH, ILLS, JEPS, JSU, LE,
MAK, MO, NAS, NY, PR, RM, RSA, TEX, UBC, US,
VDB, WTU); along Shasta-Trinity National Forest road
27, 0.6 mi. N of Squaw Creek Forest Service station,
T35N R2W Sec. 20 NENE, 1,200 ft., 13 June 1992,
Shevock & Enter 12197( ARIZ, CAS, CPU, DAV, JEPS,
NCU, NY, OBI, OSC, PE, RSA, SBBG, UC, UCR); along
Shasta-Trinity National Forest road 7H009 at Ellery Creek
on McCloud River Arm of Shasta Lake, T35N R3W Sec.
6 SE !4, 1,250 ft., 13 June 1992, Shevock & Ertter
12199 (CAS. CHR, 1BSC, LA, SD, UC).
Eponymy. Named in honor of one of the discov¬
erers, Glenn L. Clifton (1943 ), ardent collector of
the California and western United States flora. His
30,000 specimens distributed among 6,000 species,
including numerous significant collections, form a
major portion of the holdings at Pacific Union Col¬
lege (PUA). We suggest “Shasta Snow-wreath” as
a euphonious vernacular name, parallel to the es¬
tablished name of “Alabama Snow-wreath” for /V.
alabamensis.
Ecology. Lite three known occurrences are well
spaced around the eastern half of Shasta Lake north¬
east of Redding, 60-80 km south of Mount Shasta,
with an elevation range between 300 and 500 m.
All sites are on limestone substrates in shaded cool-
air canyons adjacent to creeks; similar situations on
other substrates were searched without finding any
Neviusia. Limestone, relatively rare in northern Cal¬
ifornia, is centered around Shasta Lake, where ac¬
cess to many areas is extremely limited due to the
rugged, densely forested terrain with few roads or
trails. Limestone is likewise a common substrate for
I\. alabamensis, which also occurs on sandstone,
sandy loam, and shale.
The ecological setting in which the /V. cliftonii
populations occur is typically very species-rich, in¬
cluding species at their northern or interior range
limit. Associated species include the following: Acer
macrophyllum Pursh, Aesculus californica (Spach)
Nutt., Arbutus menziesii Pursh, Aristolochia cal¬
ifornica Torrey, Asarurn hartwegii S. Watson, Her-
beris aquifolium Pursh var. dictyota (Jepson) Jep-
son, Calycanthus occidentalis Hook. & Am., Cercis
occidentalis Torrey ex A. Gray, Clematis lasiantha
Nutt., Corylus cornuta Marsh var. californica (A.
DC.) Sharp, Cornus sericea L., Cornus sessilis Du¬
rand, Holodiscus discolor (Pursh) Maxim., Tigtts-
ticum californicum L. Coulter & Rose, Lonicera
hispidula Douglas var. vacillans A. Gray, Paxis-
tima myrsinites (Pursh) RaL, Philadelphus letcisii
Pursh subsp. californicus (Bentham) Munz, Phy-
socarpus capitatus (Pursh) Kuntze, Polygala cor¬
nuta Kellogg, Prunus subcordata Bentham, Pseu-
Volume 2, Number 4
1992
Shevock et al.
Neviusia cliftonii
287
dotsuga menziesii (Mirbel) Franco, Quercus
chrysolepis Iiebm., Q. garryana Hook. var. bretv-
eri (Engelm.) Jepson, Q. kelloggii Newb., Q. wis-
lizenii A. DC., Rosa gymnocarpa Nutt., Smilax
californica (A. DC.) A. Cray, Styrax officinalis L.
var. californica (Torrey) Rehder, Symphoricarpos
albas (I,.) S. F. Blake var. laevigatas (Fernald) S.
F. Blake, Toxicodendron diversilobum (Torrey &
A. Gray) E. Greene, Trientalis latifolia Hook., Vitis
californica Bentham, and If hip plea modesta Tor¬
rey.
Although the three known sites are relatively
288
Novon
accessible and N. cliftonii occurs there as a domi¬
nant understory shrub, it is still possible that the
species may never have been previously collected.
There are no misidentified CAS, DS, JEPS, or UC
holdings of Physocarpus Maxim, (the genus it is
mostly likely to be mistaken lor, as it was in Arkansas
(Moore, 1956)), Holodiscus Maxim., or Spiraea L.
One potential cause is the prominent co-occurrence
of poison-oak, Toxicodendron diversilobum, suffi¬
cient to discourage casual exploration. In addition,
at the type locality along Highway 299 crossing
Cedar Creek during peak flows can be difficult; how¬
ever, 1992 marks the fifth year of drought in north¬
ern California.
Relationships. Kerrieae is a distinctive tribe
characterized by serrate or imbricate persistent se¬
pals, numerous stamens arranged in several series,
and 1-8 drupaceous achenes (Baillon, 1869). As
discussed by Robertson (1974), the tribe is mor¬
phologically intermediate between subfamilies Spi-
raeoideae and Rosoideae.
Neviusia alabamensis is apetalous and has 2-4
carpels; it is restricted to the southeastern United
States. Kerria japonica (1,.) DC., with five showy
yellow petals and 5-8 carpels, is native to China
and Japan. Rhodotypos scandens (Thunb.) Makino,
also native to eastern Asia, has opposite leaves, large
white petals, and 14 carpels. Hutchinson (1964)
placed Rhodotypos in its own tribe, Rhodotypeae,
based on the opposite leaves (unusual in Rosaceae)
and two ovules per carpel.
Biogeographic significance. Neviusia cliftonii
is a remarkable addition to the California Floristic
Province. The immediate interpretation is that \ev-
iusia is an old, formerly widespread genus with
relicts in forest refugia separated by the uplift of
the western Cordillera and the formation of the Creat
Plains. It is therefore immensely satisfying that, by
serendipity, Wolfe and Wehr’s (1988: 181) recent
work came to our attention, wherein a fossil “ail.
Kerria" is mentioned from early middle Eocene
montane assemblages in the Pacific Northwest.
Wolfe’s suspicion (pers. comm.) that this fossil, from
the Princeton flora of southern British Columbia,
was probably closest to Neviusia was confirmed
when he was given the opportunity to examine leaves
of N. cliftonii. This also supports previous specu¬
lations that N. alabamensis is an epibiotic, relatively
old relict species, whose current distribution follows
the former edge of the Mississippi embayment of
the old Gulf Coastal Plain (Moore, 1956; Robertson,
1974).
Similar California eastern United States disjunc¬
tions are well known (Sharp, 1951; Wood, 1970);
e.g., Torreya, Dirca, and Calycanthus. Neviusia
cliftonii occurs at the southeastern edge of Klamath-
Siskiyou province of northern California, which con¬
tains the maximum concentration of arcto-tertiary
relicts in California (Raven & Axelrod, 1978: 55-
57). Counterparts to California-eastern North
American relicts are also often present in eastern
Asia, including Torreya and the Taxodiaceae. In the
case of Neviusia, this would be the genus Kerria
of Japan and China (Fernald, 1931). Several asso¬
ciates of N. cliftonii also represent this pattern to
a greater or lesser extent.
Rarity Status. Based on the known distribution
and extent of potentially suitable habitat, we believe
that N. cliftonii will prove to be a relatively re¬
stricted and rare endemic in the Shasta Lake area.
Mining is a very real potential threat; the Hosselkus
formation is a high-quality source material for ce¬
ment production and is the only such limestone in
northern California (Bowen et al.. 1973). Many ac¬
cessible deposits throughout the region have already
been mined. In addition, a recent major forest fire
came perilously close to wiping out the type locality
before the species was even described. We therefore
recommend that N. cliftonii be considered as a
candidate for the next addition to the U.S. Fish and
Wildlife Service Notice of Review ol plant taxa for
listing as Endangered and Threatened Species in the
Federal Register. It will be included in future edi¬
tions of the California Native Plant Society’s Inven¬
tory of Rare and Endangered I dscular Plants of
California.
Neviusia alabamensis is also a candidate lor
either threatened or endangered status in the bed-
eral Register. It is known from two sites in Alabama,
three in Arkansas, three in Tennessee, one in Geor¬
gia, and one in Mississippi (element records from
The Nature Conservancy, Arlington, Virginia). An
occurrence in Missouri is presumed extirpated, as
is the type locality in Alabama; goats are a potential
threat to some extant populations (Rogers & Wilmes,
1989).
All three genera ol Kerrieae are currently in
cultivation. Neviusia alabamensis, in cultivation
since its discovery in 1858 (Howard, 1976), has
recently been promoted for native plant gardens
(Rogers & Wilmes, 1989). It is generally propagated
from softwood cuttings and is hardy as far north as
Chicago and Boston. Cuttings and seeds of N. clif¬
tonii have been given to the University of California
Botanical Garden, a member of the Center for Plant
Conservation.
Acknowledgments. The full significance of this
exciting discovery has been amplified by the con¬
tributions of many other people, all ol whom have
Volume 2, Number 4
1992
Shevock et al.
Neviusia cliftonii
289
earned our deepest appreciation. Paleobotanists Jack
Wolfe, Wesley Wehr, and Howard Shorn provided
us with a fossil history of Neviusia. Forest Service
botanist Julie Kierstead Nelson acquired maps show¬
ing limestone outcrops around Shasta Lake. David
Whetstone of JSU gave us additional information
on the morphology and ecology of /V. alabamensis,
as did The Nature Conservancy staff in Arlington,
Virginia. Kenneth Robertson provided valued re¬
viewer’s comments. Linda Vorobik was able to fit a
“need-ASAP” illustration into her crowded sched¬
ule, with no sacrifice of her usual high quality. Dieter
\\ ilken granted an exemption to the “absolutely no
more additions” barrier for including new discoveries
in the Jepson Manual. Last, but certainly not least,
Rupert Rarneby checked and corrected the Latin,
sharing our enthusiasm for this discovery.
Literature Cited
Baillon, H. 1869. Monographic des Rosacees. Hist. PI.
1: 345-483. 1869. [English translation by M. M.
Hartog, The natural history of plants 1: 335-471.
1871.]
Bowen, 0. E., C. H. Gray & J. R. Evans. 1973. The
mineral economics of carbonate rocks. Bulletin 194,
California Division of Mines and Geology, Sacra¬
mento.
Fernald, M. L. 1931. Specific segregation and identities
in some floras of eastern North America and the Old
World. Rhodora 33: 25-63.
Hitchcock, C. L., A. Cronquist, M. Ownbey & J. W.
Thompson. 1961. Vascular plants of the Pacific
Northwest. Part 3: Saxifragaceae to Ericaceae. Univ.
Wash. Publ. Biol. 17: 1-614.
Howard, R. A. 1976. In defense of the Rev. Dr. Reuben
D. Nevius and the plant called Neviusia. Arnoldia
36: 57-65.
Hutchinson, J. 1964. The Genera of Flowering Plants.
Clarendon Press, Oxford.
Moore, D. M. 1956. Neviusia alabamensis in Arkan¬
sas. Rhodora 58: 187-191.
Munz. 1959. A California Flora. U niv. California Press,
Berkeley.
Raven, P. H. & D. I. Axelrod. 1978. Origin and re¬
lationships of the California flora. Univ. Calif. Publ.
Bot. 72: 1-134.
Robertson, K. R. 1974. The genera of Rosaceae in the
southeastern United States. J. Arnold Arbor. 55:
303-332, 344-401, 611-662.
Rogers, G. & P. Wilmes. 1 989. The Alabama snow
wreath. Amer. Hort. (Alexandria) 68(4): 40-41.
Sharp, A. J. 1951. Relationships between the floras of
California and southeastern United States. Contr.
Dudley Herb. 4: 59 61.
Wolfe, J. A. & W. Wehr. 1988. Rosaceous Charnae-
batiaria-Yike foliage from the Paleogene of western
North America. Aliso 12: 177-200.
Wood, C. E. 1970. Some floristic relationships between
the southern Appalachians and western North Amer¬
ica. Pp. 331-404 in P. C. Holt (editor). The Dis¬
tributional History of the Biota of the Southern Ap¬
palachians. Part II. Flora. Virginia Polytech. Inst.
State Univ. Res. Div. Monogr. 2.
Eleven New Species, a New Variety, and a New Varietal Combination
in the Fern Genera Asplenium and Diplazium in Central America
C. Dennis Adams
British Museum (Natural History), Cromwell Road, London, SW7 5BD, United Kingdom
ABSTRACT. The following new taxa of ferns, As¬
plenium barclayanum, I. salicifolium var. aequi-
laterale, A. seileri, Diplazium atirrense var. lob-
ulatum , /). chimuense, D. ehiriquense, D.
croatianum, D. gomezianum, D. hammelianum ,
D. matudae, D. moranii, D. panamense, D. In¬
tense, have been revealed through research carried
out in the preparation of contributions lor Horn
Mesoamerieana.
Asplenium barclayanum C. 1). Adams, sp. now
TYPE: C osta Rica. Isla del Coco: “Cocos Is¬
land,” 3-7 Apr. 1838, Barclay 2196 (holo-
type, BM). Figure 1.
Filix A. eristati Lam. affinis sed lamina basin versus
sensim contracta, pinnis mediis lanceoiatis (non oblongis),
pinnula basali singulari libera 2-4-lobata, rhachidi viridi-
alata, apicem versus ad 1 mm plusve dilatata differt.
Rhizome 1-3 x 0.2-0.4 cm, suberect or shortly
creeping; scales 1-5 x 0.20.8 mm, elongate-del-
tate from a rounded auriculate base, tapered to a
short hair tip, remotely denticulate, flat or slightly
crispate, clatlirate, dark brown or blackish with nar¬
row paler margins, slightly lustrous, eventually re-
tiform, clustered over rhizome apex and extending
to base of petiole only; leaves 1 1 38 cm long, tufted
or closely approximate; petiole 2 8 cm long, pur¬
plish or grayish brown, narrowly green-winged from
near base, brittle with residual vascular shreds; blade
9-30 x 2-5.5 cm, 2-pinnate-pinnatifid, lanceolate,
gradually narrowed to base, with apex shortly acu¬
minate, nonconform; rachis green-winged adaxially,
widening at pinna bases and laterally to ca. 1 mm
or more overall below the tip, confluent with pinna-
bases on the basiscopic side, rounded and purplish
brown abaxially, becoming stramineous or green dis-
tally, with a few short glandular hairs near junctions
with pinnae; pinnae 0.5—3 x 0.4—1.2 cm, the lon¬
gest submedial, stalked to less than 1 mm long, 1 5
30 pairs, obliquely debate to lanceolate in outline,
with a distinct 2-4-lobed pinnule proximally on the
acroscopic side and up to 6 uniform pairs of simple
or shortly forked lobules; costae prominent adaxi¬
ally, winged throughout; proximal pinnules 3 5 x
2 1 mm, subflabellate; terminal segment linear, as-
eending-lobulate; ultimate segments to ca. 4 x 1
mm, linear or oblong, obtuse or emarginate, her¬
baceous, glabrescent, scarcely discolorous; veinlets
simple or 1-forked in the lobed segments, evident,
ending in linear hydathodes well short of margins;
sori 0.5-3 mm, solitary or double and sometimes
continuous around the veinlet-tip; indusia 0.4-0.5
mm wide, hyaline, colorless or light brown with
broadly rounded entire or shortly erose margin, gap¬
ing and pocketlike at maturity; spores ca. 42 x 30
jurn, light brown, reniform-ellipsoid with loose, un¬
sculptured, sparsely rugose perispore.
Paratypes. Costa Rica, isla m i coco: along brook
flowing into Wafer Bay, ca. 700 ft., abundant on walls
of cliffs, 19 Apr. 1930, Svenson 339 (MO); rivulet flowing
into Chatham Bay, Mar. 1970, Gomez 3337 (F); sin.
loc., Menzies s.n. (K).
This species has been collected several times on
the Isla del Coco and has been aligned with As-
plenium macraei Hook. & Grev. (Svenson, 1938;
Gomez, 1975). A sheet seen by Svenson at Kew
and filed with the holotype of A. macraei carries a
mixture of collections comprising a piece of A.
rnyriophyllum from Galapagos, a small withered
leaf collected by Macrae which may or may not be
f. macraei , and three leaves of another isplemum,
annotated “Asplenium Owhyhee A. M.. ’ which led
Svenson to believe that his collection from Cocos
Island was also A. macraei. It has been confirmed
by E. W. Groves that these three leaves were not
collected by Macrae, but rather by Alexander Men¬
zies on Cocos Island on 26 Jan. 1795, and he agrees
with me that they do not represent that species.
Asplenium macraei seems to be a polymorphic
species within the Hawaiian Islands, but the type
comes closer to A. sciadophilum Proctor than to
A. barclayanum, which has the ultimate segments
linear or elliptic, rather than oblanceolate-oblong,
with more open evenly spaced sinuses and a more
broadly winged upper rachis. Other species of the
affinity of A. cristatum, but with the blade narrowed
gradually to the base, are A. cladolepton fee and
I. rnyriophyllum (Sw.) C. Presl. 1 hey both dilfer
from A. barclayanum in having up to seven pairs
of pinnules per pinna rather than a single free proxi¬
mal acroscopic pinnule on the larger pinnae.
Collections from Isla del Coco, cited by Gomez
Novon 2: 290-298. 1992.
Volume 2, Number 4
1992
Adams
Asplenium and Diplazium
291
(1975: 43), but which have not been seen for this
study, are Gomez 4536 (CR, IJS), Wittier 16235
(GH), and Snodgrass & Heller 95 I (GH).
Asplenium salicifolium L. var. aequilaterale
(Christ) C. I). Adams, comb. nov. Basionym:
Asplenium auriculatum Sw. var. aequilaterale
Christ. Bull. Soc. Roy. Bot. Belgique 35: 197.
1896. TYPE: Costa Rica: Riolley 2672 (lec-
totype, selected here, BR).
This variety differs from the type variety by the
petiole drying brown rather than grayish green; the
pinnae being almost equilateral with rounded auricles
on both acroscopic and basiscopic sides rather than
strongly inequilateral with the proximal acroscopic
margin rounded-auriculate and the basiscopic mar¬
gin suhrectangular; the pinnae being more numerous
with up to 22 pairs instead of fewer than 17 pairs,
and herbaceous and discolorous rather than sub-
coriaceous anti concolorous.
Additional specimens examined. Costa Rica, san josk:
W part of montanas Jamaica, ca. 3 km NE of Bijagual
de Turrubares, Carara Reserve, 9°45.5'N, 84°33'W, 500-
600 m, 7 Aug. 1985, Grarum et al. 5834 (MO). Panama.
LOS SANTOS: Loma Prieta, Cerro Grande, 800-900 m, 8
June 1967, Lewis et al. 2238 ( MO).
Asplenium seileri C. D. Adams, sp. nov. TYPE:
El Sal vador. Laguna Verde, terrestrial in forest,
1,650 m, Seiler 957 (holotype, F).
Filix A. otitis Link affinis sed pinnis minoribus (2-4
mm latis), marginibus crenis late rotundatis interdum
emarginatis, textura submembranacea differt.
Rhizome 1 -1,5( 4.5) x 0.2-0.3 cm, erect; scales
1.5-2.5(-3) x 0.4 0.7 mm, linear-lanceolate, ta¬
pered to a hair tip, flat or subcucullate, slightly
sinuate, remotely denticulate, clathrate, dark brown
or black, lustrous, extending in smaller forms to
petiole; leaves 7.5-20 cm long, tufted, monomor-
phic; petiole 0.5-4 cm long, flexuous, green-mar-
ginate or winged to 0.25 mm wide from near base,
dark purplish brown and lustrous abaxially, with few
appressed septate hairs extending to rachis and ab-
axial surface of lamina, brittle, scarcely leaving vas¬
cular shreds; blade 7 17 x 1.4—2.3 cm, 1-pinnate,
linear-elliptic, narrowed to base, the tip long-acu¬
minate, nonconform pinnatifid, then linear-caudate,
regularly crenate-serrate; rachis with 2 patent green
adaxial wings (to ca. 0.2 mm wide) interrupted at
junctions with pinnae and confluent with their mar¬
gins, rounded and dark purplish brown abaxially;
pinnae 4— 12( 14) x 2-4 mm, the longest medial,
shortly (less than 0.3 mm) cartilaginous-stalked to
sessile only distally, 18—32 pairs, obliquely oblong,
the lowest deflexed and suhflabellately equilateral,
the others inequilateral, the upper [latent or slightly
ascending, shortly auriculate on the acroscopic mar¬
gin, the cuneate proximal margins mostly slightly
excavate, the acroscopic edge ± contiguous with
the rachis, the basiscopic extending at least halfway
to the obtuse dentate tip, the other margins with 3—
5 broadly rounded, sometimes emarginate, shallow
crenations, the costae scarcely more prominent than
the veins, submembranaceous, discolorous; veinlets
mostly simple, a few 1-forked on the acroscopic side
and 2-forked in the auricles, evident, ending in con¬
spicuous elongate-clavate hydathodes well within the
margins; sori 0.5-1.5(-2) mm, regularly disposed,
extending into the caudate leaf apex, rarely double;
indusia ca. 0.4 mm wide, hyaline, greenish or brown¬
ish, entire, folded back and often completely ob¬
scured by sporangia at maturity; spores 34-35 x
23-25 jam, light brown, reniforin-ellipsoid with loose
irregularly ridged echinulate perispore.
Paratypes. El Salvador, ahuachapan: terrestrial on
forest floor, Sierra Apaneca, SW of Apaneca village, 1,600
m, Molina R. & Montalvo 21797 (F). Costa Rica.
PUNTARENAS: terrestrial, Upper Rio Buru, Gomez et al.
21783 (MO).
292
Novon
The name Asplenium vicentinum Maxon was
used for this species by Standley & Calderon (1925:
21) and Calderon & Standley (1944: 29), based on
Standley 21574 (US), and Seiler (1 980: 49) in lists.
These uses were not accompanied by descriptions,
and I have been unable to ascertain that the binomial
was formally published elsewhere.
Diplazium atirrense (J. I). Smith) Lellinger var.
lobulatum C. I). Adams, var. nov. TYPE:
Costa Rica. Alajuela: Finca Los Ensayos, ca.
1 1 mi. NW of Zarcero, 850 m, 15 Aug. 1977,
( rant 43638 (holotype, MO; isotype, MEXU).
Pinnae per l A- l A distantiae ad costam in segmenta
rotundata cum areolis costalibus saepe consociata partitae.
Paratypes. Costa Rica, puntarenas: Canas Gordas,
Valle de Agua Buena, 1,100 in, Feb. 1897, Pittier 10968
(BR). Panama. CHIRIQUI: along road between Gualaca and
Fortuna Dam site, 10.1 mi. NW of Los Planes de Hornito,
1,260 m, 82°17'W, 8°45'N, 10 Apr. 1980, Antonio
4176 (MO); same locality, Croat 50043 (MO).
The new variety is distinguished from variety
atirrense by having the pinnae, at least at the base
of the leaf, divided l A-\A of the distance from the
margin to the costa into round-tipped segments,
which are often associated with costal areoles. The
typical variety has entire or subentire pinnae and
forms regular flattened areoles along the mid-upper
rachis-wing and elsewhere only casually.
Diplazium chimuense C. 1). Adams, sp. nov.
TYPE: Costa Rica. Limon: ridge between Cerro
Chimu and Cerro Matama, terrestrial, 1,200
m, 29 Apr. 1985, Gomez & Herrera 23544
(holotype, MO). Figure 2.
Diplazium foliis pinnatis, pinnis lateralibus subintegris
2 5-jugatis; pinna terminali libera 1-2-lobata, venulis an-
astomosantibus.
Rhizome ca. 1 cm thick, suberect; scales ca. 7
x 2.5 mm, ovate, shortly acuminate, somewhat
saccate, with irregularly toothed margins, dark
brown, clathrate, fenestrated, glossy; leaves 40 80
cm long, loosely tufted; petiole 18-40 x ca. 0.2
cm, purplish brown, scaly and darker near base,
like the rachis and costae puberulous; blade 22 40
x 14-20 cm, 1 -pinnate, ovate-deltate, acuminate;
lateral pinnae 10-14 x 3.5-4.5 cm, 2-4(-5) pairs,
oblong-lanceolate, broadly and slightly unequally cu-
neate to subtruncate or rounded at base with stalk
to 6 mm, the margins shallowly sinuate, the short -
or long-acuminate tip subentire with minute serra¬
tions obscured by downfolding; terminal pinna 16.5
18 x 4-5.5 cm (not including width of basal lobes),
lanceolate, acuminate, with 1 or 2 ovate, acuminate
basal lobes; lamina glabrous; midribs corrugate and
glabrescent abaxially; adaxial groove narrow, pu¬
berulous within, winged on both margins; veinlet-
groups about 20 on each side of costa, 2-4(-5)-
forked, with anastomoses between adjacent proximal
branches more frequent toward the margin; veinlet-
groups toward base of terminal pinna pinnate, up
to 10-forked; sori to 15 mm, often linked on prox¬
imal veinlets, mostly double; indusia 0.3-0.4 mm
wide including margin, fimbriate and ciliate with
septate hairs, brown, glossy.
Paratypes. Costa Rica, alajuela: Univ. of San Ra¬
mon’s Biological Field Station, ca. 20 km N of San Ramon,
cloud forest, 10°15'N, 84°30'W, 800-900 m, 27 Feb.
1988, Moran 4150 (MO, UC). HEREDIA: forest between
Rio Peje and Rio Sardinalito, Atlantic slope of Volcan
Barva, locally common on slopes in rich primary forest,
10°17.5'N, 84°04.5'W, 700-750 m, 2 Apr. 1986, Gra-
yum 6679 (MO, UC); ca. 30 km from San Ramon on
road to Los Angeles Norte, wet cloud forest, ca. 2,000
ft., 10 July 1972, McAlpin 1308 (CAS).
Among the Central American species with anas¬
tomosing veinlets, this is the only one with several
pairs of entire lateral pinnae and a more or less
conform terminal pinna.
Diplazium ehiriquense C. I). Adams, sp. nov.
TYPE: Panama. Chiriqui: Bajo Boquete, Finca
de T. Rios, terrestrial on bank of streamlet,
2,000 m, 7 Sep. 1982, Caballero 200 (holo¬
type, MO). Figure 3.
Filix IX franconis Liebm. affinis sed stipite rhachidique
brunnea (non straminea) et multo magis squamata, pin-
nulis vel pinnularum segmentis paribus 7-11 (non 13-
15 plusve) disposita, etiam pinnis prope bases gemmiferis
differt.
Rhizome to 7 x 2 cm, erect or ascending; scales
3-10 x 1-2.5 mm, lanceolate, acuminate, sub¬
entire or glandular-denticulate, twisted and curled,
brown or blackish brown, subclathrate; leaves 45-
120 cm, openly tufted in a short spiral; petiole 20
55 x 0.2 0.5 cm, like rachis and costae brown or
reddish brown, glabrous; scales at base of petiole
spreading, clathrate, extending in smaller ragged,
debate, or polygonal forms to costules and veins
abaxially; adaxial groove sparsely puberulous, gla¬
brescent or with a few dark hairlike scales; blade
25-75 x 10-27 cm, 2-pinnate or 2-pinnate-pin-
natifid (proximally at base), ovate-lanceolate, long-
acute; pinnae 4-13(-16) x 1.5-4( 5.5) cm, 12-
16( 18) pairs, lanceolate, usually alternate, long-
caudate acuminate, the basal slightly shorter, stalked
to 4 mm with, at least on mid and upper ones, a
small, rounded, dark, scaly bud on the acroscopic
side, incipient buds also on some adaxial surfaces
on costules or veins; pinnules (one pair proximal on
basal and mid pinnae only) 1-3 x 0.7-1.7 cm.
Volume 2, Number 4
1992
Adams
Asplenium and Diplazium
293
Figure 2. Diplazium chimuense C. D. Adams.
Figure 3. Diplazium chiriquense C. 1). Adams.
elliptical to ovate, entire or pinnatifid to halfway to
costule; other pinna-segments 0.5-2.5 x 0.5-1 cm,
8-10 pairs, ovate or oblong, obtuse or more usually
acute, often quite strongly falcate, shortly lobed,
crenate or serrulate, glabrous, herbaceous, green,
weakly discolorous; veinlets in larger segments ar-
buscular, alternate in 4-8 pairs, the proximal often
again lorked; sori to 6(-8) mm, mainly on acroscopic
hranchlets, double and single (distally); indusia 0.6
0.8 mm wide, dark brown with paler entire undulate
margins, folded back at maturity.
Paratypes. Panama. CHIRIQUI: along stream near trail
N of Cerro Punta, terrestrial, 29 May 1970, Croat 10449
(MO); Guadalupe, Cerro Punta, Finca Maduro, terrestrial,
2,000 m. Mar.-Apr. 1982, Caballero HI, 137, 138
(MO).
This fern has the affinity of D. franconis but the
stipe distally and the rachis are brown rather than
straw-colored and much more scaly; the pinnae have
fewer (7-11 instead of 13-15 or more) pairs of
pinnules or pinnular segments, and the mid and
upper pinnae bear small, dark, scaly buds near their
bases on the acroscopic side.
Diplazium croatianum C. I). Adams, sp. nov.
TYPE: Panama. Code: La Mesa, above El Valle
de Anton, ca. 2 km W of Cerro Pilon, on slopes
of steep knifelike ridge, cloud forest, terrestrial,
900 930 m, 22 July 1976, Croat 37301 (ho-
lotype, MO). Figure 4.
Affinis D. diplazioidis (Desr.) Alston sed indusiis pal-
lidis (non atrobrunneis) margine ciliis multo longioribus
(ca. I mm) quam membranae basalis latitudine (ca. 0.3
mm) instructo differt.
Rhizome unknown; scales (on petiole base) 3-6
x 1-2 mm, elongate-deltate, acuminate to slender
curled tips, denticulate-margined, black, shiny,
clathrate; leaves to 145 cm long; petiole 60 cm
long, brownish black, shiny, minutely puberulous;
blade 85 x 50 cm, 2-pinnate-pinnatifid, debate;
rachis, costae and costules dark brown to black with
numerous hair-tipped, sinuate, toothed scales, pu¬
berulous at least on major axes abaxially, glabres-
cent, shiny; adaxial groove glabrous or thinly scaly,
the wings interrupted and lobed at junctions; pinnae
to 30 x 13 cm, ca. 10 pairs, subopposite, ovate-
lanceolate, equilateral, stalked to 2 cm; pinnules to
6x2 cm, to ca. 12 pairs, shortly stalked, lanceo¬
late, the proximal slightly shorter, alternate, cut 2 A-
34 to the costule into 5 9 pairs of sessile segments;
ultimate segments to ca. 8 x 3.5 mm, the basal
basiscopic usually the largest, oblong, patent or an-
trorse, repand or crenulate, glabrous, membrana¬
ceous, scarcely discolorous; veinlets to 5 pairs in
294
Novon
the larger segments, simple, widely divaricate, black,
ending in hydathodes mostly reaching the margin;
sori very small (to ca. 3 mm) with 2 or 3 sporangia
each; indusia to 0.3 mm wide with membranous
light brown base and fimbriate margin with curled,
septate cilia to 1 mm long.
Besides its affinity with /). diplazioides, this spe¬
cies also comes close to the recently described I).
entecnum Mickel & Beitel (1983: 154), but prolif¬
erating buds at the bases of some of the pinnules
are lacking and the uncut part of the indusia is much
narrower. Diplazium laciniatum Mickel & Beitel
(1988: 156) is also close, but the pinnae of that
species have much shorter stalks, and the axes are
not puberulous.
Diplazium gomezianum C. I). Adams, sp. nov.
TYPE: Costa Rica. Puntarenas: road to Puerto
Jimenez, Osa, 40 km W of I.A. route 2, 100
m, Gomez 19533 (holotype, MO). Figure 5.
Filix D. herbacei Fee affinis sed foliorum axibus pu-
bescentibus squamatisque, pinnulis subsessilium, segmen-
tis majoribus petiolulis brevioribus insidentibus, hydathodis
a rnargine remote terminantibus, indusiis integris vel sub-
integris (nec fimbriato- nec glanduloso-marginatis) differt.
Rhizome shortly creeping; scales to 15 X 0.5
1.5 inm, linear, long-attenuate, tapered to one cell
or pair of cells, subentire, remotely denticulate,
brown, shiny, subclathrate; leaves 1.5-3 m long or
more; petiole 50-120 cm long or more, slightly
darker, pubescent and scaly at base; pneumato-
phores white; rachis and costae with small dark
brown scales; minor axes chaffy with hairlike, ragged
or stelliform scales and minute reddish glandular
hairs; blade 100 150 x 75-100 cm or larger,
3-pinnate-pinnatifid, ovate; adaxial groove puberu¬
lous proximally, glabrous distally, with wings inter¬
rupted and shortly lobed at junctions, thinly her¬
baceous, uniformly green; pinnae 30-60 X 1 2 30
cm, the basal slightly shorter, alternate from purplish
pulvini, equilateral with stalks to 3 cm, ovate to
oblong-ovate, acute; pinnules to 15 x 4 cm, 14-
20 pairs, fully pinnate with short stalks or more
usually sessile with continuous lamina, less than 2
cm apart, perpendicular to the costa, oblong-lan¬
ceolate, acuminate; divisions of the third order 10
20 x 5-8 mm, 8 12 pairs, obliquely oblong, adnate
at unequal attachment to costule, with a cleft sinus
on acroscopic base and a lobe to 3 X 2.5 mm, and
a rounded sinus at basiscopic base, with 5-7 pairs
of rounded, doubly crenate, slightly oblique lobes
Volume 2, Number 4
1992
Adams
Asplenium and Diplazium
295
cut one-third to nearly hallway to midvein; veinlets
l-2(-3)-forked in each segment-lobe; sori to 4 mm,
up to 7 pairs in each segment, double on proximal
veinlet and acroscopic branch, single distally; indusia
0.3 mm wide, oblong, brown, entire or subentire,
perpendicular at maturity.
Paratypes. Mexico, chiapas: Finca Mexiquito, July
1913 (st), Purpus 7112 (UC). Costa Rica. La Palma,
1,500 m, 24 Sep. 1905, llerckle 17103 (P). puntar-
ENAS: Canton de Buenos Aires Ujarras, headwaters of Rio
Kuiye following the ridges that overlook Olan, 9°17'50"N,
83°14'45"W, 1,450 m, 20 Sep. 1989, Herrera 3507
(CR not seen, MO).
Some Central American collections have been
identified as I), herbaceum Fee (Smith, 1981: 96).
That Brazilian plant differs in several respects, par¬
ticularly in having the stipe, rachis, and minor axes
glabrous, the pinnules longer-stalked, the ultimate
segments smaller and more distinctly stalked, the
hydathodes reaching to or very close to the margins,
and the indusia being hmbriate-ciliate and glandular-
margined.
Diplazium hammelianum C. I). Adams, sp. nov.
TYPE: Panama. Bocas del Loro: La Fortuna
area, ca. 2 mi. N of continental divide, in forest
along stream and on ridge above road, 8°46'N,
82°15'W, 950-1,100 rn, 7 Mar. 1986. Ilam-
mel et al. 11659 (holotype, MO). Figure 6.
Filix D. gomeziani C. I). Adams affinis sed laminae
textura membranacea (non herbacea), divisionibus pin-
natifidis ordinis tertii duplo longioribus sesquilatioribusque,
segmentis ohlongis (non rotundatis) fere ad costulas incisis
differt.
Rhizome shortly creeping, ascending; scales to
10x3 mm, lanceolate, long-attenuate to a hairlike
tip terminated by a knot of cells, subentire or ir¬
regularly lobed, not toothed, light tan to yellowish
brown, very shiny, subclathrate, not retiform; leaves
ca. 2 m long; petiole 85 cm long, not much darkened
at scaly base, glabrous, into rachis stramineous and
sparsely puberulous; costae and costules with a few
small, ragged, appressed scales abaxially; blade I 10
x ca. 90 cm, 3-pinnate-pinnatifid to nearly
4-pinnate, broadly ovate, openly divided, membra¬
naceous, dark green; adaxial groove minutely pu¬
berulous proximally, glabrous distally, the wings in¬
terrupted and shortly lobed at junctions; pinnae to
60 X 30 cm, to ca. 11 pairs, alternate from purplish
pulvini, to ca. 9 cm apart on rachis, equilateral with
stalks to 8 cm long, ovate, acute; pinnules to 1 8 x
8 cm, ca. 12 pairs, alternate to 2-2.5 cm apart on
the costa, stalked to ca. 8 mm or less, oblong-
lanceolate, acuminate; divisions of the third order
Figure 6. Diplazium hammelianum C. D. Adams.
14 x 0.7-1.2 cm, up to 4 — 10( 12) pairs, the
proximal stalked, oblong, acuminate, cut over half¬
way or nearly to the midrib into alternate segments,
the lamina continuous to the costa but not to its
base; ultimate segments 3-7 x 2-2.5 mm, 7-9
pairs, oblong, oblique, crenate-serrate; veinlets (2-)
4-5(-6)-forked in the segments, strongly antrorse;
sori 1-4 mm, single on the acroscopic veinlets;
indusia 0.2 mm wide, tapered at both ends, dark
yellowish brown, entire.
Diplazium hammelianum is a large fern resem¬
bling D. gomezianum C. D. Adams but with the
texture of the lamina tissue membranaceous rather
than herbaceous and the pinnatifid divisions of the
third order being about twice as long and half again
as wide with oblong rather than rounded segments
cut nearly to the midvein.
Diplazium matudae C. D. Adams, sp. nov. TYPE:
Mexico. Chiapas: Escuintla, near Col. Zinta-
lapa, in shaded brookside, 160 m, 23 July
1948, Matuda 18164 (holotype, MEXU). Fig¬
ure 7.
Affinis D. biolleyi Christ sed pinnulis basalibus tantum
penitus serrato-crenatis vel minus quam l A ad costulam
296
Novon
Figure 7. Diplazium matudae C. D. Adams.
divisis; squamis ad apicem piliformibus paucis vel absen-
tibus differt.
Rhizome unknown; scales on base of petiole 3-
7 x 2-3 mm, appressed, deltate, entire, brown,
clathrate, becoming retitorm and lacerate; leaves
150 160 cm long, slightly dimorphic; petiole 50-
65 x 0.7-0.9 cm, brownish-stramineous, pubescent
with appressed septate hairs, glabrescent; blade 90
100 X 45 50 cm. 2-pinnate-pinnatifid, ovate-lan¬
ceolate, pinnatifid and long-acuminate at tip; rachis,
costae and costules with few scattered, thin, ragged
or twisted, hair-tipped, small, paler brown scales and
numerous patent, septate, pale or brownish hairs,
light brown to stramineous, scarcely darker at junc¬
tion with pinnae; adaxial groove densely puberulous,
with narrow membranous, brownish wings inter¬
rupted and shortly lobed at junctions; pinnae 7-30
x 1.5-9 cm, the hasal slightly shorter with stalks
to 7 mm, 11-14 pairs, equilateral, lanceolate to
oblong-lanceolate, long-acuminate and serrate at tip;
pinnules 1.5-4 x 0.7-1.5 cm, larger on sterile
leaves, ca. 12-18 pairs, all sessile except 1 or 2
proximal often shorter and only narrowly adnate,
lanceolate or oblong, the larger cut at base l A or
less into 1 or 2 pairs of prominent lobes, elsewhere
crenate to serrulate, adaxially with a lew erect sep¬
Figure 8. Diplazium moranii C. D. Adams.
tate hairs along costae and near bases of sinuses,
abaxially with uniform erect, paler, septate hairs,
firmly herbaceous, green, scarcely discolorous; vein-
lets to ca. 13 pairs in the larger pinnules, each
branch again 1-4-forked; sori 2 8 mm, mostly di-
plazioid, single only distally; indusia 0.3 0.6 mm
wide, subentire, without or only very rarely with
short cilia, thin, hyaline or brown, folded upwards
and backwards and fragmenting at maturity.
Paratype. MEXICO. CUERRERO: along Highway 125
between Pinotepa Nacional and Tlaxiaco, ca. 8.4 mi. S
of Putla de Guerrero, ca. 1,000 in, 16 Jan. 1979, Croat
45808 (fertile) (MO), 45808A (almost all sterile) (MO).
Diplazium matudae is a medium-sized fern close
to I). biolleyi but with the basal pinnules at most
deeply serrate-crenate or divided less than one-third
to the costule; hair-tipped scales few or wanting.
Diplazium moranii C. D. Adams, sp. nov. TYPE:
Costa Rica. Cartago: forests near the entrance
to Parque Nacional Tapanti, 1,270 m, 3 Aug.
1983, Moran 3337 (holotype, MO). Figure 8.
Filix D. atirrensis (I. D. Smith) Lellinger affinis sed ad
basim apicis pinnatifidi sinibus magis dilatatis (latitudine
dimidium loborum contiguorum latitudinis excedentibus),
pinnis lateralibus latitudine 5 cm raro attigentibus differt.
Volume 2, Number 4
1992
Adams
Asplenium and Diplazium
297
Rhizome unknown; scales (on petiole-base) to 12
x 1.5 mm, elongate-deltate, slightly twisted, yellow-
brown, minutely forked-toothed along black sclerotic
margins; leaf 170 cm long; petiole 60 cm long,
brownish, scaly and thinly puberulous at base, be¬
coming more densely so into rachis and costae abax-
ially, this mixed indumentum extending to veinlets
and thinly, with minute forms of scales and hairs,
to lamina; blade 110 x 36 cm, 1-pinnate-pinnatifid,
oblong-lanceolate, narrowed at base, with deeply
pinnatifid acuminate apex, thinly coriaceous, dis-
colorous; adaxial midribs glabrous, without wings;
free pinnae 11-17 X ca. 4 cm, 2 pairs, ovate-
lanceolate, entire or subentire, with stalks 2-6 mm
long; sessile pinnae 16-22 x 3-5 cm, 4 pairs,
linear-lanceolate, subequilateral at narrowed cune-
ate base, crenate to shallowly pinnatifid with rounded
lobes cut one-third to costa, shortly acuminate to
an entire tip. the margin a narrow continuous band
of translucent tissue; apical segments 1.5-15 x
1.5-3 cm, ca. 9 pairs, the larger linear, entire,
widely spaced, slightly falcate, broadened at base to
an interconnecting narrow wing; veinlets in pinnate
groups, (l-)4- 10-forked, strongly divergent prox-
imally, subparallel distally, with intercostal anasto¬
moses forming flat areoles along the winged rachis
distally, otherwise generally free; sori double, con¬
tinuous along all veinlets except close to lamina
margin; indusium wanting.
This species is quite close to I). atirrense, but
the sinuses at the base of the pinnatifid apex are
wider, being more than half the width of the adjacent
lobes, and the width of the lateral pinnae is rarely
as much as 5 cm.
Diplazium panamense C. I). Adams, sp. nov.
TYPE: Panama. Chiriqui: along trail between
N fork of Rio Palo Alto and Cerro Pate Macho,
ca. 6 km NE of Boquete, 8°48'N, 82°23.5'\\ ,
montane rainforest and cloud forest, 1,600
2,000 m, 6 Feb. 1986, Smith et al. 2351
(holotype, UC). Figure 9.
Filix D. palmensis Rosenstock affinis sed laminae apice
pinnatifido, pinnis in paria 12-14 (non 2-9) dispositis,
costis venisque inferne appresso-pubescentibus differt.
Rhizome to 8 x 1.5 cm, erect; scales 6 10 x
2-2.5 mm, lanceolate, twisted, subentire, irregularly
denticulate, brown, lustrous, clathrate; leaves 70
100 cm long, loosely fasciculate, dimorphic (the
sterile with wider pinnae); petiole 25-37 cm long,
dark brown or blackish and scaly at base, dark brown
and densely puberulous with septate hairs into the
rachis; blade 45-60 x 15-20 cm, 1-pinnate, ovate-
lanceolate, scarcely narrowed at base, pinnatifid at
Figure 9. Diplazium panamense C. D. Adams.
apex; adaxial groove glabrescent; lateral pinnae 4-
10 x 1-2.8 cm, in 12-14 alternate or subopposite
pairs, 4.5-8 cm apart at base of rachis, with stalks
obsolete or 2-3 mm long on fertile leaves, oblong
to linear-lanceolate, subequal and truncate or sub-
cordate-hastate at base, entire or subentire, falcate
and abruptly acuminate and scarcely serrulate at
tip, subcoriaceous with shortly recurved margins
when dry, dark green, slightly discolorous; upper
pinnae sessile or shortly adnate, without axillary
buds; costae and veins abaxially with brown ap-
pressed hairs; veinlets branched from close to the
costae, 2—3(—4) forked, subparallel, slightly curved
close to the margin; sori 2 8 mm, irregularly di-
plazioid in both acroscopic and basiscopic branches;
indusia ca. 0.3 mm wide, dark brown, erose, ciliate,
perpendicular to the lamina at maturity.
Paratype. PANAMA. CHIRIQUI: humid forest between
Alto de las Palmas and top of Cerro de la Horqueta,
2,100-2,268 m, 18 Mar. 1911, Maxon 5520 (BM).
This species has affinity with I), patrnense but
the tip of the blade is pinnatifid rather than com¬
prising a more or less conform terminal pinna; the
lateral pinnae are more numerous; and the costae
and veins are appressed-pubescent abaxially.
298
Novon
Figure 10. Diplazium tutense C. D. Adams.
Diplazium tutense C. I). Adams, sp. nov. TYPE:
Panama. Veraguas: above Santa Fe beyond
Escuela Agricola Panamericana, 1.8 mi. be¬
yond fork in road on Pacific slope above rocky
ravine on side of Cerro l ute, 5 Apr. 1076,
Croat 34219 (holotype, MO). Figure 10.
Filix D. skutchii Lellinger affinis sed lamina axibusque
utrinque glabris vel subglabris, pinnulis basalibus minus
quam !A ad costulam partitis, laminae textura membra-
nacea viridique (non chartacea schistaceaque) differt.
Rhizome to 20 x 1 cm, erect; scales 10-15 x
2-4 mm, lanceolate, entire but soon lacerate and
ragged and eventually fenestrated and retiform, not
toothed, dark brown, very shiny, subclathrate with
elongated cells, mixed with much smaller scales;
leaves to 1 m or more long, compactly tufted; petiole
36 cm long, dark brown and covered with large and
minute, hairlike scales at base, soon stramineous
into rachis and costae, puberulous, glabrescent; blade
65 x 30 cm, 2-pinnate-pinnatifid, ovate or debate,
pinnatifid at apex to acuminate serrate tip; rachis
and costae with few somewhat irregular scales with
long hairlike tips, otherwise glabrous; adaxial groove
shortly puberulous, the wings interrupted but scarce¬
ly lobed at junctions; pinnae 6-17 x 1.5-4.5 cm,
ca. 15 pairs, stalked to 7 mm at subequal darker
base, the lower slightly shorter, the upper shortly
adnate at equally truncate base, linear- to oblong-
lanceolate, uniformly cut to within 2 3 mm of the
costa; pinnules at base of blade to ca. 2.5 x 1 cm,
free from one another except by a very narrow strip
of lamina tissue, rarely almost stalked and smaller
in proximal positions, to 12-14 pairs, oblong, patent
or antrorse, reduced toward apex of blade to ca. 5
x 5 mm, 14 to midvein or less, crenate-serrate
laterally, dentate at tip, glabrous on both surfaces,
membranaceous, green, discolorous; veinlets in larg¬
er segments 7-9 pairs, divergently 2-3-forked prox-
imally, simple distally, remote below sinuses; sori to
4 mm, to 6-7 pairs along acroscopic veinlet-branch-
es; indusia 0.5-0.7 mm wide, pale brown or hyaline,
translucent, entire or subentire, without cilia, fragile
and fragmenting at maturity.
This medium-sized fern resembles I). skutchii,
but the blade and axes are glabrous or nearly so on
both surfaces; the basal pinnules are lobed less than
one-third of the way to the costule; and the lamina
tissue is membranous and green rather than char-
taceous and grayish.
Acknowledgments. The author, who is an hon¬
orary associate of The Natural History Museum,
London, is grateful for the facilities provided and
the friendly cooperation of the botanical and library
staff. The curators of the following herbaria are also
thanked tor making their collections available for
study: BM, BR. CAS, F, K, MEXU, MO, P, UC.
Literature Cited
Calderon, S. & P. C. Standley. 1944. Lista preliminar
de plantas de El Salvador. 2nd Edition. San Salvador.
Gomez, L. D. 1975. Contribuciones a la Pteridologia
Costaricense VII. Pteridofitos de la Isla de Cocos.
Brenesia 6: 43.
Mickel, J. T. & J. M. Beitel. 1988. Pteridophyte Flora
of Oaxaca. Mem. New York Bot. Card. 46: 154-
156.
Seiler, R. 1980. Una guia taxonomica para helechos
de El Salvador. Ministerio de Educacion. San Sal¬
vador.
Smith, A. R. 1981. Pteridophytes. Pp. 1-370 in Flora
of Chiapas, Part 2. California Academy of Sciences,
San Francisco.
Standley, P. C. & S. Calderon. 1925. Lista preliminar
de las plantas de El Salvador. San Salvador.
Svenson, H. K. 1938. Pteridophyta of the Galapagos
and Cocos Islands. Bull. Torrey Bot. Club 65: 303-
333 .
Sarcoglottis caudata (Orchidaceae) and Veliozia kolbekii (Velloziaceae),
Two New Species from Minas Gerais, Brazil
Huy J. V. Alves
Botanical Institute, Czechoslovak Academy of Sciences, 25243 Pruhonice, near Praha,
Czechoslovakia
ABSTRACT. During a vegetation survey of the Sao
Jose mountain range in Minas Gerais, Brazil, the
species Sarcoglottis caudata (Orchidaceae) and
Vellozia kolbekii (Velloziaceae) were discovered;
they are here described as new to science.
Sarcoglottis caudata R. J. V. Alves, sp. nov.
TYPE: Brazil. M inas Gerais: Tiradentes, Serra
de Sao Jose D’El Rei, summits of quartzite
tabular mountains, 1,200 m, Oct. 1989, R. J.
V. Alves 959 (holotype, RB). Figure 1.
Planta S. rupestri Barb. Rodr. var. parviflorae Cogn.
aliquanto proxima sed apice labelli caudato, sepalo inter-
medio ad apicern inciso, petalis basi ad sepalum inter¬
medium connatis differt.
Geophyte, 6 8 cm tall, green herb with 1 to 3
flowers, leafless when flowering (September). Leaves
unknown. Bracts linearly lanceolate, 1 cm long, 3
mm wide at base, only at base of flowers. Flowers
erect, 2 cm long, externally sparsely hairy, green.
Lateral sepals 10 mm long, connate to about 14 ol
basal length, basally continuous with the ovary,
asymmetrically linear. Dorsal sepal 9 nun long, 3
mm wide, lanceolate, caudate with incise apex, ha-
sally adnate to petals; petals 6 mm long, 1 mm wide,
linearly asymmetric, apex rounded; labellum white,
base widely sagittate, apex long-caudate. Labellum
basally rhombic, apically caudate, 7 mm long, 3
mm wide, with undivided venation consisting of 3
main veins, 2 pairs ol thinner, lateral veins that
form a loop with the lamina (apically connected.
Fig. 1). Column ca. 2 mm long, sulcate. Anther cap
membranous. Bostellum rhombic, 1 mm long.
Habitat. Grows in white quartzitic sand deposits
on slopes and summits of tabular mountains, in areas
where the accumulated rainwater upwells from a
shallow layer of sand lying directly on the bedrock
or hardpan. The substrate is alternately wet (sum¬
mer) and dry (winter).
Distribution. Known only from the Sao Jose D EI
Rei range, Tiradentes, Minas Gerais, Brazil.
The unique labellum shape indicates that Sar¬
coglottis caudata forms a new subgroup in the
genus. It is similar to S. rupestris var. parviflora
in habit and habitat. Pabst & Dungs (1975, 1977)
recognized 29 species of Sarcoglottis K. B. Presl
from Brazil, of which ,S. aphylla (Ridley) Schltr.,
S. cogniauxiana (Barb. Rodr.) Schltr., S. neurop-
tcra (Reichb. f. & Warming) Schltr. and .S', rupestris
Barb. Rodr. are from Minas Gerais. Somewhat ear¬
lier, H oehne (1945) stated that the above species
are so closely allied that they should be joined into
one. According to the present data, ,S. caudata is
known only from the Sao Jose range, while the type
of S. rupestris var. parviflora was found in the
vicinity of Sao Joao D’El Rei.
More new species of Sarcoglottis can be expected
from the isolated campo rupestre ranges of Minas
Gerais. Most of the described species are endemic
to small ranges, and some are only known from type
collections.
Vellozia kolbekii R. J. V. Alves, sp. nov. TY PE:
Brazil. Minas Gerais: Tiradentes, Serra de Sao
Jose D* El Rei, 1,200 m, 22 Oct. 1989, R. J.
I . Alves 852 (holotype, SPF; phototype, NI1B).
Figures 2, 3.
Velloziae brachypodae Lyman B. Smith & Ayensu
proxima sed caudice multo longiore, tepalis longioribus,
ovario obscure trigonaliter que tereti, staminibus pau-
cioribus, foliis capsulaque glabris differt.
Heliophyllous phancrophyte. Gaudex sparingly
branched, usually 150 cm tall (rarely over 300 cm),
erect, 5-15 cm diam. including the golden-yellow,
persistent leaf sheaths. Leaves ca. 8 to 10 produced
in each season, ca. 12 live and 65 dry in each
rosette, 33 cm long (without the sheaths), base 14
mm wide, blades parchmentlike, narrowly triangu¬
lar, attenuate, glabrous except for the finely setose-
serrulate margins; venation of lamina consists of
median vein with 27 congruent secondary veins on
either side; scapes subterminal, 3-8 per rosette,
pedicel ca. 9 cm long, 3 mm diam., cylindric, curved
so that the flowers are naturally in horizontal posi¬
tion, reaching about !4 length of the leaves, glabrous
at the base, slightly verrucose under the ovary (i.e.,
subapieally). Ovary smooth, glabrous, in fresh flow¬
ers 7 mm diam., 12 mm long, cylindric to fusiform;
Novon 2: 299-301. 1992.
300
Novon
10 MM
Figure 1. Sarcoglottis caudata R. J. V. Alves. —A.
Dorsal sepal with attached petals. —B. Fused lateral
sepals. —C. Anther cap. —D. Column with rostellum.
— E. Labellum. —F. Hahit.
capsule terete, Bluntly trigonal with slightly sulcate
facets when green, apically constricted with a broad
crown and smooth when dry. Perigonium deep light
purple, anthers and stigma sulphury yellow, ovary
greenish to magenta, live parts covered with sticky
substance. Only flowering specimen seen. Epigynous
tube subcylindric, 4 mm high; tepals free except for
a slight basal connation (1 to 2 mm), violet-blue,
basally grading to white, 4 cm long, tepals of the
inner whorl obovate-lanceolate with slightly con¬
stricted apex, 3 main undivided and 8 lateral veins;
outer tepals broader, widely lanceolate, apical api-
cule less pronounced, 3 central and 5 lateral veins;
stamens in groups of 5 to 6, bases of filaments
connate in these groups, on base of tepals; stamens
in observed specimen 32, slightly less than Vi the
length of the tepals; anthers linear-cylindric, 1 mm
diam., 19 mm long; filaments 10 mm long, less than
l A diam. of anthers, white; style 24 mm long, 1 mm
diam., stigma 3-lobed, lobes suborbicular to reni-
form. Dorsiventral transverse section of lamina (data
from boiled herbarium specimen): no hairs observed.
Figure 2. Vellozia kolbekii R. J. V. Alves. —A. Tepal
of inner w'horl. —B. Tepal of outer whorl. —C. Insertion
of stamens. —D. Transversal section of ovary. —E.
Ovary with style. —F. Leaf apex with marginal setae.
— G. Hahit. —H. Transverse section of lamina.
adaxial surface smooth and slightly undulate; abaxial
surface furrowed to % of blade (Fig. 2H).
Habitat. On exposed quartzite outcrops and scree
with little or no soil, sometimes forming closed thick¬
ets.
Dedicated to Jiri Kolbek, who took part in the
field research.
Distribution. Presently known only from the Sao
Jose D’El Rei range near Tiradentes, Minas Gerais,
Brazil.
l ellozia kolbekii was confirmed as new by Ly¬
man B. Smith (pers. comm.). It differs from / .
Volume 2, Number 4
1992
Alves
Sarcoglottis caudata and
Vellozia kolbekii
301
Figure 3. Vellozia kolbekii R. J. V. Alves. Holotype,
detail of scapes and capsules.
brachypoda mainly by a longer caudex, longer te-
pals, a more angular ovary with the apex broadened
into a crown (Fig. 3), leaves with more linear setae
on margins, and deeper furrows on the ahaxial sur¬
face. It further differs from l . variabilis Martius
ex Schultes f. by a glabrous ovary, from I . tomeana
Lyman B. Smith & Ayensu by partly connivent
starninal filaments, a glabrous ovary and glabrous
leaves. The fact that V. kolbekii has a glabrous
ovary and that part of its scape is slightly vestite
may indicate a possible relationship with I . subsea-
bra Mikan (V. scabra Sprengel), listed under "Ex¬
cluded and Doubtful Taxa” by Smith & Ayensu
(1976); however, genus Vellozia Vand. needs to he
studied further before this can be proved. In the
diagnosis of V. tomeana , Smith & Ayensu (1976:
82) declared the leaves glabrous, and subsequently
discriminated large hair tufts on both surfaces.
Additional habitat observations. Both newly de¬
scribed species were discovered in the Sao Jose
mountain range north of Tiradentes, Minas Gerais,
Brazil. This range rises from 900 to 1,430 m, being
situated between 21°03 07'S and 44°06 13'\\ . It
consists of quartzite outcrops with irregularly eroded
surfaces, and sand deposits originated by erosion of
th is bedrock. Both substrates bear sparse stands of
herbaceous vegetation with an open shrub layer.
The local climate consists of cool, dry winters and
hot, rainy summers that culminate in January. Ab¬
solute temperatures range from 1.0° to 36.2°C and
mean values of 9.9° (July) to 25.9° (January). An¬
nual precipitation averages to 1,467 mm (8.6 mm
in July, 299.6 mm in January). Vellozia kolbekii
grows on quartzite outcrops, forming dense stands,
while Sarcoglottis caudata is restricted to the sand
deposits on summits and in lower-situated valleys,
where it thrives in moist spots that are flooded for
many days after each rainstorm.
Acknowledgments. I thank Lyman B. Smith, Na-
nuza L. de Menezes, Elsie F. Guimaraes, and Pedro
Fernandes for opinions and cooperation, Jiri Sojak
and Josef Holub for aid with the Latin diagnoses,
and the Brazilian Council for the Development of
Research (CNPq) for research support.
Literature Cited
Hoehne F. C. 1945. Flora Brasilica, (XII)Il: 311 334,
tab. 169-181, F. Lanzara Publishers, Sao Paulo.
Pabst, G. F. J. & F. Dungs. 1975, 1977. Orchidaceae
Brasilienses. Brucke Verlag, Hildesheim.
Smith, L. B. & L. S. Ayensu. 1976. A revision of the
American Velloziaceae. Smithsonian Contr. Bot. 30:
1-172.
Two New Species of Stigmaphyllon (Malpighiaceae) from Peru
('h ristiane Anderson
University of Michigan Herbarium, North University Building, Ann Arbor, Michigan
48109-1057, U.S.A.
ABSTRACT. Two new species of the neotropical wing-
fruited genus Stigmaphyllon (Malpighiaceae) are
described from Peru, S. argenteum from the eastern
lowlands and S. cuzcanum from the uplands of
Cuzco.
Stigmaphyllon, one ol the wing-fruited genera
of the Malpighiaceae, comprises nearly 100 species,
which occur from southern Mexico and the Carib¬
bean to northern Argentina, except in Chile. Most
species have large, cordate, long-petioled leaves and
umbels or pseudoracemes of yellow flowers disposed
in dichasially branched inflorescences. The androe-
cium of 10 stamens is most often heterogeneous;
the stamens opposite the lateral sepals usually bear
modified anthers consisting of an enlarged connec¬
tive bearing 0-2 reduced locules. Each of the three
styles is commonly ornamented with an apical ap¬
pendage, the foliole, for which the genus is named.
Typically, the samara bears a large, flared dorsal
wing, and the nut is often ornamented with lateral
winglets and/or spurs and crests.
Two new species are here described so that the
names will be available for inclusion in the forth¬
coming Catalogue oj the Flowering Plants and
Gymnosperms oj Peru (Brako & Zarucchi, in prep.).
With the addition of these novelties, 18 species of
Stigmaphyllon are now reported from Peru.
Stigmaphyllon argenteum C. Anderson, sp. nov.
TYPE: Peru. Huanuco: Prov. Pachitea, Dtto.
llonoria, Bosque National de Iparia, a lo largo
del Rio Pachitea cerca del campamento Miel
de Abejas, 1 km arriba del pueblo Tournavista
o unos 20 km arriba de la confluencia con el
Rio Ucayali. 300-400 m, 30 May 1967,
Schunke l . 2018 (holotype, NY; isotypes, COL,
E, G, IJS). Figure 1.
Liana. Laminae 2.5 15.3 cm longae, 5.7 14 cm latae,
triangulares, ovatae, ellipticae vel suborbiculares vel in-
terdurn 3-5-lobatae, supra glabrae vel interdum sparsim
sericeae, subtus sericeae, margine sparsim glanduloso.
Inflorescentia dichasialis constata ex umbellis, floribus in
quaque umbella ca. 15 30. Pedunculi 3-7.5 mm longi;
pedicelli 4 8.5 mm longi. Petala lateralia limbo orbiculari
vel late obovato, margine eroso; petalum posticum limbo
elliptico vel late obovato, margine eroso vel fimbriato-
denticulato. Stamina heteromorpha, antheris glabris; an-
therae sepalis antico-lateralibus oppositae 1-2 loculis re-
ductis instructae, antherae sepalis postico-lateralibus op¬
positae 1 loculo reducto instructae. Stylus anticus ca. 2.2
mm longus, glaber, utroque foliolo ca. 1.4 mm longo, ca.
1.2 mm lato, subquadrato; styli postici 2.6-3 mm longi,
glabri, lyrati, foliolo ca. 1.4 -2 mm longo latoque, subquad¬
rato. Samara ala dorsali ca. 4.5 cm longa, 1.4-11.7 cm
lata; alulae laterales absentes; nux 4-5.5 mm alta, 3.5-
4.5 mm diametro.
Vine to 14 m. Laminas 2.5-15.3 cm long, 5.7-
14 cm wide, triangular, ovate, or elliptical to su-
borbicular, or sometimes 3-5-lobed, apex acumi¬
nate, base truncate to cordate, sometimes sparsely
sericeous but usually glabrous above, sericeous be¬
low (trabecula 0.2-0.5 mm long, straight, sessile),
margin shallowly crenate to subentire and with ir¬
regularly spaced sessile glands (0.5-0.6 mm diam.)
in the sinuses and with filiform glands (up to 1.5
mm long), with a pair of prominent but sessile glands
at the apex of the petiole, each gland 1.5-3.5 nun
diam.; petioles 2-10+ cm long, sericeous; stipules
0.7-1.2 mm long and wide, triangular, eglandular.
Flowers ca. 15-30 per umbel, these borne in di-
chasia or compound dichasia. Peduncles 3-7.5 mm
long, pedicels 4-8.5 mm long; peduncles 0.6 1.2
times as long as the pedicels. Bracts 0.9-1.3 mrn
long, 0.6-1 mm wide, narrowly triangular; brac-
teoles 0.7-1.2 mm long, 0.6 1 mm wide, triangular,
eglandular. Sepals 1.8 2.3 mm long, 1.5-2 mm
wide, glands 1.6-2.3 mm long, 0.6-1.2 mm wide.
All petals glabrous, yellow; lateral petals with the
limbs orbicular or broadly obovate, margin erose;
anterior-lateral petals: claw 1.8-2.2 mm long, limb
ca. 7 mm long and wide; posterior-lateral petals:
claw 0.5-1 mm long, limb 6-6.7 mm long, 4.5-6
mm wide; posterior petal: claw 2.5-2.8 mm long,
apex strongly indented, limb 5-5.6 mm long, 3.5-
4.8 mm wide, elliptical or broadly obovate, margin
erose to fimbriate-denticulate, teeth/fimbriae up to
0.5 mm long. Stamens unequal, those opposite the
posterior styles the largest; anthers all loculate, gla¬
brous, those of stamens opposite the anterior-lateral
sepals with 1 or 2 locules, those of stamens opposite
the posterior-lateral sepals with only 1 locule. An¬
terior style ca. 2.2 mm long, shorter than the pos¬
terior two, glabrous; each foliole ca. 1.4 mm long,
Novon 2: 302-305. 1992.
Volume 2, Number 4
1992
Anderson
Stigmaphyllon argenteum and
S. cuzcanum
303
Figure 1. Stigmaphyllon argenteum C. Anderson. —a. Portion of branch with large leaf. —b. Detail of abaxial
surface of lamina. —c. Small leaf. —d. Flowering branch. —e. Posterior petal. —f. Androecium; second stamen
from left opposes posterior petal. —g. Gynoecium, anterior style to the right. —h. Samara. —i. Two views of an
embryo. Scale for a, c, d, h, bar = 1 cm; for b, bar = 0.5 mm; for e-g, bar = 1 mm. Based on: a, b, d-g, Schunke
V. 2018; c, h, i, Croat 19640.
ca. 1.2 mm wide, subsquare. Posterior styles 2.6-
3 mm long, glabrous, lyrate; foliole ca. 1.4-2 mm
long and wide, subsquare. Dorsal wing of samara
ca. 4.5 cm long, 1.4-1.7 cm wide, lateral winglets
absent, nut only prominently ribbed; nut 4-5.5 mm
high, 3.5-4.5 mm diam., areole 3-3.5 mm long,
2.5-2.8 mm wide, concave, carpophore up to 1.8
mm long. Embryo 5.8-7.3 mm long, ca. 2 times
as long as wide, ovoid, outer cotyledon 6.1-8.3 mm
long, 2.6-3.9 mm wide, the distal Z b folded over the
inner cotyledon, inner cotyledon 4-6.6 mm long,
2-3.6 mm wide, straight or the tip folded back on
itself.
Phenology. Collected in flower from April through
July, in fruit in May and from July through Sep¬
tember.
Distribution. Lowlands of eastern Peru; in forests
and thickets and at roadsides; 135-670 m.
Paratypes. Peru. HUANUCO: Prov. Pachitea, region of
Pucallpa, ca. 26 km S to 24 km SSE of Puerto Inca, N
of Rio Yuyapichis, 09°34-37'S, 74°53 56'W, If allnofer
11-31588 ( MICH); vicinity of Tingo Maria, 3-5 km from
Huanuco-Tingo Maria rd. on Monzon rd., Mathias A'
Taylor 3647 (F, UCLA). JUNIN: Puerto Bermudez, Killip
dc Smith 26630 (F, NY, US); Prov. Satipo, E bank of
Rio Ene at mouth of Rio Quipachiari, Madison 10427-
70 (F). LORETO: Quebrada Shanuce above Yurimaguas,
Croat 17999 (MICH); Isla de Ushpa-cano near mouth of
Rio Itaya, Croat 19640 (MICH); Ucayali, Bosque Na-
cional Alexander von Humboldt, between Km 90-130 of
Pucallpa Tingo Maria rd., 08°48'S, 75°20'W, Gentry et
a!. 41413 (MO); wooded banks on lower Rio Huallaga,
Killip & Smith 29004 (F, GH, NY); Prov. Maynas,
vicinity of Iquitos, Rio Momon, quebrada Momoncillo,
McKenna et al. DMK-91 (AMAZ, F, MICH, MO). PASCO:
Oxapampa, ca. 5 km up Rio Iscozacin from village of
Iscozacin, 10°12'S, 75°13'W, Knapp & Slaver 7802A
(MICH); Palcazu Valley, Rio San Jose in the Rio Chu-
churras drainage, 10°09'S, 75°20'W, D. Smith 4002
(MICH). SAN MARTIN: between Tocache Nuevo and Juan-
jui, 18.7 km S of Rio Pulcache, 07°55'S, 76°40'W, Croat
58052 (MICH); vicinity of Aguaytia, Boqueron de Padre
Abad, Mathias & Taylor 3591, 6092 (F, UCLA); Prov.
Mariscal Caceres, Dtto. Tocache Nuevo, quebrada de
Santiago, al E de Puerto Pizana, Schunke V. 6530 (GH,
MO); Prov. Mariscal Caceres, Dtto. Tocache Nuevo, que¬
brada de Cachiyaca, afluente de la quebrada de Huaquista,
al E de Puerto Pizana, Schunke V. 8528 (F, MICH, MO).
Stigmaphyllon argenteum is named for the sil¬
very, sericeous pubescence on the abaxial leaf sur-
304
Novon
Figure 2. Stigmaphyllon cuzcanum C. Anderson. —a. Large leaf. —b. Detail of abaxial surface of lamina and
individual hairs. —c. Umbel. —d. Posterior petal. —e. Androecium; second stamen from left opposes posterior petal.
— f. Gynoecium, anterior style to the right. —g. Samara. —h. Embryo. Scale for a, c, g, bar = 1 cm; for b (detail)
bar = 0.5 mm, b (hairs) bar = 0.3 mm; for d, h, bar = 2 mm; for e, f, bar = 1 mm. Based on: a, b, d. If est 6466;
c, e, f, Stafford 9; g, Tulin 1310; h, Nunez I . dv Motocanchi 3751.
faces. It is characterized by its small petals, the
limbs only up to 7 mm in diameter, the uniloculate
anthers of stamens opposite the posterior-lateral se¬
pals, and its samaras, which lack lateral wings. Col¬
lections ol this species have been assigned most
commonly to the widespread and variable ,S. sinu-
atum (IX L) Adr. Juss., though labeled with one of
its many synonyms (S. fulgens, S. hypoleucum, S.
rnartianum , 5. splendens; see Anderson, in press).
In S. sinuatum , the flowers are aggregated into
pseudoracemes instead of umbels, the petals are up
to 14 mm in diameter, the stamens opposite the
posterior-lateral sepals have unmodified biloculate
anthers, the styles are commonly pubescent (gla¬
brous in ,S. argenteum ), and the nut of the samara
usually hears lateral winglets. Stigmaphyllon ar¬
genteum is rarely mistaken for one of the three
other sympatric species with abaxially sericeous
leaves, .S', maynense Huber, ,S. puberum (Rich.)
Adr. Juss., and S. eardiophyllum Adr. Juss. Stig¬
maphyllon maynense and S. puberum differ from
most species of the genus in that the anterior style
and its opposing stamen are much larger than the
posterior styles and their opposing stamens. Idle nut
of the samara of .S', maynense bears 3-4 lateral
winglets per side, whereas the unique samara of S.
puberum is distinguished by an erect dorsal wing,
tapered from the base, and lacks a carpophore.
Stigmaphyllon eardiophyllum is easily separated
by its usually glabrate to glabrous leaves, though
sometimes very sparsely sericeous below, and its
abundantly pubescent anthers. The remaining sym¬
patric species differ in that the leaves are beset with
T-shaped hairs below.
Stigmaphyllon cuzcanum C. Anderson, sp. nov.
TYPE: Peru. Cuzco: below Machu Picchu,
2,100 m, West 6166 (holotype, MO). Figure 2.
Liana. Laminae 7-19 cm longae, 5.5-12.7 cm latae,
ovatae, supra glabrae, subtus tomentosae, margine eglan-
duloso vel sparsim glanduloso. Inflorescentia solitaria vel
dichasialis constata ex umbellis, floribus in quaque umbella
ca. 10-35. IVdunculi 3.5-15.5 nun longi; pedicelli 7-
13.5 mm longi. Petala limbo orbiculari, margine fiinbriato
vel fimbriato-denticulato. Stamina heteromorpha, antheris
glabris vel raro pubescentibus; antherae sepalis lateralibus
oppositae 2 loculis reductis instructae. Stylus anticus 4.2-
5.1 mm longus, glaber, utroque foliolo ca. 1.4 1.8 mm
longo, 1-1.5 mm lato, elliptico; styli postici 5-6 mm
Volume 2, Number 4
1992
Anderson
Stigmaphyllon argenteum and
S. cuzcanum
305
longi, glabri vel basi sparsim pubescentes, lyrati, foliolo
2- 2.5 mm longo, ca. 1.8 mm lato, subrectangulari. Sam¬
ara ala dorsali 3.8-4.5 cm longa, ca. 2 cm lata; nux ca.
10 mm alta, ca. 5.5 mm diametro, alulis lateralibus in-
structa.
Vine. Laminas 7 19 cm long, 5.5 12.7 cm wide,
ovate, apex emarginate-mucronate, base cordate to
deeply so in larger leaves and to truncate in smaller
ones, glabrous above, tomentose below (trabecula
0.9-1.5 mm long, crisped and curled, stalk 0.1-
0.3 mm long), margin eglandular or with irregularly
spaced sessile glands (0.3-0.4 mm diam.), with a
pair of prominent but sessile glands at the apex of
the petiole, each gland 1.8-2.5 mm diam.; petioles
3- 7.3 cm long, densely sericeous to glabrous in
older leaves; stipules 1-1.5 mm long, 0.8 1.3 mm
wide, triangular, eglandular. Flowers ca. 10-35 per
umbel, these borne solitary or in dichasia or in small
thyrses. Peduncles 3.5-15.5 mm long, pedicels 7
13.5 mm long; peduncles 0.6-1.4 times as long as
the pedicels. Bracts 1.2-2.5 mm long, 0.8-1.3 mm
wide, triangular; bracteoles 1.1-2.3 mm long, 0.7-
1.4 mm wide, narrowly triangular, eglandular or
each bracteole with a pair of glands (each 0.4-0.5
mm diam.). Sepals 2.5-3.5 mm long, 2.5-3.2 mm
wide, glands 2 2.5 mm long, 1-1.3 mm wide. All
petals glabrous, yellow, with the limbs orbicular,
margin fimbriate or denticulate-fimbriate, fimbriae/
teeth up to 0.5 mm long; anterior-lateral petals:
claw ca. 3-4 mm long, limb ca. 16-18 mm long
and wide; posterior-lateral petals: claw ca. 2.5-3
mm long, limb ca. 15 mm long and wide; posterior
petal: claw 3.5-4.5 mm long, apex not or only very
slightly indented, limb ca. 14 mm long and wide,
margin at the base sometimes with a stalked gland
(ca. 0.4 mm long). Stamens unequal, those opposite
the posterior styles with the longest filaments, an¬
thers of those opposite the anterior-lateral sepals
with the connective enlarged and the locules re¬
duced; anthers all loculate, glabrous or rarely sparse¬
ly pubescent. Anterior style 4.2-5.1 mm long, short¬
er than the posterior two, glabrous; each foliole 1.4
1.8 mm long, 1- 1.5 mm wide, elliptical. Posterior
styles 5-6 mm long, glabrous or with a few scattered
hairs in the basal l A, lyrate; foliole 2-2.5 mm long,
ca. 1.8 mm wide, subrectangular. Dorsal wing of
samara 3.8- 4.5 cm long, ca. 2 cm wide, nut bearing
1-2 grossly dentate rectangular lateral winglets per
side, these 3.5-8 mm long, up to 2.5 mm wide, and
also spurs; nut ca. 10 mm high, ca. 5.5 mm diam.,
areole ca. 4 mm long, ca. 3.5 mm wide, concave,
carpophore up to 5 mm long. Embryo 8.5 mm long,
ca. 2 times as long as wide, ovoid, outer cotyledon
ca. 13 mm long, ca. 4.2 mm wide, the distal l A
folded over the inner cotyledon, inner cotyledon ca.
7.3 mm long, ca. 3.3 mm wide, straight.
Phenology. Collected in flower in February, May,
June, and August, in fruit in January, February,
June, and August.
Distribution. Peru, Depto. Cuzco, Prov. Fa Con¬
vencion; in brush forests and clearings; 1,800 2,700
m.
Paratypes. Peru, CUZCO: San Miguel, Urubamba Val¬
ley, Cook & Gilbert 939 (NY, US); Prov. La Convencion,
139 km de Cuzco en Quillomayo, entre Santa Teresa y
Chaullay, 13°08'S, 72°36'W, Nunez V. & Motocanchi
8751 (MICH); Machu Picchu, Stafford 9 (K); Machu
Picchu, Urubamba Valley, Tutin 1310, 7328(BM); Prov.
La Convencion, Machu Picchu, Vargas C. 814 (F); Prov.
La Convencion, Weberbauer 4989 (G).
Stigmaphyllon cuzcanum is notable for its large,
abaxially tomentose leaves and its large flowers,
borne in umbels that are aggregated into thyrses.
The petals are among the largest in the genus; the
limb ol the anterior-lateral petal is 16-18 mm in
diameter. The stamens opposing the posterior-lateral
sepals are not modified, as in most other species.
The nut of the samara bears 1-2 grossly dentate
rectangular lateral winglets per side. This species is
readily separated from the two other species of Stig¬
maphyllon reported from Cuzco. In .S’, cardiophyl-
lum, the leaves are abaxially glabrate to glabrous
or sometimes sparsely sericeous, the flowers are very
small (the limbs of the petals only up to 6.5 mm in
diameter), and the samaras lack lateral winglets.
Stigmaphyllon strigosum Adr. Juss. differs in its
smaller long-fimbriate petals streaked with red (limbs
up to 14 mm in diameter, fimbriae up to 0.9 mm
long), modified anthers of stamens opposing the pos¬
terior-lateral sepals (the connective enlarged and the
locules reduced), and samaras with 3 to 4 lateral
winglets per side.
Acknowledgments. I thank William R. Anderson
for his comments. Karin Douthit drew the illustra¬
tions. I am grateful to the curators of the following
herbaria for making their collections of Stigma¬
phyllon available to me: AMAZ, BM, COL, F, G,
CH, K, MICH, MO, NY, UCLA, US.
Literature Cited
Anderson, C. The identities of the sericeous-leaved spe¬
cies of Stigmaphyllon (Malpighiaceae) in the Ama¬
zonian region. Contr. Univ. Michigan Herb. 19 (in
press).
Change in Status of Pinas cembroides Subsp. orizabensis (Pinaceae)
from Central Mexico
I). K. Hailey
University of Colorado Museum, Boulder, Colorado 80302, U.S.A.
Frank G. Hawksworth
USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins,
Colorado 80526, U.S.A.
ABSTRACT. The Orizaba pinyon (Pinas cembroides
subsp. orizabensis) is raised to specific status. Pinas
orizabensis differs from P. cembroides in many
characters, including needle and hark morphology
and terpene chemistry.
Orizaba pinyon, Pinas cembroides Zucc. subsp.
orizabensis I). K. Hailey, is a local taxon of east
central Mexico described by Bailey (1983). The
taxon is here raised to specific rank because of
marked differences in needle and bark morphology
and terpene chemistry between it and P. cem¬
broides.
Pinus orizabensis (1). K. Bailey) I). K. Bailey &
Hawksworth, comb, et stat. nov. Basionym:
Pinus cembroides Zucc. subsp. orizabensis D.
k. Bailey, Phytologia 54: 89. 1983. TYPE:
Mexico. Puebla: Mpio. Soltepec, ca. 10 km
SW of San Salvador el Seco on Highway 140,
19°04'N, 97°42'W, 2,370 m, 1983, D. K.
Hailey 83-01 (holotype, MEXU; isotypes,
ARIZ, CHAPA, COLO, E, ENCB, 1NIF, k,
MO, NY, RM, TEX, UC, US, UTC).
Pinas cembroides , as interpreted by Bailey (1983)
and Bailey & Hawksworth (1988), was considered
to be comprised of three allopatric subspecies:
1. Subspecies cembroides , the most widespread
taxon, ranging from northern Chihuahua and west
Texas throughout the Sierra Madre Occidental and
Sierra Madre Oriental to central Mexico (Hidalgo
and northern Veracruz).
2. Subspecies lagunae (Robert-Passini) I). K. Bai¬
ley, a local endemic restricted to the Sierra Laguna
in southern Baja California.
3. Subspecies orizabensis I), k. Bailey, a local
endemic in Puebla, Tlaxcala, and Veracruz.
Since that time there has been a trend to accept
subspecies lagunae as a distinct species, Pinas la¬
gunae M.-F. Passini (1987), e.g., in the recent
compendium of the pines of Mexico and Central
America (Perry, 1991). However, because the dif¬
ferences between subspecies cembroides and ori¬
zabensis are greater than those between subspecies
cembroides and lagunae , we suggest raising sub¬
species orizabensis to species rank as well.
Pinus orizabensis and P. cembroides are not
sympatric, and the gap between them is about 140
km (Bailey, 1983). The two pines differ in several
morphological, chemical, and ecological features
(Bailey, 1983), some of which are listed in Table 1.
Acknowledgment. We thank Teobaldo Eguiluz
Piedra, Director, Centro de Cenetica Forestal, Chap-
ingo, Mexico, for encouraging us to make this new
combination.
Table 1. Comparisons of some characters of Pinus orizabensis and P. cembroides.
Character
P. orizabensis
P. cembroides
Needle number per fascicle
Range 2-5; ca. 75-95% in 3’s,
5-20% in 4’s
Range 2-4; ca. 60% in 2’s and 40%
in 3’s
Needle color
Dorsal surface bluish-green; ventral
surface markedly glaucous
Dorsal surface yellow-green; ventral
surface slightly glaucous
Bark of mature trees
Thin, little transverse Assuring, inner
bark conspicuous, orange
Thick, strong transverse Assuring, in¬
ner bark inconspicuous, yellow
Monoterpenes
3-carene usually less than 1%
3-carene usually ca. 10%
Main elevational range
2,300-2,700 m
1,700-2,400 m
Novon 2: 306-307. 1992.
Volume 2, Number 4
1992
Bailey & Hawksworth
Pinus cembroides subsp. orizabensis
307
Literature Cited
Bailey, D. K. 1983. A new allopatric segregate from
and a new combination in Pinus cembroides Zucc.
at its southern limits. Phytologia 54: 89-100.
-& F. G. Hawksworth. 1988. Phytogeography
and taxonomy of the pinyon pines, Pinus subsection
Cembroides. Pp. 41-64 in II Simposio Nacional
sobre Pinos Pinoneros, Centre d’Etudes Mexicaines
et Centroamericaines, Mexico City, Mexico.
Passini, M.-F. 1987. The endemic pinyon of Lower
California. Pinus Ingunae M.-F. Passini. Phytologia
63: 337-338.
Perry, J. 1991. The Pines of Mexico and Central Amer¬
ica. Timber Press, Portland, Oregon.
Plat an them pallida (Orchidaceae), a New Species of Fringed Orchis
from Long Island, New York, U.S.A.
Paul Martin Hr own
15 Dresden Street, Jamaica Plain, Massachusetts 02130-4407, II.S.A.
ABSTRACT. Reexamination of the species and hy¬
brids of the simple-lipped group of the genus Pla-
tanthera sect. Rlephariglottis shows that several
colonies found on eastern Long Island constitute a
new species. Platanthera pallida is described based
upon its recurved lip, reflexed lateral sepals, pale
cream coloration, and unusual habitat—dry, inter-
dunal hollows. It was previously referred to as P.
cristata because of its superficial resemblance to
that species and lack of other similar fringed orchises
in the northeastern United States.
Platanthera pallida P. M. Brown, sp. nov. TY PE:
United States. New York: Long Island, Suffolk
County, Montauk, C. S. Bryan, 10 Aug. 1948
(holotype, AMES 65176). Figure 1.
Platantherae cristatae (Michaux) Lindley similis, sed
ab ea perianthio in alabastro perpallide aurantiaco post
anthesin eburneo, labello recurvato ligulato fimbriato, pe-
talis obovatis ad apicem fimbriatis, fimbriis plerumque
furcatis, calcari obtuso 5 6 mm longo, sepalo medio in-
tegro, sepalis lateralis valde reflexis differt.
Plants (20 )29 65( 84) cm tall, glabrous and
distinctly glaucous; lower leaves 2 3, sheathed,
strongly keeled, conduplicate, to 25(-30) cm long
and 3(-5) cm wide when flattened; upper leaves
reduced to 3 5 linear bracts below the inflorescence.
Inflorescence racemose, (18-)24 80(-1 12)-flow-
ered, (5-) 10-20(-27) cm long, 2.5-4 cm diam.,
densely flowered except in very tall individuals; lower
floral bracts usually equal to or often exceeding the
pedicellate ovary and decreasing in length upward,
about 2.2 cm long. Perianth very pale orange-
yellow in bud opening to pale cream, petals and lip
often aging to a deeper creamy yellow or fading to
a dull white; lateral sepals 3 mm long, 3 mm wide,
reflexed, widest at the middle and tapering to a
rounded apex and truncated base; dorsal sepal 3
mm long, 3 mm wide, concave, entire and arching
forward to partially enclose the fringed petals; petals
obovate, 3.25 mm long, 2 mm wide, fringed at the
tip, the fimbriae usually forked and clearly visible
to the sides of the dorsal sepal; lip ligulate, 3 mm
long, 2 mm wide (exclusive of fringe); recurved or,
rarely, descending, the margin with forked fimbriae
up to 3 mm long, the fimbriae near the base per¬
pendicular to the lip; spur 5-6 mm long, tubular.
strongly curved, either obtuse or bulbous at the tip;
nectary orifice T-shaped; column with the viscidia
2.5-3 mm apart; fruit a capsule, 1-1.6 cm long.
Illustration. Photograph in Kickett (1966: plate
20) as llabenaria cristata.
Distribution and ecology■ Currently known only
from the Town of East Hampton, Suffolk County,
Long Island, New York. Plants growing in dry, in-
terdunal hollows and dune slopes (but never swales);
usually with Pin us rigida, Quercus velutina, C.la-
donia sp., Smilax glauca, S. rotundifolia. Toxi¬
codendron radicans, Hudsonia ericoides , Dan-
thonia sp., Agrostis sp., Minuartia caroliniana,
Arctostaphylos uva-ursi, Vaccinium corytnbosurn,
and Cypripedium acaule.
Etymology. The term pallida, pale, is chosen
for its appropriate description in comparison to P.
cristata (Michaux) Lindley and other brightly col¬
ored, related species. In the past, these plants have
commonly been called the “pale cristata" or "pale
fringed orchis.” The latter is recommended for a
common name.
Paratypes. United States, new york: Long Island,
Suffolk County, Town of East Hampton-East Hampton,
23 Sep. 1928, H. Latham 5995 (NYS); East Hampton,
16 Aug. 1929, R. Latham 66 34 (NYS), G. E. Lotowycz,
J. Hoar, 4 Aug. 1985(OBPF 9782); Montauk, 11 Aug.
1928, R. Latham 5782 (NYS), 12 Aug. 1928, R. La¬
tham 5782 (NYS); Montauk Point, C. S. Bryan, Aug.
1948 (AMES 80811); Montauk, C. K. Brooks et al., 1
Aug. 1951 (AMES 66346), G. E. Lotowycz, 18 Aug.
1978 (OBPF 8985), 15 Aug. 1979, C. J. Sheviak et al.
1015 (NYS); Napeague, R. Latham, 30 July 1928 (NYS),
11 Aug. 1928, R. Latham 5781 (NYS), R Latham , 16
Aug. 1929 (NYS), 16 Aug. 1929, R. Latham 7084
(NYS), 18 Aug. 1929, R Latham 7021 (NYS), G. E.
Lotowycz, 18 Aug. 1975 (OBPF 8111), 15 Aug. 1979,
C. J. Sheviak et al. 1006 (NYS), 17 Aug. 1984, R. E.
Zaremba 1503 (NYS), G. E. Lotowycz, 4 Aug. 1985
(OBPF).
Platanthera pallida is distinguished from other
closely related taxa by the following key.
Key to the Small-flowered (Lip Less Than 9 mm Long),
Fringed Species of Pi.ataxthfr i sect. Bt.rruARH.i.oms
of North America
la. Spur less than 10 mm long; shorter than ovary.
2a. Lip recurved, lateral sepals reflexed, dorsal
sepal entire, spur 5-6 mm long, obtuse;
Novon 2: 308-31 I. 1992.
Volume 2, Number 4
1992
Brown
Platanthera pallida
309
Figure 1. Platanthera pallida P. M. Brown. —A. Habit —B. Perianth, front/side view. —C. Petal.
— E. Column, side view. —F. Dorsal sepal. —G. Lateral sepal.
5 mm
i
D. Lip.
perianth cream-colored .
. Platanthera pallida P. M. Brown
2b. Lip projecting forward, lateral sepals por-
rect, dorsal sepal emarginate, spur 7-8
mm long, acute; perianth orange to yellow
. Platanthera cristata (Michaux) Lindley
lb. Spur greater than 10 mm long; exceeding ova¬
ry; perianth pale orange, yellow, cream or near¬
ly white.
3a. Spur 10-15 mm long, exceeding ova¬
ry ... Platanthera x canbyi (Ames) Luer
3b. Spur greater than 15 mm long; great¬
ly exceeding the ovary; raceme usu¬
ally 3 cm or more in diameter ....
. Platanthera x bicolor (Raf.) Luer
The simple-lipped, fringed species of Platanthera
constitute a significant part of section Hlephang-
lottis, and are some of the showiest members of the
Orchidaceae in eastern North America. Of the five
species that comprise this group, three, P. ble-
phariglottis (Willd.) Lindley, P. ciliaris (L.) Find¬
ley, and P. cristata, are widespread throughout the
East (Luer, 1975), while two, P. chapmanii (Small)
310
Novon
Luer emend. Folsom (Folsom, 1984) and P. inte-
grilabia (Correll) Luer (Zettler & Fairey, 1990),
are of restricted and local distribution. Platanthera
hiephariglottis includes variety blephariglottis, va¬
riety conspicua (Nash) Luer, and forma holopetala
(Lindlev) P. M. Brown (Brown, 1988). Three pu¬
tative hybrids occur, P. x bicolor (Hat.) Luer (P.
blephariglottis var. conspicua x P, ciliaris), P.
x canbyi (Ames) Luer ( P. blephariglottis var. con¬
spicua x P. cristata) (Luer, 1975), and P. Xchan-
nellii Folsom (P. ciliaris x P. cristata ) (Folsom,
1981). Several other hybrids have been reported,
but none have received binomials. The description
of Platanthera pallida (Fig. 1) adds a new species
to this list.
Crowing among Pinus rigida in dry, interdunal
hollows on eastern Long Island, New York, is a
Platanthera with small flowers, short spur, and a
superficial resemblance to P. cristata. The most
obvious difference is the uniform pale cream color
ol the small flowers. Unlike typical P. cristata in
the north, the plants are locally abundant.
Detailed examination and observations of P. cris¬
tata throughout its range, including all known pop¬
ulations from Long Island, have been made to see
d they reveal morphological differences. Measure¬
ments were taken of the critical characters, i.e.,
petals, sepals, lip, spur, cilia, and column, of 327
herbarium specimens and 128 living specimens, in¬
cluding 78 plants growing on Long Island. Numerous
photographs and drawings, published and unpub¬
lished, were also reviewed. Special attention was
given to those designated as “light yellow or pale"
in coloration. With the exception of the plants in
question, all specimens and living material examined
fell well within the criteria for typical P. cristata
(Correll, 1950; Luer, 1975).
Sheviak suspected that the pale plants from east¬
ern Long Island were polyploid and of hybrid origin.
He collected material, did a chromosome count, and
determined that they are diploid with a count of 2 n
= 42 (C. Sheviak 1606, NYS).
Herbarium specimens and living plants of the
northern hybrids, P. x canbyi, P. x bicolor, and P.
ciliaris x P. blephariglottis var. blephariglottis
(reported from Michigan), were also examined to
see il they might be similar to the Long Island plants.
They were found to be distinct in all respects.
Platanthera pallida shows insufficient features
to assume its parentage is the same as P. X canbyi.
Although the small flower size and pale coloration
can be found in many plants of P. x canbyi, the
longer spur characteristic ol that hybrid is lacking
in P. pallida. The reflexed sepals and recurved lip,
which are critical features of P. pallida, are also
present in P. blephariglottis var. blephariglottis.
Comparison with plants from Michigan that ap¬
peared to he the cross between P. ciliaris and P.
blephariglottis var. blephariglottis showed no sim¬
ilarities; the putative hybrids are much larger than
P. pallida and the spur much longer, as it is in
both parents.
One of the remarkable aspects of P. pallida is
the uniformity of its floral morphology. All the crit¬
ical floral characters, i.e., perianth dimensions, col¬
or, positioning of floral parts, have little, if any,
variation throughout all populations.
Platanthera pallida occurs as three populations
in two sites in the Town of East Hampton, eastern
Long Island. The plants appear to have been first
discovered by Roy Latham in 1926 (Latham, 1940).
By 1948, and in subsequent years, the stations had
been visited by several botanists and orchid enthu¬
siasts (Lamont et al., 1988).
Latham’s initial site near Montauk supports two
distinct current populations. They are separated by
nearly 14 mile of duneland. In each of these pop¬
ulations the plants are widespread and somewhat
scattered, hut retain their habitat preference. Ad¬
jacent to the areas that support P. pallida are
numerous swales and hogs. Typical P. cristata, if
present, would he found in these wetter areas, as it
is in the Pinelands of New Jersey, a region of similar
topography. Careful searches have revealed no oilier
species of Platanthera in either the immediate area
or for several miles around.
The other population is located west of Napeague
Harbor. Several specimens collected by Latham
(NYS) in 1928 1929 and simply labeled “Na¬
peague” may he from this site. It was not until 1975
that G. E. Lotowycz found the current site and
collected her first specimen—as P. cristata —from
there. In contrast to the Montauk site, where the
plants are widely scattered, here P. pallida is con¬
centrated in a much smaller area and in larger
numbers. Again, there are adjacent swales and small
hogs, hut no other Platanthera species to he found.
Companion plants are essentially the same in both
locales. Platanthera pallida is consistently ob¬
served to he restricted to the oldest, most stable
Pmus rigida stands within the dunes.
Acknowledgments. I thank Charles J. Sheviak
(NYS), Paul M. Catling (DAO), Gustavo A. Romero
(AMES), and one anonymous reviewer for encour¬
agement and helpful suggestions; Stanley N. Folsom
for the illustrations, assistance, and patience; and
the following students and friends who accompanied
me on field trips and/or loaned slides and made
helpful suggestions: Joann and Fred Knapp (who
Volume 2, Number 4
1992
Brown
Platanthera pallida
311
first led me to the plants), Roger Bradley, Hal Hor-
witz, Philip Keenan, Eric Lamont, Juliet Perkins,
Moreno and Dorothy Tagliapietra-Cherbavaz, Nan¬
cy Webb, and Joseph Welch.
Literature Cited
Brown, P. M. 1988. Checklist of New England Orchids.
Wild Flower Notes 3(1): 12-29. New England Wild
Flower Society, Framingham, Massachusetts.
Correll, D. S. 1950. Native orchids of North America
north of Mexico. Chronica Botanica, W altham, Mas¬
sachusetts.
Folsom, J. P. 1984. A reinterpretation of the status of
relationships of taxa of the yellow-fringed orchid com¬
plex. Orquidea (Mex.) 9: 320-345.
Lamont, E. E., J. M. Beitel & R. E. Zaremba. 1988.
Current status of orchids on Long Island, New York.
Bull. Torrey Bot. Cluh 115: 113-121.
Latham, R. A. 1940. Distribution of wild orchids on
Long Island. Long Island Forum 3: 103-107.
Luer, C. A. 1975. Native orchids of the United States
and Canada (excluding Florida). New York Botanical
Garden, New York.
Rickett, H. W. 1966. Wild Flowers of the United States
1: 1. New York Botanical Garden McGraw-Hill, New
York.
Zettler, L. W. & J. E. Fairey III. 1990. The status of
Platanthera integrilabia , an endangered terrestrial
orchid. Lindleyana 5: 212-217.
Orchidaceae Dunstervillorum V: Two New Species and a New
Combination in Epidendrum from the Venezuelan Guay ana
German Carnevali
Fundacion Instituto Botanico de Venezuela, Herl)ario Nacional de Venezuela (VEN), Aptdo.
2156, Caracas 1010-A, Venezuela; present address: Missouri Botanical Garden, P.0. Box 299,
St. Louis, Missouri 63166-0299, tl.S.A.
Gustavo A. Romero
Oakes Ames Orchid Herbarium, Harvard University Herbaria, 22 Divinity Avenue, Cambridge,
Massachusetts 02138, U.S.A.
ABSTRACT. Two new species and a new combina¬
tion are required for the treatment of Epidendrum
for the Flora of the Venezuelan Guayana. The new
species are Epidendrum caurense and Epidendrum
nuriense. Nealehmannia pabstii is transferred to
Epidendrum , with the new name Epidendrum di-
chaeoides. All three species are illustrated with pre¬
viously unpublished plates by the late G. C. K.
Dunsterville. Comments on affinities and geography
are provided for all species.
Although several authors (Barros, 1982; Duns¬
terville & Garay, 1976; Garay, 1977; Pabst, 1978)
assign subtrihal rank to some groups of the suhtribe
Laeliinae (members of former subtribes Epidanthin-
ae L. 0. Williams, and Stenoglossinae Benth.) that
display well-developed viscidia, most modern orchi-
dologists (e.g.. Dressier, 1984; Hagsater, 1985) agree
that most members of Epidendrum sensu lato dis¬
play variously developed viscidia and that the dif¬
ference between the proposed subtribes Epidanthin-
ae and Stenoglossinae, and the Laeliinae is one of
degree. There is a consensus in including most mem¬
bers of the Epidanthinae and Stenoglossinae, genera
such as Epidanthus L. 0. Williams, Amblostoma
Scheidw., Nanodes Bindley, Neolehmannia Kran-
zlin, Stenoglossum Kunth, and Kalopternix Garay
& Dunsterv., in the genus Epidendrum itself. Each
of these segregate genera is apparently closer to a
different group within the genus Epidendrum , and
not to the other members of the Epidanthinae or
Stenoglossinae, rendering Dunsterville & Garay’s
(1976) circumscription of these subtribes artificial.
Another genus included in the Stenoglossinae, 67a-
dobium Bindley, containing the sole species C. vio-
laeeum Bindley, clearly belongs in the genus Sca-
phyglottis Poeppig & Endl. because of its vegetative
and floral morphology. Scaphyglottis, although a
member of the Laeliinae, is only distantly related to
Novon 2: 312-318. 1992.
Epidendrum and the other former members of the
Stenoglossinae.
The broad circumscription of the genus Epiden¬
drum followed in the Orchidaceae treatment for the
Horn oj the Venezuelan Guayana requires the in¬
clusion of the new species Epidendrum caurense
(belonging to the Amblostoma group) and the trans¬
fer of Neolehmannia pabstii P. Braga to Epiden¬
drum.
Epidendrum caurense Carnevali & G. Romero,
sp. nov. TYPE: Venezuela. Bolivar: Rio Caura,
in rainforest close to Las Pavas Falls, where
Upper Rio Caura separates from the Lower,
ca. 300 m, May 1976, collected originally by
E. B. Stevens, Dunsterville & Dunsterville
1354 (holotype, AMES). Figure 1.
Species haec Epidendro lanipes Lindley siinilis sed
ramas inflorescentia laxioribus, pubescentia rachidis lax-
ioribus, floribus minoribus pallide flavo-viridis, lobo centrali
oblongo non pandurato vel cuneato, carina centrali bre-
vioris discrepat.
Epiphytic herb; rhizome shortly repent, with the
pseudobulbs ± aggregate. Pseudobulbs 30-50 cm
long, erect, homoblastic, slenderly fusiform, 3-8-
leaved in the upper half. Leaves 15 25 x 1.5-2
cm, linear-elliptic, acute, subtended by a tubular
sheath, midnerve finely sulcate and sharply carinate,
blades generally ascending when young, tending to
droop when hilly grown. Inflorescence terminal, 25
30 cm long, an arcuate panicle with slender, some¬
what divaricate, subdensely many-flowered branch¬
es; peduncle remotely sheathed, terete, glabrous;
rachis with scattered, irregular, moniliform hairs;
floral bract ca. 3 x 2 mm, glabrous, ovate, obtuse.
Pedicellate ovary 7.5-8 mm long, densely covered
with moniliform hairs. Flowers resupinate or not,
light yellowish green, with widely spreading perianth
Volume 2, Number 4
1992
Carnevali & Romero
Epidendrum from the Venezuelan
Guayana
313
segments; sepals ra. 7 mm long, 3-nerved, apex
sharply mucronate, dorsal ca. 1.9 mm wide, nar¬
rowly elliptic, acute, acuminate, laterals ca. 2.3 mm
wide, obliquely elliptic-oblanceolate, acute; petals
slightly shorter than the sepals, ca. 0.6 mm wide.
linear-oblanceolate, obtuse, 1-nerved, set at variable
angles to the column. Labellum fused to column
throughout the length of the latter, the free portion
ca. 4.5 x 5 mm, sharply 3-lobed; lateral lobes
obliquelv ovate, ca. 2.5 x ca. 2 mm; central lobe
314
Novon
ca. 2.2 x ca. 1.5 mm, oblong, apex rounded or
obtuse, mucronate; disk of the labellum with 2 basal
calli and 3 thickened keels, the central keel shorter
or subequaling the lateral ones. Column ca. 4.5 mm
long, thickened apically; anther ca. 1.2 mm long,
consisting of a fairly thin plate bearing 2 separate,
thin, bilocular, suborbicular cells; pollinia in 2 slight¬
ly asymmetrical pairs, only slightly compressed, each
pair on a single caudicle, the caudicles arising from
a common point on a viscidium whose form is com¬
pletely distorted after removal from the split ros-
tellum.
This new species belongs to the E. amblosto-
moides group of Epidendrum (E. Hagsater, pers.
comm.), which is often included within the former
genus Amblostoma (or Amblostoma group of Epi¬
dendrum, Dressier, 1984) but is recognized here as
distinct. Both groups are characterized by their
thickened column, almost spheroid pollinia, and well-
developed viscidia. The Amblostomoides group is
characterized by membranous, mostly pubescent
flowers, while in the mostly Brazilian Amblostoma
group the flowers are glabrous and (especially the
labella) very fleshy. All the species of the Amblos¬
tomoides group have fusiform or cylindrical pseu¬
dobulbs, usually paniculate inflorescences with
densely flowered, divaricate branches, and a com¬
monly pubescent pedicellate ovary. Epidendrum
caurense seems closest to E. lanipes, from Bolivia,
Peru, Ecuador, and Colombia. From this species, E.
caurense differs by its less dense inflorescence, the
less dense pubescence of the pedicellate ovary, its
smaller sepals (ca. 7 mm long vs. 10-12 mm long),
pale yellow-green flowers (vs. white), the shape of
the labellar callus (see below), and its oblong central
lobe (vs. cuneate or obovate). The callus of Epi¬
dendrum lanipes has a central keel and two lateral
ones; these lateral calli can be entire or bicallose,
thus making the entire callosity 5-keeled. The cen¬
tral keel is always longer than the lateral keels. In
E. caurense there are always five keels but the
lateral keels are longer or subequal to the central
keel.
Epidendrum lanipes is known from elevations of
800-1,400 m (Dodson & Bennett, 1990: 64), while
E. caurense is known only from elevations below
300 m. Epidendrum purum Lindley, distributed
from Venezuela to Bolivia, is also similar but has
white flowers, a glabrous rachis, and pedicellate ova¬
ry. Other similar species are E. amblostomoides
Hoehne, from Coias, Brazil, and E. subpurum Reichb.
1. from northern Venezuela. However, both species
have a shorter, broader, emarginate central lobe
and glabrous rachis and pedicellate ovary.
A specimen from Rio Orejas, La Teta, Upper
Cauca, Colombia, Lehmann 6938 (AMES, NY),
could represent another population of Epidendrum
caurense.
Epidendrum diehaeoidcs Carnevali & G. Rom¬
ero, nom. nov. Replaced name: Neolehmannia
pabstii P. Braga, Bradea 3: 171. 1981, non
Epidendrum pabstii A. Hawkes, 1957. TYPE:
Brazil. Amazonas: Rio Marie, 16 Oct. 1978,
Madison , Hraga A- Kennedy (PEE 365) ((te¬
letype, INPA-85085). Figure 2.
I bis species, recently described from Amazonian
Brazil, has been collected several times, mainly in
the Casiquiare and Rio Negro basins in the extreme
south of Territorio Federal Amazonas of Venezuela.
Here it is locally common, always growing pendent
and low on trees in the rather open, low, humid
forests that are usually contiguous to “banas.” The
plants are frequently purple-tinged or entirely wine-
purple. An unusual feature of Epidendrum di-
chaeoides is that it frequently produces upper ax¬
illary inflorescences, besides the more normal ter¬
minal inflorescences. This character was not
mentioned in the original description of A eoleh-
mannia pabstii. Vegetatively, Epidendrum di-
chaeoides resembles Epidendrum prostratum
(Lindley) Cogn. and related species (the former
members of the genus Physinga Lindley), but it
lacks the elongated inflorescence and the basal por¬
tion of the labellum does not enfold the lateral mar¬
gins of the apex of the column. The similarities of
Epidendrum dichaeoides and E. prostratum are
more likely to be due to convergence (both occur
in the same microhabitats) than to a close phyletic
relationship, as Epidendrum dichaeoides is prob¬
ably closer to species like Epidendrum jejunum
Reichb. L, E. viridibrunneum Reichb. L, and E.
williamsii Dodson, all included in Dunsterville &
Garay’s (1976) concept of the genus Neolehmania.
The new specific epithet suggests the conspicuous
vegetative resemblance of this taxon to plants of the
genus Dichaea Lindley.
Specimens examined. Brazil. AMAZONAS: Rio Negro,
caatinga at Porto Camanus, 19 Oct. 1978, Madison et
al. 6438 (SEL); Rio Curicuriari, ca. 10 kin above the
mouth, 25 Oct. 1978, Madison et al. 6648 (SEL).
Venezuela, territorio federal amazonas: Departa-
mento Atabapo, Cano Yagua, E del Cerro Yapacana,
3°3'N, 66°40'W, F. Gudnchez 1050 (TFAV); Departa-
mento Casiquiare, Capihuara, LI. Williams 15570(\ EN);
Departamento Rio Negro, IVIC Study Area, 4 km NE of
San Carlos de Rio Negro, ca. 20 km SE of confluence of
Rio Negro and Brazo Casiquiare, 1°56'N, 67°03'W, 120
m, 10 Nov. 1977, R. Liesner 3280 (MO, SEL, YEN),
Volume 2, Number 4
1992
Carnevali & Romero
Epidendrum from the Venezuelan
Guayana
315
Figure 2. Epidendrum dichaeoides Carnevali & G. Romero. —A. Habit. —B. Flower, lateral view. —C. Detail
of column, labellum, and vesicle. —D. Floral segments, spread. —E. Anther and pollinia.
9 Apr. 1979, H. Liesner 6424 (MO, VEN), 11 Nov.
1987, R. Liesner dr G. Carnevali 22981 (MO, VEN),
5 Sep. 1988, G. Carnevali & L Ramirez 2763 (VEN),
Mar.-Apr. 1981, F. Delascio et al. 9483 (V EN), 1 ■
Sep. 1975, P. Berry 1361 (SEE, VEN); Rio Arari, 1°30'N,
65°13'W, 530-550 m, Nov. 1983, F. Gudnchez 2220.
(TFAV, VEN); Cerro La Neblina, Rio Mawarinurna, 140
m, 15-18 Mar. 1984, R. Liesner 16559 (MO, VEN),
11". Anderson 13362 (MICH, VEN); Rio Extremo Sur,
cerca de la confluencia con el Rio Siapa, 1°50'N, 64°30'W,
520 m, Dunsterville E. Dunsterville 1260 (AMES,
VEN).
316
Novon
Figure 3. Epidendrum nuriense Carnevali & Hags. A. Habit. —B. Floral segments, spread. — C. Column and
labellum. —D. Lateral view of flower, dorsal sepal and petals removed. —E. Flower. — F. Anther and pollinia. —
G. Column, ventral view.
Epidendrum nuriense Carnevali & Hags., sj>.
nov. TYPE: V enezuela. Bolivar: Altiplanicie de
Nuria, apparently common in cloud forest at
600-700 m, Dunsterville & E. Dunsterville
1333 (holotype, VEN; isotype, line drawing,
AMES). Figure 3.
Iferba epiphytica. Caules ex internodiis medianis prod-
ucti, apice 2-3(-5) foliati. Flores 1-4, terminales, non
resupinati. Ovarium incrassatum, basi incurvum. Sepala
5-nervata, lateralia basi columnae breviter connata. Petala
5-nervata, oblanceolata. Labellum 6-7 x 5-7 mm, ova-
tum vel ovato-ellipticum, basi cordatum, apice truncatum,
disco pulvinato-bicalloso. Epidendro lechleri Reichb. f.
Volume 2, Number 4
1992
Carnevali & Romero
Epidendrum from the Venezuelan
Guayana
317
similis sed labello angustiore, inflorescentia laxiore pau-
ciflora, planta floresque majore recedit.
Epiphytic herb, 7-40 cm high, with sympodial
erect growth, branched. Roots produced only from
the base of the basal stems, fleshy, thin, ca. 2.5 mm
diam. Stems reedlike, cylindric, straight, produced
from medial internodes of older stems, sympodial
elements 2-15 cm long; concealed in gray, nervose,
tubular sheaths to 5 cm long. Leaves 4 10 x 1.8
2.2(-3) cm, 2—3(5) per stem, grouped toward stem
apex, narrowly elliptic or rarely ovate, acute, margin
entire, blades articulate with their laterally com¬
pressed, striate sheaths. Inflorescence terminal from
the apex of fully mature stems, spathe absent, rac¬
emose or rarely branched, somewhat nutant; pe¬
duncle 1.5-3 cm long, laterally compressed, pe¬
duncle bracts 1 or 2, ca. 2 mm long, triangular;
rachis fractiflex, laterally compressed. Flowers not
resupinate, 1 —3(—4) in tbe raceme, opening simul¬
taneously; sepals and petals bronze-green, nerves
somewhat darker; labellum shiny, bronze-colored with
green calli; column green, apically pink; anther
cream-color. Floral bracts ca. 2 mm long, arnplex-
icaul, triangular. Pedicellate ovary 1.8 2.5 cm long,
longitudinally swollen ventrally, terete, smooth, ba-
sally incurved. Sepals 5-nerved, flesby, rugulose
without, dorsal keel poorly developed; dorsal sepal
17-18 X 4 mm, reflexed over ovary, narrowly
obovate, apex obtuse, margins not revolute; lateral
sepals 17-18 x 5-6 mm, partially fused with col¬
umn at base, then reflexed, elliptic to obovate-ellip-
tic, apex subacute to acute, slightly falcate, margins
revolute; petals 17 x 4 mm, 5-nerved, less fleshy
than sepals, spreading or reflexed, narrowly obovate-
elliptic, apex obtuse, slightly falcate, margins ex¬
tended or somewhat revolute. Labellum fused to
column throughout the length of the latter, fleshy,
blade 6—7 x 5-7 mm, ovate to ovate-elliptic, basally
cordate, apically truncate, margins finely erose, ba¬
sally provided with 2 small, pulvinate calli, without
keels, marginally thinner than the rest of the blade.
Column ca. 10 mm long, slightly bent toward base,
clinandrium very small, entire; rostellum near col¬
umn-apex, slitlike; lateral lobes of column convex-
truncate. Nectary very deep. Anther ovoid, without
ornamentation, 4-locular. Pollinia 4, subglobose,
somewhat compressed, subequal, caudiculae slightly
longer than pollinia, viscidium semi-liquid. Capsule
unknown.
This species belongs to the Epidendrum arbus-
cula Lindley group, which is characterized by the
stems branching around the middle portion and pro¬
ducing roots from the base of tbe most basal stems;
the architecture of the plant is thus termed “sym¬
podial erect growth" (Ilagsater, 1987). Epiden¬
drum nuriense does not have close allies in the
Guayana Shield. The only other species in the area
with a similar branching pattern is Epidendrum
lechleri Reichb. f. This species, however, has denser
inflorescences, smaller flowers, and a proportionally
much broader labellum. Epidendrum megagas-
trium Lindley, from Andean Peru and Ecuador,
resembles E. nuriense because of its habit and lon¬
gitudinally swollen ovary, but the Andean species
comes from much higher elevations (2,500-2,700
m), has denser inflorescences, and an acuminate or
acute labellum. Epidendrum macrugastriurn Kran-
zlin from Peru is possibly related to E. nuriense but
comes from higher elevations in the Andes (2,700-
2,900 m), has smaller flowers, and a differently
shaped (very broadly cordate), much broader la¬
bellum.
Epidendrum nuriense seems to be restricted to
the eastern portion of Estado Bolivar, Venezuela,
where it is apparently common in the Altiplanicie
de Nuria, and more scarce in La Escalera, where
the senior author and Carlos Garcia Esquivel col¬
lected living material that is currently under culti¬
vation.
Acknowledgments. Nora Dunsterville permitted
use of the previously unpublished plates by tbe late
C. C. K. Dunsterville. The Department of Graduate
Studies of the Missouri Botanical Garden provided
financial support. The Department ol Biology of the
University of Missouri-St. Louis supported the senior
author while the studies leading to this paper were
being conducted. This paper was written while the
senior author was a Visiting Scientist at the Marie
Selby Botanical Gardens. Bruce K. Holst, Francisco
Lorea 1L, and Ivon M. Ramirez carefully read and
commented on earlier drafts of the manuscript. Eric
Hagsater generously provided discussion and veri¬
fied the determinations. The curators of MO, NY,
PORT, SEL, and VEN kindly allowed us the study
of their specimens.
Literature Cited
Barros, F. 1982. Duas novas combina£6es para a sub-
tribu Stenoglossinae Benth. (Orchidaceae). Bol. Mus.
Bot. Municipal 53: 1-7.
Dodson, C. H. & D. Bennett. 1990. Orchids of Peru
1. leones Plantarum Tropicarum 2: 64. Missouri
Botanical Garden, St. Louis.
Dressier, R. L. 1984. La delimitacion de generos en el
Complejo Epidendrum. Orquidea 9: 277-298.
Dunsterville, G. C. K. & L. A. Garay. 1976. Venezuelan
Orchids Illustrated 6. Andre Deutsch, London,
Garay, L. A. 1977. The Subtribe Epidanthinae L. 0.
Wins. Orchid Digest 41(1): 19-22.
Hagsater, E. 1985. Toward an understanding of the
318
Novon
genus Epidendrum. Pp. 195-201 in K. Tan (editor),
Proc. Eleventh World Orchid Conference. Interna¬
tional Press, Singapore.
-. 1987. Epidendra nova et criticae 3: Nuevas
especies del grupo Epidendrum arbuscula de Mexico
y Centroamerica. Orquidea 10: 337-353.
Pahst, G. F. J. 1978. Noticias Orquidologicas XVIII.
Bradea 2(46): 306.
A New Species of Cyperaceae (Cryptangieae) from the
Venezuelan Guayana
Gerrit Davidse
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
ABSTRACT. Lagenocarpus amazonicus (Cypera¬
ceae), from Amazonas, Venezuela, is described and
illustrated. It differs most notably from other species
of section Junciformes in its strongly pubescent rhi¬
zome sheaths and connate stamen filaments.
Lagenocarpus Nees is a tropical American genus
of the tribe Cryptangieae that has been most recently
revised by Koyama & Maguire (1965) and Koyama
(1972), who recognized 28 species in four sections.
The genus has two centers of species diversity: the
Guayana region and southeastern Brazil in the area
between the states of Bahia, Minas Gerais, and Sao
Paulo.
A review of the genus for the Flora of the I en-
ezuelan Guayana brought to my attention a pre¬
viously unrecognized species that 1 am naming L.
amazonicus, based on its occurrence in Territorio
Federal Amazonas, Venezuela.
Lagenocarpus amazonicus Davidse, sp. nov.
TYPE: Venezuela. Territorio Federal Amazo¬
nas: Depto. Rio Negro, Cerro Aratitiyope, aprox.
70 km al SSW de Ocamo, 2°1()'N, 65°34'W,
con riachuelos afluente al rio Manipitare, piedra
ignea, 990-1,670 m, 24-28 Feb. 1984, ./.
Steyermark, P. Perry & F. Delascio 130049
(holotype, MO; isotypes, NY, VFN). Figure 1.
Herba perennis rhizomatosa, rhizomate 8-14 mm in
diametro squamis dense pubescentibus obtecto, cufmis
22-78 cm longis, basin 1.7-2.9 mm latis. Laminae fo-
liorurn 3-7.5 cm longae, 2.0-4.0 mm latae. Spiculae
masculinae 4 5.5 mm longae; gluinis 3.2 4.5 mm longis;
staminibus 2, filamentis cormatis, antheris 1.4 1.7 mm
longis. Spiculae femineae 2.5 3.0 mm longae; gluinis 3
4, 2.0-3.0 mm longis. Achenia 1.8-2.3 mm longa, laevia
vel minute papillosa apicem versus; basi styli 0.3 0.4 mm
longa, cylindrica, puberula apicem versus.
Perennial, rhizomatous, monoecious herb. Rhi¬
zomes 9 cm long, 8—14 mm thick, horizontal, pro¬
ducing numerous, slender roots, the segments short
and swollen and covered by densely appressed-pu-
bescent scales. Culms 22-78 cm long, 1.7-2.9 mm
wide at the base, erect, trigonous, glabrous; nodes
numerous; lower 4-6 internodes elongated to 15
cm, bearing somewhat loose, bladeless sheaths to¬
ward the base, the blade more fully developed dis-
tally; upper 10-20 internodes not elongated, the
sheaths closely overlapping and bearing fully de¬
veloped blades. Sheaths closed, puberulent or gla¬
brous, eligulate; contraligule 0.5 1.5 mm, nerved
to the apex; blades 3-7.5 cm long, 2.0 4.0 mm
wide, widest at the base, divergent, acuminate, abax-
ially puberulent along the midvein. Inflorescences
small, unbranched to 3-branched umbellate clusters
borne from the uppermost nodes, with ihe pedicels
or branches exserted from the sheaths. Spikelets
unisexual. Male spikelets 4-5.5 mm long, 0.7 1.1
mm wide, solitary, binary, or rarely ternate, borne
from nodes below those bearing the female spikelets,
oblong-ellipsoid, dark brown, many-flowered; male
floral bracts 3.2-4.5 mm long, 1-1.3 mm wide,
densely imbricate, lanceolate, membranous,
1 -nerved, the lower sterile and aristate to 1 mm,
the 2 or 3 distal fertile and subtending male flowers,
smaller and more delicate. Male flowers consisting
of 2 stamens; filaments slightly longer than the sub¬
tending floral bract, connate; anthers 1.4-1.7 mm
long, separate, basifixed, with a minute tuft of hairs
at the apex. Female spikelets 2.5 3.0 mm long,
solitary or rarely binary; female floral bracts 3 or
4, 2.0-3.0 mm long, 0.6-1.1 mm wide, lanceolate
to lanceolate-ovate, membranous, glabrous or pu¬
berulent on the midvein and apically ciliolate,
1 -nerved, the lowermost with an awn to 1 mm long,
the distal floral bracts shorter, thinner and with less
developed awns or acute. Female flower a solitary
pistil; styles 3. Achenes 1.8-2.3 mm long, 0.9-1.0
mm wide, about as long as the floral bracts, smooth
or minutely papillose distally, obovoid, trigonous,
each face with a basal depression; style base per¬
sistent, 0.3-0.4 mm long, cylindrical, minutely pu¬
berulent at the tip; perianth absent or rudimentary;
ovary 0.8 0.9 mm long, 0.3 mm wide, oblong in
outline, sharply trigonous.
Paratypes. VENEZUELA. AMAZONAS: Cuinbre del Cerro
Autana, 4°52'N, 67°27'W, 1,230-1,240 in, sabana y
afloramientos expuestos, 20-22 Sep. 1971, J. A. Stey¬
ermark 105151 (MO, YEN).
By virtue of its culms with many elongated in¬
ternodes, the lower leaves with no or reduced leaf
Novon 2: 319-321. 1992.
320
Novon
Figure 1. Lagenocarpus amazonicus Davidse. A. Habit with the basal sheaths enlarged (N) to show the dense
pubescence. B D. Male spikelets at anthesis. B. Three spikelets in cluster. —C. Two spikelets in cluster. —D.
Solitary spikelet. —E. Female spikelets. —F. Achene. G-M. Dissection of male spikelet. —G. Lowest sterile floral
bract. H. Lowest fertile floral bract. —I. Lowest male flower consisting of 2 connate filaments with the anthers
dehisced and fallen. —J. Middle fertile floral bract. k. Middle male flower with 2 connate filaments and 2 separate
anthers, near anthesis. —L. Upper fertile floral bract. M. Upper male flower with the connate filaments not yet
elongated. —N. Densely pubescent scales at the base of a culm. — 0. Detail of leaf showing the contraligule.
Volume 2, Number 4
1992
Davidse
Lagenocarpus amazonicus
321
Table 1. Characters
that distinguish Lagenocarpus junciformi;
> from L. amazonicus.
Character
L. junciformis
L. amazonicus
Rhizome scales
Glabrous
Densely appressed pubescent
Basal bladeless sheaths
Tight
Loose
Basal culm node width
0.6-0.9 mm
1.7-2.9 mm
Blade width
0.4- 1.8 mm
2.0-4.0 mm
Filaments
Separate
Connate
Female floral bracts
4-5
3-4
Achene texture
Papillose
Smooth but apically minutely papillose
Achene length
1.4-1.9 mm
1.8-2.3 mm
Style base shape
Conical-cylindrical
Cylindrical
Style base length
0.1-0.2 mm
0.3-0.4 mm
Style base pubescence
Apically glabrous
Apically puberulent
blades, the fully developed leaves clustered toward
the apex of the culms, the primarily terminal inflo¬
rescences (sensu Koyama, 1972), and the sharply
trigonous achenes, this species belongs to section
Junciformes Koyama. which Koyama (1972) con¬
sidered to have five species.
This species is only known from the two cited
collections, and Koyama (1972) considered one of
these, Steyermnrk 105151. to he /.. junciformis
(Kunth) Kuntze. He thus believed L. junciformis to
have a bimodal distribution, i.e., in the Planalto of
Brazil and in the Venezuelan Guayana. However,
the two cited Guayanan collections differ from the
Brazilian collections of this species in the characters
listed in Table 1.
Especially notable distinguishing characteristics
of L. amazonicus, and ones in which this species
differs from all the others in the section, are the
strongly pubescent rhizome sheaths and the connate
stamen filaments.
The fusion of the filaments is a feature previously
unknown in the genus and is known to me only in
two other genera of Cyperaceae: Scleria Bergius
and Rhynchospora Vahl. In Scleria it has been
reported in ,S. melanomphala Kunth, .S. bequaertii
De Wild., and .S’, glabroreticulata De Wild. (Bier-
art, 1951: pi. I, fig. 30, pi. 2, figs. 2, 9). and in
Rhynchospora it has been reported as “not unusu¬
al" in male flowers (Goetghebeur, 1986: 681), as,
for example, in R. confusa Ballard (Ballard, 1934:
t. 3250, fig. 4). That this feature is probably not a
simple anomalous occurrence in /,. amazonicus
seems evident from the fact that all plants from the
two known populations uniformly have it, even though
they are separated by a distance of approximately
370 km.
Acknowledgments. I thank Bruno Manara for
the excellent illustration and the curators at NY and
V EN for the loan ot specimens.
Literature Cited
Ballard, F. 1934. 4'abula 3250. Rhynchospora con¬
fusa Ballard. Hooker’s Icon. PI. 33: t. 3250, 1 3.
Goetghebeur, P. 1986. Genera Cyperacerarurn. Ph.D.
Thesis, Rijksuniversiteit Gent, Belgium.
Koyama, T. 1972. Two new species and a new' section
of Cyperaceae from Brazil. Brittonia 24: 279 -282.
- & B. Maguire. 1965. Cyperaceae tribe La-
genocarpeae. In: The Botany of the Guayana High¬
land — Part \ 1. Mem. New York Bot. Card. 12: 8
54.
Pierart, P. 1951. Les especes du genre Scleria Berg,
du Congo Beige et du Ruanda-Urundi. Lejeunia Mem.
13: 1-68, pis. 1-5.
New Species of Festuca, Sporobolus , and Eriochloa (Poaceae) from
Mesoamerica and South America
Gerrit Davidse
Missouri Botanical Garden, l\0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Richard H . Roll I
Department of Botany, 353 Bessey Hall, Iowa State University, Ames, Iowa 50011, U.S.A.
ABSTRACT. As a result of work on the Poaceae for
Flora Mesoatnericana, the following four new spe¬
cies are described, illustrated, and compared with
their closest putative relatives: Festuca talaman-
censis from the Cordillera de Talamanca in Costa
Rica, F. herrerae from the Cordillera de lalamanca
in Costa Rica and Panama, Sporobolus distichi-
vaginatus from the Peten, Guatemala, and Frioch-
loa stevensii from Nicaragua, Venezuela, and Ec¬
uador.
In addition to the new species of grasses recently
described as the result of a review ot the Poaceae
for Flora Mesoatnericana (Davidse & Pohl, 1992),
the following four species were found to he new and
are here described, discussed, and illustrated lor the
first time. As in the earlier paper, the new species
are published under the name ot the person primarily
responsible for the account ol the taxon.
POAEAF.
Twenty species ol Festuca were accepted lor
Flora Mesoatnericana , including the following two
new species. Alexeev (1981, 1982) has most re¬
cently studied the genus for Mesoamerica. Of the
five species that he described as new, three have
been accepted for the Flora Mesoatnericana treat¬
ment, and the other two were synonymized: F.
aguana Alexeev (syn. F. panamica Alexeev), /'.
cartagana Alexeev, F. swallenu Alexeev, h. tvilld-
enotciana Schultes & Schultes f. (syn. /’. guate-
malica Alexeev).
Festuca herrerae Davidse, sp. nov. TYPE: Costa
Rica. Limdn: Cordillera de'Talamanca, Atlantic
slope, Kamuk massif, paramo NE ol the Kamuk
peak, 3,000-3,300 m, 9°16'-17'N, 83°00'
02°W, Rlechnum -shrub association in Chits-
quea-Hypericum paramos, 17-18 Sep. 1981,
G. Davidse & G. Ilcrrera Ch. 2929 I (holo-
type, MO; isotypes, RM. CR, MEXU). Figure
1 A-D.
Novon 2: 322-328. 1992.
Gramen perenne caespitosa innovationibus extra et in-
travaginalibus. Culmi 30-90 cm longi. Vaginae foliorum
puberulae retrosum vel glabrae aliquantum fibrosae; ligula
0.3 2 mm longa; laminae 10-25 cm longae, 1-4 mm
latae, planae vel plicatae, infra glabrae, supra parce sca-
berullae. Panicula 6-19 cm longa, 511 cm lata, laxa.
Spiculae 6.6- 9 mm longae; gluina infra 3.1-4.5 mm
longa, 1-nervia; glurna supra 5-5.8 mm longa, 3-nervia;
tlosculi 3-5; lemmata 5.7-6.8 mm longa, 5-nervia, ac-
uminatis, glabra dorsaliter scaberulla apicem versum; an-
therae 3, 1.6-2.1 mm longae; ovarium glabrum.
Loosely caespitose, perennial herb; innovations
intra- and extravaginal; rhizomes not elongated.
Culms 30 90 cm long, hollow, sparsely scaberulous
below the nodes; elongated internodes 2 5. Sheaths
retrorsely puberulent (in the lower leaves) to gla¬
brous, stramineous to brown, slightly fibrous in age,
the margins free; auricles absent; ligule 0.3-2 mm
long, ciliate, membranous, the lateral extensions to
0.6-2 mm long; blades 10 25 cm long, 1-4 mm
wide, flat to folded, glabrous below, sparsely sca¬
berulous above; abaxial and adaxial sclerenchyma
girders well developed. Panicle 0-T9 cm long, 5-
11 cm wide, open, lax, erect; axis scaberulous; low¬
est panicle branches 4-5 cm long, solitary or paired,
spreading and drooping at the tips; pedicels scaber¬
ulous. Spikelets 6.6-9 mm long; lower glume 3.1
4.5 mm long, subulate, 1-nerved; upper glume 5-
5.8 mm long, narrowly lanceolate, 3-nerved; florets
3-5; lemmas 5.7-6.8 mm, obscurely 5-nerved, gla¬
brous on the back, scaberulous at the tip, entire,
acuminate, unawned; palea as long as the lemma to
0.7 mm shorter, scaberulous on and between the
keels near the tip, nearly glabrous toward the base,
bidentate, the margins glabrous; stamens 3, the an¬
thers 1.6 2.1 mm long; ovary glabrous; styles 2,
terminal, separate, long-plumose in the upper
Caryopsis 2.9-3 mm long, 0.8 mm wide, dark brown,
glabrous, convex on the embryo side, sulcate the
entire length on the hilum side; embryo as
long as the caryopsis; hilum ca. V l(l as long as the
caryopsis.
Paratypes. Costa Rica. LIMON: Cordillera de Tala¬
manca, Atlantic slope, Kamuk massif, main SW ridge
Volume 2, Number 4
1992
Davidse & Pohl
New Species of Festuca, Sporobolus,
and Enochloa
323
Figure 1. Festuca species. A I). F. herrerae Davidse. A. Habit. —B. Spikelet. —C. Uppermost fertile floret and
the ultimate reduced sterile floret. —D. Caryopsis. E, F. F. talamancensis Davidse. —E. Uppermost fertile floret
and the ultimate reduced, sterile floret. — F. Spikelet. (A based on Davidse Herrera 29291, B D on Davidse et
al. 25354, E, F based on Davidse 2477 6.)
from Kamuk peak to Cerro Dudu, 3,000 3,300 m, 9°15'
16'N, 83°02'-04'W, Blechnum -shrub paramo, 19 Sep.
1984, Davidse & Herrera Ch. 29399 (MO); Cordillera
de Talamanca, Cerro Kamuk massif, between Cerro Dudu
and Cerro Apri, 2,900-3,100 m, 9°14'30"-15'30"N,
83°03'30"-04'30"W, shrub paramo dominated by Di-
plostephium, Blechnum and other shrubs 1VS-2 m tall,
in open patches among shrubs, 23 & 26 Mar. 1984,
Davidse, Herrera Ch. & Warner 25879 (CR, ISC, LE,
MEXU, MO, US). LIMON-PUNTARENAS BORDER: Cordillera
324
Novon
de Talamanca, Cerro Kasir, on the continental divide,
2,950 m, 9°12'N, 83°03'W, small marshy paramo in
forest opening, 20 Sep. 1984, Davidse <fe Herrera Ch.
29405 (CR, F, MO). Panama, iiocas del toko: Cordillera
de Talamanca, 4 airline km NW of the main peak of
Cerro Fabrega along the NW ridge of the massif, 3,000-
3,150 m, 9°09'N, 82°54'W, lowest paramo slopes with
low-forested gullies, in open paramo, stems purple, Dav¬
idse el at. 25390 (MO); Cordillera de Talamanca, 2 airline
km NW of the main peak of Cerro Fabrega along the
NW ridge of the massif, 3,150-3,200 m, paramo with
low-forested gullies, around small pond, culms and spike-
lets dark purple, 7-8 Mar. 1984, Davidse et at. 25354
(CR, K. MO, PM A).
This species belongs to subgenus Festuca sect.
Festuca. It appears to be related to F. willdenowia
in that the leaf blades are not permanently convo¬
lute. Il differs from that species in sclerenchyma
girders associated with all vascular bundles (rather
than only 2), lemma essentially glabrous and un-
awned (not scabrous and awned 0.5-2 mm).
Festuca herrerae is so far known only from the
two peaks of the Cordillera de Talamanca that strad¬
dle the Costa Rican Panamanian border and which
form the highest part of the International Park La
Amistad. Its range overlaps that of F. talamacensis
in the Cerro Karnuk paramos.
The species is named in honor of the expert
Costa Rican field botanist and plant collector, Ger¬
ardo Herrera Chacon, a good friend and co-collector
of this species.
Festuca talamancensis Davidse, sp. nov. TY PE:
Costa Rica. San Jose-Cartago: Cordillera de
Talamanca, C.erros Cuerici, Parque National
Chirripo, continental divide, 9°35'N, 83°38'W,
Chusquea-Hypericum -ericad shrub paramo
with open areas w ith tussock grasses (dominated
by Cortaderia) near the summit, 3,200 3,394
m, inflorescences nodding, 17 Sep. 1983, G.
Davidse 2477 6 (holotype, MO; isotypes, AAU,
RM, COL, EAP, ISC, K, L, LE, MEXU, NY,
PM A, PRE, SI, US, USM, YEN). Figure IE, F.
Gramen perenne caespitosa innovationibus extravagin-
alibus. Culmi 45 90 cm longi. Vaginae foliorum glabrae
integrae; ligula 0.4 1 mm longa; laminae foliorum 10-
30 cm longae, 0.5-0.7 mm latae, involutae, infra glabrae,
supra scaberullae. Panicula 10-14 cm longa, 3-4 cm
lata, laxa. Spiculae 6.4-7 mm longae; gluma infra 3.4-
4.5 mm longa, 1-nervia; gluma supra 4.8 6 mm longa,
3-nervia; flosculi 3; lemmata 5-6 mm longa, 5-nervia,
glabra vel scaberulla, arista 0.3-1 mm longis; antherae
3, 1.11.4 mm longae; ovarium glabrum.
Densely caespitose, perennial herb; innovations
extravaginal; rhizomes not elongated. Culms 45-90
cm long, hollow, glabrous; elongated internodes 2(3).
Sheaths glabrous, stramineous, nonfibrous, the mar¬
gins free; auricles absent; ligule 0.4-1 mm long.
ciliolate, membranous, the lateral extensions un¬
equal, the larger extension 1-1.5 mm long; blades
10-30 cm long, 0.5-0.7 mm wide, involute, gla¬
brous below, scaberulous above; abaxial sclerenchy¬
ma girders well developed, sometimes limited to scle¬
renchyma strands below the marginal vascular
bundles; adaxial sclerenchyma absent. Panicle 10-
14 cm long, 3-4 cm wide, nodding, lax; axis gla¬
brous; lowest branches 4-6 cm long, paired, as¬
cending or the ends arching; pedicels glabrous.
Spikelets 6.4-7 mm long; lower glume 3.4-4.5 mm
long, subulate, 1-nerved; upper glume 4.8-6 mm
long, lanceolate, 3-nerved, abaxially glabrous, adax-
ially scaberulous; florets 3, the lower 2 bisexual, the
upper rudimentary; lemmas 5 6 mm long, obscurely
5-nerved, abaxially scabrous in the lower
scaberulous to nearly glabrous in the upper
adaxially scaberulous throughout, entire, the awn
0.3-1 mm long; paleas 0.5-1.5 mm shorter than
the lemmas, relatively longer in the uppermost flo¬
rets, densely scabrous on and between the keels, the
tip slightly notched, the margins glabrous on the
lower Lfc-Li, scabrous in the upper stamens
3, the anthers 1.1 1.4 mm long; ovary glabrous;
styles 2, terminal, separate, long-plumose in the
upper %. Caryopsis 2.8 3.5 mm long, 0.7 0.8 mm
wide, dark brown, glabrous, convex on the embryo
side, deeply and broadly grooved on the hilum side;
embryo 14 as long as the caryopsis; hilum ca. % as
long as the caryopsis.
Paratypes. Costa Rica. LIMON: Cordillera de Tala¬
manca, Atlantic slope, Karnuk massif, paramo NE ol the
main Karnuk peak, 9°16'-17'N, 83°0()'-02'W, Blech-
mxm-shrub association in Chusquea-Hypericum paramo,
17-18 Sep. 1984, Davidse & Herrera Ch. 29335 (RM,
CR, ISC, MEXU, MO, US).
Festuca talamancensis belongs to subgenus Fes¬
tuca sect. Festuca. It is closely related to F. tolu-
censis Kunth, a species distributed from Jalisco and
Veracruz, Mexico, south to Costa Rica, hut F. tal¬
amancensis differs in its leaves glabrous abaxially
(not scabrous), florets 3 (not 4 8), pedicels and
inflorescence axis glabrous (not scaberulous), lower
glume 3.4-4.5 mm (not 4.8 8.2 mm), upper glume
4.8-6 mm (not 6.4-9.5 mm), lemmas 5-6 mm and
less strongly scabrous (not 6.5-9.5 mm), anthers
1.1-1.4 mm (not 3.2-4.2 mm). The abaxial cuticle
is thicker than the epidermal cells, which are quite
thick-walled. The adaxial ribs are prominent.
So far it is known only in Costa Rica from the
Cordillera de Talamanca (after which it is named)
in paramos associated with the two highest peaks in
the country, Chirripo Grande (and its western ex¬
tension Cerros Cuerici) and Cerro Karnuk. Its dis¬
tribution overlaps with that of F. tolucensis in the
Chirripo paramos.
Volume 2, Number 4
1992
Davidse & Pohl
New Species of Festuca, Sporobolus,
and Eriochloa
325
Eracrostideae
Sporobolus dislichivaginalus R. Pohl, sp. nov.
TYPE: Guatemala. Peten: Dolores, en orillando
Rio Machaquilla, lado sur este de caserio, fruto
amarillo, en foresta alta, 18 Feb. 1971, R. Tun
Ortiz 1610 (holotype, ISC; isotypes, F, US).
Figure 2.
Gramen annuum 35-50 longum. Vaginae foliorum valde
distichae; laminae 2-3.5 mmlatae, papilloso-ciliatae. Pan-
icula 5-7 cm longa, 2-3 cm lata, cilindrico-pyramidalis;
rami 0.5-1.5 cm longi, verticillati. Spiculae 2.8-3.4 lon-
gae; gluma inferna 1.2-1.8 mm longa; lemma 2.5-2.7
mm longum; palea findens ubi maturum; stamina 3. Utric-
ulus 1.5-1.6 mm longus, 0.5-0.6 latus, valde complan-
atus.
Caespitose annual herb. Culms 35-50 cm tall,
erect, unbranched, glabrous, the internodes solid.
Leaves mostly basal; sheaths keeled, densely over¬
lapping, distichous; ligule 0.1-0.2 mm long, a ciliate
rim; basal blades 5-10 cm long, 2-3.5 mm wide,
folded to involute toward the apex, blunt-tipped,
glabrous except for basal and marginal pustulose-
based elongate hairs; cauline blades ca. 2 mm, much
reduced. Panicle 5-7 cm long, 2-3 cm wide, open,
cylindrical-pyramidal; rachis exposed; branches 0.5
1.5 cm long, verticillate, spreading, somewhat stiff,
spikelet-bearing on the outer Vi\ pedicels 0.2-0.6
mm long, appressed. Spikelets 2.8-3.4 mm long,
brown; glumes, lemma, and palea membranous; low¬
er glume 1.2-1.8 mm long, narrowly lanceolate,
acuminate, 1 -nerved, the margins frequently be¬
coming inrolled; upper glume 2.9-3.4 mm, lanceo¬
late, acute, l-nerved; floret solitary; lemma 2.5
2.7 mm, lanceolate, 1-nerved; palea nearly as long
as the lemma, 2-nerved, split ting as the fruit ma¬
tures; flower bisexual; lodicules 2, glabrous, truncate
to emarginate; stamens 3, the anthers 1-1.2 mm,
yellow; stigmas 2, separate; styles plumose, white.
Utricle 1.5-1.6 mm long, 0.5 0.6 mm wide, some¬
what oblong in side view, strongly flattened in cross
section, apically broadly rounded; pericarp gelati¬
nous when moistened; embryo ca. % as long as the
utricle.
Paratype. Guatemala. PETEN: La Cumbre, in Chacte
Abajo, bordering the river, 15 Feb. 1971, Contreras
10529 (MO).
This species is closely related to Sporobolus er-
ectus A. Hitchc. but differs by its apparently annual
(vs. perennial) habit, the more strongly distichous
leaf sheaths (the basis of the epithet), and smaller
inflorescences (5 7 x 2 3 cm vs. 13-17 x 5-7
cm).
Sporobolus distichivaginatus is so far known
only from river margins below 500 m in the Peten,
Guatemala. Sporobolus credits is known from pine-
oak forests in Chiapas, Oaxaca, and Veracruz, Mex¬
ico, at elevations ranging from 1,200 to 2,300 m.
Paniceae
Eriochloa stevensii Davidse, sp. nov. TYPE: Nic¬
aragua. Granada: 31 km NW of Rio Ochomogo
bridge along Hwy. 1, ca. 11°40'N, 85°59'W,
ca. 60 m, level Crescentia savanna, common,
erect, inflorescence whitish green, 20 Aug.
1981, W. I). Stevens & O. M. Montiel 20608
(holotype, MO; isotypes, BRI, CR, HNMN, ISC,
k, MEXU, SI, ITS). Figure 3A-F.
Gramen annuum 50-100 cm longum. Folia, culmi,
rami inflorescentiae et pedicelli dense velutini. Panicula
14-25 cm longa, 2-5 cm lata; racemi 17-57, 1-3.5 cm
longi. Spiculae 3.4-5.5 mm longae, plerumque binatae,
lanceolatae, acuminatae vel aristatae ad 1.0 mm, pilosae
adpressae infernae 34; flosculus superus 1.9-2.3 mm lon¬
gus, 0.8-1.0 mm latus, ellipticus, apiculo vel arista 0.1-
0.2(—0.4) mm longo.
Annual herb. Culms 50- 100 cm tall, erect, densely
velvety pubescent; internodes hollow. Leaves mostly
cauline, densely velvety pubescent in all parts; sheaths
somewhat keeled toward the apex; ligule with the
membrane 0.1 0.2 mm long and cilia 0.5-1.0 mm
long; blades 10-21 cm long, 4-14 mm wide, linear,
flat. Inflorescence 14-25 cm long, 2-5 cm wide, a
panicle of simple racemes or the racemes rarely with
a short branch bearing solitary spikelets in large
specimens; racemes 17-57, 1-3.5 cm long, as¬
cending, the rachis 0.4-0.5 mm wide; main rachis,
rachis of racemes, and pedicels densely velvety pu¬
bescent; pedicels 0.3 1.0( 2.0) mm. Spikelets 3.4
5.5 mm long, 1.2 mm wide, paired, solitary toward
the apex of a raceme, ascending, lanceolate; basal
callus 0.2-0.3 mm long; upper glume 3.4-5.2 mm,
acuminate or aristate for 1.0 mm, lanceolate,
3-nerved, appressed pilose in the lower 3 A; lower
floret sterile; lower lemma as long as to 1.0 mm
shorter than the upper glume, lanceolate, 3-nerved,
appressed pilose in lower 2 A- 3 A, acuminate; lower
palea absent; upper floret 1.9-2.3 mm long, 0.8-
1.0 mm wide, minutely papillose, elliptic, glabrous
except puberulent at the apex on the apiculum,
obtuse, apiculate or rarely short aristate to 0.2(-0.4)
mm; lodicules 2, truncate; stamens 3, the anthers
1.1-1.3 mm, yellow; stigmas 2, the styles plumose.
Caryopsis 1.1-1.2 mm long, ca. 0.7 mm wide,
broadly elliptic, ± flattened on both the hilum and
embryo sides; hilum ca. l A as long as the caryopsis,
narrowly elliptic; embryo slightly over l A as long as
the caryopsis.
Paratypes. NICARAGUA. Rio SAN JUAN: 0.5 km S of
Rio Oyote, bridge on road to San Miguelito, 11°43'N,
84°58'W, 40 m, 28 Aug. 1983, Ace & Miller 27004
326
Novon
Figure 2. Sporobolus distichivaginatus R. Pohl. —A. Habit. — B. Panicle branch with spikelets. C,
— C. End view. —D. Side view. (A, B based on Contreras 1052V: C, D based on Tun Ortiz 1610.)
D. Utricle.
5mm
Volume 2, Number 4
1992
Davidse & Pohl
New Species of Festuca, Sporobolus,
and Eriochloa
327
Figure 3. Eriochloa species. A-F. E. stevensii Davidse. —A. Flabitat. —B. Portion of leaf in the ligular area
illustrating the velvety pubescence. —C. Raceme with spikelets. —D. Portion of raceme with two spikelet pairs. E,
F. Spikelets. —E. Dorsal view. —F. Ventral view. —G. E. eggersii A. Hitchc., ventral view of spikelet. —H. E.
pacifica Mez, ventral view of spikelet. (A-F based on Stevens cV Montiel 20608; G based on Sagdstegui 2930; H
based on Asplund 5738.)
5mm
328
Novon
(BM, CANB, ISC, MO, VEN). granada: 36 km ENE of
Managua, 18 Sep. 1976, Danin 7()-4-15 (MO).
CIIINANDEGA: Coriato, ca. 0 m, 10 Nov. 1911, Hitchcock
8749 (US). Venezuela. GUARICO: Carretera Dos Cami-
nos-El Sombrero, ca. 10 km, 17 Sep. 1981, Trujillo &
Correa 17823 (MO). Ecuador. Puna, Ostkiiste, 5 m, 13
June 1934, Schimpf 1 184 (MO, US), guayas: Sabanilla,
25-30 m, 12 Apr.' 1951, Solis 20394 (US); Capeira,
km 21 Guayaquil to Daule, 20-200 in, 16 Mar. 1982,
Dodson & Dodson 13003 (MO, SEE). PERU. TlMBES:
Prov. Zarumilla, Dtto. Matapalo, Bosque Nacional de
Tumbes cerca de Campo Verde, along road to Tutumo,
ca. 20 km from Tumbes, 600-800 m, 26 Apr. 1969,
Simpson & Schunke I . 833 (US).
I bis species has often been misidentified as Er-
iochloa pacifica Mez or sometimes as K. eggersii
A. Hitchc., closely related annual species from Ec¬
uador and Peru. Both of these species are similar
to E. stevensii in having densely velvety pubescent
inflorescence branches, paired spikelets, and a sim¬
ilar ligule.
Kriochloa pacifica (Fig. 3H) is a more delicate
plant with fewer (4 17 vs. 17 57), more appressed
racemes, relatively wider and shorter leal blades
that are glabrous to pubescent (but not densely vel¬
vety), modally larger spikelets (5.0 7.3 mm vs. 3.4-
5.5 mm) with longer (1.0 3.0 mm vs. to 1.0 mm)
awns, and pedicels with noticeably elongated tri-
chomes toward the apex versus more or less the
same length.
Kriochloa eggersii (Fig. 3G) is morphologically
more variable than K. pacifica or K. stevensii. It
also tends to have fewer racemes (usually fewer than
15), but some plants may rarely have as many 24.
The leaf blades are similar to K. pacifica in tending
to be relatively shorter and broader and generally
less pubescent. Compared to E. stevensii, E. eg¬
gersii has longer spikelets (5.5-8.1 mm vs. 3.4
5.2 mm), the upper glume is widest at about or
above the middle (vs. below the middle) and strongly
clasped and enrolled over the lower lemma, the awn
of the upper floret is longer (0.3 1.0 mm vs. 0.1-
0.2 mm), and the tip of the upper glume is frequently
bidentate (vs. entire at the apex) in at least some
spikelets of a raceme. However, this may be limited
to a single indentation on one side only, and the
glume margins may he entire in other spikelets ol
the racemes.
Shaw & Webster ( 1987 ) treated Hitchcock 8< 19
collected in Nicaragua as part of E. pacifica and
considered it adventive in Nicaragua. However, it
is probably native in Nicaragua, and is another ex¬
ample of the well-known distribution pattern exhib¬
ited by species of the dry, deciduous forest zone
along the Pacific side of Mesoamerica that reoccur
in Ecuador and continue on into Peru (Gentry, 1982 ).
What is more surprising in this case is its occurrence
on the eastern side of the Andes in Venezuela, al¬
though it occupies a similar vegetation type in the
savannas of the Venezuelan llanos. In Nicaragua it
is found in the dark, heavy clays typical of the
Crescentia savannas.
The callus sometimes has a very narrowly tri¬
angular free lower glume up to 1.1 mm long.
The species is named in honor of Warren Douglas
Stevens, Director of Research of the Missouri Bo¬
tanical Garden and collector of the type collection,
in recognition of his outstanding contributions to our
knowledge of the Nicaraguan llora.
Acknowledgments. Davidse is grateful to the Na¬
tional Geographic Society for financial support, which
enabled him to explore remote areas of the Cordillera
de Talamanca in Costa Rica and Panama. We thank
Linda S. Ellis for the illustrations.
Literature Cited
Alexeev, E. B. 1981. The new taxa of the genus Festuca
(Poaceae) from Mexico and Central America. Bot.
Zhurn. (Moscow & Leningrad) 66: 1492 1501.
-. 1982. A new section and three new species
of the genus Festuca from Mexico and Central Amer¬
ica. Bot. Zhurn. (Moscow & Leningrad) 67: 1289-
1292.
Davidse, G. & R. W. Pohl. 1992. New taxa and no-
menclatural combinations of Mesoamerican grasses
(Poaceae). Novon 2: 81-110.
Gentry, A. H. 1982. Neotropical floristic diversity: phy-
togeographical connections between Central and South
America, Pleistocene climatic fluctuations, or an ac¬
cident of the Andean orogeny. Ann. Missouri Bot.
Card. 69: 557-593.
Shaw, R. B. & R. D. Webster. 1987. The genus Er-
iochloa (Poaceae: Paniceae) in North and Central
America. Sida 1: 165-207.
Podocarpus acuminatus (Podocarpaceae), a New Species from
South America
David J. deLaubenfels
Department of Geography, Syracuse University, Syracuse, New York 13244-1160, U.S.A.
ABSTRACT. A new species, Podocarpus acumi¬
natus, from southern Venezuela and nearby Brazil
is described.
A new species of Podocarpus was recognized in
material recently collected in South America.
Podocarpus acuminatus de Laubenfels, sp. now
I't PE: Brazil. Amazonas: Serra da Neblina,
barney 870 (holotype, MO).
Folia linearia vel ovata, ca. 3-4 cm longa, 8 rririi lata,
acuminata, plus rninusve revoluta; juvenilia 10 cm x 13
mm. Gemma globularis; squamae cuspidatae, 4-6 mm
longae. Tegmina carnosa ovata seminorum crista prom-
inenti, 1 mm longa, praedita; receptaculum 3 bracteis
obtectum.
Small tree 4-5 rn tall. Leaves linear to ovate, ca.
3-4 cm long x 8 nun wide, acuminate, slightly
revolute; juvenile leaves to 10 cm x 13 mm. Foliage
buds globular; bud scales erect, cuspidate, 4-6 nun
long. Fleshy covering of the seed with a prominent
apical crest 1 mm long; receptacle with at least
three bracts: with one seed a third, smaller bract
occurs below the usual sterile bract, with two seeds
one or more smaller bracts occur between the two
fertile bracts. Pollen structures unknown.
Distribution. Grows as an exposed tree in cloud
forests and among rocks, at 1,900-2,400 m.
Acuminate leaves are uncommon in Podocarpus ,
and there are no examples among closely related
species, all of which occur in South America, with
acuminate leaves anywhere near the size of those
of P. acuminatus. Among other Venezuelan species
of Podocarpus (see deLaubenfels, 1982), P. mag-
nifolius is the only closely related one with acu¬
minate leaves, and it is noted for its large leaves
more than 18 mm wide. Similar-sized leaves occur
in P. buchholzii and P. brasiliensis, where the acute
to blunt leaves show no signs of an acuminate apex
(deLaubenfels, 1982). The extra sterile bracts on
the fertile structures are unusual and have not pre¬
viously been observed on any South American spe¬
cies. The prominent crest helps to differentiate P.
acuminatus from the other South American species
with similar-sized leaves. Short, erect, cuspidate bud
scales are also distinctive.
Parntvpes. VENEZUELA. AMAZONAS: Sierra de la Neb¬
lina, .Xee 30743 (MO). BOLIVAR: Chimanta, SW Amuri-
tepui, Huber 11408 (MO).
Literature Cited
Laubenfels, D. J. de. 1982. Podocarpaceae. In: Flora
de Venezuela 11(2): 7-41.
Novon 2: 329. 1992.
A New Species of Brongniartia (Fabaceae: Faboideae) from (aierrero,
Mexico
Oscar Dorado
Rancho Santa Ana Botanic Garden, Claremont, California 91711, U.S.A. and
Universidad Autonoma del Estado de Morelos, Ave. Universidad 1001,
Cuernavaca, Morelos, Mexico 62210
Jaime Bonilla and Armando Burgos
Universidad Autonoma del Estado de Morelos, Cuernavaca
Abstract. A new species, Brongniartia ziran-
darensis (Fabaceae: Faboideae), from Guerrero,
Mexico, is described and illustrated. Its affinities to
B. mollicula Brandegee are discussed, and data are
provided on habitat and abundance.
Best MEN. Se describe y se ilustra una especie
nueva, Brongniartia zirandarensis (Fabaceae: Fa¬
boideae), del estado de Guerrero, Mexico. Se dis-
cuten sus afinidades con B. mollicula Brandegee y
se proporcionan datos de su habitat y abundancia.
Brongniartia is a neotropical genus nearly en¬
demic to Mexico. 01 the approximately 60 species
from this genus, only two, B. minutifolia S. Watson
(southern Texas) and B. ulbrichiana Harms (Boliv¬
ia), are found outside of this country. The Rio Balsas
Basin in southern Mexico is an important center ot
diversification for the genus. The state of Guerrero,
with most of its territory included in this region,
contains about ten species known from the genus;
four of these are endemic to the state. Two new
species were recently described Irom Guerrero, one
endemic, B. guerrerensis (Jimenez & Contreras,
1989), and the other distributed on the Rio Balsas
Basin, B. montalvoana (Dorado & Arias, 1992).
During a systematic study of the Podalyrioides group
of Brongniartia (Dorado, 1988), some specimens
were collected in northern Guerrero that did not
correspond with any of the previously known taxa.
These collections are proposed here as representing
a new species.
Brongniartia zirandarensis 0. Dorado, J. Bo¬
nilla & A. Burgos, sp. nov. TYPE: Mexico.
Guerrero: Mpio. Zirandaro, “El Mirador,” 17.6
mi. W of Zirandaro on dirt road to El Tarna-
rindo, 2 Sep. 1991, (). Dorado , Bonilla &
A. Burgos 2115 (holotype, MEXU; isotypes,
ENCB, HUMO, MO, RSA, XAE). Figure 1.
Brongniartia molliculae maxime similis, sed differt
ramis juvenibus glabratis, stipulis reniforinibus versus ra-
morum basem, 8 vel pluribus foliis in ramis juvenibus,
17-21 foliolis per folium, bracteolis 2-3 mm longis, et
calyce 0.9-1.3 cm longo.
Small shrub to 1.5 m tall, with grayish bark, the
younger branches yellowish green and glabrous.
Stipules (0.7-) 1-2.3 cm long, (0.4 ) I 2.2 cm wide,
texture foliaceous or slightly coriaceous in older
branches, glabrate or sparsely pilose, obliquely ovate
or elliptic at end of branches and reniform at base
of branches, persistent, venation conspicuous, es¬
pecially at the upper surface, lower lobe present and
forming a (5° )35° to 80°(-90°) angle, with a cordate
base, upper lobe present and forming a 90° to 135°
angle, the edge of the stipule adjacent to the petiole
excurvate, apex obtuse to rounded. Leaves 7.5
11(—17.5) cm long, glabrate; petiole 0.9-1.7 cm
long, terete, glabrate; rachis 8.2 1 3( 14.7) cm long,
glabrate, with 17-21 opposite or subopposite leaf¬
lets; stipels 0.6 0.8 mm long, commonly absent;
leaflets 1.4-2.4(-2.8) cm long, 0.6— 1 -6(— 1.8) cm
wide, petiolule 1 2 mm long, glabrate, lamina elliptic
or sometimes obovate, texture rather thin, glabrate,
base acute or sometimes rounded, apex acute to
rounded, with a mucro 0.5-1 mm long. Flowers
borne solitary at the base of leaf-bearing nodes;
peduncles (0.8-)1.3-2 cm long, erect, glabrate;
braeteoles 2-3 mm long, 0.2 0.3 mm wide, yel¬
lowish, subulate, normally persistent, opposite, with
some whitish hairs along the margin; calyx 0.9-1.3
cm long, 0.4-0.6 mm wide, sparsely pilose, cam-
panulate, two vexilar lobes 7-8 mm long, 4.5-5
mm wide (both lobes), connate % of their length;
lateral lobes 5.5-7 mm long, 2.5-3 mm wide, ca-
rinal lobe 6-7 mm long, 2-2.5 mm wide; standard
1.3—1.5 cm long, 2-2.3 cm wide, red, reflexed ca.
90°, widely obovate and becoming narrower at the
base, slightly auriculate, the apex cleft to 2 mm,
claw 4-4.5 mm long, 2-3 mm wide, strongly curved;
wing petals 1.7-1.8 cm long, 0.65-0.7 cm wide.
Novon 2: 330-322. 1992.
Volume 2, Number 4
1992
Dorado et al.
Brongniartia zirandarensis
331
Figure 1. Brongniartia zirandarensis 0. Dorado et al. (from O. Dorado et al. 2145). ■ —a. Branch with flower
and fruit. —b. Standard, front view. —c. Standard, face view. —d. Wing petal. —e. Keel petal. —f. Gynoecium.
Scale bar = 2 cm in a; 1 cm in b-f.
332
Novon
obliquely oblanceolate, red, claw 2-2.5 mm long,
about 1 mm wide; keel petals 1.7-1.8 cm long,
0.9 0.95 mm wide, pinkish, lunate, the ventral lace
straight, claw 2.5-3 mm long, about 1 mm wide,
auricula of the keel forming a 95°-l 10° angle with
the claw; androecium 2.1 2.2 cm long, anthers
1.8 2 mm long; ovary about 0.7 cm long, glabrous,
with ati obconical staminal sheath 0.9 1.1 mm long
at the base of the ovary, with 3-5 ovules, style 1.5
cm long, glabrous, sometimes slightly exserted from
the keel. Fruit 4.3-5 cm long, 1.7-1.9 cm wide,
oblanceolate, rather sessile, mucro about 3.5 mm
long, usually with 3 5 seeds (only immature seeds
known).
Paratypes. Mexico, cuerrero: Mpio. Zirandaro, “El
Mirador,” 17.6 mi. W of Zirandaro on dirt road to El
Tamarindo, 15 Dec. 1991, J. Bonilla , A. Burgos & J.
I tana 2134 (HOMO, MEXU, MO, RSA).
Brongniartia zirandarensis is most similar to B.
mollicula Brandegee, a species distributed in south¬
ern Mexico, in the states of Puebla and Oaxaca.
Both species share several morphological features,
such as the presence of thin leaflets, short stipels
(inconspicuous in most cases), presence ol short,
leafless, flowering branches (more commonly absent,
the flowers then occurring at leaf-bearing nodes),
and red to maroon flowers. However, the two species
can be distinguished by several morphological char¬
acters. The leaves of Brongniartia zirandarensis
bear 17-21 leaflets; the young branches are gla¬
brous and bear 8 or more leaves with reniform
stipules; the bracteoles are 2-3 mm long and the
calyx is 0.9 1.3 cm long. Conversely, the leaves ol
B. mollicula bear 5 1 3 leaflets; the young branches
are pubescent and bear fewer than 6 leaves, these
with obliquely ovate stipules; the bracteoles are 0.2
0.6 mm long, and the calyx is (1.2 ) 1.4 1.7 cm
long.
Brongniartia zirandarensis is known only from
the type locality in tropical deciduous forest at 240
m. The dominant trees are Acacia spp., Bursera
spp., Caesalpinia pulchcrrima (L.) Sw., Lysiloma
tergemina Benth., Pithcccllobium dulce (Roxb.)
Benth., Jacquinia pungens A. Cray, and Cordia
clacagnoidcs DC. The first collections ol this species
were made from about ten individuals growing on
disturbed edges of a road cut. In a second search
for more populations (J. Bonilla el al. 2134), fewer
individuals were encountered. This was apparently
due to recent road widening activity. Although the
rarity of B. zirandarensis may be explained by the
lack of explorations in the area, the low number of
individuals suggests that this species may be one of
the rarest of Brongniartia.
The specific epithet is derived from the nearby
town of Zirandaro and pays homage to the friendly,
helpful residents of that town.
Acknowledgments. We thank S. Boyd, A. Del¬
gado, T. S. Elias, C. T. Philbrick, T. Boss, M. Sousa,
I). Thompson, and J. E. Villasenor for valuable sug¬
gestions regarding the manuscript. J. L. Villasenor
kindly provided the Latin diagnosis. We also thank
.1. Viana for assistance in the field during the second
search for more populations of this species. We are
grateful to G. Avila, Director ol Research at the
University of Morelos, for providing logistic support
for fieldwork.
Literature Cited
Dorado, O. 1988. A morphological revision of the Po-
dalyrioides group of Brongniartia (Fabaceae: Fa¬
boideae). M.Sc. Thesis. The Claremont Graduate
School, Claremont, California.
- & I). M. Arias. 1992. Brongniartia montal-
voana (Fabaceae: Faboideae), una especie nueva de
la Cuenca del Rio Balsas en el sur de Mexico. Acta
Bot. Mex. 17: 13-17.
Jimenez, J. & J. L. Contreras. 1989. Una especie nueva
de Brongniartia (Leguminosae: Faboideae) de Guer¬
rero, Mexico. Anales Inst. Biol. Ser. Bot. UNAM
(Num. Unico). Vol. 58: 63-68.
New Species of Woody Plants from Amazonian Peru
Alwyn II. Gentry
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A.
Abstract. New species of Ancistrothyrsus (Pas-
sifloraceae), Spathelia (Rutaceae), and Allophylus
(Sapindaceae) from Amazonian Peru (and adjacent
regions) are described and discussed. In addition, a
new combination is made to elevate Allophylus pi-
losus to specific rank.
Miscellaneous new species from Amazonian Peru
have accumulated in my files over the past decade.
Unpublished names for some of these species have
been used in the herbarium, hut formal publication
has been hitherto postponed in hopes ot obtaining
additional material or more complete field obser¬
vations. The names are now needed for inclusion in
the Catalogue of the Flowering Plants and Gym-
nosperms of Peru (Brako & Zarucchi, in prep.).
Passifi.oraceak
Ancistrothyrsus hirtellus A. Gentry, sp. nov.
TYPE: Venezuela. Amazonas: trail S from Cer-
ro Neblina base camp on Rio Mawarinuma,
from clear water stream to top of first small
bills, 150-350 m, 0°50'N, 66°1 l'W, 3 May
1984 (fr). Gentry & Stein 47114 (holotype,
MO; isotypes, AAU, HUA, MO, NY, \ EN).
Figure 1.
Frutex scandens, ramulis hirtellis. Folia lamina obovata
vel oblongo-elliptica, 7-24 cm longa, 3.5*12 cm lata,
infra dense hirtella, supra sparsim hirtella. Inflorescentia
axillaris, in cirrhum desinens, ramulis fertilibus lateralibus.
Flores albi, sepalis lanceolatis dense puberulis, petalis lan-
ceolatis ca. 2.5 cm longis, staminibus ovarioque in an-
drogynophoro portatis. Capsula 4-valvata, dense puberula.
Canopy liana, the branchlets terete, persistently
hirtellous with somewhat crisped trichomes, with
axillary tendrils having thickened curved tips. Leaves
with blade obovate to oblong-elliptic, rounded at
apex, broadly cuneate to rounded at base, 7-24 cm
long, 3.5-12 cm wide, densely and persistently hir¬
tellous with erect or suberect trichomes below, also
with scattered peltate scales, above conspicuously
peltate-lepidote with reddish scales and sparsely hir¬
tellous over surtace and more persistently along
main veins, the petiole 0.8-1 cm long, adaxially
grooved, rather densely pubescent with subap-
pressed-rufescent trichomes, arising from a con¬
spicuously raised projection from the branchlet. In¬
florescence axillary, the short fertile branches borne
laterally near a hooked or coiled apical tendril, the
peduncle reddish hirtellous, 4 13 cm long. Flowers
white, the sepals lanceolate or lanceolate-elliptic,
acute, densely tannish puberulous, ca. 3 cm long,
7-8 mm wide, persistent in lruit, the petals lanceo¬
late, ca. 2.5 cm long, the ovary and gynophore
encircled by a corona formed from a ring of fila¬
mentous staminodia basally fused and densely pu¬
bescent, the stamens with filaments arising from base
of ovary at apex of ca. 4-mm-long androgynophore,
at least 6 mm long (anthers and filament apex eaten
in only mature flowers examined), the anthers in
bud 1.5-2 mm long, dorsifixed, not apiculate, the
ovary ovoid, densely villous, topped by four slender
style branches. Capsule yellow at maturity, 4-valved,
ellipsoid, acute at apex, conspicuously stipitate at
base, 6-9 cm long, 3.5-4 cm wide, the stipe 1 2
cm long, densely tannish puberulous with both short
crisped hairs and longer suberect ones as well as
numerous reddish peltate glands, the valves woody
but fragile, ca. 3 mm thick, pulp completely lacking,
the seeds 12-14 mm long, 1 mm wide, borne 2-5
together along the midline ol each valve, flattened
and obovate with a shallowly pitted surface, not
arillate.
Distribution. Poor soil terra firme areas of central
and northwestern Amazonia, below 200 m.
Paratypes. Venezuela. AMAZONAS: Cerro Neblina base
camp, Rio Mawarinuma, 0°50'N, 66°1 l'W, mature forest
on sandy “ultisol” and along clear water stream, 1 40 m,
2 May 1984, Gentry & Stein 47103 (MO, VEN). PERU.
I.ORETO: Prov. Loreto, Dept. Tigre, Rio Corrientes, Shi-
viyacu, camino a Forestales (38 km), 25 Nov. 1979 (fl,
fr), Ayala 2334 (AMAZ, MO). Brazil, amazonas: Es¬
trada do Rais, silva secundaria non inundabili, 23 Nov.
1936 (fl), Ducke 290 (NY); Igarape do Binda, terra firme,
solo argiloso, mata virgein, Rodrigues & Chagas 4074
(1NPA 10639) (NY).
This species is not uncommon in the vicinity of
the Cerro Neblina base camp, as judged by fallen
fruits in three different places along the limited trail
system explored. All plants were canopy lianas grow¬
ing in tangles of other lianas in the top of a canopy
tree and invisible from the ground. We were only
able to locate the leaves and attached fruits of the
Novon 2: 333-338. 1992.
334
Novon
C
Figure 1. Ancistrothyrsus fruits. A, B. A. hirtellus
(Gentry & Stein 47114). —A. Habit. —B. Fruit, close
up. —C. A. tessmannii (Gentry et al. 55911 , from
Indiana, Loreto, Peru). Shears are 21 cm long.
type, recognized in the field as a new species, after
climbing several nearby trees.
The hitherto monotypic genus Ancistrothyrsus
Harms, previously known only from a few collections
from Peru and Brazil, is of unusual phylogenetic
and phvtogeographic interest. Ancistrothyrsus is
the only neotropical representative of the otherwise
palaeotropical, mostly African, tribe Paropsieae, with
features intermediate between Flacourtiaceae and
Passifloraceae (DeWilde, 1971). Moreover, Ancis¬
trothyrsus is the only climbing member of Parop¬
sieae, and is closer to Passifloraceae sensu stricto,
with which it shares such unusual characters as
tendril-bearing inflorescences, than is any other
member of Paropsieae. Ancistrothyrsus was de¬
scribed in Flacourtiaceae and is retained under that
family in most herbaria and in the Flora oj Peru
(Macbride, 1941), although modern African floristic
authors (e.g., DeWilde, 1975; Fernandes & Fer¬
nandes, 1978) generally treat the tribe Paropsieae
under Passifloraceae. DeWilde (1971) has shown
convincingly that Ancistrothrysus , too, is best treat¬
ed under that family, a conclusion accepted by Keat¬
ing (1973) and Sleumer (1980).
Examined Peruvian material of A. tessmannii
Harms (including an isotype, Tessmann 1479 (NY)),
differs conspicuously from the plant described above
in having the mature leaves uniformly glabrous,
except for the lepidote glands. Juvenile leaves have
short, appressed hairs different in form from the
much longer, erect ones of A. hirtellus.
The mature fruit of A. tessmannii (and thus of
the genus) has never been described. Harms (1931)
merely noted in the original description that juvenile
fruits are subglobose and densely hirsute. The ma¬
ture fruit is globose or depressed-globose, 4 9 cm
long, 4 8 cm in diameter, densely and finely tannish
puberulous, with the pericarp 3 4 mm thick. The
seeds are flattened-obovoid, 1-1.5 cm long, and
0.9-1 cm wide. Thus the fruit of A. tessmannii is
quite different from that described here for A. hir¬
tellus, which differs in being much narrower and
more ellipsoid with a more acute apex, a longer
basal stipe, and more strongly tomentose valves.
There is no doubt that these very different fruits
belong to different species.
Interpretation of the limited available floral ma¬
terial is more problematic. DeWilde (1971) studied
th ree collections, one in fruit and two in flower. He
emphasized that the two flowering collections, Ducke
24385 and 35681 (neither of which I have ex¬
amined), had very different flowers, differing in flow¬
er size and in apiculate versus nonapiculate anthers,
fhe large-flowered specimen (Ducke 21385) had
tepals ca. 4 cm long and nonapiculate anthers; the
small-flowered specimen (Ducke 35681) had tepals
less than 2 cm long and long-apiculate anthers. It
would be tempting to suppose that these two flower
morphs correlate with the two different species rec¬
ognized here, except that a single stamen of the two
flowers of Klug 650 and Ducke 639 each has a
long, conspicuous apicule, while the remainder of
the stamens of these flowers are nonapiculate.
DeWilde gave no indication of what other features
correlate with his two flower types. However, since
both the glabrous-leaved collections here referred
to A. tessmannii and the pubescent-leaved ones
referred to A. hirtellus have large tepals, flower size
Volume 2, Number 4
1992
Gentry 335
New Woody Plants from Amazonian Peru
may not be a useful differentiating character. One
of the collections cited above, Rodrigues & Chaves
4074, is not included in the description of A. hir-
tellus. It is dramatically different from all other
material examined in the 3-4-mmdong trichomes
that are interspersed with the normal reddish hir-
tellous ones on its branchlets. These long trichomes
have short, reddish lateral branches and resemble
miniature bottle-brushes. The examined duplicate of
the Rodrigues collection has only buds, hut if Du eke
35681, the small-flowered specimen with apieulate
anthers illustrated by DeWilde, should prove to share
this unusual pubescence, these two collections would
merit specific recognition.
The two Ancistrothyrsus species are apparently
allopatric, but they are rarely collected and their
geographical distributions are not entirely clear. All
but one of the thirteen Peruvian Ancistrothyrsus
collections examined belong to A. tessmannii, in¬
cluding material from Loreto, Amazonas, and San
Martin departments. Roth species also occur in Bra¬
zil: Ducke 630 (MO, NY), from Sao Paulo de Oli-
vernja near the Colombian border, is typical gla¬
brous-leaved A. tessmannii, but Ducke 290 (NY),
from near Manaus, is A. hirtellus. One of the spec¬
imens illustrated by DeYi ilde (1971) (Ducke 2138 /,
provenance not indicated) has ellipsoid stipitate fruits
and hirtellous branchlets and would seem referrahle
to A. hirtellus. Since the two flowering collections
examined by DeWilde had very different floral mor¬
phologies, it is possible that his two flower morphs
correspond to the two different species of Ancis¬
trothyrsus. Apparently, A. hirtellus is restricted to
the poor soil area of central and northwest Ama¬
zonia, while A. tessmannii occurs in the lateritic soil
areas of the Solimoes and Peruvian Amazonia. In
the Iquitos area I have seen A. tessmannii only in
mature forests on relatively fertile soils.
Rutaceae
Spathelia terminalioides A. Gentry, sp. nov.
TYPE: Peru. Loreto: Mishana, Rio Nanay half¬
way between Iquitos and Santa Maria de Na¬
nay, 140 m, mature forest on upland white
sand near Campamento Uno, 73°30 , W, 3°50 , S,
25 Feb. 1981 (fl, fr). Gentry et at. 31751
(holotype, MO; isotypes, AMAZ, USM). Fig¬
ure 2.
Arbor eramosa. Folia pinnata, rhachidi in quoque latere
foliolos 14-39 vel ultra gerenti, foliolis alternis, oblongo-
lanceolatis, acutis, 2-19 cm longis, 0.6-4 cm latis, gla-
bratis. Inflorescentia terminalis, late paniculata. Flores
feminei albi, sepalis ovatis 1 mm longis puberulis, petalis
ellipticis 3-4 mm longis, staminodiis 1.5-2 mm longis
filamentis alatis, ovario complanato orbiculato. Fructus
planus, 2-alatus, transverse oblongus, 1.4-2 cm longus,
3-4 cm latus.
Tree ca. 30 m, unbranched, slender, with a ter¬
minal cluster of large pinnate leaves surrounding
the large paniculate terminal inflorescence. Leaves
multifoliolate (14-39+ leaflets (or leaflet scars) per
side), the leaflets alternate, entire, oblong-lanceolate,
asymmetrically obtuse at base, acute at apex, 2-
I 9 cm long, 0.6-4 cm wide, with scattered glandular
punctations, glabrous except for minute trichomes
on the petiole and petiolules, at base of midvein
above, and very sparsely scattered over lower sur¬
face when young, the petiole slightly flattened to
strongly grooved above, flattened at base (the en¬
larged petiole base to at least 3.5 cm across) and
forming conspicuous ridges on stem. Inflorescence
large, openly paniculate, minutely puberulous, the
bracts virtually absent, mostly reduced to tiny tri¬
angular flaps of tissue, the pedicels slender, minutely
puberulous, 3-7 mm long. Flowers (only female
seen) white, the sepals ovate, obtuse, ca. 1 mm long
and wide, somewhat puberulous; petals elliptic, 3-
4 mm long, rounded at tip, glabrous, staminodes
1.5-2 mm long, flat and incurved, the basal part
of filaments strongly winged, the wing bilobed at tip,
villous adaxially; pistil with the ovary flattened and
orbicular, ca. 3 mm long, glabrous except the pu¬
berulous, thickened, 1-mm-long basal gynophore,
the stigma sessile. Fruit flat, broadly 2-winged, trans¬
versely oblong and much broader than long, 1.4 2
cm long, 3-4 cm wide (narrower and rhombic when
immature before the wings have expanded), gla¬
brous, the central seed-hearing portion somewhat
swollen, completely surrounded by the wing, heart-
shaped if with 2 seeds or tear-drop-shaped if with
one.
Distribution. Known only from the type collec¬
tion from an ecologically distinctive area of white
sand forest on the Rio Nanay near Iquitos, Peru.
This wing-fruited genus is not typical of Rutaceae,
and at first I thought the Peruvian plant might
represent a new genus of Sapindaceae. Instead, this
tree turns out to belong to a mostly West Indian
genus otherwise known in South America only from
the Guayana Shield area and central Amazonia.
Most of the South American species are locally
endemic, small trees of the tepui summits (Cowan
& Brizicky, 1960).
Most of the West Indian species are quite dif¬
ferent from Spathelia terminalioides, with narrow¬
er, 3-winged, fruits, while the Guayana Highland
material I examined is similarly 2-winged hut with
wings that are thicker and less developed. There is
also a single lowland Amazonian species, S. excelsa
336
Novon
(K. Krause) R. Cowan & Briz., which is the nearest
relative of the* Peruvian species. That species is
known only from a few collections from central
Amazonian Brazil (Albuquerque, 1977). Spathclia
excelsa differs from S. terminahoides in having a
tannish villous inflorescence and in larger (2.5 3.5
x 4.5-6 cm) fruits with differently shaped wings.
I he fruit wings of ,S. terminnlioides are somewhat
triangular in outline, with relatively narrow tips and
a straight ventral margin, while S. excelsa has more
Volume 2, Number 4
1992
Gentry 337
New Woody Plants from Amazonian Peru
rounded wing apices and a prominently cordate ven¬
tral margin. The fruits of S. terminalioides are
similar in form to those of such Terminalia species
as T. obovata (Ruiz & Pavon) Steudel, hence the
specific epithet.
The area in which S. terminalioides w'as discov¬
ered has white sand soil and many disjuncts from
the upper Rio Negro area, where similar soils pre¬
dominate. Thus the discovery of a Guayana genus
halfway across South America from its previously
known range is perhaps not as surprising as it might
seem at first.
Sapindaceae
Allophylus multicostatus A. Gentry, sp. nov.
TYPE: Peru. Amazonas: Aintami Creek, Rio
Cenepa, 550 ft., 24 Nov. 1972 (fl), Berlin .'lib
(holotype, MO; duplicates distributed as A. di-
varicatus Radik.).
Arbor 5-15 m alta. Folia 3-foliolata, foliolis ellipticis
vel obovato-ellipticis, plus minusve serrulatis, puberulis
praecipue secus venas. Inflorescentia axillaris, pyramido-
paniculata, tornentosa, pedicellis minus quam 1 mm longis.
Flores ca. 1 mm longi, plus minusve albi, sepalis subor-
biculatis, puberulis, petalis ca. 1 mm longis, starninibus
fertilibus 6-7, exsertis. Fructus ellipsoideus vel globosus,
8-9 mm longus, 7-9 mm latus, sparsim puberulus.
Tree 5-15 m tall, with sparsely puherulous and
reddish punctate branchlets. Leaves 3-foliolate, the
leaflets with blade elliptic to obovate-elliptic, acute
to short-acuminate, broadly cuneate to obtuse at
base, 7-25 cm long, 3.5-12 cm wide, firmly mem¬
branaceous, the margin serrulate to shallowly serrate
or essentially entire, sparsely puherulous mostly along
veins below and along midvein above, with 12 23
secondary veins on a side, these rather straight and
parallel, the petiole 3.5-12 cm long, sparsely to
conspicuously tannish tomentose. Inflorescence ax¬
illary, openly pyramidal-paniculate with at least the
lowermost of the generally 5- 9 primary lateral
branches several-branched, tannish tomentose in
flower, partially glabrescent in fruit, mostly over 1 5
cm long; pedicels < 1 mm long. Flowers ca. 1 mm
long, whitish to greenish, the suborhicular sepals
somewhat appressed-puherulous, the petals ca. 1
mm long, glabrous except the inconspicuously ciliate
margins, the 6-7 fertile stamens exserted almost 1
mm beyond perianth, the anthers short. Fruits broadly
ellipsoid to globose, greenish yellow (mature?), ca.
8-9 mm long, 7-9 mm diam., sparsely and incon¬
spicuously appressed-puberulous.
Distribution. Lowland Amazonian Peru (and
probably adjacent Ecuador).
Paratypes. Peru. AMAZONAS: Rio Cenepa region, Ber¬
lin 309, 346, Tunqui 204, Kayap 115, Huashikat 1473
(all MO).
bocal names. “Suhi knum” (Berlin 300, 316),
“tsanignum” (Tunqui 204), “jimagma” (Kayap 115:
probably confused with sympatric A. floribundus,
for which this name is usually used), “wayu” (Hu¬
ashikat 1473).
The most noteworthy features of this plant are
its large, open, much-branched panicles and the
relatively close together and straight secondary ve¬
nation of its usually large and dark-drying leaflets.
Following Radlkofer (1900, 1932), this species keys
to Allophylus paniculatus because of the well-de¬
veloped paniculate inflorescence with branched lat¬
eral branches, but it differs from that species in the
puherulous tomentum of twigs, leaves, and inflores¬
cence. If the inflorescence is taken as merely 3-6-
branched, A. multicostatus keys to A. divaricatus
and I. cinnamomeus. In addition to the unbranched
inflorescence branches, A. divaricatus differs from
A. multicostatus in densely and conspicuously tan-
nish-puberulous petioles and leaf undersides (es¬
pecially the main veins), and in having generally
more membranaceous leaflets with the terminal one
tapering gradually to the base and distinctly differing
in shape from the smaller laterals; A. cinnamomeus
differs in its smaller, sharply serrulate leaflets and
much larger (3 mm wide) flowers.
Using the Flora of Peru (Macbride, 1956), Al¬
lophylus multicostatus keys to I. divaricatus be¬
cause of its 3-foliolate leaves and laxly branched
inflorescence. However, the usually larger leaves
distinguish A. multicostatus from that species; large-
leaved specimens have the largest leaflets of any
Peruvian Allophylus. The smaller (maximum 9 cm
wide) leaves of A. divaricatus also dry a different
color (± olive below vs. brownish to blackish brown
in A. multicostatus) and mostly have fewer sec¬
ondary veins (10 1 3(—15) vs. 12-23). Another dif¬
ference is in the fruits; the only fruiting collection
I have examined of A. divaricatus has smaller,
densely puherulous fruits, which are very different
from the large, glabrescent ones of A. multicostalus.
The only neotropical species published since Mac-
bride (1956) are Bolivian Allophylus steinbachii F.
Barkley and several Cuban species. I have not com¬
pared this species with the Cuban or palaeotropical
taxa, which I assume are different on phytogeo¬
graphic grounds.
Two collections with somewhat smaller leaves and
inflorescences and entire leaflet margins are included
in this circumscription and description because of
the otherwise similar aspect and branching of the
basal inflorescence branches.
338
Novon
Allophvliis pilosus(J. F. Macbr.) A. Gentry, comb,
et stat. nov. Allophylus semidentalus (Miq.)
Radik, var. pilosus .1. F. Macbr., Field Mus.
Nat. Hist., Rot. Ser. 13: 376. 1956. TYPE:
Peru. San Martin: Juan Jui, Rio Huallaga, 400
in, tree 6 m, flowers white. King 3783 (holo-
type, F; isotype, MO).
Small tree 3-10 (-20) m tall with rufous pilose
twigs, the trichomes 1 2 mm long. Leaves 3-foliolate,
the leaflets petiolulate, obovate (terminal) or asym¬
metrically oblong-ellliptic (laterals), 6 30 cm long,
2.5-13 cm wide, membranaceous, the margin ser¬
rate, densely pubescent below; petiole 5 26 cm long,
conspicuously rufous pilose. Inflorescence simply
racemose, densely rufous pilose. Flowers ca. 2 mm
across, the petals and sepals ca. 1.5 nun long. Fruits
ellipsoid, pilose, 10—13 mm long, 6-10 mm wide,
orange at maturity.
Additional specimens examined. Ecuador. NAt’o: near
Rio San Miguel, 21.2 km N of Lago Agrio, 470 in, 0°8'N,
76°50'W, (shrub, flowers in bud), Croat 50363 (MO).
Pk.RU. AMAZONAS: Rio Santiago, Quebrada Caterpiza, 200
in, 77°40'W, 3°50'S, (arbol 3 m, frutos verdes), Tunqui
642 (MO), (arbol 4 m, frutos verdes), Tunqui 078 (MO),
(arbol 4 m, flores 11-4), Huashikat 584 (MO), Huashikat
1657 (MO), huanuco: Bosque Nacional de Iparia, 1 km
arriba del pueblo de Tournavista, Prov. Pachitea, 300
400 in, ./. Schunke I . 17867 (MO). I.ORETO: Quebrada
Yanomono, Explorarna tourist camp, ca. 72°48'\V, 3°28'S,
120-130 m, (tree 10 m tall, 2" D.B.H., fruits green),
Gentry et al. 27846 (AMAZ, MO), (tree 4 m, fruits
orangish), Gentry et al. 20065 (AMAZ, MO), (arbol 20
m, frutos verdes y marrones, pubescentes), Diaz et al.
1184 (AMAZ, MO). MADRF. DE dios: Tambopata, tropical
moist forest on alluvial soil along Rio Tambopata, 280
m, 12°49'S, 89°18'W, Gentry et al. 45750 ( MO), i cayali:
Provincia Coronel Portillo, Rio Abujao, Quebrada Shesha,
73°45'W, 8°20'S, Diaz et al. 830 (AMAZ, MO).
Local name. “Tsanignum” (Aguaruna).
In his Flora of Peru Sapindaceae treatment, Mac-
bride (1956) referred only one specimen to A. sem-
identatus (Miq.) Radik., the pilose King specimen
that he designated as the type of variety pilosus.
Macbride noted that Standley had originally pro¬
posed the unpublished name A. pilosus for the Klug
collection, but “since ex char, it seems too near A.
semidentalus, I propose it myself as merely a vari¬
ant
Recent Peruvian material collected by the Flora
of Peru project includes not only the above-cited
1 1 additional specimens of this taxon with distinc¬
tive, conspicuously rufous-pilose indumentum, but
also nine collections of the real A. semidentalus,
characterized by a short puberulous indumentum.
The two taxa prove consistently distinct. In addition
to the different indumentum, A. semidentalus has
much smaller, narrower leaflets with a less pro¬
nounced reticulation of tertiary venation below,
smaller, glabrous fruits, and a more slender inflo¬
rescence axis. Its leaves usually dry a different darker
color both above and below than do those ol A.
pilosus, and the inflorescences tend to branch near
the base, whereas those of A. pilosus are uniformly
simple racemes.
Acknowledgments. The fieldwork that led to dis¬
covery of these species was funded by the National
Science Foundation, the National Geographic So¬
ciety, and the Mellon Foundation. Preparation ol
the manuscript was completed with the support of
a Pew Foundation Conservation and the Environ¬
ment Fellowship.
Literature Cited
Albuquerque, B. W. P. de. (1976) 1977. Revisao tax-
onomica das Rutaceae do Estado do Amazonas. Acta
Amaz. 6(3): Supl. 1-67.
Cowan R. S. & G. K. Brizicky. 1960. Taxonomic re¬
lationships of Diomma Engler ex Harms. Mem. New
York But. Card. 10: 58 64.
DeWilde, W. J. J. O. 1971. The systematic position
of tribe Paropsieae, in particular the genus Ancis-
trothyrsus, and a key to the genera of Passifloraceae.
Blumea 19: 99-104.
-. 1975. Passifloraceae. In: R. M. Polhill (edi¬
tor), Flora of Tropical East Africa. Crown Agents for
Overseas Governments and Administrations, London.
Fernandes, R. & A. Fernandes. 1978. Passifloraceae.
In: E. Launert (editor), Flora Zambesiaca 4: 368-
411. Robert MacLehose, Glasgow.
Harms, H. 1931. Eine neue Gattung der Flacourti-
aceae. Notizbl. Bot. Gart. Berlin-Dahlem 11: 146
149.
Keating, R. C. 1973. Pollen morphology and the re¬
lationships of the Flacourtiaceae. Ann. Missouri Bot.
Gard. 60: 273-305.
Macbride, J. F. 1941. Flacourtiaceae. In: Flora of Peru,
Field Mus. Nat. Hist., Bot. Ser. 13(4): 5 52.
-. 1956. Sapindaceae. In: Flora of Peru, Field
Mus. Nat. Hist., Bot. Ser. 13(3A): 291-391.
Radlkofer, L. 1900. Sapindaceae. In: C. Martius, Flora
Brasiliensis 13(3): 225-658.
-. 1932. Sapindaceae II. In: A. Engler, Das
Pflanzenreich IV. 165 (Heft 986): 321-640.
Sleumer, H. O. 1980. Flacourtiaceae. Flora Neotrop.
Monogr. 22: 1-499.
A New Combination in Psiguria (Cucurbitaceae) from Mesoamerica
Rachel Jane Hampshire
The Natural History Museum, Cromwell Road, London SW7 5BD, England
ABSTRACT. The new combination Psiguria dun-
lapii is made, and a key to the three Mesoamerican
species of Psiguria is given.
Psiguria dunlapii (Standley) K. J. Hampshire,
comb. nov. Basionym: Angaria dunlapii Stan¬
dley, Field Mus. Publ., Bot. Ser. 4: 298. 1929.
TYPE: Panama. Bocas del Toro; Farm Six,
Changuinola Valley, 3 Mar. 1924, 1 . (.. Dun¬
lap 490 (holotype, F not seen; isotype, F).
Jeffrey (1962) showed that the generic name
Anguria Jacq. was illegitimate. Howard (1973)
transferred three species from Anguria to Psiguria ,
and the name Psiguria is widely accepted by most
authors, e.g., Wunderlin (1978), Jeffrey (1978),
Condon & Gilbert (1990).
In preparing a treatment of the genus Psiguria
for Flora Mesoamericana, it became clear to the
author that the combination Psiguria dunlapii was
needed. Wunderlin (1978) used the combination P.
longipedunculata (Cogn.) Wunderlin (svn. Anguria
longipedunculata Cogn.) in his treatment for Horn
of Panama, including Anguria dunlapii as a syn¬
onym. Psiguria dunlapii is known only from the
type, collected in Panama, while Cogniaux (18* .)
had described Anguria longipedunculata from Ca-
temaco, in the state ol Veracruz, Mexico, based on
the specimen Galeotti s.n. (BR).
However, Psiguria dunlapii and P. longipedun¬
culata are very different. Psiguria dunlapii has
leaves that are markedly truncate to slightly cordate
at the base, while P. longipedunculata leaves are
cuneate to truncate. The leaf apex in P. dunlapii
is shortly acuminate (the acumen 0.6-2 mm, slen¬
der) while P. longipedunculata has leaves that are
long-acuminate (the acumen 7-15 mm, broad). The
peduncle of P. dunlapii is 20-21 cm, while that of
the type of P. longipedunculata is 37 cm. The type
specimen of P. longipedunculata had one few-flow¬
ered inflorescence; that ol P. dunlapii had two, one
of which was badly insect-damaged. Because both
were on loan from other institutions, no flowers were
dissected. The type specimen ol P. dunlapii is clear¬
ly distinct from P. longipedunculata and does not
appear to match material of any other species ol
Psiguria with unlobed leaves. It therefore appears
to be a distinct species, although a more detailed
study, beyond the scope of a treatment for Flora
Mesoamericana , might show P. dunlapii to be a
synonym of P. warscewiczii.
The Galeotti collection of Psiguria longipedun¬
culata, which is clearly different from P. dunlapii ,
matches material of P. triphylla with unlobed leaves.
Psiguria longipedunculata should be considered a
synonym of P. triphylla.
Wunderlin (1978) included under Psiguria lon¬
gipedunculata a specimen from Llanuras de Santa
Clara, Gosta Rica (Smith 6519, BM, BR, K), which
has a remotely dentate leaf margin and obtuse to
truncate leaf base. This specimen appears different
from P. dunlapii and, although similar to some
unlobed P. triphylla material, it may represent a
new species.
Wunderlin (1978) also included Psiguria pedala
in bis Flora of Panama treatment, following the
report by Cogniaux (1916) of the species from Pan¬
ama. As no material of P. pedala has been seen
by the author, the species is not included in the key
below. Therefore, only three species of Psiguria are
recognized from Mesoamerica (from the Mexican
states of Tabasco, Yucatan, and Chiapas to the
Panama-Colombia border). A key to these species
follows:
la. Leaves simple, less than 4 cm across, the base
truncate to cordate, not decurrent (Panama)
. P. dunlapii
lb. Leaves trifoliolate, lobate or rarely simple, more
than 4 cm across, the simple leaves cuneate
and decurrent at the base (Tabasco to Panama).
2a. Leaves thick and fleshy; calyx tube not
spotted dark green . P. triphylla
2b. Leaves membranaceous; calyx tube spot¬
ted with dark green. P. warscewiczii
Acknowledgments. The author thanks Sandy
Knapp (BM) for comments on the manuscript.
Literature Cited
Cogniaux, A. 1877. Diagnoses de Cucurbitacees nou-
velles et observations sur les especes critiques. Mem.
Couronnes Autres Mem. Acad. Roy. Sci. Belgique
27: 3-44.
-. 1916. Cucurbitaceae — Fevilleae et Meloth-
rieae. In: A. Engler, Pflanzenreich, IV.275.1. Verlag
von Wilhelm Engelmann, Leipzig.
Condon, M. A. & L. E. Gilbert. 1990. Reproductive
biology and natural history of the neotropical vines
Gurania and Psiguria. Pp. 150-166 in D. M. Bates,
Novon 2: 339-340. 1992.
340
Novon
R. W. Robinson & C. Jeffrey (editors), Biology and
Utilization of the Cucurbitaceae. Comstock Publish¬
ing Associates, Cornell Univ. Press, Ithaca and Lon¬
don.
Howard, R. A. 1973. The enumeratio and selectarum
of Nicolaus von Jacquin. J. Arnold Arbor. 54: 435-
470.
Jeffrey, C. 1962. The application of the generic names
Anguria and Elaterium (Cucurbitaceae). Kew Bull.
16: 197-198.
-. 1978. Further notes on Cucurbitaceae: IV.
Some New-World taxa. Kew Bull. 33: 347-380.
Wunderlin, R. P. 1978. Family 182. Cucurbitaceae.
In: Flora of Panama, Part IX. Ann. Missouri Bot.
Card. 65: 285 366.
Five New Species of Solarium Section Geminata (Solanaceae) from
Peru and Ecuador
Sandr(i Knapp
Department of Botany, The Natural History Museum, Cromwell Hoad, London SW7 5BD,
United Kingdom
ABSTRACT. Five new species of Solatium sect.
Geminata are described from Peru and Ecuador:
S. clivorum, S. goniocaulon, S. malletii, S. smithii,
and S. tovarii. Relationships within section Gemi¬
nata are discussed, and an illustration and distri¬
bution map are provided for each species.
Solarium is one of the largest genera of flowering
plants, with some 1,500-2,000 valid species. Most
of these species are neotropical, with taxa occurring
in all habitats throughout the tropics and subtropics.
Peru and Ecuador are particularly rich in Solanum
species, probably due to their great diversity in hab¬
itat (Weberbauer, 1945). Section Geminata is one
of the largest of the subdivisions of the nonspiny
solanums, with about 145 species, all but one of
these neotropical. Species of section Geminata are
distinguished by their small white flowers, woody
habit, usually leaf-opposed inflorescences, and hard
green fruits at maturity (in all except the .S', pseu¬
docapsicum alliance).
Solanum clivorum S. Knapp, sp. nov. TYPE:
Peru. La Eibertad: entre Huamachuco y Ca-
jabamba, 3,000-3,200 m, 16 Mar. 1948, Fer-
reyra 3059 (holotype, USM; isotypes, IBE,
MO, US). Figure 1.
Frutex; caules dense pubescentes; sympodia difoliata
geminata vel plurifoliata; folia elliptica supra glabra subtus
dense dendritico-pubescentia; inflorescentiae foliis oppos-
itae vel terminales dendriticae; corolla alba; bacca globosa
in pedicello deflexo inserta; semina cornplanata reniformia.
Bushy shrubs, 2-3 m tall; young stems and leaves
densely pubescent with dendritic trichomes ca. 0.5
mm long; bark of older stems reddish or green,
glabrous. Sympodial units difoliate, geminate, or
appearing plurifoliate due to rapid inflorescence
growth. Leaves elliptic, commonly with several ax¬
illary shoots, glabrous (occasionally with scattered
simple uniseriate trichomes on the lamina) and with
a few dendritic trichomes along the main veins adax-
ially, densely pubescent with dendritic trichomes
abaxially; major leaves 13-21 cm long, 6-9 cm
wide, with 7-9 pairs of primary veins, these drying
yellowish, the apex acute, the base acute; petiole
2-3 cm long; minor leaves differing from the majors
only in size, 7-10 cm long, 3-4 cm wide, the apex
acute, the base acute; petiole 1-2 cm long. Inflo¬
rescences opposite the leaves or appearing shoot-
terminal, 3-7 cm long, many-times branched, 20-
40-flowered, densely pubescent with dendritic tri¬
chomes; pedicel scars unevenly spaced 1-2 mm
apart. Puds globose, the corolla only halfway ex-
serted from the calyx tube. Pedicels at anthesis
erect, tapering, 6-8 mm long, ca. 0.5 mm diam.
at the base, densely pubescent with soft dendritic
trichomes like those of the rest of the inflorescence.
Flowers with the calyx tube conical, 1-1.5 mm long,
the lobes deflate, 0.5-1 mm long, densely pubescent
with dendritic trichomes; corolla white, 1 1.1 cm
diam., lobed halfway to the base, the lobes planar
at anthesis, densely pubescent on abaxial lobe sur¬
face with minute, dendritic trichomes; anthers 1.5
2 mm long, 1-1.5 mm wide, poricidal at the tips,
the pores not opening to slits; free portion of the
filaments ca. 0.5 mm long, the filament tube ca.
0.5 mm long, with 5 triangular projections ca. 0.5
mm long alternating with the anthers and closely
investing the style; ovary glabrous; style 3-4 mm
long, glabrous; stigma capitate, the surface minutely
papillose. Fruit a globose, green berry, 1 1.2 cm
diam.; fruiting pedicels woody, erect, 1.5-1.7 cm
long, ca. 1.5 mm diam. at the base. Seeds dark
reddish brown, flattened-reniform with thickened
margins, 3-4 mm long, 2-2.5 mm wide, the sur¬
faces minutely pitted.
Distribution. In the Cajabamba valley, depart¬
ment of Cajamarca, Peru, on steep rocky slopes
from 2,000 to 2,800 m. Figure 5.
Paratypes. PERU. CAJAMARCA: Cutervo, Lagunac, cam-
ino a Cochabamba, 2,000 in, 26 Feb. 1985, Llatas
Quiroz 1178 (F); Cajabamba, Finca Zul, 2,500 m, 22
Aug. 1985, Mostacero L. & Guerra L. 63 (HUT, MO,
NY); Cajamarca, entre Matara y Namora, 2,600 m, 16
Aug. 1973, Sanchez Vega et al. 1217 (NY); Namora-
Matara, 2,600 m, 16 Aug. 1973, Sagastegui A. 7755
(HUT, MO, NY); Huamachuco-Cajabamba, 2,800 m,
16 Nov. 1983, Sagastegui A. 11144 (HUT, NY).
Novon 2; 341-350. 1992.
342
Novon
Figure 1. Solarium clivorum S. Knapp (from Ferreyra 3059, USM). Scale bar = 1 cm.
Local name. Cajamarca: shirac.
Solarium clivorum is closely related to the com¬
mon and widely distributed species ,S. oblongifolium
Dunal. It differs from S. oblongifolium in its smaller
flowers, denser pubescence, and in the projections
from the filament tube. Solarium clivorum is unusual
in section Geminata in not having the terminal pores
of the anthers elongate to slits with age or drying.
This condition is also found in .S. reitzii and .S’.
pseudoquina, two species not closely related to .S’.
Volume 2, Number 4
1992
Knapp
Solanum sect. Geminata
343
Figure 2.
Solarium goniocaulon S. Knapp (from Young 1213, N5). Scale bar — 1 cm.
clivorum and found in southern Brazil. 1 he restrict¬
ed range of S. clivorum is within the range oi ,S\
obIongifolium, hut S. clivorum appears to occur in
drier rnicrohabitats.
This species is named for its habitat (clivorum—
growing on slopes).
Solatium goniocaulon S. Knapp, sp. nov. T\ PE:
Peru. San Martin: Hio Abiseo National Park,
hill E of Gran Pajaten ruins, 2,350 m, ca. 7°S.
77°W, 13 Aug. 1986, K. Young 4213 (ho-
lotype, HUT; isotypes, K, NY). Figure 2.
Frutex vel arbor; caules glabri valde quadrangulati;
sympodia difoliata geminata; folia elliptica supra glabra
nitida subtus glabra nitida in status sicco aurea; corolla
ignota; bacca globosa glabra in pedicello deflexo inserts;
semina ovoideo-reniformia.
Trees, 7 m tall; young stems and leaves minutely
pubescent with golden uniseriate trichomes ca. 0.5
mm long; stems distinctly four-angled, erect; bark
of older stems yellow-gray, glabrate. Sympodia! units
difoliate, geminate. Leaves elliptic, thick anil leath¬
ery, glabrous and shiny adaxially, drying dark green,
glabrous or pubescent with uniseriate golden tri¬
chomes ca. 0.5 mm long along the midrib and pri¬
mary veins abaxially, drying dark gold; major leaves
15-28 cm long, 5-8 cm wide, with 8-10 pairs ol
primary veins, these obscure abaxially, the apex
acute, the base attenuate, somewhat winged onto
the petiole; petioles 1-1.5 cm long; minor leaves
differing from the majors only in size, 3-10 cm long,
1.5-5 cm wide, the apex acute, the base attenuate,
somewhat winged onto the petiole; petiole 0.7-1 cm
long. Inflorescence opposite the leaves, simple, 1-
344
Novon
Figure 3. Solarium malletii S. Knapp (from Knapp & Mallei 6564, US). Scale bar = 1 cm.
3 cm long, sparsely puberulent at the tip, 20-30-
flowered; pedicel scars beginning ca. halfway from
the base, closely spaced, not overlapping. Burls elon¬
gate when very small, sparsely pubescent with uni-
seriate trichomes on the tips of the calyx lobes, larger
buds not known. Flowers not known. Fruit a globose,
green berry, 1 1.2 cm diam.; fruiting pedicels woody,
deHexed, 2-2.2 cm long, ca. 2 mm diam. at the
base. Seeds pale tan, ovoid-reniform, 3-4 mm long,
2.5-3 mm wide, the surfaces minutely pitted.
Distribution. In rainforest in northern Peru, Ca-
jamarca and San Martin departments, from 2,000
to 2,500 m. Figure 5.
Paratypes. PERU. CAJAMARCA: Jaen, Rio Agua Blanca
near Rio Jeronga, 8,000 ft., 30 July 1943, Fvinger 461
(US).
Solarium goniocaulon is a species of uncertain
relationships in section Geminata. It is probably
related to species in the S. arboreum species group
(Knapp, 1986), a complex of glabrous species with
usually erect fruiting pedicels and small white flow¬
ers. 'I’he Mowers of S. goniocaulon are not known,
but the buds (on Fvinger 161) are similar to those
of ,S. anisophyllum van Heurck & Muell.-Arg., a
primary forest species of the eastern Andean slope
in Peru and Ecuador. The strongly 4-winged stem
and the distinctive yellow color of the undersides of
dried leaves readily differentiate ,S. goniocaulon from
any other species of section Geminata.
This species is named for its distinctly angled
stems (gonio—angled; caulon—stem).
Solanum malletii S. Knapp, sp. nov. TYPE: Peru.
Amazonas: Bagua, Quebrada Chinganza, 8 km
N of Muyo, km 472 of Oleoducto Norperuano,
350-400 m, 5°25'S, 78°28'W, 5 July 1984,
Knapp & Mallet 6 561 (holotype, USM; iso¬
types, BH, F, K, US). Figures 3, 4.
Frutex; caules dense pubescentes; sympodia difoliata
geminata anisophylla; folia supra glabra nitida subtus dense
dendritico-pubescentia; folia inajora obovata minora or-
biculata; inflorescentiae foliis oppositae elongatae dense
pubescentes; corolla viridis lobis subtus dense pubescentia;
bacca globosa pubescentia; semina complanata reniformia
margine incrassata.
Volume 2, Number 4
1992
Knapp
Solanum sect. Geminata
345
Figure 4. Solarium malletii S. Knapp (Knapp & Mallet 6564, Quebrada Chinganza, Depto. Amazonas, Peru).
346
Novon
Slender shrubs to small trees, 1-7 m tall; young
stems and leaves densely pubescent with dendritic
trichomes, these 0.5-1 nun long, pale golden when
dry, beige when fresh; bark of older stems reddish
brown, sparsely pubescent with dendritic trichomes.
Sympodial units difoliate, geminate, anisophyllous.
Leaves glabrous and shining adaxially, with a few
dendritic trichomes along the main veins, pubescent
with dendritic trichomes abaxially, the pubescence
denser along the veins; major leaves elliptic-obovate
to obovate, 17-26 cm long, 8-15 cm wide, with
9-13 pairs of primary veins, the apex acute to
acuminate, the base sessile and truncate, usually
somewhat auriculate; petiole if present ca. 2 mm
long; minor leaves differing from the majors in both
size and shape, orbicular, 3-7.5 cm long, 3 7 cm
wide, the apex rounded, the base rounded; petioles
1 2 mm long. Inflorescences opposite the leaves,
simple, 1-15 cm long, 10 — 15-flowered, but with up
to 100 scars, densely pubescent with dendritic tri¬
chomes; pedicel scars densely spaced, not overlap¬
ping. liuds globose, the corolla strongly exserted
from the calyx tube, densely pubescent with den¬
dritic trichomes. Pedicels at anthesis somewhat de-
ilexed, strongly contracted beneath the calyx tube,
ca. 1 mm diam. at the apex, 0.5 mm diam. at ihe
base, 0.7-1 cm long, densely pubescent with den¬
dritic trichomes like those of the rest of the inflo¬
rescence. Flowers with the calyx tube broadly con¬
ical, 1.5-3 mm long, the lobes triangular, 1.5 3
mm long, densely pubescent with dendritic tri¬
chomes; corolla pale green, 1-1.2 cm diam., lobed
ca. 3 A of the way to the base, the lobes planar at
anthesis, the abaxial surface ol the lobes densely
dendritic pubescent; anthers 2 2.5 mm long, 1.5
2 mm wide, in the type with a few dendritic tri¬
chomes on the abaxial surface, poricidal at the tips,
the pores opening to slits with age; free portion of
the filaments ca. 0.25 mm long, the filament tube
ca. 0.5 mm long; ovary densely pubescent with
dendritic trichomes ca. 0.5 mm long; styles heter-
omorphic, in short-styled flowers 11.5 mm long,
in long-styled flowers 5-6 mm long; stigma clavate,
the surface minutely papillose. Fruit a globose, green
berry, 1-1.2 cm diam.; fruiting pedicels woody,
erect or deflexed, 0.7-1 cm long, ca. 1.5 mm diam.
at the base. Seeds reddish brown, flattened-reniform
with the margins incrassate and paler, 1 3 mm long,
1.5-2.5 mm w'ide, the surfaces minutely pitted.
Distribution. On the western margin of the Am¬
azon basin in Ecuador and Peru, in tropical wet
forest, from 200 to 500 m. Figure 5.
Paratypes. ECUADOR. NAI’O: Reserva Biologica Jatun
Sacha, Rio Napo, 8 km E of Misahualli, 450 m, 1°04'S,
77°36'W, 21-25 May 1987, Ceron M. 1394 (MO, NY);
Parque Nacional Yasuni, Pozo Amo 2, trochas de Amosur,
230 m, 0°52'S, 76°05'W, 9-13 Jan. 1988, Ceron M.
& Coello 3182, 3298 (MO, NY). Pkku. amazonas: 400
in behind La Poza, Rio Santiago, 180 m, 22 Aug. 1979,
Huashikat 130 (MO). LORETO: Balsapuerto, lower Rio
Huallaga, 150-350 in, 28-30 Aug. 1929, Killip &
Smith 28636 (NY, US).
Solarium malletii is related to Solarium xarit/w-
phaeum Hitter ol Peru and Solarium rnicroleprodes
Bitter of Central and northern South America, but
differs from both those species in its almost auric¬
ulate leaf bases and pubescent fruits. Solarium xan-
thophaeum shares with S. malletii loose dendritic
trichomes, while S. rnicroleprodes has congested,
echinoid trichomes (see Roe, 1971; Knapp, 1989).
Solarium malletii is found in primary rainforest as
w'ell as secondary forest. The species appears to be
andromonoecious, as many flowers in the type spec¬
imen were found to be short-styled (see Fig. 4). This
breeding system is probably common in section Gem-
inata, but no species have yet been tested experi¬
mentally.
I bis species is named in honor of James Mallet,
who found the type specimen and who for many
years has been a patient companion.
Solnmim smithii S. Knapp, sp. nov. TYPE: Ec¬
uador. Loja: ca. 30 km S ol Catamayo on road
to Cariamanga, 1,900-2,000 m, 4°10'S,
79°20'W, 6 Eeb. 1984, Knapp & Mallet 0232
Volume 2, Number 4
1992
Knapp
Solanum sect. Geminata
347
Figure 6. Solanum smithii S. Knapp (from Knapp &: Mallei 6 252, NY). Scale bar — 1 cm.
(holotype, NY; isotypes, AI), BH, F, G, GH,
K, MEXU, MO, QCA, QCNF, US). Figure 6.
Frutex; caules glabrae; sympodia difoliata vel plurifol-
iata; folia elliptica coriacea supra glabra nitida subtus
pubescentia trichomatibus dendriticis in axillis venarum
ferentia; corolla ignota; bacca globosa alboviridis in ped-
icello deflexo inserta; semina ochracea ovoidea reniformia;
testa foveolata.
Shrubs to 1 m; young stems and leaves glabrous
to densely red papillose, the branches erect; bark
of older stems dark reddish brown. Sympodial units
difoliate or plurifoliate, not geminate, with many
short shoots. Leaves elliptic, widest at the middle.
occasionally thick and somewhat coriaceous, gla¬
brous adaxially, pubescent with tufts of dendritic
trichomes ca. 0.5 mm long in the axils of the primary
veins abaxially, with 4-6 pairs of primary veins, the
blades 5-14 cm long, 1.5-5.5 cm wide, the apex
acute to rounded, the base attenuate, minutely winged
onto the petiole; petiole 0.5-1.5 cm long. Inflores¬
cences internodal or on short shoots, simple, gla¬
brous, 1-4 cm long, 6-7-flowered; pedicel scars
unevenly spaced 3-5 mm apart. Buds elongate and
somewhat pointed, only very small buds known.
Pedicels at anthesis not known. Flowers not known.
Fruit a globose, green to greenish white berry, 1-
348
Novon
1.2 cm diain.; fruiting pedicels somewhat deflexed,
woody. 1 1.3 cm long, 1-1.5 mm diam. at the
base, 3-4 mm diam. at the apex, the calyx lohes
reflexed in fruit. Seeds pale tan, ovoid reniform, 4-
5 mm long, 3-4 mm wide, the surfaces minutely
pitted.
Distribution. In dry forests and scrublands in the
Huancabamba depression of northern Peru and
southern Ecuador, from 1,900 to 2,600 m. f ig¬
ure 5.
Paratype. Peru, cajamarca: Condebamba valley, Ca-
jabamba-Cajamarca road, 2,100-2,600 in, 7°33'S,
78°09'W, 15 Feb. 1983, Smith A I 'asquez 3409 (MO,
NY, USM).
Solanum smithii is similar to S. barbulntum
Zahlbr., also of highland Ecuador and Peru, but
differs from that species in its ovoid reniform seeds
and axillary tufts of dendritic trichomes. The closest
relative of S. smithii is S. tunariense Kuntze of
Andean Bolivia. Solarium smithii differs from .S.
tunariense in having trichomes confined to small
tufts in the axils of the primary veins on the ahaxial
leaf surfaces, its somewhat larger leaves, and its
longer fruiting pedicels. The flowers of ,S. smithii
are not known, but from the shape of the immature
buds I expect them to have elongate calyx lobes.
The leaves on the type specimen are much larger
than and not as coriaceous as those on Smith &
I asquez 3409. The type was collected in a shady,
wet quebrada, while the Peruvian collection is prob¬
ably from a more open habitat.
This species is named in honor of the late 1). N.
Smith, an indefatiguable collector of Peruvian plants
and the collector of the paratype.
Solanum tovarii S. Knapp, sp. nov. TYPE: Peru.
I luancavelica: Hacienda Tocas entre Colca-
bamha y Pauearbamba, 3,000-3,100 m, 19
Apr. 1954, Tovar 1932 (holotype, USM; iso¬
type, 1BE). f igure 7.
Frutex; caules dense pubescentes trichomatibus sim-
plicibus uniseriatis; sympodia difoliata geminata; folia el-
liptica eoriacea supra sparse subtus dense pubescentia
margine revoluta; inflorescentiae foliis oppositae simplices
sense pubescentes; calycis lobi quadratus; corolla alba
carnosa; bacca matura et semina ignotae.
Shrubs 1-2 m tall; young stems and leaves dense¬
ly pubescent with soft, uniseriate, simple (occasion¬
ally branched) trichomes 1-2 mm long; bark of older
stems greenish brown, glabrate. Sympodia! units
difoliate, geminate, heaves elliptic, thick and some¬
what leathery with revolute margins, sparsely pu¬
bescent with uniseriate trichomes like those of the
young growth adaxially, the trichomes denser along
the main veins, the adaxial epidermis large-celled
and crystalline, more densely pubescent with uni¬
seriate, simple or branched trichomes abaxially; ma¬
jor leaves 4-8.5 cm long, 1.5-3.5 cm wide, with
7-10 pairs of primary veins, these drying yellowish,
the apex acute, the base acute; petiole 0.3 1 cm
long; minor leaves differing from the majors only in
size, 1.5-4.5 cm long, 0.5-2.1 cm wide, the apex
acute, the base acute; petiole 2 5 mm long. Inflo¬
rescences opposite the leaves, simple, 2 4 mm long,
densely pubescent with uniseriate trichomes like those
of the stems and leaves, 3-5-flowered; pedicel scars
densely packed, overlapping. Buds globose, the co¬
rolla exserted from the calyx tube. Pedicels at an-
thesis tapering, 0.7-1 cm long, deflexed, sparsely
pubescent with uniseriate, simple trichomes like those
of the inflorescence, ca. 0.5 mm diam. at the base,
1 mm diam. at the apex. Flowers with the calyx
tube broadly conical, 1.5-2.5 mm long, the lobes
quadrate with minute apiculae, sparsely pubescent
with simple, uniseriate trichomes, the apiculae with
tufts of simple trichomes; corolla white, waxy, 1.3-
1.5 cm diam., lobed V\ of the way to the base, the
lobes planar at anthesis, the tips and margins of the
lobes densely papillose; anthers 4-4.5 mm long,
l. 5-2 mm wide, poricidal at the tips, the pores
becoming slitlike with age; free portion of the fila¬
ments ca. 0.25 mm long, the filament tube minute,
often not present, glabrous; ovary sparsely pubes¬
cent with simple, uniseriate trichomes ca. 0.5 mm
long; style ca. 6 mm long, pubescent at the base;
stigma capitate, the surface minutely papillose. Fruit
a globose, green berry (immature), sparsely pubes¬
cent; fruiting pedicels woody, erect, ca. 1.5 cm long.
Seeds not known.
Distribution. In high-elevation savanna areas in
the Peruvian departments of Huancaveliea and Ju-
nin, from 2,700 to 3,300 m. Figure 5.
Paratypes. Peru, huancavelica: Tayacaja, Salcabain-
ba, 3,250 m, 7 Jan. 1939, Stork & Horton 10273 (F,
K). JUNIN: Carpapata above Huacapistana, 2,700-3,200
m, 7 June 1929, Killip & Smith 30080 (NY, IIS).
Solanum tovarii is superficially similar to S. ma-
tureealvans Bitter (an earlier name for S. aurei-
foliurn Rusby), but lacks the arachnoid trichomes
of that species. The trichomes of 5. tovarii are
exclusively uniseriate and can he simple or dendritic.
Solanum tovarii is probably most closely related to
S. barbulntum Zahlbr. of Andean Colombia to north¬
ern Peru and differs from that species in its quadrate
calyx lobes, larger flowers, and in its evenly distrib¬
uted ahaxial leaf pubescence. In ,S. tovarii trichomes
are found over the entire lower leaf surface, while
Volume 2, Number 4
1992
Knapp
Solanum sect. Geminata
349
Figure 7. Solarium tovarii S. Knapp (from Tovar 1932, IBE). Scale bar = 1 cm.
in S. barbulatum they are generally confined to the
axils of the main veins.
This species is named in honor of Oscar Tovar,
Peruvian graminologist, collector of the type spec¬
imen, and a great help to botanists visiting Peru.
Acknowledgments. I thank Blanca Leon, Emma
Cerrate de Ferreyra, Ramon Ferreyra, and Oscar
Tovar for help at USM and for the loan of specimens;
Jaime Jaramillo for help at QCA; Abundio Sagastegui
for help at HIJT; James Mallet for field assistance;
Blanca Leon, Ken Young, and Oscar Tovar for
information about field characteristics of some of
these species; Micky Luck for the illustrations; Sid¬
ney McDaniel for space at IBE; and the curators
350
Novon
of F, HUT, I BE, MO, NY, US, USM for the loan
of specimens.
Literature Cited
Knapp, S. 1986. A revision of Solarium section Gem-
inata (G. Don) Walpers. Ph.D. Thesis, Cornell Uni¬
versity, Ithaca, New York.
-. 1989. A revision of the Solarium riitidum
group (section Holophylla pro parte): Solanaceae.
Bull. Brit. Mus. (Nat. Hist.), Bot. 19: 63-102.
Roe, K. E. 1971. Terminology of hairs in the genus
Solarium. Taxon 20: 501-508.
Weberbauer, A. 1945. El Mundo Vegetal de los Andes
Peruanos. (Nueva edicion revisada de Die Pflanzen-
welt der peruanischen Anden. Leipzig, 1911). Lima.
Two New Species of Cladocolea (Loranthaceae) from
Mexico and Surinam
Job Kuijt
Missouri Botanical Garden, and (mailing address) Department of Biology, University of Victoria,
Victoria, V8W 2Y2, Canada
ABSTRACT. Cladocolea racemosa and (.. elliptica
are illustrated and described as new. The former,
Mexican species is unique in the genus in having
progressive indeterminate inflorescences, and highly
unusual in having pedicellate flowers. The latter,
Suriname species also has an indeterminate inflo¬
rescence, hut is closely related to C. micrantha.
Since my monograph of Cladocolea (Lorantha¬
ceae) (Kuijt, 1975), the circumscription of the genus
has changed through the publication of new entities
(Kuijt, 1981, 1987a, 1987b), the removal of some
to the genus Ixocactus (Kuijt, 1991b), and the in¬
clusion of species from the genus Phthirusa (Kuijt,
1991a). I originally visualized the genus as being
characterized by determinate inflorescences, though
even then a few clearly derived species had inflo¬
rescences that aborted terminally; and by flowers
that were arranged in monads on inflorescences and
were ebracteolate.
The two new species violate both of these major
generic characters: P. racemosa has a truly inde¬
terminate inflorescence, and C. elliptica has at least
some bracteolate triads at the base of an indeter¬
minate inflorescence. In both cases, however, the
alliances to the main body of Cladocolea seem be¬
yond question, and these modified characters are
regarded as derivative.
Cladocolea racemosa Kuijt, sp. nov. TYPE:
Mexico, Cuerrero: 2.5 km al WNW dc Tlaxco,
camino de Tototepec a San Miguel Amoltepec,
bosque de pino, ladera de exposicion S. colgante
parasito de Pinas, 2,100 m, 28 Mar. 1982,
Lorea 055 (holotype, FCME; isotypes, LEA,
WIS). Figure 1.
Plantae delicatissimae, graciles, pendulae, probabiliter
dioeciae, solum masculae visae. Folia usque ad 60 x 4
mm, angustissime lanceolata. Inflorescentiae staminatae
axillares, singulae, indeterininatae, racemosae, prophyllis
binis crassis suffultae, floribus binis, pedicellatis, ebrac-
teolatis 10-14; bractea suffulciens cum pedicello non
coalita. Flores hexameri: petala staminaque dimorpha;
antherae sessiles; sacculi polliniferi 4, connectivum in
cornu prominens protractum.
Extremely delicate and slender, glabrous plants,
the leaf-bearing stems 1 mm or less thick, terete,
sparsely branched, possibly sympodial. Leaves paired,
to 6 cm long and 4 nun wide, very narrowly lan¬
ceolate, tapering into a slender petiole ca. 7 mm
long, venation not evident except for the midrib,
which runs into the apiculate apex. Probably dioe¬
cious, only the staminate seen. Inflorescence one
per axil, indeterminate, subtended by two distinctive,
thick, blunt, pink to purplish prophylls less than 0.5
mm long, and becoming at least 3-4 cm long in
anthesis, racemose, bearing 10-14 paired, pedicel¬
late flowers, the bracts 1 mm long, ovate, with
rounded apex, caducous, completely free from the
pedicels, which become 2.5 mm long and are deeply
constricted just below the ovary in anthesis. Flower
bud at least 5 mm long, one-third of which is an
ovary with inconspicuous, smooth calyculus; petals
6, dimorphic, ca. 3 mm long; anthers 2 mm long,
in 2 series, both series completely sessile, attached
to the petal in the middle of the abaxial surface, a
median ridge present below each anther; pollen sacs
4, the connective extended into a minute terminal
horn; style 1.5 mm long, slender, straight, somewhat
expanded just above the ovary, stigma poorly dif¬
ferentiated.
This is perhaps the most anomalous species of
Cladocolea, and it could be argued that a distinct
genus should be erected for it. 1 prefer to maintain
it in Cladocolea, even though this placement clashes
with one major element of the current circumscrip¬
tion of the genus. This divergent feature is the pro¬
gressive indeterminacy of the inflorescence. There
exist several other species in the genus which, tech¬
nically, also have indeterminate inflorescences, but
the situation is different there in that all flowers
mature more or less simultaneously, e.g., C. cori-
acea Kuijt (Kuijt, 1987a); C. cupulata Kuijt, C.
rncvaughii Kuijt (Kuijt, 1975); C. biflora Kuijt
(Kuijt, 1981). Considering that the terminal flower
in most Cladocolea species tends to be the first to
mature, the simultaneous maturation of all flowers
may be seen as a transitional feature to true inde¬
terminacy. In C. racemosa, progressive indetermi-
Novon 2: 351-354. 1992.
352
Novon
Figure 1. Cladocolea racemosa kuijt. —a. Habit. b. Mature inflorescence. —c. Young inflorescence with
caducous bracts and (arrow) prophylls. —d. Mature flower bud. —e. Ovary and style. —f. Petals and anthers. (Loren
1955, LEA.)
nacy is lully realized: the lowest flowers mature while
the apex of 1 he inflorescence is still forming new
flowers. There is no other Cladocolea species for
which this is true; in several related genera such as
Struthanlhus, Oryctanthus, Oryctina, and Phthi-
rusa, this condition is the rule (kuijt, 1980).
Curiously, there is entomological support lor
placement in Cladocolea: the plant bears a couple
Volume 2, Number 4
1992
Kuijt
Cladocolea racemosa and C. elliptica
353
Figure 2. Cladocolea elliptica Kuijt. —a. Habit. —b. Mature inflorescence, with young superposed inflorescence
bud (arrow). —c. Petals and style. (Irwin et al. 57535, P.)
of small, galled flowers exactly like those I noted
years ago for a male plant of C. loniceroides (Knijt,
1975, fig. 22d), also a Mexican endemic.
Two more, highly unusual features are the ped¬
icellate flowers and the placement of their bracts
directly on the inflorescence axis, rather than fused
on the pedicel, thus the long pedicels are left com¬
pletely free. Cladocolea is otherwise characterized
by sessile flowers in all but C. coriacea Kuijt (Kuijt,
1987a), C. harlingii Kuijt, and C. pedicellata Kuijt
(Kuijt, 1975). In the last two of these species, the
pedicels also are free from the bracts, even though
those of the lowest flowers in C. pedicellata are
fused with the inflorescence axis, instead. None of
these species appears to be closely related to C.
racemosa. Beyond Cladocolea, free pedicels are
rare in Loranthaceae (Kuijt, 1980), probably being
limited to several New 7 Zealand species and the me¬
dian triad flowers of some American inflorescences
in Struthanthus, Tripodanthus, and Psittaranthus.
354
Novon
Notwithstanding its unusual features and the large
geographical distance involved, C. racemosa ap¬
pears to have some affinity to (.. harlingii kuijt ol
southern Ecuador and northern Peru (the latter range
extension is based on a recent new record from
I'rov. Lambayeque, Dept. Lambayeque, Proceden-
cia Colaya, 1,400 m, S. I Jala Quiroz 2002 , LEA).
That species has evolved a remarkable, leaf-medi¬
ated mode of parasitism (Kuijt, 1975), of which
there is no evidence in C. racemosa; and its inflo¬
rescence is determinate. Other inflorescence fea¬
tures, however, are very similar. Cladocolea har¬
lingii thus far has not been considered closely related
to any other species of Cladocolea.
My suggestion of dioecy is based on the unusually
large anthers and, conversely, the insignificant style
and stigma. Neither criterion is reliable, however,
and further material is needed to confirm the point.
Surprisingly, the pollen is not mature in these large
anthers.
Clmiocolca elliptic;! Kuijt, sp. nov. TYPE: Su¬
riname. Wilhelmina Gebergte, Zuid Rivier, in
savanna and disturbed areas, Kayser Airstrip,
45 km above confluence with Lucie Rivier, 270
m, on shrubs 1-4 m above ground. Irwin,
Prance, Soderstrom A Holmgren 57535 (ho-
lotype, P; isotypes, BBS, NY). Figure 2.
Partes novellae leviter furfuraceae; phyllotaxis irre-
gulariter decussata-alterna; folia usque ad 30 x 8 nun,
anguste lanceolato-elliptica, apice plus minusve rotundata.
Planta dioecia, typus pistillatus. Inflorescentiae minus-
culae (4-5 mm), sessiles, in glomerulis, quoque paribus
triadum singulis vel duabus et monadis singulis vel duabus
ebracteolatis, indeterminatae; apex abortivus parvus, ap-
planatus, cuneoliformis. Flores tetrameri; petala 1 mm
ionga, dimorpha; flos pistillatus ut videtur antheram ster-
ilem carens.
Plants lightly scurvy on young parts and inflo¬
rescences, phyllotaxy irregularly decussate-alter¬
nate, leaves from 4 X 1.5 cm (lower part ol in¬
novations) to 20 x 8 mm (upper part ol innovations),
narrowly lance-elliptic, apex rounded to mucronate-
apiculate, only the lower midrib evident but venation
probably pinnate; dried leaf surface strongly gran¬
ular, perhaps indicating sclereid clusters in meso-
phy 11. Dioecious, the type female. Inflorescences
extremely small (4-5 mm), sessile, 1-3 [ter leaf axil,
the 2nd and 3rd superposed slightly gray-furfura-
ceous, made up of 1 or 2 pairs of triads followed
by 1 or 2 pairs of ebracteolate monads, indeter¬
minate, the aborted apex forming a flat wedge be¬
tween the two most distal monads. Flowers 4-merous,
petals 1 mm long, dimorphic, each with oblong me¬
dian cushion but apparently without sterile anther;
style 0.5 mm, clavate, stigma weakly differentiated.
Cladocolea elliptica is closely related to C. mi-
crantha, but very different in leaf appearance and
furfuraceous surfaces of young [tarts and inflores¬
cences, especially the indeterminate inflorescence.
It shares with C. micrantha the aberrant feature of
having bracteolate triads in the lower [tart of the
inflorescence, tetramery, and a very similar general
appearance.
Acknowledgments. 1 am indebted to Karel U.
Kramer for the Latin diagnoses, and to the Natural
Sciences and Engineering Research Council of Can¬
ada for continuing financial support.
Literature Cited
Kuijt, J. 1975. The genus Cladocolea (Loranthaceae).
J. Arnold Arbor. 56: 265-335.
-. 1980. Miscellaneous mistletoe notes, 1-9.
Brittonia 32: 518-529.
-. 1981. Inflorescence morphology of Loran¬
thaceae— an evolutionary synthesis. Blume 27: 1-
73.
-. 1987a. Miscellaneous mistletoe notes, 10-
19. Brittonia 39: 447-459.
-. 1987b. Novelties in Mesoainerican mistletoes
(Loranthaceae and Viscaceae). Ann. Missouri Bot.
Card. 74: 511-532.
-. 1991a. Inflorescence structure and generic
placement of some small-flowered species of Phthi-
rusa (Loranthaceae). Syst. Bot. 16: 283-291.
-. 1991b. Two new species of Ixocactus (Lor¬
anthaceae) and a reformulation of the genus. Syst.
Bot. 16: 292-298.
Hypnella ambrosia (Bryopsida: Hookeriaceae),
a New Species from Bolivia
Marko Lewis
Hert )ario Nacional de Bolivia, Casilla 10077, La Paz, Bolivia
Bruce Allen
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Abstract. The existence of a Hypnella species
(Hypnella ambrosia, sp. nov.) with short marginal
leaf cells, cells with walls projecting at the upper
ends, occasionally doubly serrate leaf margins, cos¬
tae serrate above, and costal tips strongly projecting
as spines is suggestive of a close generic relationship
to Hookeriopsis sensu lato.
Cerro Amboro, within Amboro National Park, is
an area with deep sandstone canyons lying between
the high tropical Andes of the Cordillera Real, the
sub-Andean ranges of Chuquisaca and Tarija, and
the sandstone ranges of the Chiquitos. Clusia shrub-
lands, grading into isolated grasslands, dominate the
upper slopes of Cerro Amboro. The lower canyons
are densely forested, and isolated riverine forests
follow the deeper canyons nearly to the summit.
During a botanical expedition to the summit of Cerro
Amboro (September 1990), the senior author col¬
lected extensively in the upper canyon forests where
an unusual species was encountered along rills and
creeks from 900 to 1,250 m. The species has fea¬
tures found in Hypnella, Hookeriopsis sensu lato,
and Callicostella, but the presence of pluripapillose
cells indicates the first-named genus. This taxon is
unlike any known species of Hypnella.
Hypnella ambrosia M. Lewis & B. Allen, sp. nov.
TYPE: Bolivia. Santa Cruz: Prov. Ichilo, Cerro
Amboro, between upper Rio Isama and summit,
ca. 35 km S of Buena Vista, Parque Nacional
de Amboro, Cerro Amboro, 17°45'S, 63°39'W,
ca. 900 m, Lewis 37820 (holotype, LPB; is¬
otypes, MO, NY). Figures 1-7.
Species nova Hypnellae piliferae proxima foliis gra-
datim angustatis, acuminatis, longis differt.
Plants in thin green mats, occasionally purple
tinted. Stems red, 2-3 cm long, 0.1-0.15 cm wide,
central strand absent, epidermal cells somewhat dif¬
ferentiated, thin-walled, enlarged, at times appearing
as a hyaloderm, not fluted; axillary hairs 2-celled,
the basal cell quadrate, red-brown, the upper cell
swollen, oblong, hyaline; pseudoparaphyllia absent;
paraphyllia absent; irregularly branched, branches
± parallel. Leaves complanate wet or dry, erect
when dry, spreading to wide-spreading when wet,
lanceolate to oblong-lanceolate, gradually narrowed
to long, slender, flexuose acumina, 1.0-1.8 mm x
0.15-0.30 mm, margins weakly serrate at middle,
strongly serrate at apex, teeth mostly single, oc¬
casionally double, each tooth often ending in a pa¬
pilla; costae double, ± parallel, projecting as 1-2-
celled spines near the base of the acumen, with multi-
celled serrations along the upper margins, serrations
often ending in 1 or 2 papillae, in cross section
rounded, with 3-4 layers of homogeneous, uniform¬
ly thick-walled cells; leaf cells thin-walled through¬
out, at insertion reddish or hyaline, rectangular, 30-
40 p.m x 6-8 /am, basal cells long rectangular, 25-
47 /am x 5-6 /um, upper cells elongate fusiform to
vermicular, 23 53 /am x 6-8 /am, upper marginal
cells somewhat wider and shorter than laminal cells,
cells with single papillae variously placed over the
surface and often projecting at the upper cell ends,
or 2-4 papillae scattered, at times in irregular hor¬
izontal rows, over the surface of the cell lumina.
Sporophytes unknown.
Paratypes. Bolivia. Santa cruz: Prov. Ichilo, Cerro
Amboro, ca. 35 km S of Buena Vista in Parque Nacional
de Amboro, 17°45'S, 63°39'W, Lewis 37823, 37859,
37860 (all LPB, MO).
Hypnella ambrosia is common on the banks of
creeks and rills in the upper canyons of Cerro Am¬
boro between 900 and 1,250 m. It appears to be
semi-aquatic and is especially abundant in areas near
creek high-water levels and seepy zones near springs
and rills. It grows on soil and bare rock. Its soft,
pale green, occasionally red-tinted, flattened mats
make it a distinctive element of the upper canyon
forests of Amboro.
Hypnella ambrosia is similar to Callicostella in
having some cells with single papillae; however, its
Novon 2: 355 357. 1992.
356
Novon
Figures 1-7. Hypnella ambrosia M. Lewis & It. Allen. —1. Habit. —2. Median leaf cells. —3. Marginal leaf
cells below apex. —4. Apex of costa. —5. Leaves. —6. Stem cross section. —7. Marginal leaf cells. Figures 1, 4-
7 from holotype; Figures 2, 3 from Lewis 37860. Top scale bar = 40 gm, leaf cells; middle scale bar = 400 ^m,
leaves and stem cross section; bottom scale bar = 1 mm, habit.
Volume 2, Number 4
1992
Lewis & Allen
Hypnella ambrosia
357
long, vermicular, frequently pluripapillose leaf cells
remove it from that genus. Long leaf cells (occa¬
sionally with projecting end-walls), short, broad, dou¬
ble-toothed marginal cells, and strongly serrate cos¬
tae that may end in distinct spines are all features
of Hookeriopsis sensu lato. The long, pluripapillose
leaf cells of this species place it in Hypnella. Hyp¬
nella and Hookeriopsis sensu lato are closely re¬
lated, as demonstrated by several shared features.
For example, leaves ol Hypnella pilifera (Hook. &
Wils.) A. Jaeger occasionally have weakly spined
costae, and the marginal cells of Hypnella leptor-
rhyncha (Hook. & Grev.) A. Jaeger frequently have
projecting ends. Hypnella ambrosia is close to II.
pilifera (Hook & Wils.) A. Jaeger. The latter species
has oblong leaves, often wider above the middle,
that are abruptly narrowed to the acumen. Although
the lower marginal leaf cells in //. pilifera are long
and narrow, the marginal cells in the upper one-
third can be short and broad. There are now seven
species in Hypnella (Crosby et al., 1985; Allen,
1986; Buck, 1990). They are separated in the fol¬
lowing key.
Key to the Species of Hypnella
1. Leaves long acuminate, subpiliferous or pili¬
ferous . 2
1. Leaves long-acute, subacute or obtuse . 4
2(1). Leaves rounded at the apex, abruptly pilifer¬
ous . H. pilfer (Hook. & Wils.) A. Jaeger
2. Leaves gradually and evenly long acuminate
to a piliferous apex . 3
3(2). Marginal leaf cells long, narrowly fusiform;
costae smooth above, not or very weakly pro¬
jecting by a single cell .
. . . H. leptorrhyncha (Hook. & Grev.) A. Jaeger
3. Marginal leaf cells short, broadly rectangular;
costae serrate above, strongly projecting by
two or more cells .
. H. ambrosia M. Lewis & B. Allen
4(1). Leaves ecostate; leaf cells all papillose or en¬
tirely to partly smooth .
. H. guayanensis B. Allen & Buck in Buck
4. Leaves costate; nearly all leaf cells papillose
. 5
5(4). Leaves long-acute, apex serrate by projecting
cell wall ends, ± bordered by long narrow ±
smooth cells .//. punctata Broth.
5. Leaves subacute or obtuse, apex serrulate due
to lateral, projecting papillae, not bordered ... 6
6(5). Leaf papillae mostly multifid; leaves obtuse,
not apiculate, 0.5-1.0 x 0.2-0.4 mm; mo-
noicous; exostome furrowed; capsule neck tu¬
bercular to scabrous .
. H. pallescens (Hook.) A. Jaeger
6. Leaf papillae simple; leaves subacute, apicu¬
late, 1.0-1.2 x 0.3-0.6 mm; dioicous; ex¬
ostome not furrowed; capsule neck papillose
. H. diversifolia (Mitt.) A. Jaeger
Acknowledgments. The senior author thanks
Bobin Clark for the opportunity to participate in the
expedition to Amboro, and the National Geographic
Society and the Missouri Botanical Garden for fi¬
nancial support.
Literature Cited
Allen, B. H. 1986. The taxonomic status of Hypnella
punctata. Bryologist 89: 224-226.
Buck, W. R. 1990. Contributions to the moss flora of
Guyana. Mem. New York Bot. Card. 64: 184-196.
Crosby, M. R„ B. H. Allen & R. E. Magill. 1985. A
review of the moss genus Hypnella. Bryologist 88:
121-129.
New Species of Passiflora Subgenus Plectostemma (Passifloraceae)
John M. MacDougal
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Abstract. Four new species are described, one
with two distinct subspecies: Passijiora (sect. Circa)
jitliana, P. (sect. Circa) xiikzodz subsp. xiikzodz
and subsp. itzrnsis , P. (sect. Pscudodysosmia) oax-
accnsis , and P. (sect. Xerogona) escobariana. Pas¬
siflora Juliana appears to be the closest relative of
P. viridflora Cavanilles, a species that traditionally
has been segregated as a distinct section or subgenus.
Revision of Mexican, Central American, and Co¬
lombian collections of passionflowers is yielding nu¬
merous undescribed species, especially among the
small-flowered subgenus Plectostemma Masters.
Apetalous passionflowers, found only in subgenus
Plectostemma, form a group of about 18 closely
related species called section Circa (Medikus)
DeCandolle. This section is most diverse and com¬
mon in northern Mesoamerica and southern Mexico,
with a radiation of relatives of P. suberosa Linnaeus
in the Caribbean. There are at least two other apet¬
alous species in other sections, hut these occur in¬
dependently elsewhere in the subgenus. The follow¬
ing two new species are in section Circa.
Passiflora juliana J. MacDougal, sp. nov. TYPE:
Mexico. Michoacan: Mun. Coahuayana, high
point on coastal road (Hwy. 200) between San
Telmo and San Juan de Lima, 70 m, 2 Nov.
1079,./. M. MacDougal i l )2 (holotype, DUKE;
isotypes, CAS, CHAPA, DUKE, ENCB, F, (ill.
IBUG, MICH, MO, MEXU, NY, TEX, US,
XAL). Figure I.
Passijiora ad sectionem Ciecarn pertinens, scandens;
stipulae (3—)5— 12(- 15) mm latae; petioli in parte media
biglandulosi; folia peltata glandulosa trilobata, lobis acutis
vel plerumque obtusis vel subrotundatis, marginibus in-
tegris; pedunculi ebracteati; stipes floralis 2-4 mm longus;
petala nulla; coronae filainenta 2-seriata, filamentis ex-
terioribus 6-8 mm longis, viridiflavis; operculum plicatum;
androgynophorum 4-5 mm longum; ovarium glabrum;
fructus baccatus atropurpureus; semina 3.7-4.1 mm lon-
ga, reticulatim foveata.
Vine (1.5—)2—3 m, minutely to microscopically
puberulent throughout at the shoot tip with an-
trorsely appressed curved trichomes 0.08-0.1 2 mm
long, except adaxial surface oi laminas and stipules
glabrous; plant glabrescent below. Stems ca. terete
to elliptical, often reddish, with minute appressed
trichomes. Stipules (6—)10—20(—23) x (3—)5 -12
(- 15) mm, loliose, asymmetrically obovate (to ovate),
acute; petioles 1-4 cm, 2-glandular (0.25-)0.3-0.6
the distance to the apex, the nectaries 2 x 1.0-
1.5 mm, saucerlike, fleshy, subsessile or narrowed
at the base; laminas 3.5-10 x 6-16 cm, peltate
(3—)5— 15 mm from margin, entire, sometimes var¬
iegated as juveniles, the variegations scattered as
small patches of white, this rarely retained at re¬
production, depressed obovate to widely depressed
obovate in general outline, 3-lobed 0.40-0.72 the
distance to the peltate base, 5-7-veined, glabrous
and often matte adaxially, glabrescent and glaucous
abaxially, the lobes elliptic or more usually obovate
(or the central widely obovate), apices obtuse or
sometimes rounded, the central lobe narrowed at
base, the angle between the lateral lobes (80° )1 10°
160°; juvenile leaves more deeply lobed with narrow
segments; laminar nectaries present as 2-6(- 10)
sessile submarginal glands distal to lateral veins, (one
juvenile plant seen with 1 gland per lateral lobe also
proximal to lateral veins), glands 1 mm or less in
diameter, (2 )3 5 mm from margins. Peduncles
(1 )2 per node, 7-20(-27) mm long, uniflorous, green
(one individual seen with branched peduncles, 1
2(-3)-flowered); bracts absent; conspicuous inflo¬
rescences 5 19 cm long often present as condensed
terminal shoots with aborted laminas, the petioles
and their nectaries still present. Flowers borne facing
above the horizontal to nearly vertical, yellowish
green, with a hardly detectable slightly sweet odor,
open and receptive from late morning past midday;
stipe 2.0-4.0 mm; hypanthium ca. 7-8 mm diam.;
sepals (9—) 10— 12 x 4.0-6.0 mm, light green or
yellowish green, with no apical projection; petals
absent; coronal filaments in 2 series, the outer ca.
28 48, 6 8 mm long, tapering to points at tips,
slightly laterally compressed, opening to or slightly
more than 180°, light greenish yellow to light yellow
apically, unmarked or with a mere flush of purplish
or reddish spots adaxially at base, the inner 3.2
3.7 mm, erecl to slightly spreading, greenish yellow
and sometimes with a tinge of purplish at very base,
capitellate; operculum 2 mm long, membranous,
plicate, greenish yellow; limen floor violet to dark
Novon 2: 358-367. 1992.
Volume 2, Number 4
1992
MacDougal
Passiflora subgenus Plectostemma
359
Figures 1-3. —1. Passiflora Juliana J. MacDougal, with beginning of terminal inflorescence ( MacDougal 492GR).
Scale bar = 1 cm. —2. Passiflora xiikzodz subsp. xiikzodz (MacDougal 4677). Scale bar = 1 cm. —3. Passiflora
xiikzodz subsp. itzensis (MacDougal 4633). Scale bar = 1 cm.
purple, its raised edge white or purplish and slightly
inclined toward androgynophore; androgynophore
(4.0-)4.5-5.0 mm long, whitish or pale green with
the purple coloring of the limen ascending only a
few mm at base, or purple nearly to the apex;
staminal filaments free 3.0 -3.5 mm, light greenish,
anthers 3.3-4.0 mm long, light greenish, not spotted
or marked; pollen light yellow; ovary 2.4-3.0 x
1.5-2.0 mm, ellipsoid, glabrous, green; styles 4.0
5.0 mm total length, including stigmas, the stigmas
1.5 mm diam., light green. Fruit 1.3-2.5 x 1.3-
2.0 cm, depressed globose to slightly ellipsoid, some¬
times somewhat trigonal-round in cross section, es-
tipitate, bluish black with a white bloom, thus ap¬
pearing dark gray-blue, exocarp mildly fruity, neither
sweet nor sour; arils clear, with a distinct white line
(funiculus) on the side, close fitting, ± tasteless;
seeds 3.7-4.1 x (2.2-)2.5-2.9 mm, slightly cam-
pylotropous, the chalazal beak slightly inclined to¬
ward the raphe, reticulate-foveate with 11 — 15( 18)
foveae per side. Germination epigeal. Chromosome
number 2 n = 12.
Distribution and habitat. Passiflora juliana is
restricted to the coast and coastal plain of south¬
western Mexico. It is not uncommon near rocky
cliffs overlooking the Pacific Ocean and in the coast¬
al forest in the hills and away from the coast. Col¬
lectors’ notes give the following habitats: common
on road-cut banks and cliffs in sun; on rocky ledge
of arid slope with low scattered trees; in shrubs;
wooded plain of Crescentia—Juliania savanna; selva
baja caducifolia; bosque tropical caducifolia; abun¬
dant on shrubs on low mountain summits with de¬
ciduous woodlands; thickets and rocks at cliff-tops;
roadside thickets; grows best in sunny areas; dappled
shade of secondary forest of low stature. In Mi-
choacan I found it associated with or near Passiflora
holosericea Linnaeus, P. goniosperma Killip, P.
mexicana A. L. Jussieu, and P. filipes Bentham.
Paratypes. MEXICO. COLIMA; Mun. Tecoman, N of
Tecoman, 3.9 mi. NE on Hwy. 110 from jet. of road to
Tecoman (Hwy. 200), 30 Oct. 1979 (H), MacDougal &
Miley 486 (CAS, CHAPA, DUKE, ENCB, IBUG, MEXU,
MICH); 15 mi. SE of Tecoman, near Cerro de Ortega,
0-50 m, 9 Dec. 1959 (fl), McVaugh & Koelz 1615
(MICH); 7 mi. N of Santiago, rd. to Durazno, Jalisco, via
bridge over Rio Cihuatlan, 200 m, 30 July 1957 (fl),
McVaugh 15924 (MICH). Jalisco: 2 km by road E of
Cihuatlan, 0-50 m, 8 Mar. 1970 (fl), Anderson & An¬
derson 6154 (MICH); 19°30'N, 105°03'W, 6 Jan. 1983
(fl), Lott 1701 (DUKE, MEXU); Mun. La Huerta, entrada
a la verada a las Piletas cerca de Rincon de Ixtlan, viejo
camino a Nacastillo, 12 Aug. 1983 (fr), Lott 1832 (DUKE,
MEXU); Mun. Cihuatlan, 2.8 mi. W on Hwy. 200, 4.2
mi. E of road to Barra Navidad, 10 Nov. 1979 (fl),
MacDougal <& Miley 507 (MO-spirit); Mun. La Huerta,
Estacion de Biologia Chamela, 10 Dec. 1976 (fr), Ma-
gallanes 344 (DUKE, MEXU); 13 Dec. 1976 (fl, fr),
Magallanes 364 (MEXU); high rocks above the Pacific
Ocean, Rancho Paraiso, 10 km SE of Chamela, 20 m,
14 Feb. 1975 (fr), McVaugh 26277 (MICH), michoacan:
Mun. Apatzingan, Tancitaro region, Mt. Apatzingan, 2,000
360
Novon
Figure 4. Diagram of floral cross sections of Passiflora
xiikzodz J. MacDougal. Scale bar = 5 mm. —a. P.
xiikzodz subsp. xiikzodz (MacDougal 4677). —b. P.
xiikzodz subsp. itzensis (MacDougal 4633).
ft., 20 Aug. 1941, Leavenworth & Hoogstraal 1717
(F); Mun. Coahuayana, 7.7 mi. SE from Rio Coahuyana
on Hwy. 200, 31 Oct. 1979, MacDougal & Miley 486
(DUKE); Mun. Huetamo, 3 km al NW de Erendira, sobre
el camino a Caracuaro, 800 m, 6 Nov. 1977, Rzedowski
35506 (ENCB).
Passiflora juliana very closely resembles P. vir-
idiflora Cavanilles vegetatively, except that it has
notably wide and foliose stipules and the stem is not
as bright red. The length of the androgynophore is
the main difference between the two: P. viridijlora
has an elongate tubular flower, while P. juliana has
the shallow-dish shape that is usual in the section
and subgenus. Both inhabit similar habitats along
the Pacific Coast of Mexico, with P. juliana to the
north and P. viridijlora southward down the coast,
from Acapulco to the Isthmus of Tehuantepec. The
two species apparently are not sympatric, but the
possible zone of contact has not been sampled. They
are undoubtedly sister species.
Passiflora viridijlora is pollinated by humming¬
birds in coastal Oaxaca (A. Lau, pers. comm.). The
usual pollinator of P. juliana is unknown; my one
morning of observation yielded only a single hurried
visitor—a hummingbird. The floral structure of P.
juliana suggests adaptation to a small or medium¬
sized insect.
The discovery of a passionflower that is clearly
referable to section Cieca , but that very closely
resembles P. viridijlora (except for the bird-adapted
flower), reinforces the placement ol P. viridijlora
in section Cieca, a position previously suggested by
MacDougal (1983).
Passiflora viridijlora has been recognized as a
distinct monotypic genus, Synactila Rafinesque, and
at various infrageneric ranks within Passiflora, but
the discovery of this new species with intermediate
morphology confirms that P. viridijlora is not as
distinct as once thought. The elongate flower prob¬
ably merely represents a shift in pollinators.
Passijlora juliana has a conspicuous terminal
infloresence on older plants that consists of a con¬
densed shoot with extremely reduced or aborted
laminas. A short petiole remains at each node and
the petiolar nectaries are well developed and active.
The shoot of the inflorescence is not determinate,
however, and vegetative growth with normal leaves
and longer internodes often continues after weeks
or months of blooming. This inflorescence is identical
to those seen in the related species, e.g., P. viri-
diflora and P. coriacea A. E. Jussieu. Laminar
nectaries are distal to the lateral veins except in one
specimen, Rzedowski 35506, where single glands
occur proximal to the main veins of each lateral
lobe. Plants from Jalisco and Colima tend to have
more laminar nectaries than the forms from Mi-
choacan, with up to 8 or 10 per leaf. The type
gathering consisted of several individuals growing
adjacently. Branched peduncles, seen only in one
individual of the type gathering (cf. isotype, DUKE),
are to be considered an abnormal condition in this
species. This character state is not shared with any
other collection of P. juliana, nor with other species
in section Cieca, except that it has been observed
in certain races of P. suberosa (cf. MacDougal
421).
A few seeds from fruits on the type gathering
were stored dry and clean for 2'/5 months, then
sown. Germination was excellent within 10 days,
and the seedlings were numbered MacDougal
492GR. In 1983 several of these seedlings were
distributed to the horticultural trade in California
and made available at the retail level in limited
quantities. Less than 20 cuttings were distributed in
the next decade, but clones are still available from
several growers. The chromosome count was from
MacDougal 492GR (Snow & MacDougal, in press).
Passiflora juliana appears to be self-incompat¬
ible, as no fruits were produced from over 50 con¬
trolled self-pollinations performed on MacDougal
492GR in the greenhouse. No fruits have been
produced by autogamy during ten years of culti¬
vation. Hybrids were produced from 492GR on an
extremely limited basis by MacDougal in 1982 by
Volume 2, Number 4
1992
MacDougal
Passiflora subgenus Plectostemma
361
crossing P. juliarui with P. trinifolia Masters and
P. tenuiloba Engelmann. The several plants grown
oi these two crosses were variegated (chimera-type)
and dwarfed like their smaller parents, but otherwise
were essentially intermediate to the parental mor¬
phology. Hybrids flowered with normal-looking, in¬
termediate flowers with abundant pollen; pollen vi¬
ability or stainahility has not been examined. Fertile
vouchers of examples of these crosses are deposited
at DUKE.
Dried leaf samples of MacDougal 492GR (as
492) were chromatigraphically screened by Mc¬
Cormick (1982) for flavonoids. Flavonol 3-O-gly-
cosides but not C-glycosylflavones were detected.
Ibis is an unusual pattern in Passifloraceae, and
was found in all of the species of section Circa that
contained any flavonoids, including P. viridiflora
(McCormick. 1982).
Lacebugs and the butterfly Hcliconius chariton-
ius were important herbivores on MacDougal &
Miley 486.
The species is named for Julianna Miley, my
former wife and beloved companion during the Mex¬
ican and Guatemalan fieldwork of 1978 to 1980.
Her enthusiastic assistance resulted in the discovery
or re-collection of many species of passionflowers.
Passiflora xiikzodz .1. MacDougal, sp. nov. TYPE:
Mexico. Campeche: Tuxpena, [18°26'N,
90°06'W], 19 Jan. 1932, Lundell 1210 ( ho-
lotype, MICH; isotypes, AHIZ, DS, F, GH,
MICH, MO, NY, US, WIS—2 sheets, U). Fig¬
ures 2 4.
Passiflora ad sectionem Ciecarn pertinens, ad rupes
calcareas scandens; stipulae 0.4-0.7 mm latae; petioli ad
apicem biglandulosi; folia peltata glandulosa bilobata vel
truncata vel raro trilobata, lobis lateralibus acutis vel raro
acuminatis, marginibus integris; pedunculi ebracteati, sti¬
pes floralis 12-30 mm longus; petala nulla; coronae fi-
lamenta 5-7(-8)-seriata, filamentis exterioribus filifor-
mibus 8-12 mm longis, pro parte maxima atropurpuris,
ad apicem flavidis; operculum ad basim in denticulos lac-
eratum; androgynophorum nullus vel 0.5-4 mm longuin;
ovarium glabrum; semina ca. 4.3-5.5 mm longa, reti-
culatim foveata.
Vine ca. l-3(-6) m, minutely to microscopically
puberulent throughout with antrorsely appressed,
curved blunt trichomes 0.08-0.12 mm long, and
sometimes also straight, erect, capillary trichomes
to 0.3 mm long, except the laminas glabrous or
nearly so; plant glabrescent below. Stems subterete
to obtusely subangular, ca. scabrous, minutely pu¬
berulent. Stipules (2.8-)3.5 6.5 x 0.4-0.7 mm,
linear to linear-lanceolate; petioles 1—2(—3) cm,
2-glandular at or near the apex (or at least on the
distal 14), 0-5(-10) mm from the apex, the nectaries
0.7-1 x 1.3-2 mm, saucerlike, fleshy; laminas
(1.2-)1.5-7.5 x (3.5-)5- 19 cm, peltate (2.5 )3-
5(-10) mm from margin, entire, usually strongly
variegated along main veins as juveniles, often var¬
iegated at reproductive nodes, depressed obovate to
shallowly obtriangular to narrowly transversely el¬
liptic, 2(-3)-lobed or ca. truncate, 5(-7)-veined at
base, the lateral lobes (acuminate to) acute, often
rounded at the very apex, the central lobe retuse
to obsolete (or obtuse), the angle between the lateral
lobes (75°-)90°-130°(-150°); laminar nectaries
(6-)8-16(-28) per leaf, borne 1-12 mm from mar¬
gin, both proximal (exmedial) to the primary lateral
veins and between the primary lateral and central
veins. Inflorescence often present to 20 cm long as
a condensed terminal shoot with aborted laminas,
the petioles and their nectaries still present. Pedun¬
cles (1)2 per node, 1—7(—10) mm, uniflorous; bracts
absent. Flowers borne vertically, with no detectable
odor, light yellowish green, the corona dark violet
or purple, apically yellowish; stipe 12 30 mm; hy-
panthium ca. 6 9 mm diam.; sepals (8—)10— 1 4.5
x 3.5 6.0 mm, narrowly triangular to oblong tri¬
angular, outermost two slightly cucullate, with no
apical or subapical corni, greenish yellow inside and
out, or greenish outside and lightly flushed with
reddish purple, reflexed at anthesis so that calyx is
open ca. 240°; petals absent; coronal filaments in
ca. 5—7(—8) series, the outermost series 8 12 mm
long, subequal to sepals, very slender-filiform, re¬
flexed slightly more than the sepals at anthesis, the
inner ca. 3-7 series 0.5-3 mm, the three innermost
series greatly apically dilated-capitate; operculum
represented by vestigial fan-shaped denticles ca. 0.3
mm long; limen 5-6 mm diam., the floor convex,
conspicuously glossy brown, dark purple, or reddish
purple, with no raised edge; nectary apparently ab¬
sent; androgynophore absent or present, when pres¬
ent light greenish yellow and unmarked, or purplish
at the base or lower !4; stamina] filaments not con¬
nate, or connate 0.5-4.5 mm along androgynop¬
hore, the free portions 2.5 3.5 mm long, greenish
yellow and not marked, or dark reddish purple;
anthers 1.8 3.0 mm long, greenish yellow, not
marked, dehiscence extrorse or distal; pollen yellow;
ovary 1.9-2.5 x 1.3-2.0 mm, ellipsoid, glabrous;
styles 1.5-5.5 mm long including stigmas, light
greenish yellow or green, unmarked; stigmas capi¬
tate, 1.2-1.9 mm diam. Fruit ovoid. Arils and ger¬
mination unknown. Chromosome number 2 n =12.
Distribution and habitat. Edges, thickets, sec¬
ondary and primary growth on limestone outcrops
or on walls of cenotes, most usually collected on
362
Novon
Mayan ruins or small cliffs, in tropical humid or
subdeciduous forest, Yucatan peninsula and adja¬
cent Guatemala and Belize, 100-300 m. It has been
found growing with the similar P. coriacea at some
sites.
Ethnobotany. Vernacular names are “Bat wing”
(Belize), “Yerba del cuate” (Campeche), “Zodz’ ak“
(Yucatan), and most commonly “Xiik zodz” (vari¬
ously transcribed as “Xig-Sodz,” “Xi ik so’otz,” “Xiik
zotz,” or “Shig sots”) (Yucatan, Campeche). In Be¬
lize, “used in a mix to treat pain” {Hodge .s & Klassi
20 ).
Passiftora xiikzodz presents a fundamentally dif¬
ferent floral corona from other species in section
Cieca; furthermore, there is no floral nectary. I he
corona is highly proliferated, 5-7-seriate instead of
2-seriate, not counting the operculum. Even the
operculum is different. While almost all species in
the genus have a nectary-cover called an operculum
formed by the innermost corona (fused-membranous
or not), this protection against rain, useless visitors,
and desiccation is lacking in P. xiikzodz , the rem¬
nants existing only as minute teeth. A floral nectary
is absent, and no nectar is formed. The reward for
the pollinators is unknown. Pollen might be the re¬
ward, but the shiny convex liinen is deceptively
similar to glistening nectar, and the possibility of
attraction by deception should be investigated.
The floral stipe (that portion of the pedicel distal
to the articulation) is diagnostically long, 12-30 mm,
longer than most other species in the genus. The
seeds are unusually elongate for species in section
Circa or subgenus Plectostemma, being more than
two times longer than wide.
The leaves of P. xiikzodz are very similar to
those of the closely related P. coriacea; both are
transversely oblong, peltate, with laminar nectaries
proximal to the lateral veins, and variegated or not.
They are easily distinguished by the position of the
large, paired petiolar nectaries. In P. coriacea the
glands are near the middle of the petiole. In contrast,
P. xiikzodz has the glands distal or apical on the
petiole. The glands are often not visible from above
the leaf, being hidden under the base of the blade.
Phis batwing plant has in its northern populations
a variant with a truly unusual flower. The popula¬
tions around Chichen Itza, Yucatan, have been sam¬
pled and grown in the greenhouses; careful study
of fresh flowers revealed that the central reproduc¬
tive structures are shortened and modified so that
the flower must dust the pollen on the ventral surface
of the pollinators, not on the dorsum as does the
southern form. Other differences combine to suggest
a cohesive unit recognized here as a separate sub¬
species.
Key to the Subspecies of Passifloka xiikzodz
la. Androgynophore 2.6-4.5 mm long; free por¬
tions of staminal filaments spreading perpen¬
dicular to androgynophore at floral anthesis, the
anthers dehiscing proximally, toward corona;
styles (including stigmas) 3.5-5.5 mm long
. P. xiikzodz subsp. xiikzodz
lb. Androgynophore absent or to 1.0 mm long, the
ovary sessile or nearly so; free staminal fila¬
ments suberect at anthesis, the anthers dehisc¬
ing distally, away from the corona; styles (in¬
cluding stigmas) 1.5-2.0 mm long .
. P. xiikzodz subsp. itzensis
Passiflora xiikzodz J. MacDougal subsp.
xiikzodz
Hypanthium 7-9 mm diam.; staminal filaments
connate along androgynophore 2.6-4.5 mm, the
androgynophore [tale greenish yellow or sometimes
purplish brown at base, the free staminal filaments
greenish yellow, not marked with purple, reflexing
to become perpendicular to the androgynophore at
anthesis; anthers extrorse; styles (including stigmas)
3.5-5.5 mm long, f ruit 2.0-3.2 x 1.5-2.2 cm,
ovoid, slightly 3-sided, purplish black, estipitate; seeds
(4.3-)4.5-5.5 x 1.9-2.3, obcampylotropous, the
chalazal beak inclined toward raphe, reticulate-fo-
veate with 13—20( 22) foveae per side. Chromo¬
some number 2 n = 12. Figure 2, 4a.
Paratypes. BELIZE. COROZAL: Cerros Maya ruins, Low¬
ry's Bight, coastal area, 8 Apr. 1983, (.rune 513 (LL);
without locality, 1931-1932, Gentle 255 (MICH, US).
el cayo: El Cayo, river bluffs, 15 Feb. 1931 (juvenile
and seedling), Bartlett 11496(1 (MICH); El Cayo, 5-13
Mar. 1931, llarlletl 12012 (MICH); hillside near Camp
6, 17 Mar. 1938 (fl), Gentle 2377 (MICH); Macal Hiver,
20-25 Aug. 1991, Hodges & Klassi 20 (MO); ruins of
Xunantunich, coll, by J. Turner, MacDougal 4677 (MO);
ruins of Xunantunich, 17°05'N, 89°08'W, 600-700 ft.,
2 Dec. 1968 (fl), Proctor 29617 (BM); Panti Trail, lx
Chel Farm, Chaa Creek, 21 Mar. 1990 (fr), J. Turner
s.n. (MO); 1 mi. NE of Benque Viejo on rd. to Xunan¬
tunich, 25 Mar. 1990 (fr), J. Turner s.n. (MO). TOLEDO:
cliff face, Jacinto Hills, 16°09'N, 88°51'W, 400 ft., 4
Nov. 1933 (fr), Schipp 603(F). Guatemala, peten: ruins
of Uaxactun, 22 Mar. 1931 (fl), Bartlett 12270{ MICH);
ruins of Uaxactun, 26 Apr. 1931 (fl), Bartlett 12755
(MICH); ruins of Temple 1, Tikal Nacional Park, 13 Mar.
1966 (fl), Contreras 5578 (F); Santa Elena, on La Lib-
ertad road, km 5, 12 Sep. 1966, Contreras 6083 (LL);
Dos Lagunas, 5 km W on Carmelita road, 6 May 1969,
Contreras 8478 (LL, MO); Lake Peten Itza, cliff along
shore E of San Jose, 22 Jan. 1962, I.undell 17235 (LL);
apotal on hilltop, La Cumbre, San Luis area, bordering
Arroyo Quebrada Seca, km 122, 28 Mar. 1977, Lundell
& Contreras 20711 (LL); Plaza Mayor T ikal Ruins, 300
m, 11 Nov. 1967, Molina 21075 (EAP, F); Tikal, Parque
Nacional, km 16 camino Zococzal a Tikal, 12 Sep. 1969
(fl). Tun Ortiz 271 (EAP, F). Mexico, campeche: Campo
Experimental Forestal Tropical “El Tormento," km 5
carretera Escarcega a Candelaria, 24 Dec. 1965, Chav-
Volume 2, Number 4
1992
MacDougal
Passiflora subgenus Plectostemma
363
elas et al. ES-153 (MEXU); ruins of Becan, coll. E.
Leiter, cult, at MO 1990-1992, MacDougal 4690( MO);
Mun. Champoton, a 30 km al N de Calakmul a Escarsega,
220 m, 4 Feb. 1983 (fl), Martinez S. & Tellez 2910
(MEXU). QUINTANA H(K): Kohinlich, July 1974, Boege
3243bis (MEXU); 16 km al S de San Jose de la Montana,
sobre el camino a Tomas Garrido, 26 Sep. 1983, Cabrera
& Cabrera 5565 (MEXU, MO); cleared ruin, near Xpujil,
18°10'N, 88°45'W, 300-400 m, 18 Aug. 1974 (fl),
Shepherd 210 (WIS).
The specific epithet, xiikzodz, is a compound
form of the transliteration of the undeclinable Mayan
vernacular name for the plant, meaning “bat-wing.”
The epithet is pronounced (more or less) “sheik
zoats.” Several other species of plectostemmad pas-
sifloras (notably P. biflora Lamarck, P. sexflora A.
L. Jussieu, P. rovirosae Killip, and P. coriacea) are
locally called “ala-murcielago” (Spanish for bat-wing)
in northern Mesoamerica, but the shape, texture,
and venation of the leaves of this new species par¬
ticularly suggest a comparison with a bat’s wings.
The species is endemic to the region of the Old
Mayan Empire, and the vernacular name is well
known there (E. Leiter, pers. comm.). Most her¬
barium specimens examined early by me were in¬
scribed with a different manuscript name which may
persist on some sheets; corrected labels have been
distributed. The chromosome count was from
MacDougal 4677 (Snow & MacDougal, in press).
Passiflora xiikzodz subsp. itzensis J. Mac¬
Dougal, subsp. nov. TYPE: Mexico. Yucatan:
Chichen Itza, [20°40'N, 88°34'W], June-July
1938, Lundell & Lundel-l 7470 (holotype, LL;
isotypes, LL—2 sheets, MEXU, MICH, US).
Figure 3, 4b.
Androgynophorum nullus vel 1.0 mm longum; sty 1 i
breves, fere sessiles, plus minusve immobiles.
Hypanthium 5.8 7 mm diam.; petals almost al¬
ways absent (but rarely recorded from occasional
sports on cultivated plants as (3.5-)5-7 mm x
(l-)2.5 mm); androgynophore absent to ca. 1 mm
long, the ovary subsessile within the basally fused
or free staminal filaments, the filaments dark reddish
purple throughout, suberect at anthesis; anthers de¬
hiscing distally; styles (including stigmas) 1.5-2.0
mm long; immature fruit ovoid, otherwise unknown;
seeds unknown. Chromosome number 2 n = 12.
Additional specimens examined. Mexico. Yucatan:
Progreso, 1934, Flores 2 (F); Lake Chichankanab,
[19°54'N, 88°46'W], Apr. 1917, Gaunter 23714 (F);
without further locality, 1917 1921, Gaumer 24415 in
part (F, G, MO); Chichen Itza, near Piste, [20°40'N,
88°34'W], June-July 1938 (fl), Lundell & Lundell 7375
(MICH); Chichen Itza, June-July 1938 (fl), Lundell 7439
(MICH); Chichen Itza, coll, by Dzib Dzib & Leiter, cul¬
tivated at MO 1989-1992, MacDougal 4633 (MO).
I hc androgynophore in this distinct subspecies is
reduced to nearly nothing, the ovary thus subsessile
and surrounded by five nearly free stamen filaments
thrusting upwards. The presence of a distinct an¬
drogynophore is usually considered a generic char¬
acter in this family, but other species of Passiflora
have been described with a similarly reduced or
absent column (cf. P. apoda Harms, P. multiflora
Linnaeus, P. ursina Killip & Cuatrecasas, and P.
monadelpha Jorgensen & Holm-Nielsen). The an¬
thers in P. xiikzodz subsp. itzensis differ by not
flipping over to an extrorse position after the flower
buds open, but instead moving only slightly from
the original introrse position to dehisce distally (up¬
wards), away from the corona. The stigmas are
nearly sessile on short nonmoving styles, with their
receptive surfaces also directed distally. The pollen
is thus dusted on the ventral surface of any insect
visitor that might walk on the flower. Although pol¬
linators of neither subspecies are known, relatives
are hymenopteran-pollinated (sect. Circa , Lindman,
1906; P. coriacea , J. Turner, pers. comm.). The
difference in placement of pollen between the two
subspecies may indicate differences in pollinators,
but this remains unknown.
Petals appeared in a few flowers of the cultivated
MacDougal 4633 in October of 1989 and 1991.
These were well formed but small, and their occur¬
rence is an anomaly in the otherwise apetalous sec¬
tion Cieca.
Both subspecies appear to be self-incompatible.
More than 60 artificial self-pollinations of clones of
both subspecies proved unsuccessful in the green¬
houses, and no fruits have been produced by au¬
togamy in two years of cultivation. Viable seeds were
produced easily between two clones of subspecies
xiikzodz (MacDougal 4690, 4677) and even sub¬
species xiikzodz and P. coriacea, with hybrids grown
to flowering. The hybrids with P. coriacea
(MacDougal 4678) had normal-looking, interme¬
diate flowers with abundant pollen; pollen viability
or stainability has not been examined.
Artificial cross-pollinations in the greenhouse be¬
tween the two subspecies has proved unsuccessful.
About 15 crosses in both directions between
MacDougal 4633 and 4677 have been tried, with
aborted fruits with no viable seed formed with 4677
as the pollen donor, and with no fruits formed w ith
1633 as the pollen donor. The lack of fertility be¬
tween the two subspecies underscores the divergence
of the taxa, but more information would be needed
to support the recognition of two separate species.
364
Novon
Figure 5. Passiflora oaxacensis J. MacDougal, with immature fruit (MacDougal 329). White ruler is 16.2 cm
long.
The flowers ol both variants of P. xiikzodz look
similar from a distance, although the northern type
is smaller, arid both have the same unique, convex,
shiny limen with modified operculum and prolifer¬
ation of coronal rows.
The specimens collected by both Flores and Gau¬
nter cited above are tentatively included in this sub¬
species due to their geographic position at the north
of the range, but the specimens are sterile. The
chromosome count was from MacDougal 1633
(Snow & MacDougal, in press).
Passiflora oaxacensis J. MacDougal, sp. nov.
TYPE: Mexico. Oaxaca: Distr. Miahuatlan, 1
km N of [San Jose de] El Pacifico, or 34 km
S of Miahuatlan on Hwy. 175 at Km 132,
16°05'N, 96°30'W, ca. 2,500 m, 26 July 1965,
Roe, Roe & Mori 616 (holotype, WIS; pho¬
tographs, DUKE, MEXU; isotypes, DUKE,
ENCB, F). Figure 5.
Passiflora ad sectionem Pseudodysosmiam pertinens,
scandens, puhescens trichomatis uncinatis; petioli duo
glandulis subsessilis infra medium instruct!; folia trilohata,
basi cordata et denticulata, lobis acute vel acuminate
triangularis; bracteae lineares vel setaceae; coronae fila-
menta uniseriata vel biseriata, filamentis exterioribus 11 -
14 mm longis; operculum plicatum; ovarium glabrum;
arillus rubroaurantiacus; semina reticulatim foveata.
Climbing vine 1.5-3 m with stems apparently
annual from slightly thickened perennial roots,
sparsely pubescent throughout with 0.25 0.60-mm-
long uncinate trichomes; stem subterete and slightly
striate to subangulate. Stipules 8 14 x 4- 8 mm,
ovate-auriculate and apiculate to long caudate, the
margins entire and ciliolate; petioles 2.8-6 cm,
2- glandular 0.25 0.50 the distance to the apex, the
nectaries 1.0 2.0 mm total length, 1.0-1.8 mm
diam., paired or subopposite, pyriform or hemi¬
spherical, subsessile to short-stipitate; laminas
(5.5—)7— 14 x (6.5 )8 15.5 cm, never variegated,
3- lobed 0.30-0.40 (-0.45) the distance to the cor¬
date base, the lobes triangular and acute to nearly
acuminate, the central lobe not narrowed at base,
the angle between the lateral lobes 85°-l 10°(-11 7)°,
ratio of lateral to central lobe lengths (0.65-)0.70-
0.83, margins proximal to lateral lobes denticulate
with 5-8(-10) teeth per lobe, margins distal to lat¬
eral lobes entire; laminar nectaries 0.5-0.9 mm
diam., 2-3 per sinus, very rarely 1 -2 also proximal
to lateral lobes, 1.5-6(-9.0) mm from margin, ses¬
sile; juvenile leaves more shallowly lobed, 0.10 0.30
the distance to the base, the lobes with convex
margins forming a terminal angle of ca. 90°. Pe¬
duncles (1)2 per node, 2.0-2.5 cm; bracts (2.6-)
3.0-6.5 x 0.15-0.5 mm, linear-triangular to cap¬
illary, entire. Flowers 4-5 cm diam., oriented sub-
laterally but below horizontal, with a pleasant flow¬
ery-sweet odor, open and receptive from morning
to midday; stipe 0.8-2.5 mm; hypanthium 12 15
(-18) mm diam., conspicuously concave abaxially
at junction of stipe; sepals 15-21 x 7-10 mm,
ovate-triangular and acuminate, ecorniculate or cor-
ni 1-2 mm long, pale yellowish green to greenish
white and puberulent abaxially, pale yellowish green
Volume 2, Number 4
1992
MacDougal
Passiflora subgenus Plectostemma
365
to whitish adaxially; petals 6 7 x 4.5-5.0 min,
(widely) ovate, white; coronal filaments in 1 main
series, sometimes with a vestigial second series, the
outer filaments ca. 52 56, 11-14 mm long, whitish
to cream and sometimes becoming light yellow dis-
tally, violet or purplish at base and with 13 violet
bands 4-9 mm from base, giving the appearance
of 2-3 violet rings, or the upper bands sometimes
splotchy and rnn together as one pale indistinct
band; inner coronal filaments (if present) 0.25-0.75
mm long, submicroscopic and inconspicuous, cap-
illiform to davate, generally occurring alternate with
the outer filaments, expressed mainly in the sepal
sectors, usually absent in the petal sectors, white,
purplish at base; operculum 1.6-2.2 mm, whitish
or usually whitish with purple streaks near base;
limen white or white with purple speckling; staminal
filaments connate 6.0-7.0 mm along androgyno-
phore, the free portions 5.5 mm long, very lightly
speckled w'ith reddish brown (dried); anthers 3.9
5.0 mm long, the connective speckled with reddish
brown at its abaxial proximal end (dried); ovary 4.5
6.3 x 2.1-2.8 mm, ellipsoid, borne on a 0.8-2.0
mm long stipe, glabrous; styles 7-8 mm including
stigmas; stigmas 2.0-2.5 mm diam. Fruit 6.0-8.0
cm long including 1.5 2 cm stipe, 2.8-3.0 cm
diam., ovoid-ellipsoid, ellipsoid, or slightly obovoid,
apically abruptly and bluntly conical, sometimes also
with an abrupt nipple, purplish black with white
bloom, often light greenish yellow proximally and
on stipe; arils ca. 8 9 mm long, medium to very
orange, fimbriate at apex, fruity-sour tasting; seeds
4.5-5.0 x 3.0-3.5 mm, obovate to widely obovate,
coarsely reticulate-foveate with (15—)16—20 loveae
per side; germination epigeal. Chromosome number
2 n = 12.
Distribution and habitat. Known only from sev¬
eral populations near the road over the crest of the
Sierra de Miahuatlan of Oaxaca, Mexico, in moist
to wet forest at the lower edge ol the pine zone,
2,400-2,500 m. Passiflora oaxacensis is found on
steep slopes in open pine woods, forest edges, dis¬
turbed areas, or roadside thickets, climbing herbs,
shrubs, and dead branches of tree falls. The forest
contains a small amount of oak, and herbs such as
Maianthemum Wiggers are common. This locality
is in an area classified as “humedo, semifrio,” with
1,200-1,500 mm of rain annually and an average
annual temperature of ca. 12°C (CETENAL, 1970).
This climate type has a restricted distribution in
Oaxaca hut extends roughly 40 km eastward from
San Jose del Pacifico, with two or three small isolated
patches to the west. Passiflora oaxacensis, or a
close relative, may thus be on other nearby peaks.
Paratypes. Mexico, oaxaca: W-facing slope, 1.0 mi.
N of San Jose del Pacifico, Distr. Miahuatlan, ca. 7,800
ft., MacDougal 329 (CHAPA, DUKE); rootstocks of
MacDougal 329 cultivated at Duke University 1979
1980, MacDougal 329GR (CHAPA, DI KE, MEXU);
Puerto del Pacifico, Lau s.n. in 1987 (photographs:
DUKE); cultivated at the Missouri Botanical Garden 1988
1992 from seeds from Lau s.n. in 1987, MacDougal
3009 (BM, MEXU, MO).
Passiflora oaxacensis belongs in subgenus Plec¬
tostemma sect. Pseudodysosmia (Harms) Killip on
account of the plicate operculum, basally serrate
leaves, one main filamentous coronal row, large stip-
itate purple fruit with orange pulp, circinnate de¬
velopment of the tendrils at the shoot apex, and the
uncinate trichomes. The species is similar to P.
sicyoides Schlechtendal & Chamisso, from which it
may be distinguished vegetatively by having larger
stipules, subsessile petiolar nectaries, angles between
the lateral laminar lobes of usually less than 110°
compared to usually more than 1 10°, ratios of lateral
to central lobe lengths usually 0.70-0.80 compared
to 0.53-0.75, and 10-20 laminar marginal teeth
compared to less than 12. Fertile material differs
notably by the longer outer coronal filaments 11-
14 mm long compared to 4-8(-10) mm long, which
number 52-55 compared to 59-65(-70), by having
fruits more ovoid than obovoid, and seeds with fewer
testal foveae. Additionally, P. sicyoides often has
peduncles that exceed 30 mm.
A second, vestigial but numerous inner row of
coronal filaments is seen in two of the three collec¬
tions. Other species of this section sometimes have
flowers with a few vestigial inner filaments, but here
the filaments are numerous although diminuitive and
doubtless nonfunctional in the reproduction of the
plant.
The chromosome count was made from Mac¬
Dougal 3009 (Snow & MacDougal, in press). Clones
MacDougal 320GR and MacDougal 3000 were
self-incompatible in the greenhouse, based on more
than 20 controlled self-pollinations.
This species is named for the State of Oaxaca,
Mexico, where it appears to be restricted to a small
area.
Passiflora escobariana J. MacDougal, sp. nov.
TYPE: Colombia. Antioquia: Mun. de Frontino,
road between Nutibarra and La Blanquita, re¬
gion of Murri, ca. 24.5 km from Nutibarra,
1,090 m, 8 Feb. 1991, J. MacDougal, D.
Restrepo & D. S. Sylva 3823 (holotype, HUA;
isotypes BM, COL, CR, HUA, MEDEL, MO,
US). Figure 6.
Passiflora ad sectionem Xerogonam pertinens; caulis
sub3-5-angularis, vinosus; folia 7-25 cm longa, 6-12
366
Novon
Figure 6. Passiflora escobariana J. MacDougal, flowers closing (MacDougal pI al. 3823GR). Scale bar on midrib
of leaf is 1 cm.
lata, eglandulosa nonprofunda bilobata, lobis acutis vel
obtusis, marginibus integris, trichomatibus adaxialibus mi¬
nus quam 0.8 mm longis; pedunculi ebracteati; sepala
15-23 mm longa, 4.5-9 mm lata; coronae filamenta 1
2-seriata, filamentis exterioribus 10-13 mm longis, can-
didis vel eburneis, parte inferiore roseis; operculum pli-
catum; androgynophorum 7.0-9.5 mm longum; ovarium
minute dense puberulens; capsula fusiformis 6-valvata,
carinis rubris.
Vine 5-10 m, minutely puberulent throughout
at the shoot tip with 0.04 0.25 mm long antrorsely
appressed trichomes. Stems ca. 3 5-angular above,
obscurely so below, reddish, puberulent. Stipules 5
7 x 0.5-0.7 mm, linear-triangular or slightly fal¬
cate; petioles eglandular, 2.5-6 cm long; laminas
7-25 x 6-12 cm, 2(3)-lobed 0.11-0.20(-0.27)
the distance to the shallowly cordate base, with
the central lobe always shortest, entire, not varie¬
gated, somewhat bullate, adaxially sparsely but evenly
pubescent or hirsutellous with straight 0.10 0.5
(-0.8) mm long trichomes, abaxially lightly and
evenly pubescent with 0.2-0.3 mm trichomes, the
lateral lobes acute to acuminate, the central lobe
obtuse, the angle between the lateral lobes 33° 50°;
laminar nectaries absent. Older stems cauliflorous.
the inflorescence a condensed leafless axillary shoot
1.5-3.5 cm. Peduncles (1)2 per node, (5-)7-26
mm, uniflorous, red; bracts absent. Flowers creamy-
white tinged pink, with a reddish center, borne ±
upwards, with a mild sweet odor, open from mid¬
morning to just after midday; stipe (2.5-)4-8 mm
(to 12 mm in fruit); hypanthium 10-13 mm diam.,
very pale greenish or nearly white, sometimes slight¬
ly tinged with pink. Sepals (1 5—)1 8 23 x (4.5 )6
8(-9.0) mm, narrowly triangular or oblong-trian¬
gular, acute, strongly 3-nerved, with no apical pro¬
jection, whitish, cream, or pale green abaxially, whit¬
ish adaxially and sometimes tinged pink or red on
the basal part and edges; petals (1 1 —) 13— 1 7 x 4.0-
5.5 mm, narrowly ovate to broadly lanceolate, acute
but the very tip rounded, (whitish to) pale pink;
coronal filaments in 1 or 2 series, the outer ca. 27-
29 in number (N = 2), 10-13 mm long, filiform,
the basal portion forming a bowl, the tips spread
about 180°, the filaments white in the lower half,
pale yellow in the upper half, marked reddish purple
adaxially in the lower %; inner corona present and
numerous in the flowers of some branches of the
type clone but absent in others, absent or only a
Volume 2, Number 4
1992
MacDougal
Passiflora subgenus Plectostemma
367
few vestigial filaments seen in paratype, 2-3 mm
long, capillary, purplish distallv, with paler apex;
operculum 2.0-3.0 mm long, membranous, plicate,
light purple; limen 6-7 mm diarn., concave-shal-
lowly funnelform at insertion of androgynophore,
very pale greenish yellow', not marked with purple;
nectar ring 1 mm high, conspicuously closer to limen
edge than base of operculum; staminal filaments
connate 7.0 9.5 mm along androgynophore, light
greenish yellow at base, paler distally, not marked
with purple, the free portions ca. 4 mm long; anthers
3.0-4.8 mm long, not marked with purple, with pale
to light yellow pollen; ovary 2.0 4.5 x 1.2—3.0
mm, ellipsoid, 6-ridged, finely and densely (micro¬
scopically) puberulent, pink or pale purplish red, or
pale yellowish green; styles 5-6 mm long including
stigmas, reddish (especially distally); stigmas ca. 2
mm diam., capitate, green, fruit 7.0 x 2.5 cm,
fusiform, 6-ridged, dehiscent, red on ridges, cream
between ridges; arils, seeds, and germination type
unknown. Chromosome number 2 n =12.
Distribution anil habitat. Known only from the
Colombian tvpe and one collection in adjacent Pan¬
ama, Passiflora escobariana is from primary low¬
land and premontane rainforest, in small to medium¬
sized trees at edges of light gaps.
Paratypes. Colombia, antioquia: cuttings from type
collection grown 1989-1992 at Missouri Botanical Gar¬
den, MacDougal el al. 3823GR (HUA, MO). Panama.
DAKIF.N: Parque Nacional Darien, Estacion Pirre, a orillas
del Rio Perresenico entre la Estacion Pirre y la cascada,
8°00'N, 77°45'W, 150 m, 10 Oct. 1990 (fl), II. Herrera
720 (BM, MO, PM A).
Passiflora escobariana is assigned to subgenus
Plectostemma sect. Xerogona (Rafinesque) killip
on account of the plicate (loral operculum, lack of
extrafloral nectaries, lack ol bracts, and dehiscent
carinate fruit. It is most similar to P. rovirosae and
P. costaricensis Killip, anti shares with them the
tendency to form leafless flowering axillary short-
shoots. Ehe stem ol P. costaricensis is always sharp¬
ly triangular, even 3-winged, and the upper surface
of the leaf is hirsute with most trichomes longer than
1 mm; the new species has sub3-, 4-, or 5-angular
stems, and the leaves are not hirsute. Passiflora
rovirosae, from southeastern Mexico to northern
Guatemala and Belize, is most similar but has nar¬
rower, more densely pubescent, nonbullate leaves
with a lateral lobe angle of 22°-3() o (-40°), and yel¬
lower flowers with a well-developed inner coronal
row.
A clone from the type collection was grown in
the greenhouses where it flowered a few' times a
year (mainly in the winter). The species may be sell -
incompatible: ten controlled sell-pollinations failed
to set fruit, and no fruit has been produced by
autogamy in four years of cultivation. The chro¬
mosome count was from MacDougal et al. 3823(711
(Snow & MacDougal, in press).
The paratype, from Panama, differs from the type
in its 4-5-angular stem instead of sub3-4-angular,
in having notably longer petioles (4-6 cm vs. 2.5-
3.5 cm), and longer trichomes on the upper surface
of the leaf (0.2-0.8 mm vs. 0.10-0.25 mm).
It is a privilege to name this passionflower in
honor of Linda Albert de Escobar, eminent student
of passionflowers, whose valuable work in the genus
and on the flora of Colombia is well known.
Acknowledgments. 1 thank Eldon Leiter lor his
dogged pursuit and collection of live material of P.
xiikzodz; his help alone is responsible for most of
our understanding of this species. 1 am also indebted
to Joanna Turner for her photographs, observations,
and live material of that species from Belize. Victoria
Bricker and Anne Bradburn at I ulane University
guided the transliteration of the Mayan epithet. Al¬
fred Lau generously collected seed of P. oaxacensis,
Melvin Turner assisted with P. juliana, and Kathy
Upton cared for all four of the new taxa in the
greenhouse. I thank my Colombian colleagues Diego
Restrepo and D. Stella Sylva for guidance during
fieldwork in Antioquia, and the curators ol the listed
herbaria. The Latin diagnoses were graciously pol¬
ished by Henk van der Werff and William L. Cul¬
berson. The initial research on three of these species
and fieldwork in Mexico was accomplished during
graduate studies at Duke University under a Na¬
tional Science Foundation Fellowship, further sup¬
ported by NSF grant DEB-7912607.
Literature Cited
CETENAL e Instituto de Geographia, UNAM. 1970.
Carta de climas. Hoja San Pedro Pochutla 14P-(I1)
Oaxaca 14Q-V11I. Secretaria de la Presidencia, Go-
bierno de Mexico, Mexico, D.F.
Killip, E. P. 1938. The American species of Passiflor-
aceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 1-
613.
Lindman, C. ,A. M. 1906. Zur Kenntnis der Corona
der einiger Passifloren. Pp. 55-79 in F. R. Kjellman
(editor), Botaniska Studier. Upsala.
MacDougal, J. M. 1983. Revision of Passiflora L.
section Pseudodysosmia (Harms) Killiip emend. J.
MacDougal, the Hooked Trichome Group. Ph.D. Dis¬
sertation, Duke University, Durham, North Carolina.
McCormick, S. P. 1982. Flavonoid Chemistry of Pas¬
siflora subgenus Plectostemma. Ph.D. Dissertation,
The University of Texas at Austin, Austin, Texas.
Snow, N. & J. M. M acDougal. New chromosome numbers
in Passiflora (Passifloraceae). Syst. Bot. 18: in press.
New Species ol the Fern Genus Elaphoglossum from Mesoamerica
John T. Mickel
New York Botanical Garden, Bronx, New York 10458, II.S.A.
Abstract. 01 the I 18 species of Elaphoglossum
treated in the flora of Mesoamerica, 33 were de¬
termined to be new and are described here. In ad¬
dition, one new variety is described and one variety
is raised to species rank.
Elaphoglossum is a large anil taxonomically dif-
ficult genus of wet tropical regions. Fee (1845) and
Christ (1899) published major treatments of the
genus, treating 105 and 142 species, respectively.
Today there are thought to be ca. 600 species. Since
the turn of the century many additional collections
have been made, but until recently there have been
no usable treatments for their identification. While
preparing a treatment of Elaphoglossum for E/ora
Mesoamericana, a number of previously unde¬
scribed species were distinguished. Of I 18 species
in the flora, 33 species and one variety are new, a
figure consistent with the recent regional treatments
of Mexico (19 new r of 45; Mickel, 1980, 1981,
1988, 1992; Smith, 1975), Venezuela (35 new of
101; Mickel, 1985, 1990b, 1991b), anil Peru (51
new ol 122; Mickel, 1990a, 1991a). Gradually a
clearer understanding is emerging of the genus, the
species and species groups, and overall character
diversity. Several species complexes remain unre¬
solved. and more collections and detailed studies are
needed. Most of the species in Flora Mesoameri¬
cana (all but 1 I of the 1 18) will be treated also in
Lellinger’s pteridoflora of Costa Rica-Panama Cho-
co, together with illustrations.
Eluphoglossum andersonii Mickel, sp. nov.
TYPE: Costa Rica. San Jose: near Quebradillas,
about 7 km N of Santa Maria de Dota, Standley
42932 (holotype, US).
Ah E. hieracioide frondibus majoribus longius stipitatis
textura tenuioribus necnon hydathodiis manifestis abstat.
Epiphytic; rhizome 2 3 mm diarn., compact; rhi¬
zome scales 3-5 mm long, linear, orange-tan, lus¬
trous, entire; leaves 8 17 cm long, approximate;
phyllopodia lacking; petiole the sterile leaf length,
tan, flattened; petiole scales 2.5 4 mm long, linear,
entire, orange-tan, subulate, spreading, mixed with
scattered to dense glandular hairs at petiole base;
sterile lamina 4-8 cm long, 1.3—1.7 cm wide, el¬
liptic, chartaceous, base cuneate, apex rounded to
acute; veins evident, 2 mm apart, at 45-55° angle
to costa; hydathodes evident as orange dots; lamina
scales 3-4 mm long, scattered, spreading, subulate,
hairlike, orange-tan, entire; fertile leaves shorter
than the sterile; petiole Vfc-34 the fertile leaf length;
fertile lamina elliptic, ca. 2.5 times longer than wide,
base and apex rounded; intersporangial scales lack¬
ing.
Distribution. Wet forests; 1,500-2,400 m.
Earatypes. CoSTA RlCA. CARTACO: Cerro de La Car-
pintera, Standley 34327 (US). SAN JOSE: Finca La Cima,
above Los Lotes, N of El Copey, Standley 42659 (IIS).
Panama. CHIRiptU: Bajo Grande, Cerro Punta, Caballero
178 (US).
This species differs from E. hieracioides Mickel
in having evident hydathodes, but resembles it in
the elongate, longitudinally folded fertile lamina.
1 here seems to be considerable variation in the
abundance of glandular hairs, from dense to lacking.
The species is named in honor of William R.
Anderson, who completed outstanding studies of
hlaphoglossum in Hawaii and Jamaica.
Elnphoglossum atrobarbatum Mickel, sp. nov.
TYPE: Panama. Panama: Cerro Jefe, Valdes-
pino & Aranda 167 (holotype, NY; isotype,
UC).
Frondium squamis longissimis capilliformibus nigris
necnon frondibus fertilibus abbreviatis praestans.
Epiphytic; rhizome 3-4 mm diam., compact, hor¬
izontal; rhizome scales 4-7 mm long, linear, orange-
tan, dull, entire, spreading; leaves 7-20 cm long,
approximate; phyllopodia lacking; petiole '/-%( '/>)
the sterile leaf length, tan; petiole scales 4-7 mm
long, dense, subulate, hairlike, black, lustrous, den¬
ticulate, spreading; sterile lamina 5-14 cm long,
1.4-2.3 cm wide, elliptic to lanceolate, chartaceous,
base rounded to broadly cuneate, apex acute; veins
obscure, at 65° angle to midrib; hydathodes present;
lamina scales 4 8 mm long, black, lustrous, dentic¬
ulate, spreading; fertile leaf shorter than the sterile;
petiole % ol fertile leaf length; lamina narrowly el¬
liptic, 6-9 mm wide, broadly cuneate, apex acute
to obtuse; intersporangial scales 3-8 mm long, liair-
Novon 2: 368 382. 1992.
Volume 2, Number 4
1992
Mickel
Elaphoglossum
369
like, subulate, black, lustrous, inconspicuously
sparsely denticulate.
Distribution. Wet forests; 100-1,000 m.
Paratypes. Panama. CHIR1QUI: ca. 0.5 km E of Cerro
Pate Macho, headwaters of Rio Palo Alto, Knapp et al.
2109 (CR). cocle: Cerro Gaitel above F.I Valle, Miller et
al. 788 (MO); Comarca de San Bias, Cerro Obu, Nevers
et al. 8082 (MO). PANAMA: Cerro Jefe, Stimson 5418
(NY), Tyson 3599 (MO, TTC), van der Werff & van
Hardeveld 6950 (MO, NY, UC), Porter et al. 5079 ( MO),
Folsom 3836 (MO), Dressier 3164 (TTC), l aldespino
& Aranda 122 (MO), 211 (MO, NY), 220 (MO, NY),
Churchill 3951A (MO); headwaters of Rio Chagres, Rio
Esperanza, and Rio Piedras, de Nevers 4073 (MO).
Elaphoglossum atrobarbatum is distinguished by
the long, black, hairlike lamina scales and short
fertile leaves. It resembles E. lanceiforrnc somewhat
in lamina shape, but in E. atrobarbatum the scales
are longer and are black, rather than orange-brown.
The specific epithet comes from the Latin, ater,
black, and barba, beard, referring to the long, black,
subulate laminar scales.
Elaphoglossum auripilum Christ var. longi-
pilosuin Atehortua, var. nov. TYPE: Costa
Rica. Puntarenas: 5 km S of San Vito de Java,
Finca Wilson, Lloyd 4242 (holotype, NY).
A var. auripilo fronde sterili lineari-lanceolato (nec
oblanceolato), stipite dense (nec dissite) squamoso, squam-
is 3-6 mm longis aurantiacis vel rubro-aurantiacis (nec
1-3 mm brunneo-aurantiaco) divisa.
Phis variety is most readily distinguished from
variety auripilum by the more abundant, longer
scales on petiole (3—6 mm vs. 1-3 mm long) and
lamina (2-3 mm vs. 1.5-2 mm long).
Distribution. Wet forests; 1,000 1,800 m.
Paratypes. Costa Rica. COCOS ISLAND: Fournier 351
(NY), Gomez 3347 (NY), puntarenas: San Vito, Finca
Wilson, Lloyd 4226 (NY), 4227 (NY), Mickel 3098
(NY), Seidenschnur 131, 132, 135, 161, 185 (all NY),
McAlpin 85-71 (F), Raven 21966 (F). san josk: Gomez
et al. 22964 (MO, NY).
Elaphoglossum boquetense Mickel, sp. nov.
TYPE: Panama. Chiriqui: Boquete, Cerro Hor-
queta, Dwyer & Hayden 7706 (holotype, NY;
isotype, MO).
Ab E. papilloso laminae stipitisque squamis aurantiacis
distat.
Epiphytic; rhizome 4-5 mm diam., compact, hor¬
izontal; rhizome scales 1-2 mm long, linear, black,
lustrous, slightly denticulate toward apex; leaves 28-
40 cm long, approximate; phyllopodia lacking; pet¬
iole l A- l A of sterile leaf length, tan to brown; petiole
scales 1.5-2.5 mm long, sparse, tan (to black at
petiole base), entire; sterile lamina 15 25 cm long,
3.4-5 cm wide, elliptic, chartaceous, base attenuate,
apex acuminate; veins evident, 2 mm apart, at 60°
angle to costa; hydathodes present; lamina scales
2-3 mm long on surface, 3 mm long on margin,
scattered on both surfaces, linear, orange-tan, dull,
entire; fertile leaf slightly shorter than the sterile;
petiole % of fertile leaf length; fertile lamina elliptic,
base attenuate, apex acute; intersporangial scales
ca. 1 mm long, rare, orange-tan, linear, dull.
Distribution. Wet forests; 1,500-1,900 m.
Paratypes. Panama. CHIRIQUI: vicinity of Fortuna Darn
in valley of Rio Chiriqui, Croat 66563 (MO), l aldespino
1179 (NY).
This species resembles E. papillosum (Baker)
Christ in lamina shape and white patches, but is
distinct in the scattered, orange, petiolar and laminar-
scales (rather than sparse, minute, black scales).
Elaphoglossum brenesii Mickel, sp. nov. TYPE:
Costa Rica. Puntarenas: 5 km S of San Vito
de Java, vicinity of Finca Wilson, Seiden¬
schnur 180 (holotype, NY).
Ab F. cuspidato laminae squamis lanceolatis laxe dis-
positis, costae squamis angustis pallidis et lamina apice
acuta divergens.
Epiphytic; rhizome 3-6 mm diam., compact; rhi¬
zome scales 3-6 mm long, linear, dark reddish brown,
lustrous, ciliate-denticulate; leaves (10—)1 7—48 cm
long, approximate; phyllopodia present, indistinct;
petiole % ~Vi of sterile leaf length, tan to orange;
petiole scales 3-7 mm long, dense to scattered, some
linear, 3-7 mm long, dark, spreading, most ap-
pressed, to 0.5-1 mm long, pale with dark teeth
and central streak, lanceolate to orbicular; sterile
lamina (8-)13-30 cm long, 1.4-2.5 cm wide, lin¬
ear-elliptic, chartaceous, base cuneate, apex acu¬
minate; veins obscure, ca. 1 nun apart, at 70° angle
to costa; hydathodes lacking; abaxial lamina scales
1 -2 mm long, lightly imbricate, some lamina visible,
linear-lanceolate, light orange, ciliate-denticulate,
with darker point of attachment, costal scales 2-3
mm long, linear, ciliate-denticulate, paler than other
laminar scales, with dark teeth and point of attach¬
ment, often with dark red-brown lustrous teeth; ad-
axial scales linear-lanceolate to orbicular, pale or¬
ange, costal scales as abaxially; fertile leaves slightly
longer than the sterile; petiole % of fertile leaf length,
usually dark; fertile lamina linear-elliptic, base broadly
cuneate to rounded, apex acuminate; costal scales
1-1.5 mm long, pale with reddish brown to black
370
Novon
lustrous point of attachment, linear-lanceolate, ap-
pressed; intersporangial scales lacking.
Distribution. Wet forests; 500-2,500 m.
Paratypes. Costa Rica, alajuela: ca. 20 km N of
San Ramon, Moran 3235 (MO); NW end of Finca Los
Ensayos, Buena Vista de Zarcero, Judziewicz 4391 (WIS).
caktaco: La Estrella, Standley 39261 (LJS); Santa Maria
tie Dota, 50 mi. S of Cartago, Stork 2970 (LJS). guan-
ACASTE: Parque Rincon de la Vieja, Hacienda Santa Maria,
llcrrcra 733 (F, MO, UC); Rio Chiquito de Tilaran, llaber
& Hello 5158 (MO). HEREDIA: Virgen del Socorro Rio
Sarapiqui-Cariblanco, Chacon & Herrera 1162 (LJC).
PLNTARENAS; foothills of the Cordillera de Talamanca,
between Agua Caliente and the Rio Canasta, Davidse et
al. 28348 (MO). SAN JOSF.: ca. 7 km N of San Rafael de
Heredia on Volcan Barba, Lellinger 662 (MO); Pan Amer.
llwy. to San Isidro de General, Lloyd 4248 (NY). Pan¬
ama. CHIR1QUI: vie. of El Boquete, Maxon 5034 (US);
Distrito de Bugaba, road Volcan Santa Clara, van der
II erff t£ Herrera 7058 (MO). COCLE: El Valle Mesa, 5.6
km along mesa road from main road in El Valle, Folsom
3887 (MO), panama: Distrito Capira, Cerro Campana,
van der IVerff& Herrera 6214 (MO), veracuas: between
Escuela Agricola Alto Piedra and Rio Dos Bocas, Croat
2589a (MO).
Elaphoglossum brenesii is distinct from E. pa-
leaceum (Hook. & Grev.) Sledge by the dark red
central point of attachment of linear-lanceolate scales
on costa and often by round, fringed abaxial scales,
rhe former is similar to E. cuspidatum (Willd.) T.
Moore in tbe dark-centered laminar scales, but there
are some to many abaxial lanceolate scales, the
costal ones often very pale with dark teeth.
The species is named in honor of Alberto M.
Brenes, collector of plants in the vicinity of San
Ramon, Costa Rica.
Elaphoglossum brevissimum Mickel, sp. nov.
TYPE: Panama. Panama: Cerro Jefe, I aldes-
pino & Aranda 130 (holotype, NY; isotype,
MO).
Ab E. doanensi frondium fertilium stipite brevissimo
differt.
Epiphytic; rhizome 4-5 mm diam., short-creep¬
ing; rhizome scales ca. 5 mm long, linear-lanceolate,
tan to dark brown mixed together, lustrous, sub¬
entire, with an occasional process; leaves 24-37 cm
long, 12 mm apart; phyllopodia lacking; petiole
5 of sterile leaf length, tan; petiole scales 1-3
mm long, sparse, dark, sclerotic, with processes;
sterile lamina 22-35 cm long, 3.5-6.2 cm wide,
narrowly elliptic, coriaceous, attenuate at both ends;
veins obscure, ca. 1 mm apart, at 80° angle to costa;
hydathodes lacking; abaxial lamina surface with
scattered minute stellate trichomidia, abaxial costa
base with a few scattered scales as on petiole, adax-
ially glabrous; fertile leaves much shorter than the
sterile ('/-*/, the length); petiole V u - l / s the fertile leaf
length; fertile lamina 1.2-1.8 cm wide, narrowly
elliptic, attenuate at both ends, sparse dark scales
on petiole and costa; intersporangial scales lacking.
Distribution. Wet forests; 800-1,000 m.
Paratypes. Panama. PANAMA: Cerro Jefe, Dressier
3163a (TTC), 3163b (TTC), Churchill 3952 (MO), Ty¬
son 1001 (TTC), Porter et al. 5078 (MO), Folsom 3833
(MO), Moran 3285 (MO), Croat 43572 (MO), l aldes-
pirio et al. 194 (NY, UC).
This species is similar to E. doanense Gomez in
the sterile leaf size and short petiole, but that species
has large, pale, ovate, appressed petiole scales, pale
rhizome scales, longer fertile leaves, and no tri-
chomidia.
1 he name alludes to the very short petiole.
Elaphoglossum caricifolium Mickel, sp. nov.
TYPE: Panama. Chiriqui: Cerro Colorado, 50
km N of San Felix on the Continental Divide,
Mori & Dressier 7890 (holotype, NY).
Ab h. ciliato (C. Presl) T. Moore lamina lineari, stipite
gracili, phyllopodiis brevibus abstans.
Epiphytic, epipetric, or terrestrial; rhizome 1.2
2 mm diam., short- to moderately creeping, resinous,
black, or with scales ca. 1 mm long, linear-lanceo¬
late, sclerotic, lustrous brown, entire; leaves 12 42
cm long, approximate to 2 mm apart; phyllopodia
0.3-0.6 cm long; petiole / w ~'/\ of sterile leaf length,
orange to brown; petiole scales ovate, brown, scle¬
rotic proximally, substellate, orange-tan distallv, or
lacking, or reduced to resinous dots; sterile lamina
1 1-35 cm long, 0.5-1.2 cm wide, linear, charta-
ceous, base and apex attenuate; veins obscure, ca.
1 mm apart, at 65-70° angle to costa; hydathodes
lacking; lamina indument adaxially of scales ca. 0.5
mm long, substellate, orange-tan to white with age,
more abundant near the margin, abaxial surface with
resinous dots, abaxial costa with scattered ovate
scales ca. 0.5 mm long; fertile leaves slightly shorter
than or equaling the sterile; petiole V 3 -V 2 the fertile
leaf length; fertile lamina 4-7 mm wide, linear,
attenuate at base and apex, with scattered orange
to black, minute (0.2 0.3 mm long) scales adaxially;
intersporangial scales lacking.
Distribution. Wet forests; 1,000 2,000 m.
Paratypes. Costa Rica, alajuela: Monteverde Re¬
serve, Rio Penas Blancas, Bello 979 (MO). GUANACASTE-
ALAjliELA: slopes of Miravalles, above Bijagua, Gomez et
al. 19159 (MO, UC). puntarenas: valley of Rio Coton
between Sitio Coton and Sitio Tablas, Davidse 24472
Volume 2, Number 4
1992
Mickel
Elaphoglossum
371
(MO); upper slopes of Cerro Buru, Davidse 23806 (MO).
san JOSE: vie. of Santa Maria de Dota, Standley 43452
(US). Panama, bocas dei. toro: Oleoducto Road, 2 km
NE of Continental Divide, Churchill et nl. 4934 (MO).
CHIRIQUI: El Respingo, Cerro Punta, Chavarria 38 (US);
Fortuna, Correa et al. 2179 (US); Cerro Colorado, 50
km N of San Felix on Continental Divide, Mori & Dressier
7890 (US); El Boquete, Common 853 (UC, US); end of
road past Palo Alto, ca. 4.5 mi. NE of Boquete, Hammel
7358 (MO); Cerro Hornito, Croat 6 7973 (UC); E of main
camp at dam site, Folsom et al. 5461 (UC); Distrito de
Bugaba, Santa Clara, van der Werff & Herrera 7172
(MO, NY, UC). VERACUAS: Cerro Tute, Thompson et al.
8850 (MO).
Elaphoglossum caricifolium is distinguished by
the slender, naked to resinous-scaly rhizome, linear
lamina, and short phyllopodia, but is variable in the
scaliness of the rhizome and leaf.
The name refers to the sedgelike leaves.
Elaphoglossum caroliae M ickel, sp. nov. TY PE:
Costa Rica. Volcan Barba, 7,800 ft., Idoyd
4138 (holotype, NY).
Ab E. smithii (Baker) Christ lamina latiori basi rotun-
data differt.
Epiphytic; rhizome 4 6 mm diam., compact; rhi¬
zome scales 4-5 mm long, lanceolate, reddish brown
to red-orange, lustrous, denticulate, spreading; leaves
10-33 cm long, approximate; phyllopodia lacking;
petiole Vi-Vs the sterile leaf length, stramineous;
petiole scales 3-4 mm long, scattered to dense,
lanceolate to subulate, orange, dull, denticulate,
spreading; sterile lamina (4-)7-14 cm long, 2-5
cm wide, lanceolate, chartaceous, base rounded, apex
caudate; lamina scales ca. 5 mm long on costa, 2-
3 mm long on margin, sparse to rare, subulate,
orange, dull, spreading; veins evident, 1.5 2 mm
apart, at 70° angle to costa; hydathodes present;
fertile leaves shorter than the sterile; petiole 3 A the
fertile leaf length; fertile lamina lanceolate, base
rounded, apex acute to mucronate; abaxial costa
glabrous or with scales 3-5 mm long, scattered,
linear, hairlike; intersporangial scales 2 mm, orange-
tan, hairlike, or lacking.
Distribution. Wet forests; 1,200-2,600 m.
Paratypes. Costa Rica, cartaco: La Carpintera, Tor¬
res 120 ( US), Standley 34433 (US); Santa Clara de
Cartago, Lankester 898 (US); San Cristobal-El Tejar
road. Stork 2236 (UC). HEREDIA: Cerros de Zurqui, NE
of San Isidro, Standley & I alerio 50361 (US). SAN JOSE:
ca. 10 km N of San Rafael de Heredia on Volcan Barba,
Mickel 2656 (NY), 2677 (NY), 3023 (NY); Volcan Bar¬
ba, Brade 545 (NY), Lloyd 4140b (NY); San Cristobal
road, Danielson (Stork no. 2545) (UC, MICH). LIMON:
Cordillera de Talamanca, Cerros Tararia (Tres Picos),
Davidse et al. 28790.4 (MO). Panama. Cerro Tute,
Thompson et al. 8850 (MO).
Elaphoglossum caroliae is distinguished by the
orange scales of the rhizome and leaf, lustrous lam¬
inar surface, and rounded lamina base. There is
considerable variation in the abundance of scales on
the petiole and lamina, and even among the spo¬
rangia.
The species is named in honor of my wife, Carol
.1. Mickel.
Elaphoglossum christianeae Mickel, sp. nov.
TYPE: Panama. Panama: 4.5 km N of Lago
Cerro Azul by road, Nee 7010 (holotype, NY;
isotypes, MO, UC).
Rhizomatis squamis elongatis aurantiacis, laminae
squamis sparsis dissectis aurantiacis praestans.
Epiphytic, rarely epipetric; rhizome 4-6 mm
diam., compact; rhizome scales 7-13 mm long, lin¬
ear-lanceolate, orange-tan, dull, entire with occa¬
sional hairlike processes; leaves 10-32 cm long,
approximate; phyllopodia present; petiole 0“
3 cm) of sterile leaf length, stramineous to tan;
petiole scales 1-2 mm long, scattered, spreading,
linear, lacerated with narrow processes, orange to
brownish black, dull; sterile lamina 10-29 cm long,
1.4 1.8 cm wide, narrowly elliptic, coriaceous, base
attenuate, apex acuminate; veins obscure, ca. 0.8
mm apart, at 75° angle to costa; hydathodes lacking;
lamina scales 1 mm, scattered (dense at base) on
costa and lamina, deciduous on adaxial surface, highly
dissected, substellate, orange, dull; fertile leaves lon¬
ger or shorter than the sterile; petiole l A the fertile
leaf length; fertile lamina linear-elliptic, Vi as broad
as the sterile; intersporangial scales lacking.
Distribution. Wet forests; 0 700 m.
Paratypes. Costa Rica, puntarenas: Osa Peninsula,
9 km N of Rincon de Osa lumber camp, Lloyd 4154
(NY), 4256 (NY); top of ridge just N of OTS station,
Lloyd 4164 (NY): ridge W of Rincon de Osa, Seiden-
schnur 57b (NY), 60 (NY), 61 (NY); at OTS field station,
Seidenschnur 103 (NY), 117 (NY); near airfield, ca. 4
mi. W of Rincon de Osa, Burger & Stolze 5484 (F).
Panama, canai. zone: near Madden Dam, Alston 8903
(CR). DARIEN: Parque Nacional del Darien, Estacion Ran¬
cho Frio at N base of Cerro Pirre, de Nevers et al. 8262
(MO). PANAMA: Rio La Maestra, Allen 39 (LIS).
This species is distingished by the long, orange
rhizome scales and the slender lamina with scattered,
orange, highly dissected abaxial scales.
The species is named in honor of Christiane S.
Anderson, who made many valuable collections of
Elaphoglossum m Costa Rica and who co-authored
the treatment of that genus for Jamaica.
372
Novon
Elaphoglossum cocosense Mickel, sp. nov.
TYPE: Costa Rica, lies Cocos, Pittier 12359
(holotype, US; isotype, US).
Ab E. paleaceo frondibus magnis basi longe attenuatis
necnon squamis rhizomatis, stipitis laminaeque magnis,
tenuibus et sparsirn longidentatis distat.
Epiphytic; rhizome ca. 5 mm diam., compact;
rhizome scales 7-12 mm long, linear-lanceolate,
orange-tan to blackish, thin, margin with sparse,
very long (to 1 mm long) hair-teeth; leaves 37-65
cm long, approximate; phyllopodia present; petiole
the sterile leaf length, orange-tan, densely
clothed with orange-tan scales 4-8 mm long, sparse¬
ly long-hair-toothed; sterile lamina 36-50 cm long,
(2.4-)3.0-4.0 cm wide, narrowly oblanceolate,
chartaceous, base gradually long-attenuate, apex
cuspidate; veins obscure, ca. 1 mm apart, at ca.
70° angle to costa; hydathodes lacking; lamina scales
abaxially 3-6 mm long, abundant, imbricate, linear-
lanceolate, long hair-toothed, adaxially mostly 2 3
mm long, to glabrescent; fertile leaves not seen.
Distribution. Wet forests; “middle elevation.”
Paratypes. Costa Rica, cocos isi.and: Gomez 3327
(CR, F, US), 3362 (CR, NY, US), Klawe, 3 Sep. 1967
(US), Valerio 1095 (CR).
This resembles E. paleaceum in the linear-lan¬
ceolate scales on both laminar surfaces, but is dis¬
tinct in the more dissected blade and petiole scales,
tbe large leal size, the long-attenuate lamina base,
the very long, slender petiole scales, and the rhizome
scales thin, only sparsely toothed.
Elaphoglossum coriifolium Mickel, sp. nov.
TY PE: Panama. Chiriqui: Distrito Bugaba, Cer-
ro Punta, van der Werff & Herrera 6 317
(holotype, MO; isotypes, NY, UC).
Ab E. lingua foliis magnis et lamina basi truncata
differt.
Epiphytic or terrestrial; rhizome 4-5 mm diam.,
creeping; rhizome scales 4-6 mm long, lanceolate,
dark reddish brown, lustrous, entire; leaves 49 60
cm long, 1-2.5 cm distant; phyllopodia present;
petiole l A the leal length, brown; scales at petiole
base 5- 10 mm long, scattered, orange, spreading,
with hairlike processes, scales distally reduced, ob¬
scure, linely skeletonized, appressed; sterile lamina
22-29 cm long, 7 10 cm wide, broadly lanceolate,
coriaceous, base truncate to broadly cuneate, apex
acute to rounded; veins obscure, 0.7-0.9 mm apart,
at i 5-80° angle to costa; hydathodes lacking; lamina
scales lacking except for minute, stellate, black to
dark orange trichomidia; fertile leaves slightly longer
than the sterile; petiole % the fertile leaf length;
fertile lamina similar in shape to the sterile but l A~
as wide; intersporangial scales lacking.
Distribution. Wet forests; 1,800 2,300 m.
Paratypes. Costa Rica, san josk: Desamparados,
Frailes, C. Abejonales, Chavarria <£• Herrera 25 (CR,
MO). PUNTARENAS: upper Rio Buru, Gomez et al. 21527
(CR, MO); San Isidro, La Guaba, Jimenez 303 (CR);
Cerro Chompipe, near Ermita Sta. Cruz, Lems 650202
(NY). Panama, chiriqui: Monte Azul, 1.4 mi. N of Entre
Rios on E slopes of Cerro Punta, Antonio 2724 (MO);
above Boquete, on trail to Pate Macho, McPherson 123IF
(MO); vie. of Guadalupe, between Rio Chiriqui Viejo and
the Continental Divide, Churchill <V de Nevers 5054
(MO); Distrito Bugaba, Cerro Punta, van der Werff&
Herrera 6274 (MO).
Elaphoglossum coriifolium is distinguished by
the creeping rhizome, large, strongly coriaceous,
truncate blade, and stellate trichomidia. It is related
to h. lingua (C. Presl) Brackenr. but is the largest
form in that complex, resembling it in the roughly
ovate or lanceolate, heavily coriaceous lamina,
rounded blade apex, fertile leaf longer than the
sterile, and small, black, semideciduous rhizome
scales.
Antonio 2724 is similar in rhizome diameter and
habit and the broadly cuneate blade base but has
thinner blade texture, more pointed narrower fertile
lamina, and the rhizome scales are not sclerotic.
This may represent a distinct taxon.
The specific epithet comes from the Latin, eo-
rium , leather, and folia, leaf, referring to the highly
coriaceous leaves.
Elaphoglossum eorreae Mickel, sp. nov. TYPE:
Costa Rica. Finca La Selva, Lloyd 4284 (ho¬
lotype, NY).
Elaphoglosso producto Rosenstock affinis, sed stipite
gracili, lamina lineari-elliptica, fronde fertili iis sterilibus
multo breviori, lamina fertili lanceolata, ulterius patria in
regione inferiori abstans.
Epiphytic; rhizome 2-4 mm diam., compact; rhi¬
zome scales 4-5 mm long, linear, tan at base, black¬
ish at tip, lustrous, subentire, with occasional pro¬
cesses; leaves 18-48 cm long, approximate;
phyllopodia present; petiole %-% tbe sterile leaf
length, tan; petiole scales 2 mm long, sparse, black¬
ish, lustrous, linear-lanceolate, spreading, and ap¬
pressed, scattered, orange trichomidia; sterile lamina
13-36 cm long, 1.2-2.1 cm wide, linear-elliptic,
subcoriaceous, attenuate at both ends; veins ob¬
scure, 1 mm apart, at 70° angle to costa; hydathodes
lacking; laminar scales 0.5-1 mm long, abaxially
scattered, linear, hastate or with irregular processes,
brown to black, lustrous, especially toward the lam-
Volume 2, Number 4
1992
Mickel
Elaphoglossum
373
ina base and costa, adaxially glabrous; fertile leaves
much shorter than the sterile (V s -V 2 the sterile), pet¬
iole ca. % the fertile leaf length; fertile lamina 1.0-
2.1 cm wide, lanceolate, base rounded to truncate,
apex acuminate, black scales on petiole and costa
as on sterile leaves linear to arachnidoid (spiderlike);
intersporangial scales lacking.
Distribution. Wet forests; 80-350 m.
Paratypes. Costa Rica, alajuela: Cordillera de Ti-
laran, road between San Ramon and Bajo Rodriguez,
Croat 68167 (MO), 68183 (MO); Upala, Dos Bios, 5 km
al sur de Brasilia, Herrera 1023 (UC). HEREDIA: Cerros
Sardinal, Smith 1790 (CR, MO, UC). Panama, colon:
Santa Rita Ridge Rd., Sytsma 1347 (CR). san blas:
Continental Divide along El Llano Carti Road, Nusagandi,
de Nevers Perez 3711 (MO). PANAMA: Cerro Jefe,
Dressier 3072 (TTC), 3073 (TTC), Correa & Dressier
189 (TTC), 190 (TTC). san jose ISLAND: Johnston 205
(GH).
Elaphoglossum correae is distinguished by the
slender sterile leaves and broader, much shorter
fertile leaves, tan-based black-tipped rhizome scales,
and occurrence at low elevation.
The species is named in honor of Mireya I).
Correa A., outstanding taxonomist and teacher in
Panama.
Elaphoglossum croalii Mickel, sp. nov. TYPE:
Panama. Chiriqui: Distrito Boquete, Fortuna
dam site, van der ff erff & van Hardeveld
6728 (holotype, NY; isotypes, MO, UC).
Ab E. biolleyi stipite gracili, rhizomatis compactioris
squamis linearibus abstans.
Epiphytic; rhizome 4-6 mm diam., compact to
short-creeping; rhizome scales 3-4 mm long, lan-
ceolate-deltate, dark brown, lustrous, entire; leaves
25-38 cm long, approximate; phyllopodia present;
petiole V ut -'A of sterile leaf length, tan to brown;
petiole scales 1-3 mm long, sparse, spreading, dark
brown, lustrous as on rhizome, and squamules/tri-
chomidia less than 0.5 mm long, round to stellate,
appressed, orange to brown; sterile lamina 21-32
cm long, 3.3-4.5 cm wide, elliptic, chartaceous,
attenuate at both ends; veins evident, 1 1.5 mm
apart, at 80° angle to costa, indistinctly united at
margin to form a commissural vein; hydathodes
lacking; lamina abaxially with scattered stellate tri-
chomidia, 0.1 0.3 mm long, brown, adaxially gla¬
brous; fertile leaves about equal to or slightly longer
than the sterile; petiole ca. !4> of fertile leaf length;
fertile lamina 1 2.7 cm wide, narrowly elliptic, at¬
tenuate at both ends; intersporangial scales lacking.
Distribution. Wet forests; 800-1,200(-1,700) m.
Paratypes. COSTA RlCA. CARTAGO: above San Isidro,
Weber 6013 (GH). san jose: ca. 15 km N of Tres Rios,
Lellinger 1384 (MICH). Panama, bocas del toko: For¬
tuna Dam area, Churchill 5862 (MO). CHIRIQUI: Distrito
Boquete, Fortuna Dam site, van der Werff & van Har¬
develd 6693 (UC), van der Werff 6888 (MO); NO de
Fortuna, Correa et al. 2579 (US). COCLE: Fa Mesa above
El Valle, junction with road to Cerro Pilon, Croat 25337
(MO).
Elaphoglossum croatii is distinct from E. lad-
folium (Sw.) J. Smith by the presence of dark, entire
rhizome scales (also scattered on petiole), the pres¬
ence of laminar trichomidia, and thinner texture.
This species is named in honor of Thomas B.
Croat, exceptional collector of plants in tropical
America.
Elaphoglossum davidsei Mickel, sp. nov. TYPE:
Costa Rica. Limon: Valle de Silencio, N of Cerro
Hoffman, Davidse et al. 28653 (holotype, NY;
isotype, MO).
Ab E. alfredii rhizomatis squamis deltatis appressis
sparsis differt.
Terrestrial, rarely epiphytic; rhizome 2.5-4 mm
diam., creeping; rhizome scales mostly 1-1.5 mm
long, brown, debate, dull to somewhat lustrous, en¬
tire to short-toothed, appressed; leaves 11 28 cm
long, 2-10 mm apart; phyllopodia lacking; petiole
the sterile leaf length, tan to brown; petiole
scales 0.5-1.5 mm long, deltate to lanceolate, brown,
dull, entire, appressed or tips spreading; sterile lam¬
ina 9 25 cm long, 1.1-2.1 cm wide, narrowly el¬
liptic, chartaceous; base cuneate, apex attenuate;
veins evident, ca. 2 mm apart, at 60° angle to costa;
hydathodes present; lamina scales evenly and sparsely
distributed on surfaces and margin, lamina also with
round punctae 0.1 mm or lanceolate scales 0.8 mm,
these dull brown, the longer ones with darker lus¬
trous apex, entire or minutely and sparsely toothed;
fertile leaves slightly shorter to slightly longer than
the sterile; petiole % % the fertile leaf length; fertile
lamina 5.5-9 cm long, 1-1.5 cm wide, base broadly
cuneate, apex acute to acuminate; adaxial scales
0.1 0.2 mm long, black, lustrous, deltate, inter¬
sporangial scales lacking.
Distribution. Wet forests; 2,300-3,100 m.
Paratypes. Costa Rica. CARTAGO: 16 km SE of El
Empalme, Smith 2103 (CR, MO, NY, UC); near La
Chonta, Scamman & Holdridge 8022 (GH, US), limon:
between Rio Terbi and Rio Sini, Davidse et al. 28936
(MO, UC). PUNTARENAS-LIMON BORDER: between Cerro
Kasir and Cerro Nai, Davidse et al. 25843 (CR). SAN
JOSE/CARTAGO: 22 km SE of Empalme, Burger & Liesner
6489 (US). Panama, bocas del toko: 2-5 km NW of
peak of Cerro Echandi, Davidse et al. 25085 (MO),
25479 (MO, UC), Gomez et al. 22259 (MO, UC).
374
Novon
Elaphoglossum davidsei is a good match for E.
alfredii Rosenstock in all dimensions but has rhi¬
zome scales that are debate, appressed, and scat¬
tered.
The species is named in honor of Gerrit Davidse,
who has made many interesting collections of Ela¬
phoglossum.
Elaphoglossum eximiiforme Mickel, sp. nov.
TYPE: Panama. SE slopes of Cerro Echandi,
Bocas del Toro, Gomez et at. 22281 (holotype,
NY; isotype, MO).
Ab E. eximio stipite longiori et laminae squainis lon-
gioribus crebrioribus diversa.
Epiphytic; rhizome 2-3 mm diam., compact; rhi¬
zome scales 4 7 mm long, linear, reddish brown,
lustrous, denticulate; leaves 25-33 cm long, ap¬
proximate; phyllopodia lacking; petiole l A~ x k of ster¬
ile leaf length, tan; petiole scales 2 2.5 mm long,
scattered, spreading, subulate, orange-tan, lustrous,
denticulate, many 3-5-celled hairs also present;
sterile lamina 15-22 cm long, 1.1-2.1 cm wide,
linear-lanceolate, chartaceous, base broadly cune-
ate, apex acuminate; veins evident, mostly simple,
a few once-forked, 1.2-1.8 mm apart, at 65° angle
to costa; hydathodes present; blade scales 2 3 mm
long, scattered, ascending, subulate, orange-tan, dull,
denticulate; fertile leaves shorter than to nearly
equaling the sterile; petiole 2 A the fertile leaf length;
lamina linear-lanceolate, base broadly cuneate, apex
acuminate; intersporangial scales 1.5 2.5 mm long,
dense, linear-lanceolate, dark reddish brown, lus¬
trous, denticulate.
Distribution. Wet forests; 2,600-2,800 m.
Paratypes. Costa Rica. CARTACO: Volcan de Turrialba,
Pittier 7492 (NY, US).
I bis species differs from E. eximium (Mett.) Christ
in the longer petiole and longer laminar scales.
Elaphoglossum fuliginosum Mickel, sp. nov.
TYPE: Costa Rica. San Jose: ca. 3 km NW of
Cascajal near the union of the Cascajal and
Bajo Maquina rivers, Taylor & Taylor 11329
(holotype, NY).
Ab E. muscoso laminae squamis parvis sparsis nigris
absimilis.
Epiphytic; rhizome 4-6 mm diam., compact; rhi¬
zome scales 5-10 mm long, linear, black to brown,
lustrous, subentire, occasionally with slight teeth;
leaves 10 30 cm long, approximate; phyllopodia
present, obscured by dense scales at petiole base;
petiole 2 /s the sterile leaf length, tan-orange; petiole
scales 2 5 mm long, scattered to dense, lanceolate,
black, especially the center, distally the petiole scales
all black, lustrous, erose, spreading, smaller ones
0.6-1.5 mm long, short-ciliate-denticulate, ap¬
pressed; sterile lamina 6-17 cm long, 1.8-3.8 cm
wide, oblong, coriaceous, base broadly cuneate, apex
obtuse; veins obscure, ca. 1 mm apart, at 75° angle
to costa; hydathodes lacking; lamina scales 0.7-1.5
mm long, abaxially scattered, appressed, lanceolate,
black to mottled black especially near lamina mar¬
gin, with black cell contents and clear walls, den¬
ticulate to short-ciliate, slightly arched, adaxially
orbicular, peltate, white, scattered, entire; fertile
leaves longer than the sterile; petiole V\ the fertile
leaf length; fertile lamina narrowly elliptic, base
broadly cuneate, apex acute to obtuse; costal scales
ca. 1 mm long, black, appressed, dense; interspor¬
angial scales ca. 1 mm long, scattered, black, lus¬
trous, denticulate.
Distribution. Wet forests; 1,700-2,400 m.
Paratypes. Costa Rica, cartago: near Empalme,
Brown CR-64 (US). LIMON: Valle de Silencio, along the
Rio Terbi, Davidse et al. 28783 (CR, MO); ca. 1 kin S
of El Empalme along Interamerican Hwy., Lellinger 1200
(F). san jose: ca. 3 km NW of Cascajal, Taylor & Taylor
11338 (US); S of Cartago, ca. 4 km S of El Empalme,
near La Chonta, Lellinger 1200 (US), 1004 (US). PROV.
UNKNOWN: vie. of Coliblanco, Maxon 281 (NY, US).
Elaphoglossum fuliginosum differs from E. mus-
cosum (Sw.) T. Moore by the small, sparse, black
laminar scales.
The specific epithet derives from the Latin, fu-
ligio, soot, alluding to the black lamina scales.
Elaphoglossum gloeorrhizum Mickel, sp. nov.
TYPE: Costa Rica. Volcan Barba, Lloyd 1140a
(holotype, NY).
Rhizomate crasso nudo glutinoso, stipite forti, necnon
lamina eximie coriacea notabilis.
Epiphytic, rarely terrestrial; rhizome 5-9 mm
diam., short-creeping, resinous, black; rhizome scales
lacking; leaves 35-53 cm long, approximate to 5
mm apart; phyllopodia 1.5—3( 4) cm long; petiole
Vi-Vfc of sterile leaf length, orange-tan (to brown),
lacking scales but with resinous dots at base, rarely
with a few minute (1-mm-long) brown scales near
base; sterile lamina 21-40 cm long, 2.3-4.6 cm
wide, narrowly elliptic, strongly coriaceous, base
cuneate, apex attenuate; veins obscure, 1 mm apart,
at ca. 75° angle to costa; hydathodes lacking; lamina
glabrous or with very obscure minute dots on abaxial
surface; fertile leaves about same length as the sterile
Volume 2, Number 4
1992
Mickel
Elaphoglossum
375
leaves; petiole V 3 -V 2 of fertile leaf length; fertile
lamina 1-3.4 cm wide, narrowly elliptic, base cu-
neate, apex attenuate, lacking indument.
Distribution. Wet forests; 1,800—2,700 m.
Pnratypes. Honduras, ocoteit.oue: Aldea F.1 Portia,
Agua Caliente-Sta. Rosa de Copaii, Croat & Hannon
6 3800 (MO). Costa Rica, i.imon: Cordillera de Tala-
rnanca, Valle de Silencio, along Rio Terbi, Davidse et al.
28972 (MO); Cordillera de Talamanca, S side ot cordillera
between Rio Terbi and Rio Sini, Davidse et al. 28972
(MO); Varablanca, Lloyd 4124 (NY); region of Zarcero,
Smith H104 (F, MO); Werckle , in 1910 (NY, UC).
This species is distinguished by the thick, naked,
resinous rhizome, stout petiole, and coriaceous lam¬
ina.
'I’he specific epithet derives from the Creek, gloios,
glue, and rhiza , root, for the black, resinous rhi¬
zome.
Elaphoglossum grayumii Mickel, sp. nov. PE:
Costa Rica. Puntarenas: 5 km S of San Vito,
Finca Wilson, Mickel 3066 (holotype, NY).
Ab E. exirnio hydathodiis nullis, costae squamis fuse is
processibus provisis, phyllopodiis, necnon lamina fertili
basin versus attenuata absimilis.
Epiphytic; rhizome 3-4 mm diam., compact to
short-creeping; rhizome scales 1-2 mm long, linear,
black, rarely orange-tan, touched with black, lus¬
trous, entire; leaves 32-40 cm long, approximate
to 6 mm apart; phyllopodia present; petiole less than
yj 5 the sterile leaf length (12 cm long), essentially
lacking, stramineous to tan; petiole scales at petiole
base 1-2 mm long, sparse, appressed, lanceolate to
linear, brown to black, lustrous, entire to occasion¬
ally toothed, distally with stellate scales to trichom-
idia less than 0.5 mm long; sterile lamina 30 38
cm long, 1.7-3.0 cm wide, linear-elliptic, coria¬
ceous, base attenuate, apex acuminate; veins ob¬
scure, ca. 1 mm apart, at 75° angle to costa, uniting
at margin to form commissural vein; hydathodes
lacking; scales on costa sparse, linear to lanceolate,
black, lustrous, ca. 1 mm long, irregularly lobulate,
laminar scales reduced to 0.3 0.8 mm stellate hairs
or 0.1 mm punctae; fertile leaves slightly shorter
than the sterile; petiole %-'/ 2 the lertile leaf length;
fertile lamina narrowly elliptic, attenuate at both
ends; intersporangial scales lacking.
Distribution. Wet forests; 100-2,000 m.
Paratypes. Honduras. Santa Cruz de Yojoa, Cortes,
Edwards 721 (F). Nicaragua, zelaya: Cerro La Pi-
mienta, Pipolv 5243 (CR. UC); Cano F.l Hormiguerro,
Pipoly 5928 (CR, MO); Costado SW of Cerro El Hor¬
miguerro, Grijalva 488 (UC, US); ca. 6 km N of Colonia
San Antonio on new road to Colonia San Martin, Stevens
9015 (UC). Costa Rica, alajuela: ca. 20 km SW of San
Ramon, Smith 2254 (CR, UC); between San Ramon and
Bajo Rodriguez, Croat 68181 (MO, UC). CARTAGO: SE
of bridge over Rio Pacuare, Lent 670(F). HEREDIA: Cerros
Sardinal, ca. 2-2.5 km N of Chilamate de Sarapiqui,
Smith 1791 (CR, NY, UC); Finca La Selva, Grayum
1964 (NY). LIMON: vie. of Guapiles, Stand-ley 37070
(US). PUNTARENAS: San Vito, W of Finca Wilson, Lloyd
4223 (NY); Osa Peninsula, Tropical Science Center field
station, Seidenschnur 119 ( NY); Boruca, Ocampo 1340
(CR). san jose: basin of El General, Skutch 2158 (MO,
NY, UC). Panama, canal ZONE: Barro Colorado Island,
Croat 10996 (MO). CH1RIQUI: 2.5 km from Questu Piedra
along Rio Monte Road, Folsom 3981 (MO, L'C). gogee:
foot of Cerro Pilon, above El Valle de Anton, Porter et
al. 4666 (MO, NY).
Elaphoglossum grayumii differs from E. exi-
mium (Mett.) Christ in lacking hydathodes, and by
the presence of dark costal scales (with lateral pro¬
cesses), phyllopodia, and fertile lamina attenuate at
the base.
The species is named in honor of Michael II.
Grayum, collector and student ol the Costa Rican
flora.
Elaphoglossum heterochroum Mickel, sp. nov.
TYPE: Costa Rica. Region sureste del Lago
Dabagri, Gomez et al. 23217 (holotype, NY;
isotype, MO).
Inter E. cuspidatum et E. paleaceum quasi intermedia,
a prima stipitis costaeque squamis ad mediam nigro-pur-
pureis, laminae squamis lanceolatis arcuatis aurantiacis
laxe dispositis diversa.
Epiphytic; rhizome 3-4 mm diam., compact; rhi¬
zome scales 4-5 mm long, linear, castaneous, lus¬
trous, margin with sparse stiff cilia often abraded so
as to appear entire; leaves 24-35 cm long, ap¬
proximate; phyllopodia present; petiole, the sterile
leaf length, tan to orange; petiole scales 1-3 mm
long, dense, spreading, linear-lanceolate, castaneous
to black-lustrous, distally on petiole scales with dark
center and pale margin, ciliate-denticulate; sterile
lamina 18-24 cm long, 1.2-2.1 cm wide, linear-
elliptic, coriaceous, base cuneate, apex acuminate;
veins obscure, ca. 1 mm apart, at 80-85° angle to
costa; hydathodes lacking; lamina scales 1-2 mm
long, dense, appressed, lanceolate, pale, dull, ciliate-
denticulate, costal scales black-centered, adaxially
subimbricate to very sparse or lacking, lanceolate
to suborbicular, whitish; fertile leaves longer than
the sterile; petiole V 3 ~% the fertile leaf length; fertile
lamina linear, base broadly cuneate, apex acumi¬
nate; costal scales black, appressed. suborbicular,
ciliate-denticulate, adaxially denser than on sterile
lamina, suborbicular; intersporangial scales lacking.
376
Novon
Distribution. Wet forests; 600-2,900 m.
Paratypes. Costa Rica, alajuela: N side of Volcan
Arenal, Lent 2921 (F); 8 mi. NW of San Ramon on road
to Tilaran, Lloyd 4194 (NY). CARTACO: near Rio Pejibaye,
2 km SW of Taus, Lent 2971 (F). LIMON: ridge between
Quebrada Camagre and Rio Barbilla, Grayum 8904 (MO);
between headwaters of Rio Madre de Dios and Quebrada
Barreal, Grayum 8802 (MO). HEREDIA; Virgen del So¬
corro Rio Sarapiqui Cariblanco, Chacon & llerrera 1162
(MO). PliNTARENAS: just N of Santa Elena on Fila Coton,
S of Agua Caliente, Davidse et al. 28273 (MO). Panama.
coci.E: La Mesa above El Valle, Folsom 5629 (MO, UC);
foot of Cerro Pilon, above El Valle de Anton, Porter el
al. 4662 (MO, NY); 9.4 km above El Cope, Croat 44657
(MO). DARIEN: Pirre Massif, Alturas de Nique, McPherson
I2202C (MO), panam a: Distrito Capira, Cerro Campana,
van der Werff & Herrera 6214 (NY, UC), Porter et al.
4939 (MO, UC).
Elaphoglossum heterochroum resembles both K.
cuspidatum and K. paleaceum, but the scales of
the petiole and costa are blackish at the center, the
lamina scales are imbricate, lanceolate, orange, and
the rhizome scales castaneous. It is close to K. ni-
grocostatum Mickel of Venezuela, but differs in the
bicolorous costal and petiolar scales.
The specific epithet comes from the Creek, ket-
eros, other, chroma , color, referring to tlie bicolo¬
rous petiolar and costal scales.
Elaphoglossum killipii Mickel, sp. nov. TYPE:
Costa Rica. San Jose: vie. of Santa Maria de
Dota, Standley 11625 (holotype, US).
Proxime ad E. longicrurem accedens, sed frondibus
majoribus rhizoinatisque squamis bicoloribus absimilis.
Epiphytic; rhizome 2.5-4 mm diam., compact to
short-creeping; rhizome scales 2-4 mm long, linear-
lanceolate, bicolorous, castaneous to black-centered
with orange-tan margin, entire; leaves (14 )26 46
cm long, approximate to 2 mm apart; phyllopodia
distinct; petiole the sterile leaf length, glabrous
to sparsely scaly; petiole scales 3-5 mm long, lan¬
ceolate, lustrous, bicolorous as on rhizome, entire;
lamina (15—)20-32 cm long, (1.5—)2.0-3.4 cm wide,
narrowly oblanceolate to elliptic, coriaceous, grad¬
ually attenuate at base, apex acute to acuminate;
veins obscure, ca. 1 mm apart, at 60 70° angle to
costa; hydathodes lacking; blade surfaces glabrous
or with very sparse minute stellate trichomidia on
abaxial surface and often with a few lanceolate scales
on base of abaxial costa; fertile leaves slightly longer
than the sterile; petiole of fertile leaf length;
fertile lamina slightly narrower than the sterile, base
attenuate, acute to rounded; intersporangial scales
lacking.
Distribution. Wet forests; 1,000-2,000 m.
Paratypes. Costa Rica. PDNTARENAs: valley of the Rio
Coton between Sitio Coton and Sitio Tablas, Davidse
24452 (CR, MO); Upper Rio Burn, Gomez et al. 21487
(MO), san JOSE: Quebrada de los Yases, entre La Guaria
y Palmilera, Dota, Jimenez 1455 (CR, NY); Alto del
Sacatal, Pittier (herb. no. 10536) (CR). Panama, chi-
RIQUI: slopes of Cerro de Lino, above El Boquete, Maxon
5213 (LJS); vie. of Boquete, Common 853 (ARIZ), 895
(ARIZ), 897 (ARIZ), 1014 (ARIZ, US), Killip 5180
(US), 5203 (US), Maxon 5032 ( US); lower slop >es of Cerro
Pelota, Knapp 1491 (CR).
Elaphoglossum killipii closely resembles E. Ion-
gicrure Christ, differing in its larger size and bico¬
lorous rhizome scales.
The species is named in honor of Ellsworth P.
Killi [>, whose avid collecting has contributed greatly
to our knowledge of Central American pterido-
phvtes.
Elaphoglossum lanceiforme Mickel, sp. nov.
1 Y PE: Costa Rica. Los Angeles de San Ramon,
IIrenes 16141 (holotype, NY).
Ab k. setoso (Liebrn.) T. Moore lamina lanceolata ma-
jori, stipitis laminaeque squamis longioribus, necnon lam¬
ina fertili angusta diversa.
Epiphytic; rhizome 2-4 mm diam., compact; rhi¬
zome scales 3-5 mm long, linear-lanceolate, dark
reddish brown, lustrous, entire, appressed; leaves
11-30 cm long, approximate; phyllopodia present;
petiole A the sterile leaf length, tan; petiole scales
2-4 mm long, abundant, subulate, dark reddish
brown, lustrous, denticulate, spreading; sterile lam¬
ina 8-19 cm long, 1.8 3.2 cm wide, linear-lanceo¬
late, chartaceous, base broadly cuneate to rounded,
apex acuminate; lamina scales 1-5 mm, scattered,
hairlike, subulate, dark reddish brown, lustrous, the
longer ones on costa and margin, denticulate,
spreading; veins evident, 1.5-2 nun apart, at ca.
70° angle to costa; hydathodes present; fertile leaves
shorter than the sterile; petiole % the fertile leaf
length; fertile lamina 6.5-9 cm long, 0.8-11 cm
wide, linear to linear-lanceolate, base cuneate, apex
acuminate; intersporangial scales 3-5 mm long,
abundant, hairlike, orange-tan to dark reddish brown,
lustrous, remotely and indistinctly denticulate.
Distribution. Wet forests; 800- 1,5()()( 2000) m.
Paratypes. Costa Rica, alajuela: ca. 23 km NE of
San Ramon, Taylor 1 7846 (NY); 8 mi. NW of San Ramon
on road to Tilaran, Lloyd 4193 (NY). CARTACO: Reserva
de Tapanti, Gomez 18919 (MO, UC). HEREDIA: between
Rio Peje and Rio Sardinalito, Grayum 6666 (MO); Laguna
del Barva and summit of Volcan Barva, Grayum & Que-
sada 7432 (UC). SAN JOSE: near km 50 along Pan Amer¬
ican Hwy., Cerro de la Muerte, Seidenschnur 232 (NY);
along Rio Zurqui, Smith 1709 (VC); Alto La Palma, NW
of San Jose, Moran 2352 (MO); vie. of La Palina, Maxon
Volume 2, Number 4
1992
Mickel
Elaphoglossum
377
Harvey 7907 (US); La Hondura, Standley 3791 (US);
Parque Nacional Braulio-Carrillo, Moran 3265 (NY).
PUNTARENAS: 5 km S of San V ito, Finca Wilson, Mickel
3132 ( NY); Monteverde, Palmer 62 (NY). Panama, bogas
DEL TORO: Fortuna Dam Chiriqui Grande road, McPher¬
son 6792 (MO), chiriqui: Distrito Boquete, Fortuna Dam
site, van der If erfj & van Hardeveld 6819 (MO); 0.5
km E of Cerro Pate Macho, Knapp et al. 2109 (MO).
PANAMA: Cerro Jefe, Witherspoon 8539 (MO). VERAGUAS:
Cerro Tute, Moran 4033 (NY).
Elaphoglossum lanceiforme is closely allied to
E. atrobarbalum of Panama but the latter has lon¬
ger, black intersporangial scales rather than subulate
orange-tan ones and the lamina apex of the latter
tends to be broadly acute rather than narrowly so.
Elaphoglossum lankesteri Mickel, sp. nov.
TYPE: Costa Rica. Heredia: Volcan Barba,
Lloyd 1201 (holotype, NY).
Ab E. latifolio lamina majori distat.
Epiphytic, less commonly terrestrial; rhizome 5
12 mm diam., compact to short-creeping; rhizome
scales 4-8 mm long, linear-lanceolate, dull tan to
brown, with occasional lateral processes; leaves 55
90 cm long, approximate to ca. 5 mm apart; phyl-
lopodia distinct; petiole usually 14-14 the sterile leaf
length; petiole scales present only at the base, to 4
mm long, spreading, lanceolate, with occasional pro¬
cesses; sterile lamina 37-60 cm long, 7-1 1.5 cm
wide, elliptic, chartaceous to subcoriaceous, base
cuneate, apex acuminate; veins obscure, ca. 1 mm
apart, at ca. 80° angle to costa; hydathodes lacking;
lamina surfaces glabrous or with brown stellate Iri-
chomidia; fertile leaves slightly shorter to slightly
longer than the sterile, somewhat narrower; petiole
ca. !4 the fertile leaf length; intersporangial scales
lacking.
Distribution . Wet forests; (300-)800 2,600 m.
Paratypes. Costa Riga, alajuela: ca. 20 km NW of
San Ramon, Smith 2272 (CR, MO, NY, UC). cartago:
El Zanjon, Lankester 645 (US). HEREDIA: Cerro Chorri-
pipe, N of San Rafael, Lems 64082908 (NY), union:
Reserva Indigena Talamanca Sukut, Hummel 17576
(MO). Panama, chiriqui: Distrito Boquete, Fortuna Dam
site, van der If erfj A' van Hardeveld 6617 (MO); 2.2
km SW of Cerro Punta, Croat 26307 (MO). PANAMA,
COMARCA DE SAN BI AS: Nusagandi, van der II erjf 7005
(MO, NY).
This species is distinct from E. latifolium by being
considerably larger. It is not to be confused with h.
coriifolium because of E. lankesteri s greater rhi¬
zome diameter, cuneate laminar base, more pointed
apex, and thinner texture.
The species is named in honor of C. 11. Lankester,
who collected many fine plant specimens in Costa
Rica.
Elaphoglossum latum (Mickel) Atehortua ex
Mickel, comb, et stat. nov. Elaphoglossum
apodum (Kaulf.) Schott ex J. Smith var. latum
Mickel, Amer. Fern J. 69: 100 (1979). TYPE:
Guatemala: Steyermark 46369 (holotype, E).
Elaphoglossum lepidothri.x Mickel, sp. nov. TYPE: Pan¬
ama. Code: La Mesa above El Valle, Croat 25402
(holotype, NY).
Ab E. muscoso stipitis squamis linearibus ciliato-den-
ticulatis patulis, laminaeque squamis linearibus ciliato-den-
ticulatis sparsis diversa.
Epiphytic; rhizome 4-8 mm diam., compact; rhi¬
zome scales 3-6 mm long, linear with hairlike at¬
tenuate apex, reddish brown, lustrous, entire; leaves
12-31 cm long, approximate; phyllopodia incon¬
spicuous, covered by scales; petiole 14-14 the sterile
leaf length, orange-tan; petiole scales 5-8 mm long,
spreading, linear, brown to blackish, lustrous, ciliate-
denticulate, mixed with 0.5-1 mm understory of
orange-tan scales; sterile lamina 8 1 6 cm long, 3.0
4.3 cm wide, lanceolate-oblong, coriaceous, base
broadly cuneate to rounded, apex rounded; veins
obscure, ca. 1 mm apart, at 80° angle to costa;
hydathodes lacking; lamina scales abaxially 2 3 mm
long, linear-lanceolate, orange-tan, ciliate-denticu-
late, adaxially mostly glabrous, costa occasionally
scaly toward base; fertile leaves longer than the
sterile; petiole 14-14 the fertile leal length, usually
longer than the entire sterile leaf; fertile lamina
linear-lanceolate, base rounded, apex obtuse; petiole
scales blacker than on sterile leaf, especially distally,
costal scales 2 mm long, linear, ciliate-denticulate,
intersporangial scales 1 mm long, lanceolate, black-
brown, lustrous, ciliate-denticulate, teeth longer than
the scale body width.
Distribution. Wet lorests; 500-1,300 in.
Paratypes. PANAMA. BOCAS DEL TORO: Oleoducto road,
between Rios Guabo and Guabito, Churchill 5037 (MO).
GHIRIQL1: Fortuna Dam area, Churchill et al. 4843 (MO),
coc IE: Cerro Pilon, Liesner 782 (MO); 8 km above El
Cope, llammel 787 (CR). DARIEN: Cerro de Garagara,
Pittier 5649 (US); SW ridge leading to Alturas de Nique,
Hartman 12389 (MO). PANAMA: Valle de Anton, Ocampo
87t> (CR); summit of Cerro Campana, Porter et al. 4910
(MO); Cerro Jefe, ITitherspoon 8537 (MO).
This species is in the E. muscosum complex, but
is distinguished by the linear, ciliate-denticulate,
spreading petiolar scales.
The specific epithet comes from the Greek, lepis,
scale, and trichos, hair, referring to the very slender
scales on petiole and lamina.
378
Novon
Elaphoglossum maculatum Mickel, sp. nov.
TYPE: Panama. Code: Cerro Pilon, summit,
Dwyer & Lallathin 8959 (holotype, NY).
AL> E. pygmaeo frondibus majoribus margine integris,
lamina fertili baud decurrenti sporisque cristatis diversa.
Epiphytic; rhizome 1 2 mm diam., horizontal,
compact; rhizome scales 1 mm long, linear, hairlike,
hlack, or mixed orange and black scales, strongly
indurated, lustrous, grading to dull tan, entire; leaves
4.515(-20) cm long, approximate; phyllopodia
lacking; petiole l A- l A of sterile leaf length, tan;
petiole scales 1.5—2 mm long, scattered, ascending
to spreading, linear to subulate, orange-tan, to scle¬
rotic, lustrous, dark brown, entire; sterile lamina 3
12 cm long, 0.6-1.9 cm wide, narrowly elliptic,
chartaceous, base attenuate, apex acuminate; veins
free, 1.5-2 mm apart, at 60° angle to costa; hy-
dathodes present; lamina scales 1-1.5 mm long,
sparse on costa, veins, and margin, spreading, su¬
bulate, dull, tan to sclerotic but not dark brown,
entire to crenulate; fertile leaves longer than the
sterile, the petiole %-% °f fertile leaf length, as long
as the sterile leal; fertile lamina narrowly elliptic,
base cuneate, apex acute to acuminate; scales pres¬
ent on costa and margin but lacking among spo¬
rangia.
Distribution. Wet forests; 0-1,400 m.
Paratypes. Panama, cocle: Alto Calvario, 5.5 mi. N
ol El Cope, Croat 97549 (MO); vie. of La Mesa, N of
El Valle de Anton, Croat 57282 (UC). DAMIEN: SW ridge
leading to Alturas de Nique, Hartman 12380 (MO); Cerro
Pirre, Duke A Elias E-13706 (MO, NY). Panama: Ran¬
cho Chorro, mts. above Torti Arriba, Folsom et al. 5695
(CR, MO); Cerro Jefe, Churchill 395 HI (MO). SAN Ml. as:
Cerro Brewster, de Nevers 4033 (MO). VERAGUAS: Cerro
l ute, Moran 4041 (MO); mts. W of Alto de Piedras Siclo
Basico school N of Santa Fe, Hummel 4693 (MO).
Colommia. EL VAI.I.E: El Silencio, Yanaconas, Fillip A
Garcia 33790 (US); Sabaletas, Killip A Cuatrecasas
38756 (US). INTENDENGi.A DEL CHOCO: La Concepcion, 15
km E of Quibdo, Archer 2083 (US); Rio Nuqui, Haught
5499 (US).
This species is closest to E. pygmaeum (Mett. ex
Kuhn) Christ of Ecuador, which has smaller (4-8
cm long) leaves with crenulate margins, decurrent
fertile lamina, and spores spiculate (vs. crested in
Panama). Elaphoglossum maculatum is variable in
the petiole—mostly short to sessile, some stipitate.
Elaphoglossum pygmaeum has a rhizome less than
1 mm diameter and orange rhizome scales.
The specific epithet comes from the Latin, ma-
culosus , spotted, alluding to the white patches on
the leaves.
Elaphoglossum micropogon Mickel, sp. nov.
TYPE: Costa Rica. Cartago: Tapanti, ca. 15
km S of Paraiso, Mickel 2369 (holotype, NY).
Ab E. paleaceo stipite brevi et lamina fertili lineari
differt.
Epiphytic or epipetric; rhizome 3 4 mm diam.,
compact; rhizome scales 4-6 mm long, linear-lan¬
ceolate, dark reddish brown, lustrous, ciliate-dentic-
ulate; leaves 30-39 cm long, approximate; phyl¬
lopodia present; petiole ca. '/[„ of sterile leaf length,
tan; petiole scales 3-4 mm, dense, spreading, linear-
lanceolate, reddish brown to light orange, lustrous
to dull, ciliate-denticulate; sterile lamina 24 30 cm
long, 1.4 2.7 cm wide, linear-lanceolate, charta¬
ceous, base attenuate to narrowly rounded, apex
acuminate; veins obscure, 1 mm apart, at 70° angle
to the costa; hydathodes lacking; lamina scales dense,
imbricate, abaxially 2 3 mm long, linear-lanceolate,
ciliate-denticulate, pale orange; costal scales 2-4 mm
long, dark-tipped; adaxially scales 1-2 mm long,
linear with 2 4 strongly ascending teeth, scales erect,
reddish brown; fertile leaves slightly longer than the
sterile; petiole x A~Vi of the fertile leal length; fertile
lamina 1 1 — 14 mm wide, linear; adaxial scales and
costal scales dark, arachnidoid (spiderlike), ca. 1
mm long, intersporangial scales 0.1 0.3 mm long,
orange-tan, substellate, among the sporangia and
hidden by them.
Distribution. Wet forests; 1,000-1,400 m.
Paratypes. Costa Riga, alajuela: Reserva Monte-
verde, Rio Penas Blancas, Bello 134 (CR). cartago: along
tributary of Quebrada Casa Blanca Tapanti, Grayum A
Sleeper 3673 (MO); Canon del Rio Grande de Orosi,
Chacon et al. 1520 (CR). PUNTAREN AS: Monteverde re¬
serve along Rio Penas Blancas to small lagoon, llammel
et al. 15408 (MO); Navarro, Werckle, in 1905 (P). san
JOSE: 1-4 km N of La Hondura, Lloyd 4111 (NY).
Panama. CHIRIQUI: between Gualaca and Fortuna Dam
site on Rio Cbiriqui, Croat 49977 (MO); Distr. cle Bugaba,
along road Volcan-Santa Clara, van der Werj)'& Herrera
7070 (MO, NY, UC).
I bis species is similar to E. auricomum (Kunze)
T. Moore in the short petiole and linear adaxial
laminar scales, but that species has skeletonized
scales on both laminar surfaces whereas E. micro-
pogon has linear-lanceolate scales on the abaxial
surface. It is closely allied to E. paleaceum, but
differs in the linear fertile lamina, short petiole, and
linear adaxial scales.
Lloyd 4245 (Costa Rica, San Vito, Finca Wilson,
NY) has a rounded lamina base, sterile lamina 4
4.5 cm wide, and fertile lamina ca. 2 cm wide, and
may represent a distinct taxon.
Volume 2, Number 4
1992
Mickel
Elaphoglossum
379
The specific epithet comes from the Creek, mik-
ros, small, and pogon , beard, referring to the short,
scaly petiole.
Elaphoglossum mitorrhizum Mickel, sp. nov.
TYPE: Costa Rica. Alajuela: 1 1 km N of San
Ramon, Mickel 296/ (holotype, NY).
Ab E. proximo rhizomatis squamis bicoloribus (parte
centrali nigra saepissime sola persistenti) laminisque utr-
inque nigropunctatis diversa.
Epiphytic; rhizome 1-1.8 mm diain., long-creep¬
ing; rhizome scales 0.5 1 mm long, appressed, lan¬
ceolate to debate, black, lustrous, entire; leaves (5-)
8-16 cm long, 1.5-5 cm distant; phyllopodia pres¬
ent; petiole V 2 -V 2 of sterile leaf length, tan to brown;
petiole scales 0.2-1 mm long, lanceolate to round,
black to brown, appressed; sterile lamina 3.5-11
cm long, 1.4 4.2 cm wide, ovate-lanceolate, cori¬
aceous, base broadly cuneate, apex acuminate; veins
obscure, ca. 1 mm apart, at 75-80° angle to costa;
hydathodes lacking; lamina punctate abaxiallv with
round to substellate, black punctae 0.1 0.2 mm
long, fewer punctae adaxially; abaxial costa occa¬
sionally with sparse, black, reduced, lanceolate, has¬
tate scales; fertile leaves longer than the sterile;
petiole { A~V\ of fertile leaf length; fertile lamina
narrowly lanceolate, base broadly cuneate, apex
acuminate; scales few, small, black on costa, inter-
sporangial scales lacking.
Distribution. Wet forests; 800 1,500 m.
Paratypes. Costa Rica, alajuela: ca. 1 km SE of La
Balsa de San Ramon, ca. 16 km NW of San Ramon,
Smith 2297 (MO, UC), ca. 20 km SW of San Ramon,
Smith 2215 (MO, UC). guanacaste: Fila del Volcan
Cacao, Chacon dr Chacon 2291 (MO): Parque Rincon
de La Vieja, Herrera 1469 (NY). SAN JOSE: Estacion
Carrillo, Parque Nacional Braulio Carrillo, Zamora &
Elizondo 677 (CR). Panama. CHIRIQIT: Distrito Boquete,
Fortuna Dam site, van der Werff& van Hardeveld 6811
(MO, UC).
The very slender, long-creeping rhizome with
sparse, small, black scales and the abaxial lamina
with dark punctae or trichomidia distinguish this
species. It has been misinterpreted in most herbaria
as E. proximurn Christ (synonym, E. bradeorum
Christ), hut the type specimen of E. proximurn has
scattered, lustrous, golden scales on the abaxial lam¬
ina surlace.
The specific epithet comes from the Greek, mitos,
thread, and rhiza , root, referring to the very slender
rhizome.
Elaphoglossum montgomeryi Mickel, sp. nov.
TYPE: Costa Rica. Cartago: ca. 22 km E of
Turrialba, above Platanillo, Mickel 3406 (ho¬
lotype, NY).
Ab E. lingua frondium stipite atque costa squamis
nigris obsitis differt.
Epiphytic; rhizome 2 mm diam., creeping; rhi¬
zome scales ca. 1.5 mm long, black, sclerotic, lus¬
trous, lanceolate, with weak irregular hair-teeth, es¬
pecially at scale base; leaves 11-18 cm long, ca. 1
cm apart; phyllopodia present; petiole ca. V b the
sterile leaf length, light brown; petiole scales 2.5-
4 mm long, black, lanceolate to linear-lanceolate,
lustrous to dull, with irregular lateral processes; ster¬
ile lamina 8 15 long, (2-)3-4.5 cm wide, oblong,
coriaceous, base narrowly cuneate, apex rounded;
veins obscure, ca. 1 mm apart, at 80° angle to costa;
hydathodes lacking; lamina with abaxial costal scales
to 3 mm long, as on petiole but reduced, scattered,
still more reduced (ca. 1 mm long) on adaxial costa
and abaxial lamina surface; fertile leaves far ex¬
ceeding the sterile; petiole ca. V 2 the fertile leaf
length; fertile lamina 17 cm long, 1.1 cm wide, base
abruptly narrowed, apex rounded; lamina scales in¬
cluding intersporangial scales lacking.
Distribution. Wet forests; 700-1,500 m.
Paratypes. Costa Riga. CARTAGO: beside Rio Pejibaye,
2 km SW of Taus, Lent 2962 (F). L1MON: along road
233 between Turrialba and Siquirres, Crosby 2548
(MICH). Panama, panama: summit of Cerro Campana,
Porter et al. 4944 (MO).
This species is close to E. lingua in the creeping
rhizome, very dark, sclerotic, small, deltate rhizome
scales, and rounded lamina apex, hut is distinct from
that species by the black scales on the stipe and
costa. The petiole scales are linear-lanceolate and
conspicuous, 3-5 mm long, with slender lateral pro¬
cesses. The costal scales are slender with propor¬
tionally longer processes; abaxially on costa they
are 1-2 mm long, becoming as small as 0.2 mm
on the lamina surface, with fewer adaxially.
The species is named in honor of James I). Mont¬
gomery, collector, friend, and student of llryopteris.
Elaphoglossum moranii Mickel, sp. nov. TYPE:
Costa Rica. Heredia: end of rte. 113 where
road crosses Rio Patria, Moran 3075 (holo¬
type, NY).
Ab E. lingua lamina angustiori differt.
Epiphytic; rhizome 2-3 mm diam., creeping; rhi¬
zome scales 0.4-1 mm long, deltate, black, sclerotic,
lustrous, entire, appressed, scattered; leaves 13-
26(-32) cm long, 2-8 mm apart; phyllopodia pres¬
ent; petiole V\- l A the sterile leaf length, stramineous
to brown, glabrous or with dark stellate trichomidia;
380
Novon
sterile lamina 8-18 cm long, 1.2-2.7(-3.4) cm
wide, narrowly oblong, coriaceous, base narrowly
cuneate, apex rounded; veins obscure, ea. 1 mm
apart, at 60 70° angle to costa; hydathodes lacking;
fertile leaves far exceeding the sterile; petiole Vz-
V\ the fertile leaf length; fertile lamina 7-12(-30)
mm wide, narrower than the sterile; lamina scales
lacking.
Distribution. Wet forests; 800 2,500 m.
Paratypes. Costa Rica, a la jit. la: along upper Rio
Sarapiqui, near Cariblanco, Burger & Antonio 11128
(F); road between San Ramon and Bajo Rodriguez, Croat
6 8099 (MO); Upala, Bijagua Camino de El Pilon a Rio
Celeste, Herrera 2081 (MO); ca. 20 km N of San Ramon,
Moran 3220 (NY), guanacaste: slopes of Miravalles,
above Bijagua, Gomez et al. 19153. HEREDIA/SAN JOSE:
along Rio Para Blanca, Cerro de Zurqui, Burger et al.
9308 (F). HEREDIA: Volcan Barba, Lloyd 4212 (NY).
Panama, chiriqui: Ei of the Fortuna main campsite, Fol¬
som et al. 5283 (MO); NO de Fortuna, Correa et al.
2234 (US), 2244 (US). coci.E: 8 km above F,1 Cope,
Hammel 789 (MO).
Elaphoglossum nwrnnii is similar to E. lingua
in the creeping rhizome and small black sclerotic
rhizome scales, but the lamina is more slender (1.2-
2.7(-3.4) cm long vs. 2.7-6.5 cm) and the petiole
is more slender (ca. 1 mm diam. vs. (1-)1.5-3 mm).
It is also similar to E. longicrure in lamina shape,
but E. moranii has a creeping rhizome with small
black sclerotic rhizome scales.
The species is named in honor of Bobbin C. Mor¬
an, fern scholar and coodinator of the Mesoamerican
pteridoflora.
Klnphoglossum ornatiforme Mickel, sp. nov.
TYPE: Panama. Veraguas: above Rio Primero
Rrazo, 5 mi. NW of Santa Fe, Liesner 900
(holotype, MO).
Elaphoglosso ornato affinis, sed cum ea squamis cos-
talibus multo minoribus grossius dentatis, laminae stipi-
tisque pilis erectis glandulosis, necnon lamina apice obtusa
hasi cuneata incongrua.
Epiphytic; rhizome ca. 5 mm diam., compact;
rhizome scales 5-9 mm long, linear-lanceolate to
lanceolate, red-orange, dull, with weak hair-teeth;
sterile leaves 15-18 cm long, approximate; phyl-
lopodia present; petiole x A~ x k the sterile leaf length,
orange-brown; petiole scales 2 3 mm long, dense,
lanceolate, orange-brown, dull, coarsely toothed, of
uneven lengths and with weak hair tips, spreading,
the petiole also bearing erect, glandular hairs; sterile
lamina 14-15 cm long, 2.8 3.6 cm wide, narrowly
oblanceolate, coriaceous, base cuneate, apex obtuse;
veins obscure, 1-13 mm apart, at 70° angle to the
costa; hydathodes lacking; lamina scales ca. 2 mm
long, dense on and limited to costa and margin,
lanceolate, orange-tan, dull, coarsely toothed,
spreading, lamina surface bearing erect glandular
hairs; fertile leaves longer than the sterile; petiole
ca. V 2 the fertile leaf length; petiole nearly as long
as the sterile leaves; lamina linear-oblanceolate, base
narrowly cuneate, apex obtuse; scales scattered on
costa; intersporangial scales lacking.
Distribution. Wet forests; 700-1,300 m.
Paratypes. Panama, darif.n from (’ana up the Cerro
Pirre Massif, Moran 5050 (US); ca. 6 km S from gold
mining camp at Cana up the fila leading W to Alturas de
Nique, bellinger 1905 (US).
I bis species is closely allied to E. ornatum but
differs in the following: the costal scales are much
smaller (ca. 2 mm long, rather than 5-6 mm) and
with coarser teeth; lamina and petiole have erect
glandular hairs (lacking in E. ornatum)'- the lamina
apex is obtuse, not acute, the base is cuneate, not
truncate.
Klaphoglossum pallidiforme Mickel, sp. nov.
TYPE: Panama. Panama: 2.4 mi. beyond Cer¬
ro Jefe on road to Altos de Pacora, Croat 22050
(holotype, NY; isotypes, MO, UC).
Ab E. pallido lamina basi rotundata apice caudata
necnon rhizomatis squamis brunneis diversa.
Epiphytic; rhizome 5-8 mm diam., compact; rhi¬
zome scales 7-10 mm long, 1.2 mm wide, linear-
lanceolate, brown, lustrous, entire; leaves 28-50
cm long, approximate; phyllopodia present but short
(2-3 mm long), obscured by scales; petiole Vz-Vz of
sterile leaf length, orange-brown; petiole totally gla¬
brous; sterile lamina 15-27 cm long, 5.0-6.7 cm
wide, lanceolate, chartaceous, base rounded, apex
caudate; costa keeled below; lamina glabrous; veins
evident, 1.5-2 mm apart, at 70-80° angle to the
costa, the vein ends often extended laterally to form
a commissural vein; hydathodes lacking; fertile leaves
nearly as long as the sterile; petiole nearly l A the
fertile leaf length; fertile lamina linear-lanceolate;
intersporangial scales lacking.
Distribution. Wet forests; 600 1,000 m.
Paratypes. Costa RlCA. ALAJUELA: San Carlos, Jime¬
nez 979(F). PUNTARF.NAS: Boruca, Ocampo 1341 (CR).
Panama, chiriqui: Palo Alto, 4.5 mi. NE of Boquete,
Hammel 7503 (UC). COCLE: foot of Cerro Pilon, above
El Valle de Anton, Porter et al. 4070 (MO); Cerro Caital
Caracora, Dwyer 8922 (MO), panama: trail to top of
Cerro Pelado, Antonio 1094 (MO); P. Grande, Santa Fe,
Ocampo 800 (CR). VERAGUAS: shoulder of Cerro Tuti,
Folsom <V Edwards 3307 (MO).
Elaphoglossum pallidiformc resembles E. pal¬
lidum (Baker) C. Chr. in the rounded lamina base
and obscured phyllopodia, but it totally lacks lamina
Volume 2, Number 4
1992
Mickel
Elaphoglossum
381
scales and petiole glands and lias laterally extended
vein tips. It may he more closely related to E.
albomarginatum A. R. Smith in the lack of glands
and scales and the vein tip extensions, but differs
from it in the caudate rather than acuminate lamina
apex.
Elaphoglossum pardalinum Mickel, sp. nov.
TYPE: Panama. Chiriqui: above El Boquete,
Maxon 5718 (holotype, US).
Ab E. boragineo lamina angustiori, stipite breviori,
squamisque paucioribus, integris, iis laminae et stipitis
fuscis diversa.
Epiphytic; rhizome ca. 6 mm diam., horizontal,
compact; rhizome scales 3-5 mm long, linear-del-
tate, dark reddish brown, lustrous, entire, spreading;
leaves 21-35 cm long, approximate; phyllopodia
lacking; petiole of sterile leal length, light brown;
petiole scales 3-4 mm long, scattered, linear-deltate,
dark reddish brown or with paler margin, lustrous,
entire, ascending to spreading; sterile lamina 18-
31 cm long, 2.4-4.4 cm wide, narrowly elliptic,
chartaceous, base narrowly cuneate, apex acumi¬
nate; lamina scales 1-3 mm long, sparse, more on
costa, few on margin, linear, castaneous to tan,
lustrous to dull, entire, appressed; veins obscure, at
ca. 70° angle to costa; hydathodes present; fertile
leaf not seen.
Distribution. Wet forests; 1,400-1,700 m.
Known only from the type collection.
Elaphoglossum pardalinum resembles E. oblan-
ceolatum in the oblanceolate lamina with white
patches, but is distinct in the smaller, narrower
leaves, slightly longer petiole, and more random
white patches (not just between the veins).
The specific epithet comes from the Greek, par-
dos, leopard, alluding to the white spots on the
lamina.
Elaphoglossum standleyi Mickel, sp. nov. TYPE:
Costa Rica. San Jose: La Palma, Standley
33224 (holotype, LJS).
Elaphoglossum omissum simulans, sed stipitis longioris
squamis concoloribus laininaeque fertili conduplicata div¬
ersa.
Epiphytic; rhizome 3-4 mm diam., compact; rhi¬
zome scales 5-7 mm long, linear-lanceolate, orange-
tan, lustrous, entire, spreading; leaves 15-35 cm
long, approximate; phyllopodia lacking; petiole % to
nearly half the sterile leaf length, stramineous to
tan; petiole scales 2-3 mm long, dense, linear-del¬
tate, orange-tan, dull, denticulate, appressed to as¬
cending; sterile lamina 8-18 cm long, 1.2-2.6 cm
wide, lanceolate, chartaceous, base broadly cuneate
to rounded, apex acuminate; lamina scales 1 -2 mm
long, scattered, linear-deltate, tan-orange, lustrous,
denticulate, appressed, spreading at margin; veins
mostly obscure, 1.5-2 mm apart, at 65° angle to
costa; hydathodes present; fertile leaf slightly shorter
than the sterile; petiole % the fertile leaf length;
fertile lamina elliptic, base broadly cuneate, apex
obtuse to acute, folded lengthwise; intersporangial
scales lacking.
Distribution. Wet forests; 1,500-2,400 m.
Paratypes. Costa Rica, heredia: Cerro de las Caricias,
N of San Isidro, Standley & l alerio 52191 (US), 52198
(US). PUNTARENAS-ALAJUELA BORDER: Monteverde Re¬
serve, Cordillera de Tilaran, Grayum & Sleeper 3840
(CR, MO), san JOSE: N fork of Rio Zurqui, Cordillera
Central, Smith 1652 (UC).
This species resembles E. omissum Mickel but
has concolorous petiole scales, longer petiole, and
longitudinally folded fertile lamina.
The species is named in honor of Paul C. Standley,
whose extensive collecting has contributed greatly
to our knowledge of Central American pterido-
phytes.
Elaphoglossum teleglottis Mickel, sp. nov.
TYPE: Panama. Chiriqui: Distrito Bugaba, Cer¬
ro Punta, t'an der Werff & Herrera 6 503
(holotype, NY; isotypes, MO, UC).
Ab E. lloensi laminae squamis lineari-hastatis diversa.
Epiphytic; rhizome 1-1.3 mm diam., long-creep¬
ing; rhizome scales 3-4 mm long, ovate-lanceolate,
tan, dull or occasionally with lustrous center, entire,
ascending; leaves 13-19 cm long, 3-6 cm apart;
phyllopodia present; petiole Vs-V'i the sterile leal
length, tan; petiole scales 3-4 mm, scattered,
spreading, lanceolate to ovate-lanceolate, tan, dull,
entire or with a few processes at base; sterile lamina
8 12 cm long, 2.2-3 cm wide, lanceolate, char¬
taceous, base narrowly cuneate, apex acuminate;
lamina scales orange-brown, abaxially linear-has¬
tate, 4 mm long, with basal processes, to hastate,
1-2 mm long, to linear, 1 mm long, or reduced to
punctae; lamina adaxially glabrous; veins evident,
1-1.5 mm apart, at 65-70° angle to costa; byda-
thodes lacking; fertile leaves longer than the sterile;
petiole 2 A of fertile leaf length; fertile lamina nar¬
rowly lanceolate; intersporangial scales lacking.
Distribution. Cloud forests; 2,200 m.
Known only from the type collection.
This species resembles E. proximum Christ but
has thinner texture, veins more evident, lamina base
more narrowly cuneate, rhizome scales entire (rather
than fimbriate) and narrower; lamina scales are has¬
tate rather than fimbriate. Elaphoglossum mitor-
382
Novon
rhizurn differs in having black, appressed rhizome
scales 0.5-1 mm long (or larger, tan scales with
sclerotic black center) rather than concolorous tan
ascending scales 3-4 mm long.
The specific epithet comes from the Greek, tele ,
far, and glottis , tongue, referring to the distant
leaves on the rhizome.
Elaphoglossum valdespinoi Miekel, sp. nov.
TYPE: Panama. Panama: Cerro Jefe, I aides-
pino HO I (holotype, NY).
Stipite abbreviato, laminae squamis angustis nigris, nec-
non lamina fertili lineari notabilis.
Epiphytic; rhizome 4-6 mm diam., horizontal,
compact; rhizome scales 5-7 mm long, linear, red¬
dish brown, lustrous, entire, spreading; leaves 12
22 cm long, approximate; phyllopodia lacking; pet¬
iole the sterile leaf length, brown; petiole scales
2-3 mm long, sparse, hairlike-subulate, blackish
brown, lustrous, denticulate, spreading; sterile lam¬
ina 10-20 cm long, 1.4-2.3 cm wide, narrowly
elliptic, chartaceous, base cuneate, apex acute to
acuminate; veins obscure, ca. 1.5 mm apart, at 65°
angle to costa; hydathodes present; lamina scales
scattered, as on petiole, lustrous, 3 6 mm long on
costa, 2-4 mm long elsewhere, minutely denticulate,
spreading; fertile leaves equal to or slightly longer
than the sterile; petiole x /i the fertile leal length;
fertile lamina 5 mm wide, linear, base attenuate,
apex obtuse; intersporangial scales 4 6 mm long,
dark reddish brown to black.
Distribution. Cloud forests; 100 900 m.
Paratypes. Panama, chiriqui: Distrito Boquete, For-
tuna Darn site, van tier If erff & van Hardeveld 6 808
(UC). PANAMA: Cerro Jefe, Sytsma et al. 142b (CK. MO),
245b (MO), Porter, et al. 5081 (MO), Tyson 3404 (MO).
This species is distinguished by its short petiole,
long, narrow, black laminar scales, and linear fertile
lamina.
The species is named in honor of Ivan A. Val-
despino, outstanding collector of Panamanian pte-
ridophytes and student of American Selaginella.
Acknowledgments. I am grateful to Rupert Bar-
neby for assistance with the Latin diagnoses and to
Ruth Russell for her careful reading of the manu¬
script and many valuable suggestions.
Literature Cited
Christ, H. 1899. Monographic des Genus Elaphoglos¬
sum. Denkschr. Schweiz. Naturf. Ges. 36: 1-159.
Fee, A. L. A. 1845. Memoires sur les families des
fougeres. 2: 1-114, pi. 1-64.
Miekel, J. T. 1980. Nine new species of Elaphoglossum
(Elaphoglossaceae) from Mexico. Brittonia 32: 334
339.
-. 1981. Elaphoglossum. Pp. 104-116 in D.
E. Breedlove (editor), Flora of Chiapas. 2. Pterido-
phytes, by A. R. Smith. Calif. Acad. Sci.
-. 1985. Elaphoglossum. Pp. 78 107 in A. R.
Smith, Pteridophytes of Venezuela, an annotated list.
Dept, of Botany, Univ. of California, Berkeley.
-. 1988. Elaphoglossum. In: J. T. Miekel & J.
M. Beitel, Pteridophyte Flora of Oaxaca, Mexico.
Mem. New York Bot. Card. 46: 168 186.
-. 1990a. Three new species of Elaphoglossum
from Peru. Amer. Fern J. 80: 110-112.
-. 1990b. Two new species of Elaphoglossum
(Elaphoglossaceae) from Amazonas, Venezuela. Arm.
Missouri Bot. Card. 77: 594 595.
-. 1991a. Elaphoglossum. In: R. M. Tryon &
R. G. Stolze, Pteridophyta of Peru, part IV. Fieldiana,
Bot. n.s., no. 27: 110-166.
-. 1991b. Four new species of Elaphoglossum
(Elaphoglossaceae) from Venezuela. Ann. Missouri
Bot. Gard. 78: 259-261.
-. 1992. Pteridophytes. In: R. McVaugh & W.
R. Anderson (editors), Flora Novo-Galiciana. Univ.
of Michigan, Ann Arbor, Michigan 17: 120 431.
Smith, A. R. 1975. New species and new combinations
of ferns for Chiapas, Mexico. Proc. Calif. Acad. Sci.
4th ser. 40: 209-230.
Two New Combinations in Peruvian Jaltomata (Solanaceae)
Thomas Mi one
Biological Sciences, Copernicus Hall, Central Connecticut State University, New Britain,
Connecticut 06050-4010, II.S.A.
helix G. Coe
Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut
06269-3043, IJ.S.A.
ABSTRACT. The new combination Jaltomata we-
berbaueri (Dammer) Mione is based on Saracha
weberbaueri Dammer; this species resides in Peru,
Dept. Ancash, between 3,300 and 3,700 m ele¬
vation. The new combination Jaltomata aspera (Ruiz
& Pavon) Mione is based on Atropa aspera Ruiz
& Pavon; Hebecladus asperus (Ruiz & Pavon)
Miers is a synonym. This species resides in Peru,
Dept. Lima, from near sea-level to 1,800 m, and
is rare, likely due to habitat destruction by humans.
Jaltomata (tribe Solaneae) is a genus of about
28 species ol tropical and subtropical perennial herbs
and small shrubs that are distributed from Arizona,
U.S.A., to southern Bolivia, and with one species
each in the Greater Antilles and on the Galapagos
Islands.
During study of the genus Jaltomata we have
found the following new combinations to be neces¬
sary. The species treated here are similar but are
easily distinguished. Jaltomata weberbaueri resides
above 3,000 m and is glabrate, while ./. aspera
resides below 1.800 m and is pubescent and ciliate.
Jaltomata weberbaueri (Dammer) Mione, comb,
nov. Basionym: Saracha iveberbaueri Dam¬
mer. Bot. Jahrb. Syst. 37: 638. 1906. TV PE:
Peru. Ancash: Prov. Cajatambo, Ocros, A. II e-
berbauer 2657 (holotype, B destroyed, photo
of B sheet F neg. 2558, W IS).
Saracha weberbaueri Dammer subsp. pallascana Bitter,
Repert. Spec. Nov. Regni Veg. 19: 267. 1924.
Saracha pallascana (Bitter) Macbride, Publ. Field
Columbian Mus., Bot. Ser. 8: 110. 1930. TYPE:
Peru. Ancash: Prov. Pallasca, valley of the river
Pelagatos, near Yungabal, A. Weberbauer 7225
(holotype, B destroyed, photo of B sheet F neg.
2559, WIS; isotype, F, photo of F sheet F neg.
49345, WIS).
See Macbride (1962, as Saracha weberbaueri)
for an English description of this taxon.
Additional specimens examined. PERU. ANCASH: Prov.
Recuay, km 107 between Recuay and Pativilca, (fl), M.
Dillon et al. 3174 (BH, GH, MO); Prov. Aija, cuchu
cuchu, entre Huamanwilca y Sequipoquian, C. Ochoa &
A. Salas 15160 (NY).
Habitat. Grasslands with scattered bushes, A.
Weberbauer 2657, 7225; moist ravines, M. Dillon
et al. 3174.
Altitude. 3,300-3,700 m.
Phenology. Flowering and fruiting January
through March.
Local name. “Tomentito,” M. Dillon et al. 3174.
Jaltomata aspera (Ruiz & Pavon) Mione, comb,
nov. Basionym: Atropa aspera Ruiz & Pavon,
FI. Peruv., et Chilensis 2: 45. 1799. I Iticona
aspera (Ruiz & Pavon) Raf., Sylva Telluriana
55. 1838. Hebecladus asperus (Ruiz & Pavon)
Miers, London J. Bot. 4: 322. 1845. TYPE:
Peru. In collibus Limae moninatis Amancaes,
Ruiz Lopez & Pavon s.n. (holotype, presum¬
ably MA not seen).
Saracha ciliata Miers, Ann. Mag. Nat. Hist., ser. 2, 3:
446. 1849, syn. nov. TYPE: Peru. Valley of Lima,
Mathews 834 (holotype, K).
Saracha lacrima-virginis Bitter, Repert. Spec. Nov. Reg¬
ni Veg. 17: 341. 1921, syn. nov. TYPE: Peru.
Amancaes, A. Weberbauer 1587 (holotype, B de¬
stroyed, photo of B specimen F neg. 2551, WIS).
Saracha urbaniana Bitter & Dammer, Repert. Spec.
Nov. Regni Veg. 17: 342. 1921, syn. nov. TYPE:
Peru. Above San Bartolome, Lima- Oroya Railroad,
A. Weberbauer 5262 (5262 = 5305 according to
label on photo at WIS) (holotype, B destroyed, photo
of B specimen F neg. 2557, GH, W IS; isotype, GH).
See Macbride (1962, as Hebecladus asperus,
Saracha ciliata, and S. urbaniana) for English
description of this taxon.
Additional specimens examined. PERU. LIMA: Prov.
Huarochiri, valley of Rio Rimac, near Lima Oroya high¬
way at km 70 E of Lima, T. H. Goods peed & R. D.
Novon 2: 383-384. 1992.
384
Novon
Metcalf 30248 (MO, US); near Lima, IF. Nation s.n.
(K , on same sheet as holotype of ,S. ciliata); hills of
Mongomarca, .4. Weberbauer 5235 (US). LIMA: Aman-
caes, A. Weberbauer 524ba (F, US), A. FT eberbauer
5305 (F); San Geronimo, J. F. Macbride 5912 (F), I).
Stafford 1932 (K).
Habitat. Loma-formation, A. H eberbauer 1587,
5235, 5246a; rocky hillside, .Macbride 5012; dry
open hillside, granite soil, with cacti, Goodspeed &
Metcalf 30248; sandy hills of Lima, shady places,
between stones, rare, II . Nation s.n.
All specimens examined were collected prior to
1943; perhaps endangered, due to habitat destruc¬
tion by humans.
Altitude. Near sea-level to 1.800 in.
Phenology. Flowering and fruiting July through
October, and March.
Loral name. “Lacrima de la virgen,” A. II e-
berbauer 1587.
Acknowledgments. We thank Gregory J. An¬
derson, Tilton Davis IV, William D’Arcy, Michael
Nee, and the herbarium staffs at B1L BM, C, COLO,
F, GH, K, LD. MKRF, MO, NY, US, and WIS for
loan of specimens and/or accommodating visits,
Thomas Stark (or assistance with German, and Ar¬
mando T. Hunziker and Luis Bernardello, who sug¬
gested this group for study. Support was from fel¬
lowships from the Graduate School and the Research
Foundation of the University of Connecticut to T.
Mione, and an N.S.F. grant to T. Mione and Gregory
J. Anderson.
Literature Cited
Macbride, J. F. 1962. Solanaceae. Field Mus. Nat.
Hist., But. Ser. 13, part V-B, No. 1. [.Jaltomata as
Saracha and licheeIndus.]
Two New Species of Simsia (Asteraceae: Heliantheae) from
Southern Mexico
Jose L. Panero
Department of Botany, University of Texas, Austin, Texas 78713-7640, IJ.S.A.
Edward E. Schilling
Department of Botany, University of Tennessee, Knoxville, Tennessee 37996-1100, U.S.A.
ABSTRACT. Two new species from Mexico are de¬
scribed, Simsia sylvicola from Oaxaca and S. spoo-
neri from Guerrero. The two species are placed in
Simsia because they exhibit most of the features
that characterize the genus, although they differ in
possessing a biconvex achene shape that is more
typical of the related genus I iguiera. Chromosome
counts of n = 17 are reported for both species.
In the course of our revisionary studies of l i-
guiera Kunth ser. Grammatoglossae S. F. Blake,
we have discovered two previously undescribed spe¬
cies. There is some question whether these should
be placed in l iguiera or in the related Simsia Per-
soon, because the distinction between these genera
is still problematic. In practice, Spooner (1990) con¬
sidered Simsia to be delimited by features of the
disk achenes, which are laterally very flattened and
usually have a pappus of two awns but typically lack
squamellae. In contrast, the typical disk achene in
Viguiera is biconvex and has a pappus that includes
both two awns and also two or more squamellae.
There is, however, a larger suite of traits that, to¬
gether with the achene features, characterize Simsia
(Spooner, 1990), including nodal disks, herbaceous
involucres, long, narrow ray ovaries, long, tapering
style branches, and setose anther connectives. Many
of these traits are also exhibited in combination by
some of the members of Viguiera ser. Gramma¬
toglossae. The two new species are particularly
striking in differing from typical members of Simsia
only in features of the achene, and thus further
bridge the morphological distinction between the two
groups. Further support indicating that Simsia and
some members of Viguiera ser. Grammatoglossae
are related phylogenetically has been revealed from
restriction site analysis of chloroplast DNA (Schilling
& Jansen, 1989; Schilling & Panero, unpublished
data). Resolution of the exact boundaries between
Simsia and Viguiera ser. Grammatoglossae will
await further information from DNA-based studies.
We argue at this point that the concept of Simsia
should be broadened relative to that of Spooner
(1990) to include some species that have biconvex
achenes (there is, in fact, variation within one species
of Simsia , S. ghiesbreghtii (A. Gray) S. F. Blake,
for this trait). For this reason, we suggest placement
in Simsia of the two species newly described here,
.S. sylvicola and S. spooneri.
Simsia sylvicola Panero & E. Schilling, sp. nov.
'EY PE: Mexico. Oaxaca: Km 65 of the road
Oaxaca-Sola de Vega, occasional in pine-oak
forest, 1,950 m, 10 Nov. 1990, Panero 21 Ob
(holotype, MEXU; isotypes, ENCB, SI. TENN,
TEX, US). Figure 1.
A Viguiera rhombifolia (Robinson & Greenrnan) S.
F. Blake foliis perfoliatis et late deltatis et petiolis pler-
umque non alatis differt.
Perennial herbs 4 10 dm tall, sometimes pros¬
trate. Stems terete, purplish brown, sparsely to mod¬
erately puberulent, hispid distally toward the capi-
tulescence. Leaves opposite, triplinerved; petioles
10-23 mm long, terete to slightly canaliculate,
sparsely to moderately puberulent to hispid; blades
4-7 cm long, 12-55 mm wide, gradually decreasing
in size distally toward the capitulescence, deflate to
ovate, base rounded to truncate, apex acuminate,
margins serrate to dentate, adaxial surface shiny
dark green drying dull brownish green, sparsely to
moderately strigose to hirsute, abaxial surface slight¬
ly paler in color than adaxial surface, moderately
hirsute, pubescence denser on veins outlining them
against a green background. Capitulescences dicha-
sial, thyrsoid, open or sometimes heads congested
at distal end of peduncles with 3-25+ heads; pe¬
duncles 2-30 cm long, pubescence like that of the
stem. Heads 6.5-7.5 mm diam. (ligules excluded),
12-13 mm high, campanulate to cylindrical; re¬
ceptacles 4.5-5.5 mm diam. in fruit, flat to slightly
convex. Phyllaries 19 23 in 3 graduated series;
phyllaries of first series 3-5.5 mm long, 1.2-1.5
mm wide, ovate to lanceolate, herbaceous, dark
Novon 2: 385-388. 1992.
386
Novon
Figure 1. Simsia sylvicola Panero & E. Schilling (Panero 210 6). —a. Habit. —b. Head. —c. Ligule. —d. Disk
Hower. —e. Athene. Scale bars = 1 cm (a); I min (h e).
green drying blackish green, sparsely puberulent,
hispid distally toward the apex, margins eiliate; phyl-
laries of second and third series 9 11 mm long,
1.7 2.3 mm wide, lanceolate to narrowly lanceolate,
herbaceous, dark green drying blackish green, pu¬
bescence as first series. Pales 7-7.8 mm long, 2.8
3.2 mm wide, lanceolate to ovate in outline, con-
duplicate, shallowly keeled, not exceeding phyllaries,
Volume 2, Number 4
1992
Panero & Schilling
Simsia sylvicola and Simsia spooneri
387
chartaceous, stramineous turning hyaline whitish with
age, essentially glabrous ahaxially, apex acuminate
to slightly cuspidate, erect, dark green drying black.
Ray flowers 7-ll(-13); corollas lemon yellow or
pale yellow; tube 1.5-1.7 mm long, essentially gla¬
brous; ligule 6.5-9 mm long, 3.5-4.5 mm wide,
oval-oblong to suborbicular, sparsely puberulent, apex
rounded and minutely bifid; ovaries 5 5.5 mm long,
linear, triquetrous, glabrous, pappus of 2-3 minute
scales. Oisk flowers 25-35, flowers protruding 2-
3 mm beyond pales; corollas lemon yellow or green¬
ish yellow turning purplish with age, cylindrical; tube
0.6 0.8 mm long, glabrous or minutely puberulent;
throat 4 mm long, 1 mm wide, veins pale orange
and discernible the entire length of the throat, gla¬
brous or sparsely puberulent; lobes 0.5 mm long,
sparsely puberulent; anthers 2.5 mm long, thecae
black; styles 6.5 mm long; style branches 1.8-2.2
mm long. Achenes 3.5 3.7 mm long, 1.5 mm wide,
biconvex, oval-obovate, sometimes edged with a min¬
ute flat rim, shiny black or mottled, sparsely seri¬
ceous, pappus of 2 awns and 0-2 squamellae, awns
slender, stramineous, subequal, 1.5-2 mm long,
squamellae 0.5 mm long. Chromosome number: n
= 17.
Paratypes. Mexico, oaxaca: 15.1 km N of Sola de
Vega on road to Oaxaca, 1,800 in, 21 Oct. 1985, Bar-
tholomew et al. 3251 (CAS, TENN); ca. 10-15 km N
Sola de Vega, road to Oaxaca, 1,880 in. 17 Sep. 1988
(in bud), Schilling & Panero 88-35 (MEXU, TENN;
voucher for chromosome count of n = 17).
Simsia sylvicola shares a similar habit and floral
morphology with Viguiera rhomhifolia , but the two
species can he separated by vegetative character¬
istics. Simsia sylvicola has petiolate leaves with
perfoliate bases, whereas V. rhomhifolia has leaves
that appear to be sessile because of the completely
winged petioles but are not perfoliate.
The species epithet refers to the characteristic
occurrence of the species in open oak-pine woods.
Simsia spooneri Panero & E. Schilling, sp. nov.
TYPE: Mexico. Guerrero: 14 km E of Ayot-
zinapa, ca. km 30 on the road Chilpancingo-
Chilapa, 1,900 m, 7 Nov. 1990, Panero 2032
(holotype, MEXU; isotypes, ENCB, SI, TENN,
TEX, US).
A Viguiera rhomhifolia (Robinson & Greenman) S.
F. Blake foliis magnis et late deltatis cum petiolis non
alatis, et Simsia sylvicola Panero & E. Schilling foliis
non perfoliatis differt.
Perennial herbs 4-15 dm tall. Stems terete, pur¬
plish brown, sparsely to moderately puberulent to
strigose and scabrous, hispid distally toward the eap-
itulescence. Leaves opposite, triplinerved; petioles
7-15 mm long, terete to slightly canaliculate, sparsely
puberulent; blades 4-9 cm long, 17-65 mm wide,
gradually decreasing in size distally toward the cap-
itulescence, debate to ovate, base truncate to round¬
ed, apex acuminate to rounded, margins serrate,
adaxial surface shiny green drying dull green, sparsely
to moderately strigose to hirsute, abaxial surface
slightly paler in color than adaxial surface, sparsely
hirsute, veins raised and clearly outlined against
background, essentially glabrous. Capitulescences
dichasial, thyrsoid, open or sometimes heads con¬
gested at distal end of peduncles with 3-25+ heads;
peduncles 0.5-35 cm long, pubescence like that of
the stem. Heads 9- I 1 mm diam. (ligules excluded),
12-13 mm high, campanulate to cylindrical; re¬
ceptacles 0.6 0.7 mm diam. in fruit, flat to slightly
convex. Phyllaries 16-20 in 2-3 graduated series;
phyllaries of first series 2.5-5.5 mm long, 1.5-2.2
mm wide, ovate, herbaceous, dark green drying
blackish green, moderately puberulent, margins cil-
iate; phyllaries of second and third series 8-13 mm
long, 2.2-2.7 mm wide, lanceolate to narrowly ob¬
long. herbaceous, dark green drying blackish green,
sparsely puberulent especially along mid costae. Pales
9 mm long, 2.5 mm wide, lanceolate to ovate in
outline, conduplicate, shallowly keeled, not exceed¬
ing phyllaries, chartaceous, stramineous turning hy¬
aline whitish with age, essentially glabrous ahaxially,
apex acuminate to slightly cuspidate, erect, dark
green drying black. Ray flowers 9 13(— 15); corollas
lemon yellow or pale yellow; tube 1.5 mm long,
essentially glabrous; ligule 11-13 mm long, 5-5.5
mm wide, oval oblong to ovate, essentially glabrous,
apex acuminate to rounded and minutely bifid; ova¬
ries 5.5 mm long, linear, triquetrous, glabrous, pap¬
pus of 2-3 minute scales. Disk flowers 15—30, flow¬
ers protruding 2-3 mm beyond pales; corollas lemon
yellow or greenish yellow turning deep purple with
age, cylindrical; tube 0.6-0.8 mm long, glabrous or
minutely puberulent; throat 5.8-6.2 mm long, 1.3-
1.6 mm wide, veins pale orange and discernible the
entire length of the throat, glabrous or sparsely
puberulent; lobes 1 mm long, sparsely puberulent;
anthers 3 mm long, thecae black; styles 7-7.5 mm
long; style branches 1.8-2.2 mm long. Achenes
3.4-4.2 mm long, 1.4-2 mm wide, biconvex, oblong
to obovate, shiny black or mottled, moderately se¬
riceous, pappus ol 2 awns and 5-7 squamellae, awns
slender, stramineous, subequal, 2.7-3.2 mm long,
squamellae 0.5 0.7 mm long. Chromosome number:
n = 17.
Paratypes. Mexico. GUERRERO: 9 mi. E of Ayotzinapa,
13 Oct. 1984, Spooner & Burgos 2610 (OS, TENN);
388
Novon
road to Chichihualco ca. 20 km W from jet. with Hwy.
95, 1,600 m, 19 Sep. 1988 (in bud), Schilling & Panero
88-43 (TENN; voucher for chromosome count of n =
17); Km 20 on the road Chilpancingo-Chichihualco, 1,900
m, 16 Nov. 1990, Panero 2027 (COL, K, MA, MEXU,
TENN, UC, US).
The species epithet honors David M. Spooner,
potato specialist at the University ol Wisconsin and
monographer of Simsia, as well as first collector of
this taxon.
Simsia spooneri closely resembles S. sylvicola,
but plants of the two species have a clearly different
aspect when grown together in the University of
Tennessee greenhouse. Simsia spooneri also re¬
sembles S. villasenorii Spooner in its habit and
general morphology. Both species tend to grow on
vertical, eroded hillsides where they produce a mul¬
titude of flowering shoots of various lengths arising
from a central mass of tuberous roots. The two
species can be separated by their involucre and disk
achene morphologies: S. spooneri has a campan-
ulate, green involucre and achenes without flat rims,
whereas S. villasenorii has an imbricate, purplish
urceolate involucre and achenes with a flat rim.
Simsia spooneri can be confused with Viguiera
rhombifolia, because they have similar head and
achene morphologies, but they differ in leaf and
habit characteristics.
Acknowledgments. This study was supported by
NSF grant BSR-8806513 (to E. Schilling). We thank
D. Spooner and J. Strother for helpful comments.
Literature Cited
Schilling, E. E. & R. K. Jansen. 1989. Restriction
fragment analysis of chloroplast DNA and the sys-
tematics of Viguiera and related genera. Ainer. J.
Bot. 76: 1769-1788.
Spooner, D. M. 1990. Systematics of Simsia (Com-
positae Heliantheae). Syst. Bot. Monogr. 30: 1-90.
Ardisia callejasii (Myrsinaceae): A New Species from
the Antioquian Choco of Colombia
John J. Pi poly III
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Abstract. Continuing studies in Myrsinaceae for
Flora de Colombia revealed a heretofore unde¬
scribed species of Ardisia closely related to Ardisia
pittieri Mez of Costa Rica and Panama.
While determining specimens in Colombian her¬
baria for my treatment of the Myrsinaceae for Flora
de Colombia , I discovered a fine series of specimens
representing an unknown taxon, described herewith.
Ardisia callejasii Pipoly, sp. nov. TYPE: Colom¬
bia. Antioquia: Mpio. Mutata, N of Hacienda
El Darien, right margin of Rio Chontadural, 28
July 1978 (fl, fr), R. Fonnegra & F. Renteria
953 (holotype, HUA; isotypes, COL, MEDEL,
MO). Figure 1.
Quoad ramulos furfuraceo-lepidotos, folia magna cor-
iacea oblanceolata brevipetiolata, panicularum ramulos
corymbosos, stamina in tubum connata, ad A. pittieri
valde arete affinis, sed ab ea laminis ad apicem longi-
attenuatis (non acutis), sepalis chartaceis (non coriaceis)
2-2.8 (nec 3-3.5) mm longis, symmetricis (nec asym-
metricis) ad apices obtusis (nec emarginatis), petalis ovatis
(nec ellipticis), 5.3-5.9 (nec 10-11) mm longis, manifeste
(nec obscure) punctatis, staminibus 4.9-5.4 (non 8.7-
9.1) mm longis, tubo staminali 1.2-1.3 (nec 1.8-2) mm
longo, antheris ad basem inter se liberis (nec conniven-
tibus), denique fructibus luteis (non ruberis) statim separ-
abilis.
Tree to 7 m; branchlets terete, 0.7 1 cm diam.,
densely and minutely furfuraceous lepidote. Leaves
coriaceous, oblanceolate, (25-)31-45.5 cm long,
(5-)7-12.2 cm wide, apex long-attenuate, gradually
tapering to a cuneate base, costa slightly raised
above, prominently raised below, secondary veins
16-20 pairs, slightly depressed or not visible above,
prominently raised below, glabrous and dull above,
sordid and sparsely and minutely sessile rufous fur-
furaceous-lepidote below, glabrescent, the margin
regular, entire, flat; petioles marginate (0.6—) 1 1.2
cm long, glabrous above, densely and minutely ru¬
fous furfuraceous-lepidote below. Inflorescence sub¬
terminal, pinnately or bipinnately paniculate, (7.5 )
12.5-24.5 cm long, pyramidal, primary and sec¬
ondary bracts unknown, secondary branches 3 12
cm long, the rachis minutely densely furfuraceous-
lepidote, glabrescent; floral bracts unknown (pre¬
sumably early caducous); pedicels cylindrical, 0.7-
1.8 cm long, densely furfuraceous-lepidote, persis¬
tent. 1 lowers 5-merous, corymbose, the corymb with
7-12 flowers, pink; sepals quincuncial, almost free,
chartaceous, ovate, 2-2.8 mm long, 1-1.3 mm
wide, symmetrical, apex obtuse, thickened and
prominently black punctate medially, densely and
minutely furfuraceous-lepidote, the margin irregu¬
lar, scarious, hyaline, sparsely glandular-ciliolate,
entire; petals quincuncial, basally connivent, cori¬
aceous, ovate, 5.3-5.9 mm long, apex acute, highly
reflexed in anthesis, medially thickened, prominently
pellucid punctate without, somewhat verruculose
medially within, the margin scarious, hyaline, entire,
glabrous; stamens 4.9-5.4 mm long, the filaments
connate into a coriaceous elobate tube 1.2-1.3 mm
long, epunctate, glabrous, the anthers free, lanceo¬
late, 3.5-4.2 mm long, 1.3-1.6 mm wide, sessile
on the staminal tube, apex attenuate, base cordate,
longitudinally dehiscent by narrow, sublatrorse slits,
the connective inconspicuously pellucid punctate;
pistil obclaviform, 5.3-5.6 mm long, 1-1.2 mm
diam., the ovary 1.5-1.7 mm long, glabrous, the
style 3.8-4 mm long, erect, inconspicuously pellucid
punctate, glabrous, the placenta ellipsoid, 0.8 1 mm
long, 0.3-0.4 mm diam., apex apiculate, the ovules
12 15, pluriseriate, stigma punctiform. Fruit glo¬
bose, ca. 1 cm diam., green, then yellow at maturity,
densely and prominently black punctate, the style
base persistent.
Distribution. Ardisia callejasii is endemic to the
Municipality of Mutata, located in the northwestern
corner of the Department of Antioquia, Colombia,
at 100 180 m elevation.
Ecology and Conservation Status. This new spe¬
cies occurs in lowland pluvial forests, in a narrow
valley belonging to the Rio Sucio drainage basin,
located in the Department of Antioquia between the
southwestern corner of the Department of Cordoba
and the northwestern corner of the Department of
Choco. Label data from the Fonnegra et al. collec¬
tions indicate it is a common species in forests pre¬
served on Hacienda El Darien. However, because
the region is traversed by the Medellin-Turbo high-
Novon 2: 389-391. 1992.
390
Novon
Figure 1. Ardisia callejasii Pipoly. —A. Habit, showing pyramidal panicle with corymbose branches. — B. Abaxial
leaf surface, showing minutely furfuraceous-lepidote tomentum. —C. Longisection of pistil and calyx, showing ovate
sepals, elongate style, and punctiform stigma. —D. Longisection of pistil, showing ellipsoid, apiculate placenta with
pluriseriate ovules. —E. Opened corolla, showing staminal tube, sessile, lanceolate anthers and narrow, sublatrorse
longitudinal slits. —F. Fruit, showing prominent punctations and persistent style base.
way, A. callejasii should he considered a threatened
species.
Etymology. It gives me great pleasure to dedicate
this new species to Ricardo Callejas Posada, friend,
curator of the herbarium of the University of An-
tioquia (HUA), and pre-eminent authority on the
morphogenesis, anatomy, and systematics of the Pi-
peraceae.
Paratypes. Colombia, antioquia: Mpio. Mutata, mar¬
gin of Rio Leon (Bucaba), Villa Arteaga, Las Caucheras,
Volume 2, Number 4
1992
Pipoly
Ardisia callejasii
391
hill above water tap, 100-180 m, 2 Oct. 1961 (bud),
Cuatrecasas 26156 (US); Hacienda El Darien, right bank
of Rio Chontadural, 20 June 1979 (fr), Fonnegra el at.
1228 , (fl), 1249 (COL, HUA, MO), 13 Sep. 1979 (fr),
Fonnegra 1343 (COL, HUA, MO).
Ardisia callejasii is most closely related to the
vicariant Ardisia pittieri Mez of Costa Rica and
Panama, owing to the furfuraceous-lepidote bran-
chlets, large, coriaceous, ohlanceolate leaves, pan¬
icles with corymbose branchlets, and stamens with
filaments connate to form a tube. However, the long-
attenuate leaf apices, much shorter, obtuse and
chartaceous sepals, much shorter, ovate, and prom¬
inently punctate petals, shorter stamens and sta-
minal tube, free anthers, and yellow fruit allow for
immediate recognition. Within neotropical Ardisia ,
formation of a stamina] tube is a synapomorphic
character shared by only these two taxa, and the
yellow fruit is known only in A. callejasii. The
vestiture, corolla, and anther morphology indicate
placement in the group of Ardisia species segregated
in the genus Auriculardisia Lundell. However, be¬
cause the majority of the diagnostic characters which
define that genus frequently occur in Malesian Ar¬
disia, I am not recognizing that entity as distinct.
Acknowledgments. My studies in the documen¬
tation of neotropical plant species diversity are sup¬
ported by a grant irom the John D. and Catherine
T. M acArthur Foundation. 1 thank the curators of
the herbaria cited for loans of material and for
hospitality during my visits. I also thank Juan Carlos
Pinzdn of Santafe de Bogota, Colombia, for his out¬
standing illustration. Paul Berry read the manuscript
and provided useful suggestions.
Notes on the Genus Myrsine (Myrsinaceae) in Peru
John J. Pipoly III
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
Abstract. Continuing studies in Peruvian Myr¬
sine have revealed eight species new to science.
These species are described and illustrated, and their
systematic relationships are discussed. Treatment of
Rapanea as a taxonomic synonym of Myrsine ne¬
cessitates six new combinations for Peruvian species
and another four combinations for related taxa from
Colombia, Ecuador, Bolivia, and Brazil.
In preparation for publication of the Myrsinaceae
for the Catalogue of the Flowering Plants and
Gymnosperms of Peru, it has become necessary to
publish a number of novelties and new combinations.
The genus Myrsine L., as here recognized, includes
all neotropical members of the genus Rapanea Au-
blet, formerly recognized as separate from it based
on tbe sessile anthers and lobed stigmas. A prelim¬
inary discussion of reasons for reducing Rapanea
to synonymy under Myrsine has been presented
elsewhere (Pipoly, 1992). Myrsine is a pantropical
genus of perhaps upwards of 150 species, of which
80, many as yet undescribed, are neotropical.
Myrsine microdonta Pipoly, sp. nov. TYPE: Peru.
La Libertad: Prov. Santiago de Chuco, Huacas,
Cachicadan, 2,800 m, 15 June 1984 (pist. II,
fr), A. Sagdstegui ,,/. Mostacero & M. Diestra
11903 (holotype, MO; isotypes, BISH, CAS,
F, HUT, US). Figure 1.
Quoad ramulos graciles ad apices dense ferrugineo-
tomentosos, folia alterna, plana, superne nitida, inferne
tristia, ad bases rotundata, necnon lobos calycinos cori-
aceos ovatos, M. brevis valde affinis sed ab ea ramulis
adpresse (non floccose) tomentosis lamina secus margines
apicem versus manifeste dentata (non integerrima), mu-
cronulata (nec emarginata), minute ciliolata (nec glabra),
lobis calycinis ad apices minutissime ciliolati (non glabris),
lobis corollinis ellipticis (non ovatis) secus margines in-
volutis (nec planis) praeclare distat.
Shrub, height unknown; branchlets prominently
ridged, 2-3 mm diam., adpressed ferrugineous to-
mentose apically, glabrescent. Leaves coriaceous,
elliptic, 1-2 cm long, 0.7-1.3 cm wide, apex broad¬
ly rounded, mucronulate, base rounded, decurrent
on the petiole, smooth and nitid above, dull and
prominently black perpuncticulose below, costa
deeply impressed above, prominently raised below,
the secondary veins not visible above or below, the
margin very minutely ciliate in bud, the cilia gla¬
brescent, prominently dentate from tbe middle to
tbe apex, subrevolute basally, glabrous; petioles
deeply canaliculate, 4 6 mm long, densely tomen-
tose above, glabrous below. Staminate inflores¬
cence: unknown. Pistillate inflorescence: a subses-
sile umbel, 3-5-flowered; floral bracts lanceolate,
1-1.2 mm long, 0.3-0.4 mm wide, apex acute, the
margin entire, minutely glandular-ciliate; pedicels
ridged, 1.4-1.6 mm long, glabrous. Flowers
5-merous, green; calyx coriaceous, cotyliform, 1.3-
1.4 mm long, the tube 0.5-0.6 mm long, the lobe
ovate, 0.8-0.9 mm long, 0.6-0.7 mm wide, apex
acute, sparsely but prominently black punctate with¬
out, very minutely ciliolate apically, the margin en¬
tire; corolla coriaceous, rotate, 3-3.6 mm long, the
tube 0.9-1.2 mm long, the lobes elliptic, 2.1-2.4
mm long, 1.1-1.2 mm wide, reflexed at antbesis,
apex rounded, very minutely puberulent near the
margin within and without, sparsely but prominently
short black punctate-lineate without, the margin en¬
tire, involute; sterile anthers inserted 0.8 1.1 mm
above tbe base of the corolla tube, the filaments 0-
0.3 mm long, flat, epunctate, the anthers elliptic,
1-1.7 mm long, apex rounded, base cordate, the
connective epunctate; pistil terete, 2.4-2.5 mm long,
the ovary globose, 1 — 1.2 mm long and in diam.,
prominently short black punctate-lineate, the stigma
sessile, sinuate-capitate, prominently vertically lobed,
1 1.1 mm long, 0.9-1 mm wide, persistent in fruit,
placenta globose, 2-ovulate. Fruit subglobose, 4.5—
5.5 mm long, 4-5 mm diam. sparsely but promi¬
nently short black punctate-lineate.
Distribution. Known only from the type, col¬
lected on rocky hillsides, at 2,800 m.
Myrsine microdonta is most closely related to
M. brevis (J. F. Macbride) Pipoly, but is easily
recognized by tbe appressed tomentum of the bran¬
chlets, leaves with dentate and ciliolate margins,
mucronulate apices, apically minutely ciliolate calyx
lobes, and elliptic corolla lobes with revolute mar¬
gins. The specific epithet refers to the dentate leaf
margins, a character unknown in any other tropical
American species.
Novon 2: 392-407. 1992.
Volume 2, Number 4
1992
Pipoly
Notes on the Genus Myrsine
393
Figure 1. Myrsine microdonta Pipoly. —A. Habit. —B. Pistillate flower habit, showing rotate corolla with involute
lobes. —C. Pistillate inflorescences, showing flower viewed from above and fruit. —D. Branchlet apex, showing
glandular-ciliate leaf bud margin. —E. Abaxial leaf surface, showing dentate, ciliolate margin, mucronulate apex, and
prominently raised costa. —F. Adaxial leaf surface, showing densely pilose, impressed costa and canaliculate petiole.
— G. Open pistillate flower, showing elliptic corolla lobes and sinuate-capitate stigma. AG, drawn from holotype.
394
Novon
Myrsine fosteri Pipoly, sp. nov. TYPE: Peru.
Pasco: Prov. Oxapampa, Oxapampa-Pasco
boundary, Abra San Guttardo, divide between
Ghontabamba and Hio Paucartambo Valley,
2,500-3,000 m, 29 Dec. 1983 (stam. 11), Fos¬
ter, Chanco & Alban 7636 (holotype, MO;
isotypes, AMAZ, F, NY, USM). Figure 2.
Ob ramulos dense ferrugineo-piloso-tomentosos, folia
ad apices obtusa vel emarginata atque minutissime mu-
cronulata, lobos corollinos manifeste atropunctatos atque
punctatolineatos, lobos calycinos manifeste atropunctatos
M. dependenti valde affinis sed ab ea foliis alternis (non
distichis) ad bases obtusis vel subcordatis (non sueto ro-
tundatis), lobis calycinis sparse glandulari-pilosis (non gla-
bris) secus margines dense longe-glandulari-pilosis (nec
parce glandulari-ciliolatis) manifeste atro-punctatis (nec
punctato-lineatis), lobis corollinis ad apices acutis (non
rotundatis) secus margines involutis (nec planis), antheris
oblongis (non ovatis) ad apices apiculatis (nec rotundatis)
facile cogniscitur.
Shrub to 2( 3) m tall; branchlets terete, 2-3 mm
diam., densely ferrugineous pilose. Leaves coria¬
ceous, ovate to obovate (1.3—)2—3 cm long, 1.4 2
cm wide, apex obtuse to emarginate, mucronulate,
base obtuse to subcordate, decurrent on the petiole,
sparsely pilose above, densely pilose below, the costa
impressed and densely pilose above, prominently
raised and densely pilose below, the secondary veins
6-8, visible but not raised above and below, incon¬
spicuously pellucid punctate, the margin densely
pilose in bud, persistent, not ciliate, revolute, epunc-
tate, glabrous; petioles canaliculate, 3-5 mm long,
densely pilose above and below. Staminate inflo¬
rescence: a subsessile umbel, (l-)3-flowered; floral
bracts ovate, ca. 0.5 mm long, densely pilose, the
apex rounded, the margin revolute, glandular-pilose;
pedicels terete, 0.7 0.9 mm long, glabrous. Flowers
5-merous, pinkish green; calyx chartaceous, cotyl-
iform, ca. 1 mm long, the tube ca. 0.1 mm long,
the lobes ovate, 0.8-0.9 mm long, 0.4-0.5 mm
wide, apex obtuse, prominently black punctate,
sparsely glandular-pilose without, glabrous within,
the margin entire, densely glandular-pilose; corolla
chartaceous, campanulate, 2.1 2.9 mm long, the
tube 0.4-0.5 mm long, the lobes elliptic, 1.7-2.4
mm long, 1-1.1 mm wide, apex acute, glabrous
except minutely pilose apically, prominently short
black punctate-lineate without, the margins entire,
involute, glabrous; stamens inserted at junction of
the corolla tube and lobes, the anthers sessile, ob¬
long, 1.6—1.8 mm long, 0.6-0.8 mm wide, apex
apiculate, base cordate, the connectives epunctate;
pistillode cylindrical, 1.2-1.3 mm long, 0.5-0.6 mm
diam., the pistil ellipsoid, 0.6 0.7 mm long, the
stigma irregularly lobed, 0.30.4 mm long. Pistil¬
late inflorescence: a subsessile umbel, 1 2-flowered;
floral bracts ovate, ca. 0.3 mm long, densely pilose,
the apex rounded, the margin revolute, glandular-
pilose; pedicels terete, 0.5-0.7 mm long, glabrous.
Flowers 5-merous, pinkish green; calyx chartaceous,
cotyliform, ca. 1 mm long, the tube ca. 0.1 mm
long, the lobes ovate, 0.8-0.9 mm long, 0.4 0.5
mm wide, apex obtuse, prominently black punctate,
sparsely glandular-pilose without, glabrous within,
the margin entire, densely glandular-pilose; corolla,
staminodes and pistil unknown. Fruit globose, 3
3.5 mm long and in diam., with a few scattered,
prominent short, black punctate-lineations.
Distribution. Endemic to the short shrubby “pa-
jonal” formations above cloud forest areas in Huan-
ueo and Pasco, Peru, at 2,500-3,600 m elevation.
Paratypes. Peru. HUANUCO: Prov. Huanuco, Carpish
Pass, above Chinchao, 9°42'S, 76°04'W, 24 Oct. 1947
(fr), Fosberg 28856 (MO, NA, US, USM). PASCO: Prov.
Oxapampa, Cordillera Yanachaga, Cerro Pajonal, 12 km
SE of Oxapampa, 20 D 35'S, 75°20'W, 2,700-2,800 m,
7 Oct. 1982 (bud), Foster 9011 (AMAZ, F, MO, USM);
San Gotardo, 36 km W of Oxapampa, 10°39'S, 75°37’W,
2,850 m, 21 Nov. 1982 (stam. fl), I). Smith 2762 (MO,
NY, USM); Oxapampa-Cerro de Pasco Kd., 38 km W
of Oxapampa, 10°39'S, 75°37'W, 2,750 m, 19 May
1983 (fr), I). Smith 4121 (MO, USM).
Myrsine fosteri may be confused with M. de¬
pendent (Ruiz & Pavon) A. Sprengel, but is easily
separated by its alternate leaves with obtuse or sub-
cordate bases, sparsely glandular-pilose and prom¬
inently black punctate calyx lobes with densely long
glandular-pilose margins, acute, involute corolla lobes,
and oblong anthers with apiculate apices. This spe¬
cies is dedicated to Robin Foster, ecologist and Ho-
ristic systematist, and one of the experts in the study
of Peruvian flora.
Myrsine (lilloniana Pipoly, sp. nov. TYPE: Peru.
Amazonas: Prov. Bagua, Cordillera Golan, E of
La Peca, 3,200 m, 28 Aug. 1978 (pist. fl, fr).
Harbour 3257 (holotype, MO; isotypes, AMAZ,
F, LL-TEX, USM). Figure 3.
Quoad ramulos angulatos, folia alterna, cartilaginea,
elliptica, ad apices obtusa vel rotundata, bases late ro-
tundata, petiolos usque ad 6 mm longos marginatosque,
pedicellos acute angulatos, lobos calycinos ovatos, species
haec M. lehmanii arete similans, sed ab ea laminis 4-6
(non 2-3.5) cm longis atque, 2.5 3.2 (nec 1-1.5) cm
latis, lobis calycinis carinatis (non planis) secus margines
crenulatis (nec integerrimis), lobis corollinis lineari-lan-
ceolatis (non oblongis) inconspicue pellucido-punctatis (nec
epunctatis) secus margines minutissime ciliolatis (nec gla-
bris) praeclare diagnoscenda.
Shrub to 2.5 m; branchlets angulate, 2.5-3.5
mm diam., glabrous. Leaves cartilaginous, elliptic,
4-6 cm long, 2.5-3.2 cm wide, apex obtuse, base
Volume 2, Number 4
1992
Pipoly
Notes on the Genus Myrsine
395
5mm
Figure 2. Myrsine fosteri Pipoly. —A. Habit. —B. Staminate flower habit, showing prominently punctate-lineate
corolla lobes. —C. Calyx and fruit, showing prominent punctations on calyx lobes and few, scattered punctate lineations
on fruit. —D. Branchlet apex, showing pilose indumentum. —E. Abaxial leaf surface, showing densely pilose,
prominently raised costa, mucronulate apex, and revolute margin. —F. Adaxial leaf surface, showing deeply impressed,
densely pilose costa. —G. Opened pistillate flower and calyx with pistillode, showing densely glandular-pilose calyx
lobe margins and ellipsoid pistillode. —H. Adaxial calyx lobe surface, showing prominently raised punctations. —I.
Abaxial calyx lobe surface. A, B, D-I, drawn from holotype; C, drawn from Fosberg 28856.
396
Novon
Figure 3. Myrsine dilloniana Pipoly. A. Habit. —B. Pistillate flower habit. —C. Fruit, showing persistent
prismatic stigma and globose fruit. —D. Branchlet apex, showing leaf bud with revolute, glabrous margin. —E.
Abaxial leaf surface, showing revolute leaf base and prominently raised costa. —F. Adaxial leaf surface, showing
slightly impressed costa and canaliculate petioles. —G. Opened pistillate flower, showing linear-lanceolate corolla
lobes, apiculate anthers, crenulate calyx lobes and prismatic stigma. A-G, drawn from holotype.
Volume 2, Number 4
1992
Pipoly
Notes on the Genus Myrsine
397
rounded, decurrent on the petiole, dull above and
below, the costa deeply impressed above, promi¬
nently raised below, the secondary veins scarcely
visible from above, 12-15 pairs, densely and very
minutely scrobiculate above, smooth below, the
punctations small, pellucid, inconspicuous, the mar¬
gin revolute, entire, glabrous in bud and at maturity;
petioles marginate, 3-7 mm long, glabrous. Sta-
minate inflorescence: unknown. Pistillate inflores¬
cence: a subsessile 15—20-flowered umbel; floral
bracts ovate, 1.3-1.5 mm long, 0.6-0.7 mm wide,
apex acute, early caducous, glabrous, the margins
entire, glabrous; pedicels ungulate, 2.6-2.8 mm long,
glabrous. Flowers 5-merous; calyx coriaceous, co-
tyliform, 2-2.1 mm long, the tube ca. 0.1 mm long,
the lobes ovate, 1.8-2 mm long, 1-1.1 mm wide,
apex acute, medially thickened, carinate, incon¬
spicuously pellucid punctate, the margin minutely
crenulate; corolla coriaceous, rotate, 3.9-4.1 mm
long, the tube 0.5-0.7 mm long, the lobes linear-
lanceolate, 3.2-3.4 mm long, 0.7-0.8 mm wide,
apex subulate, inconspicuously pellucid punctate,
the margins very minutely ciliolate; pistil 2.6-2.8
mm long, the ovary ellipsoid, 1.7 mm long, 1.2 mm
diarn., the ovary costulate, the stigma sessile, pris¬
matic, 3-angled, ca. 1 mm long, 0.7 mm diarn., the
placenta subglobose, 2-ovulate. Fruit globose, 3.5
5 mm long and in diarn., smooth, inconspicuously
punctate.
Distribution. Known only from the type, in elfin
forest, at 3,200 m.
Mr rsine dilloniana is closely related to M. leh-
mannii (Standley) Pipoly, but may be recognized
by its longer and wider leaves, carinate calyx lobes
with crenulate margins, and linear-lanceolate, in¬
conspicuously punctate corolla lobes with very mi¬
nutely ciliolate margins. I dedicate this species to
Michael Dillon (F), co-principal investigator for the
American side of the Flora of Peru Project.
Myrsine diazii Pipoly, sp. nov. TYPE: Peru. Ama¬
zonas: Prov. Luya, Camporedondo-Tullanya,
path to Cerro Huicsocunga, 2,350 m, (stam.
fl). Diaz A Campos 3681 (holotype, MO; is¬
otypes, AMAZ, BISH, F, US, USM). Figure 4.
Species haec ob ramulos glabros, folia membranacea,
anguste oblanceolata vel elliptica, punctata atque punc-
tato-lineata, ad apices basesque acuta, petiolos margina-
tos, lobo.s calycinos ovato-triangulares ciliolatos necnon
minute atro-punctatos, stigmata carnosa M. pellucido-
punctatae valde arete affinis, sed ab ea ramulis angulatis
(non teretibus) 2.5-3.5 (nec 6-8) mm diametro, inflo-
rescentiis subessilibus (non ramulis brevibus insidentibus),
umbellatis (nec fasciculatis), pedicellis angulatis (non ter¬
etibus), tubo corollino quadrato (non cylindrico), lobis cor-
ollinis manifeste atropunctatis atque punctato-lineatis (non
epunctatis), antheris apiculatis (non acutis) facile separ-
abilis.
Tree 6-8 m tall; branchlets angulate, 2.5 3.5
mm diam., glabrous. Leaves membranaceous, ellip¬
tic to narrowly oblanceolate, (3.6-)5-6.2 cm long,
1.6-2 cm wide, apex and base acute, decurrent on
the petiole, the costa slightly impressed above, prom¬
inently raised below, the secondary nerves not prom¬
inent, 13-18 pairs, nitid above, dull below, densely
pellucid punctate, the margin entire, flat or subre¬
volute, opaque, minutely glandular-ciliolate in bud,
glabrous at maturity; petioles marginate, 3-6 mm
long, glabrous. Staminate inflorescence: a subsessile
umbel, 4-6-flowered; floral bracts very widely ovate,
ca. 1 mm long, 1.4 mm wide, the apex obtuse, the
margin glandular-ciliolate; pedicels angulate, 0.7-1
mm long, glabrous. Flowers 5-merous; calyx char-
taceous, cotyliform, 0.8 1 mm long, the tube ca.
0.1 mm long, the lobes widely ovate, 0.7-0.9 mm
long, 0.7 0.8 mm wide, apex acute, sparsely black
punctate, the margin entire, glandular-ciliolate; co¬
rolla cotyliform, 2.7-2.8 mm long, the tube square,
0.2-0.3 mm long, the lobes lanceolate, 2.4-2.6 mm
long, 0.9 1 mm wide, apex acute, densely and
prominently black punctate and punctate-lineate
without, glabrous within, the margin entire, minutely
glandular-ciliolate; stamens inserted at junction of
corolla tube and lobe, the anthers sessile, oblong,
2-2.1 mm long, 0.9-1 mm wide, apex apiculate,
the apiculum proximally curved, base deeply cor¬
date, the connective epunctate; pistillode obturbi-
nate, 1.5-1.6 mm long, 0.5-0.6 mm diarn., the
ovary hollow, the stigma sessile, carnose. Pistillate
inflorescence: a subsessile umbel, 4-6-flowered; flo¬
ral bracts very widely ovate, ca. 1 mm long, 1.4
mm wide, the apex obtuse, the margin glandular-
ciliolate; pedicels angulate, 0.5-1 mm long, gla¬
brous. Flowers 5-merous; calyx chartaceous, cotyl¬
iform, 0.8-1 mm long, the tube ca. 0.1 mm long,
the lobes widely ovate, 0.7-0.9 mm long, 0.7-0.8
mm wide, apex acute, sparsely black punctate, the
margin entire, glandular-ciliolate; corolla cotyliform,
2.3- 2.5 mm long, the tube square, 0.6-0.8 mm
long, the lobes lanceolate, 1.7 1.9 mm long, 0.5-
0.6 mm wide, apex acute, densely and prominently
black punctate and punctate-lineate without, gla¬
brous within, the margin entire, minutely glandular-
ciliolate; staminodes inserted at junction of corolla
tube and lobe, the sterile anthers sessile, oblong,
1.3- 1.5 mm long, 0.9-1 mm wide, apex apiculate,
the apiculum proximally curved, base deeply cor¬
date, the connective epunctate; pistil obturbinate,
1.8-2 mm long, 0.9-1 mm diam., the ovary subglo-
398
Novon
5cm
Figure 4. My rsine diazii Pipoly. A. Habit. —B. Staminate flower habit, showing corolla lobes with prominent
pellucid punctations and punctate lineations, apiculate anthers. — C. Pistillate flower habit, showing same features as
in staminate flower and conical, carnose stigma. —D. Abaxial leaf surface, showing basally revolute margin, leaf
reticulation, and prominently raised costa. —E. Adaxial leaf surface, showing slightly impressed costa and decurrent
base on rnarginate petiole. —F. Leaf apex detail, showing dense pellucid punctations. — G. Sterile anther and corolla
lobe in proximal view. —H. Pistillate corolla, showing square corolla tube and reflexed corolla lobes. —I. Pistil,
showing conical, carnose stigma. — J. Pistillate calyx, showing cotyliforrn habit, sparse black punctations, and glandular-
ciliolate margins. —K. Branchlet apex, showing glandular-ciliate leaf bud margin. A, B, D F, K, drawn from holotype;
C, G-J, drawn from Diaz & Campos 3726.
Volume 2, Number 4
1992
Pipoly
Notes on the Genus Myrsine
399
bose, 0.9-1 mm long, prominently black punctate-
lineate, the placenta subglobose, 2-4-ovulate, the
stigma conical, carnose, 0.9 1 mm long. Fruit un¬
known.
Distribution. Known only from the type and
paratype, presumably endemic to the cloud forests
of Cerro Huicsocunga, 2,350 m elevation.
Paratype. PERU. AMAZONAS: Prov. Luya, Campore-
dondo-Tullanya, path to Cerro Huicsocunga, 2,350 m,
3 Sep. 1989 (pist. fl), Diaz & Campos 3726 (AMAZ,
F, US, USM).
Myrsine diazii is most closely related to M. pel-
lucido-punctata Oersted of Costa Rica and Panama,
but is easily distinguished from it by the thinner,
angulate branchlets, subsessile, umbellate inflores¬
cences, angulate pedicels, square corolla tube, prom¬
inently black punctate and punctate-lineate corolla
lobes, and apiculate anthers. It is dedicated to Camilo
Diaz Santibanez, Peruvian national coordinator of
the field research associates for the Flora of Peru
project and specialist in Peruvian Araliaceae.
Myrsine reynelii Pipoly, sp. nov. TYPE: Peru.
Piura: Prov. Huancabamba, Dtto. Canchaque,
between Chorro Blanco and War War, 2,000—
2,500 m, 18 Jan. 1989 (fr), Diaz, Pennington
& Reynel 3180 (holotype, MO; isotypes,
AMAZ, USM). Figure 5.
Propter laminam coriaceam ellipticam vel ohlanceola-
tam insuper scrobiculatam, petiolos marginatos, lobos ca-
lycinis coriaceos inconspicue pellucido-punctatos, M. so-
diroanae affinis sed ab ea laminis atropunctato-lineatis
(non inconspicue punctatis) secus margines revolutis (nec
planis), pedicellis cylindricis (non angulatis) 2-3.3 (nec
6-8) mm longis, lobis calycinis ovatis (non late triangu-
laribus) secus margines glabris (nec ciliolatis) praeclare
distat.
Tree to 8 m tall; branchlets terete, 4 6 mm diam.,
glabrous. Leaves coriaceous, elliptic to oblanceolate,
6-10 cm long, 2 3 cm wide, apex acute, base
cuneate, decurrent on the petiole, the costa im¬
pressed above, prominently raised below, secondary
veins not prominent, 15-19 pairs, nitid, scrobiculate
and somewhat rugose above, pallid and conspicu¬
ously black punctate and punctate-lineate below, the
margin entire, opaque, revolute, prominently black
punctate, minutely glandular-ciliate in bud, glabrous
at maturity; petioles marginate, 0.8-1 (-1.4) cm
long, glabrous. Staminate inflorescence: not seen.
Pistillate inflorescence: a sessile (aside, 5-9-flow-
ered; floral bracts coriaceous, very widely ovate, ca.
0.9-1.2 mm long, 0.9-1.3 mm wide, the apex acu¬
minate, the margin opaque, entire, glabrous; pedicels
cylindrical, 2 3.3 mm long, glabrous. Fruiting calyx
5-merous, coriaceous, cotyliform, 1-1.4 mm long,
the tube ca. 0.2-0.3 mm long, the lobes ovate, 0.7-
1.1 mm long, 0.5-0.7 mm wide, apex subacumi¬
nate. inconspicuously pellucid punctate, the margin
entire, glabrous; corolla, androecium, and gynoe-
cium unknown. Fruit slightly depressed-globose, 2.7-
3 mm long, 3-3.2 mm diam., inconspicuously pel¬
lucid punctate.
Distribution. Endemic to the cloud forests of
Piura Province, 1,900-3,000 m elevation.
Paratypes. PERU. PIURA: Prov. Ayabaca, Frias, Las
Pircas, 3,000 m, 17 Oct. 1988 (fr), Diaz & Vdsquez
3006 (AMAZ, MO, USM); Prov. Huancabamba, Dtto.
Canchaque, road between Canchaque and Huancabamba,
km 16 toward the left, Quehrada La Mina, 1,900 m, 17
Apr. 1987 (fr), Diaz & Baldeon 2380 (AMAZ. MO,
USM).
Myrsine reynelii is most closely related to M.
sodiroana (Mez) Pipoly, but is easily recognized by
its black punctate-lineate, revolute leaves, shorter
cylindrical pedicels, and ovate, glabrous calyx lobes.
Myrsine sodiroana is endemic to the cloud forests
of Pichincha in Ecuador, while M. reynelii is en¬
demic to the cloud forests of Piura in Peru. This
species is dedicated to Carlos Reynel, professor ot
dendrology at the Universidad Nacional Agraria at
La Molina, and currently a graduate student at the
Missouri Botanical Garden.
Myrsine bill lata Pipoly, sp. nov. TYPE: Peru.
San Martin: Prov. Mariscal Caceres, Rio Abiseo
National Park. nr. Puerta del Monte, 7°27'S,
77°21'W, 3,200 m, 7 Sep. 1985 (fr), k. )oung
1572 (holotype, MO; isotypes, F, USM). Fig¬
ure 6.
Quoad laminas coriaceas ellipticas vel oblanceolatas
bullatas insuper nitidas secus margines revolutas, lobos
calycinis coriaceos ovatosque species haec ad M. perre-
ticulata affinis, sed ab ea laminis subter pellucido-punc-
tatis (non atro-punctato-lineatis) minute rufo glandulari-
papillosis (nec adpresso-lepidotis) secus margines coriaceis
opacis epunctatisque (nec cartaceis hyalinis aurantiaco-
punctatisque), pedicellis 5-7 (non 2-4) mm longis, lobis
calycinis planis (non carinatis) 1.4-1.7 (nec 2.5-2.8) mm
longis ad apices acuminatis integerrimisque (nec obtusis
erosisque) facile cognoscitur.
Tree to 13 m tall; branchlets terete, 3.5-5 mm
diam., glabrous. Leaves coriaceous, elliptic to ob¬
lanceolate, (5.5-)9 15 cm long, (2-)3.5-4 cm wide,
apex acute, base cuneate, decurrent on the petiole,
costa impressed above, prominently raised below,
secondary veins 12-15 pairs, impressed above,
prominently raised below, bnllate, nitid and glabrous
above, minutely rufous glandular-papillate and in¬
conspicuously pellucid-punctate below, the margin
400
Novon
1 cm
Figure 5. Myrsine reynelii Pipoly. —A. Habit. —B. Inflorescence, showing cylindrical pedicels and ovate, glabrous
calyx lobes. C. Adaxial leaf surface, showing scrobiculate, rugose surface. —D. Abaxial leaf surface, showing
punctate lineations, revolute margin and marginate petiole. —E. Branchlet apex, showing bud texture and glandular-
granulose margin. A E, drawn from holotype.
Volume 2, Number 4
1992
Pipoly
Notes on the Genus Myrsine
401
Figure 6. Myrsine bullata Pipoly. —A. Habit. —B. lufructescence, showing cylindrical pedicels and glandular-
ciliate floral bract margins. —C. Abaxial leaf surface, showing minute rufous glandular papillae and revolute margin.
— D. Adaxial leaf surface. —E. Branchlet apex, showing glandular-ciliate leaf bud margin. A E, drawn from holotype.
402
Novon
opaque, epunctate, entire, revolute, and glabrous in
bud and at maturity; petioles marginate, 0.5-0.8
cm long, glabrous. Staminate inflorescence: un¬
known. Pistillate inflorescence: a sessile umbel, 7—
10-flowered; floral bracts coriaceous, ovate, 2.2
2.0 mm long, 0.8-1 mm wide, the apex acuminate,
the margin opaque, entire, densely glandular-ciliate;
pedicels cylindrical, 5-7 mm long, glabrous. Fruiting
calyx 5-merous, coriaceous, cotyliform, 1.4 1.7 mm
long, the tube ca. 0.2-0.3 mm long, the lobes ovate,
1.2-1.4 mm long, 0.9 1.2 mm wide, apex acu¬
minate, inconspicuously pellucid punctate, sparsely
red-papillate, the margin entire, glabrous; corolla,
androecium and gynoecium unknown. Fruit de¬
pressed-globose, 4.8 5.1 mm long, 5.1-0 mm diam.,
purple, inconspicuously pellucid-punctate.
Distribution. Endemic to the Mariscal Caceres
Province, of San Martin Department, Peru, at the
upper limit of montane rainforest with cloud forest,
3,200-3,400 in.
Paratype. PERU, san MARTIN: Prov. Mariscal Caceres,
Rio Abiseo National Park, 7°0()'S, 77°00'W, NW corner
of Park, Chochos, S side of river, 3,400 m, 7 June 1986
(fr), Young 3721 (F, US, USM).
Myrsine bullata is the sister species of M. per-
reticulata Pipoly, but is easily separated from it by
the pellucid-punctate and minutely rufous glandular-
papillate abaxial leaf surface, the coriaceous, opaque
and epunctate leaf margins, longer pedicels and flat,
acuminate calyx lobes with entire margins. The spe¬
cific epithet refers to the prominently bullate char¬
acter of the leal blades.
Myrsine youngii Pipoly, sp. nov. TYPE: Peru.
San Martin: Prov. Mariscal Caceres, Rio Abiseo
National Park, NW corner, Puerta del Monte,
trail to el Mirador, 7°00'S, 77°00'W, 3,100-
3,300 m, I 1 July 1987 (fr). Young & Leon
1 PIS (holotype, F; isotypes, US, USM). Fig¬
ure 7.
Ob laminas anguste oblanceolatas vel ellipticas, punc-
tatas atque punctato-lineatas, ad apices acutas, etiam pe-
tiolos marginatos, M. pellucido-punctatae et M. diazii
simulans, sed species haec ab ambabus raniulis translu-
cido-papillatis (non glabris), laininis coriaceis (non mem-
branaceis) secus rnargines manifeste revolutis (nec planis
nec subrevolutis) parce et minute translucido-papillatis
(nec glabris), lobis calycinis coriaceis (non chartaceis) ova-
tis (nec ovato-triangularibus) ad apices acuminatis (nec
acutis) atque epunctatis (nec punctatis), stylo 0.2-0.3
mm longo (non obsoleto), ulterius a prima ramulis 2.5-
3.5 (non 6-8) mm diametro, inflorescentiis subsessilibus
(non ramulis brevibus insidentibus), stigmatibus 0.5-0.7
(non 1 1.4) mm longis, a secunda ramulis teretibus (non
angulatis) petiolis 0.7-1 (non 0.3 -0.6) cm longis, pedi-
cellis teretibus (non angulatis) 2-3 (nec 0.5 1) mm longis,
atque stigmatibus sinuato-lobulatis (non conicis) praeclare
distinguitur.
Free to 12 m; branchlets terete, 2.5-3.5 mm
diam., translucent glandular-papillate apically. Feuves
coriaceous, elliptic to oblanceolate, 3.5-5 cm long,
1.5-2 cm wide, apex acute, base cuneate, decurrent
on the petiole, costa impressed above, prominently
raised below, secondary veins inconspicuous, 8 1 2
pairs, sparsely translucent glandular-papillate above,
densely at first, then sparsely minutely translucent
glandular-papillate and black punctate and punctate-
lineate below, the margin opaque, epunctate, entire,
revolute, densely filiate in bud, glabrous at maturity;
petioles marginate, 0.7-1 cm long, glabrous. Sla¬
minate inflorescence: unknown. Pistillate inflores¬
cence: a subsessile fascicle, 5-8-flowered; floral bracts
coriaceous, very widely ovate, 1-1.5 mm long, 1.3-
1.5 mm wide, the apex acuminate, the margin
opaque, entire, densely glandular-ciliate; pedicels
cylindrical, 2-3 mm long, glabrous. Fruiting calyx
5-merous, coriaceous, cotyliform, 1.2-1.5 mm long,
the tube ca. 0.1 -0.2 mm long, the lobes ovate, 1.2
1.3 mm long, 0.6-0.9 mm wide, apex acuminate,
inconspicuously pellucid punctate, glabrous, the
margin entire, glandular-ciliate; corolla, androecium
and gynoecium unknown except persistent style 0.2
0.3 mm long, the stigma sinuate-lobed, 0.5-0.7 mm
long. Fruit globose, 2.7-3 mm long, 2.5 2.8 mm
diam., prominently black punctate.
Distribution. Known only from the type.
Myrsine youngii is closely related to M. pellu-
cido-punctata and M. diazii, but is readily sepa¬
rated from both by the translucent papillate bran¬
chlets, the coriaceous, revolute, and sparsely
translucent papillate leaves, the coriaceous, ovate
and acuminate calyx lobes, and the lobed stigma on
a style. Furthermore, M. youngii is distinguished
from M. pellueido-punctata by the thinner bran¬
chlets, subsessile inflorescence, and smaller stigma,
while from M. diazii it is clearly separated by the
terete branchlets, longer petioles, terete and longer
pedicels, and sinuate-lobulate stigma. The presence
of a style in Myrsine youngii once again reinforces
the concept that maintenance of Rapanea Aublet
as distinct from Myrsine F. is untenable. This species
is dedicated to Ken Young, phytogeographer and
floristician, whose excellent collections and papers
on Peruvian “ceja de selva’' have proven very valu¬
able in furthering its study and conservation.
Myrsine minutiflora Pipoly, sp. nov. TYPE: Peru.
Fambayeque: Prov. Ferrenafe, Bosque de Chin-
ama, 2,300-2,700 m, 15 Aug. 1988 (stain.
fl), I alencia 237V (holotype, US; isotype,
USM). Figure 8.
Volume 2, Number 4
1992
Pipoly
Notes on the Genus Myrsine
403
Figure 7. Mvrsinc y'oungii Pipoly. —A. Habit, showing subsucculent branchlets. B. Branchlet apex, showing
glandular-papillate leaf buds with margins glandular-ciliate. —C. Adaxial leaf surface. D. Abaxial leaf surface,
showing translucent glandular-papillae. —E. Pistillate infructescence, showing fruit, persistent style and sinuate-lobed
stigma. A E, drawn from holotype.
Quoad ramulos glabros, folia coriacea elliptica vel ob-
lanceolata ad apices acuta vel obtusa, infiorescentia sessilia
fasciculataque, pedicellos breves, species haec cum M.
pellucida saepenumero confusa est, sed ab ea floris stam-
inatae corollis 1.8-2 (non 2.5-2.7) mm longis, pedicellis
1-1.2 (non minus quam 1) mm longis, pistillodio costato
(non tereti), floris pistillatae calycis lobis acuminatis (non
rotundatis) secus margines glandulari-ciliolatis (nec gla-
bris), pedicellis 1.5- 1.7 mm longis (non obsoletis), denique
stiginate conico (nec lobulado) statim separabilis.
Tree, height unknown; branchlets ungulate, 3.5-
4.5 mm diam., prominently red punctate, glabrous.
Leaves coriaceous, elliptic to oblanceolate, (4.2 )5
404
Novon
Figure 8. Myrsine minutiflora Pipoly. A-D, F-G drawn from holotype, E drawn from Valencia 2248. — A. Habit,
showing angulate branchlets. —B. Branchlet apex, showing punctations on leaf hud and stem. —C. Ahaxial leaf
surface, showing black punctate lineations and marginate petiole. —D. Adaxial leaf surface. —E. Infructescence,
showing calyx and fruit. — F. Staminate inflorescence. — G. Staminate flower, showing costate pistillode.
Volume 2, Number 4
1992
Pipoly
Notes on the Genus Myrsine
405
7(-9) cm long, (1.9-)2.5-3 cm wide, apex acute
to obtuse, base cuneate, decurrent on the petiole,
costa impressed above, prominently raised below,
secondary veins inconspicuous, 13 15 pairs, red
perpuncticulose and black punctate-lineate below,
the margin opaque, epunctate, entire, flat, densely
ciliate in bud, glabrous at maturity; petioles mar-
ginate, 0.7-1 cm long, glabrous. Staminate inflo¬
rescence: a sessile fasicle, 3-6-flowered; floral bracts
coriaceous, widely ovate, 0.7-0.8 mm long, 0.7-
0.9 mm wide, apex obtuse to subacute, the margin
entire, glabrous; pedicels angulate, 1-1.2 mm long,
glabrous. Flowers 5-merous, green; calyx coria¬
ceous, cotyliform, 0.7 1 mm long, the tube ca. 0.2
mm long, the lobes ovate, 0.5-0.8 mm long, 0.3-
0.5 mm wide, apex acuminate, somewhat cucullate,
densely and prominently red punctate, the margin
densely glandular-ciliolate; corolla coriaceous, sub¬
rotate, 1.8-2 mm long, the tube ca. 0.6 mm long,
the lobes ovate, 1.2-1.4 mm long, 0.7-0.9 mm
wide, apex subacuminate and proximally recurved,
inconspicuously pellucid-punctate and glabrous with¬
out, densely glandular-granulose throughout within,
the margin opaque, densely glandular-ciliate; sta¬
mens 1-1.2 mm long, the filaments conspicuous,
flat, 0.2-0.4 mm long, the anthers ovate, 0.7-0.9
mm long, 0.4 0.6 mm wide, apex apiculate, base
deeply cordate, the connective epunctate, glabrous;
pistillode ovoid, 1-1.2 mm long, 0.3-0.4 mm diam.,
prominently costate, hollow, the stigma punctiform.
Pistillate inflorescence: a sessile fasicle, 3-5-flow-
ered; floral bracts coriaceous, widely ovate, 0.7-1
mm long, 0.8-1 mm wide, apex acute, the margin
entire, glabrous; pedicels angulate, 1.5-1.7 mm long,
glabrous. Fruiting calyx 5-merous, coriaceous, co¬
tyliform, 1.3 1.5 mm long, the tube ca. 0.2 mm
long, the lobes ovate, 1.1-1.3 mm long, 1.1-1.2
mm wide, apex acuminate, somewhat cucullate,
densely and prominently red punctate, the margin
densely glandular-ciliolate; corolla, androecium and
gynoecium unknown. Fruit subglobose, 3 3.5 mm
long and in diarn., densely and prominently red-
punctate; persistent stigma conical, ca. 0.1 0.2 mm
long.
Distribution. Endemic to the western slopes of
the Peruvian Andes in the departments of Lamba-
yeque, Piura, and La Libertad, at 2,300 2,950 in.
Paratypes. Peri:, la LIBERTAD: Prov. Otuzco, Bosque
de Llaguen, 2,500-2,800 in, 5 Nov. 1988 (ster.), Cano
2378 (US, USM); Prov. Ferrenafe, Bosque de Chinama,
2,300-2,700 in, 15 Aug. 1988 (pist. fr), Valencia 2248
(US, USM). piifra: Prov. Ayabaca, Bosque de Huamba,
2,850-2,950 m, 15 Oct. 1987 (ster.), Cano 1514 (US,
USM), (bud), Valencia 2027 (US, USM).
Myrsine minutiflora has often been confused with
M. pellucida (Ruiz & Pavon) A. Sprengel, but can
at once be recognized by its staminate flowers with
costate pistillode, with shorter corollas and pedicels.
In addition, the longer pistillate pedicels, conic stig¬
ma and acuminate, glandular-ciliolate calyx lobes of
M. minutiflora are distinctive. Both Myrsine min¬
utiflora and M. pellucida have been confused with
M. guianensis (Aublet) Kuntze and M. latifolia
(Ruiz & Pavon) A. Sprengel, but bear only superficial
vegetative resemblance to both. Aside from differ¬
ences in floral morphology, Myrsine minutiflora is
separated from M. guianensis by the sessile inflo¬
rescences (not on short shoots) on branchlets with
thin (not corky) bark, and from M. latifolia by the
fasciculate (not umbellate) inflorescences, smaller
leaves, and shorter pedicels.
In addition to the novelties described above, treat¬
ment of Rapanea as a taxonomic synonym of Myr¬
sine necessitates the following new combinations.
Because Peruvian Myrsinaceae have long been poor¬
ly understood, I include citations of representative
specimens when possible.
Myrsine sprucei (Mez) Pipoly, comb. nov. Bas-
ionym: Rapanea sprucei Mez in Engl., Pflan-
zenr. IV. 236(9): 388. 1902. TYPE: Peru.
San Martin: Tarapoto, Spruce 4251 (holotype,
B destroyed; isotypes, K, G, G-DEL, OXF).
Distribution. Endemic to the lower eastern slopes
of the Andes in Peruvian Amazonia, 2,400-2,500
m elevation.
Representative specimen examined. Peri . AMAZONAS:
Prov. C.hachapoyas, Cerros de Leimebamba, 2,400-2,500
m, 20 Apr. 1964 (fr), Ferreyra 15594 (MO, USM).
Myrsine brevis (J. F. Macbr.) Pipoly, comb. nov.
Basionym: Rapanea brevis J. F. Macbr., Can-
dollea 5: 396. 1934. TYPE: Peru. La Libertad:
Prov. Pataz, Tayabamba, 1900-1914 (fr), H e-
berbauer 7025 (holotype, F).
Distribution. Endemic to the north-central region
of the Peruvian Andes, 2,850-3,600 m.
Representative specimens examined. Peru, la liber¬
tad: Prov. Pataz, Retamas-La Paccha, rd. to Tayabamba,
3,600 m, 24 June 1974 (stam. fl), Lopez <£- Sagdstegui
8172 (HUT, MO).
Myrsine lechleri (Mez) Pipoly, comb. nov. Bas¬
ionym: Rapanea lechleri Mez, Pflanzenr. IV.
236(9): 380. 1902. TYPE: Peru. Puno: Ta-
tanara, Lechler 2647 (holotype, K).
Distribution. Known only from the type.
406
Novon
lYlyrsine rivularis (Mez) Pipoly, comb. mov. Bas-
ionym: Rapanea rivularis Mez, Bull. Herb.
Boissier 2 ser. 5: 536. 1905. TYPE: Peru.
Puno: between Sandia and Tambo Azalayo,
1,500 rn, Weberbauer 1061 (holotype, B de¬
stroyed; fragment, F).
Distribution. Known only from the type.
lYlyrsine sessilillora (Mez) Pipoly, comb. nov.
Basionym: Rapanea sessiliflora Mez, Reperl.
Spec. Nov. Regni Veg. 3: 103. 1906. SYN¬
TYPES: Peru. Amazonas: near Cuelap, near
Chachapoyas, II eberbauer 1321 (syntype, B
destroyed; G-DEL, as photo at F, Neg. 26699).
Cajamarca: Prov. Cajamarca, above San Pablo,
2,400 m, If eberbauer 3810 (syntype, B de¬
stroyed; fragment, F); Prov. Chota, between
Huambos and Montan, 2,500-2,600 rn, If e-
berbauer 1216 , 1217 (syntypes, B destroyed).
I have not seen the isosyntype of II eberbauer
3810 at G-DEL, but the type photo leaves no doubt
as to the identity ol this species.
Distribution. Endemic to the eastern slopes of
the eastern Andean cordillera of Cajamarca, lluan-
uco, San Martin and Amazonas, Peru, in cloud and
elfin forests, 1,960-3,500 m.
This species has been poorly understood, but is
at once recognized by it sessile, few-flowered, fas¬
ciculate inflorescences, flexuous, ridged branchlets,
and patelliform calyces.
Representative specimens examined. Pkki . AMAZONAS:
Leimebamba, 2,300 in, 20 Dec. 1962 (fr), II oytkoicski
7780 (MO). CAJAMARCA: Prov. Cajamarca, road between
Cajamarca and Celendin, 2,950-3,500 m (pist. H, fr),
Diaz 2161 (('PUN, MO, USM); below Paso Gavilan be¬
fore Choten, 2,800 2,850 m, 11 Feb. 1987 (fr), Sanchez
4210 (('PUN, MO); 50-60 km E of bridge over rio
Maichil, 6°28'S, 79°00'W, 2,500-2,600 in, 9 Feb. 1988
(pist. H, fr), Gentry et al. 61437 (CPUN, MO, USM);
Prov. Celendin, Celendin-Balsas Rd., 3-10 km from Ce¬
lendin, 6°52'S, 78°07'W, 2,700-2,900 m, 16 July 1983
(fr), I). Smith Sanchez 4303 (MO, USM); Prov. Chota,
Llama, caserio Callampampa, tributary of Rio Huarimar-
ca, 2,800 3,000 m, 19 Jan. 1990 (pist. ft, fr), Diaz &
Osores 4013 (AMAZ, CPUN, F, MO, US, USM); Prov.
Contumaza, 10-11 km from Contumaza on rd. to Cascas,
7°25'S, 78°50'W, 2,550 m, 7 Fell. 1987 (pist. II, fr),
Stein et al. 4045 (CPUN, MO, USM); Lleden, 2,400 m,
28 June 1983 (fr), Sagdstegui et al. 10844 (HUT, MO,
NY, USM); San Martin, between Singarran and Lleden,
2,900 m, 1 June 1979 (pist. H), Sagdstegui et al. 0347
(HUT, MO); Prov. Hualgayoc, Tahona, between Bam-
barnarca and Hualgayoc, 2,900-3,000 m, 16 Aug. 1952
(bud), Ferreyra 8500 (MO, LJSM).
lYlyrsine weberbaueri (Mez) Pipoly, comb. nov.
Basionym: Rapanea weberbaueri Mez, Re¬
port. Spec. Nov. Regni Veg. 3: 103. 1906.
SYNTYPES: Peru. Huanuco: Prov. Huamal-
ies, nr. Monzon, 1,000-1,500 m, (stain, fl),
IT eberbauer 3457, (pist. fl), 3162 (syntypes,
B destroyed, F Neg. 4895).
Leetotypification is postponed until isosyntypes
are located.
Distribution. Endemic to the Huanuco area of
the north-central Andean cordillera of Peru, 800
1,500 m.
Representative specimen examined. PERI 1 . HCANlico:
Prov. Leoncio Prado, Dtto. Rupa Rupa, E of Tingo Maria,
nr. Cerro Quemado, 800 850 m, 15 Aug. 1978 (fr),
Schunke 10488 (F, MO).
Upon examining specimens of non-Peruvian Myr-
sirie species, the following other new combinations
have come to light.
lYlyrsine sodiroana (Mez) Pipoly, comb. nov. Bas¬
ionym: Rapanea sodiroana Mez in Engl.,
Pflanzenr. IV. 236(9): 385. 1902. TYPE: Ec¬
uador. Pichincha: without locality, Sodiro 1003
(holotype, B destroyed; F Neg. 4892).
Despite the fact that the holotype of this species
was destroyed in Berlin, the photograph at F permits
positive identification.
Distribution. At present known only from Pi¬
chincha, Ecuador, at 1,500 3,500 m.
Representative specimens examined. Ecuador. I.OJa/
ZAMORA—CHINCH I PE: Parque Nacional Podocarpus, Yan-
gana Valladolid, Nudo de Sabanilla, 4°27'S, 79°08'W,
2,750-2,900 in, 28 Feb. 1985 (fr), (dllgaard et id.
58407 (AAU, QCA). pichincha: Canton Quito, Parroquia
Calacali, Reserva Geobotanica Pululahua, Road to Moras
Pungo, 00°05'N, 78°30'W, 1,800-3,356 m, 23 Aug.
1987 (fr), Ceron & Benavidez 1910 (MO, QCA).
Myrsine pseudocrenata (Mez) Pipoly, comb. nov.
Basionym: Rapanea pseudocrenata Mez in
Engl., Pflanzenr. IV. 236(9): 393. 1902. TYPE:
Bolivia. Sorata, Cochipata on Rio l leumauni,
3,000-3,200 m, Mandon 515 (holotype, B
destroyed; F Neg. 4891).
Distribution. Known only from Bolivia, but ex¬
pected in Puno, 2,500-3,500 m.
Representative specimens examined. B()LI\ IA. COCHA¬
BAMBA: Prov. Carrasco, 5 km E of bridge over Rio Lopez
Mendoza, on Cochabamba-Santa Cruz Rd., 19 kin W of
Epizana, I 7°32'S, 65°22'W, 2,900 in, 11 Feb. 1987
(fr), Solomon & Nee 16050 (LPB, MO, NY, IIS), la
paz: Prov. Sud Yungas, below Unduavi, Cerroinarca Val¬
ley, 3,450 m, 28 Aug. 1988 (stain, fl). Beck 14682
(EBP, MO, US).
Volume 2, Number 4
1992
Pipoly
Notes on the Genus Myrsine
407
Myrsine congesta (Schwacke ex Mez) Pipoly,
comb. nov. Basionym: Rapanea congesta
Schwacke ex Mez in Engl., Pflanzenr. IV.
236(9): 383. 1902. SYNTYPES: Brazil. Minas
Gerais: Serra do Ouro-Preto, Gomes 1273 (syn-
type, M—as Photo F, Neg. 20074). Rio de
Janeiro: near Petropolis on Mono Carangola,
Glaziou 11011 (syntype, C—as Photo, F Neg.
22973; isosyntype, RB).
I defer lectotypification until all syntypes have
been studied.
Distribution. Minas Gerais and Rio de Janeiro,
Brazil.
Myrsine lehmannii (Standley) Pipoly, comb. nov.
Basionym: Rapanea lehmnannii Standley,
Publ. Field Mus. Nat. Hist., Bot. Ser. 11:1 70.
1936. TYPE: Colombia. Cauca: Paramo Fas
Delicias, Central Andes of Popayan, Lehmann
8459 (holotype, F).
Distribution. Endemic to the western slopes ol
the Central Cordillera of Colombia, in the general
area of the Macizo Colombiano.
Ibis species is apparently endemic to the slopes
of the western Andean cordillera in Colombia, hut
is very closely related to Myrsine dilloniana , a
Peruvian species described herein.
Acknowledgments. Support lor my studies on the
Flora of Peru is provided by a grant Irom the
Andrew W. Mellon Foundation, which is gratefully
acknowleged. 1 thank the John I). and Catherine T.
MacArthur Foundation for support of fieldwork in
Peru as part of the Documentation of Neotropical
Plant Species Diversity program. Thanks are also
due to Michael Dillon (F) and Al Gentry (MO) for
continued collaboration with the Flora of Peru Pro¬
ject. Alan Brant and Jon Ricketson, project coor¬
dinators (MO), to the curators of herbaria cited for
loaning material, the Peruvian field stafF of the Mis¬
souri Botanical Carden, and my volunteers, Cath¬
erine Mayo, Valerie Levi, David Belt, and John
Skinner. Finally, I thank John Myers and Linda Ellis
for the skillfull illustrations of new taxa described
herein.
Literature Cited
Pipoly, J., III. 1992. Estudios del genero Myrsine L.
(Myrsinaceae) en Colombia. Caldasia 17(1): 1-5.
Adenophorus Xcarsonii hyb. nov. (Grammitidaceae): A New Endemic
Hybrid from the Island of Hawaii
Tom A. Ranker
University Museum and Department of Environmental, Population, and Organismic Biology,
University of Colorado, Boulder, Colorado 80309-0350, U.S.A.
ABSTRACT. A new hybrid between two species of
the endemic Hawaiian genus Adenophorus is de¬
scribed. On morphological grounds it appears to be
a hybrid between . 1 . hymenophylloides and A. Iri-
pinnatifidus. That hypothesis is supported by pre¬
liminary isozyme data, which also demonstrate that
different individuals were produced by separate hy¬
bridization events. The irregular size and shape of
spores and lack of complete pairing of meiotic chro¬
mosomes indicate that the hybrid is infertile.
Adenophorus Gaudich. is an endemic Hawaiian
genus comprising two subgenera, Adenophorus (6
species) and Oligadenus I.. E. Bishop (4 species;
Bishop, 1974). Most members of the genus are
widespread on the main Hawaiian Islands, although
A. haalilioanus (Brack.) K. A. Wilson is restricted
to Kauai and Oahu, and A. oahuensis (Copel.) L.
E. Bishop only occurs on Oahu. Adenophorus per-
iens L E. Bishop is known only from a few locations
on Kauai, Hawaii, and Molokai. Most species are
obligate epiphytes occurring in habitats ranging from
montane mesic to wet forests. Although Bishop (1974)
described a hybrid between A. oahuensis and A.
pinnatifidus Gaudich., both in subgenus Oligaden¬
us , the hybrid I present here is the first report of a
naturally occurring hybrid between two species in
subgenus Adenophorus.
Adenophorus xcarsonii Hanker, hyb. nov.
TYPE: U.S.A. Hawaii: Island of Hawaii, Puna
District, Kahaualea Natural Area Reserve, 5.6
km S of Hwy. 1 1 at end of South Glenwood
Rd., 700 m, 6 June 1990, Ranker 1115 (ho-
lotype, BM; isotype, COLO). Figure 1.
Rhizomata breviter reptantia; paleae elongato-lanceo-
latae; folia 12-35 cm longa, pendula sed juvenilia erecta;
lamina bipinnatifida; pinnae 5-15 cm longae; pinnulae
basales et acroscopicae simplices vel pinnatifidae; sporae
abortivae; planta inter A. hymenophylloides et A. tri-
pinnatijidus intermedia.
Plants epiphytic; rhizome short-creeping, 2-4 mm
diam.; rhizome scales 3 4 mm long, 0.3-0.5 mm
wide, amber-colored, long-lanceolate, rounded to au-
riculate at base, subulate to slightly rounded at tip
often bearing a glandular apical cell, occasionally
branched near tip with an additional glandular cell
at apex of branch; cells of rhizome scales rhomboid
to fusiform, several 1o many times longer than wide;
leaves mostly 12-25 cm long, occasionally up to
35 cm, 1.5-2.0 cm wide, erect when young but
soon becoming pendulous as they mature, appearing
tufted; stipe 0.3-0.5 mm diam., 1.0-2.0 cm long,
slightly alate when young but not apparent on older
stipes, sparsely to densely clothed with appressed
glandular trichomes; blades bipinnatifid, linear, with
appressed, glandular trichomes on both surfaces and
rachis, tip long-tapering or abrupt; pinnae 5 15 mm
long with 6-10 pinnules; pinnules mostly simple,
spathulate, apex slightly rounded to acute, basal
pinnules 2.0-7.0 mm long, 0.8-1.0 mm wide, be¬
coming smaller toward tip of pinna, basal acroscopic
pinnule often divided again; mature sori abaxial,
about 1.0 mm diam., equaling the width of the fertile
pinnule; sporangia with 12-15 annular cells; spores
irregular and aborted.
Plants discovered to date only grow epiphytically
on bryophyte-covered trunks of Metrosideros poly-
rnorpha Gaudich. (Myrtaceae) in forests dominated
by a mixture of that species and the tree fern 67-
botium glaucum (Smith) Hook. & Arn.
The epithet is in honor of Hampton L. Carson,
University of Hawaii, who first recommended that
I visit the Kahaualea Natural Area Reserve where
the hybrid was first discovered.
Paratypes. U.S.A. HAWAII: Island of Hawaii, Puna
District, same as Ranker 1115 but collected on 27 May
1991, Ranker 1202 (MICH, PTBG, US); forest NW of
Volcano refuse transfer site and SE of Olaa tract of HVNP,
870-900 m, Ranker (1315) & Trapp (COLO).
The possible hybrid nature of A. Xcarsonii was
suggested on morphological grounds because of its
similarity to both A. h ymenophylloides and A. tri-
pinnatifidus , its putative parents. Adenophorus
Xcarsonii differs from these two species in several
morphological features and is usually approximately
intermediate between them (Table 1, Fig. 1). The
hybrid is most easily confused at first sight with A.
Novon 2: 408-410. 1992.
Volume 2, Number 4
1992
Ranker
Adenophorus x carsonii
409
Figure 1. A C. Adenophorus x carsonii Ranker. —A. Entire plant. —B. Detail of abaxial surface of pinna.
— C. Detail of abaxial surface of pinnule. —D. Abaxial surface of pinna of A. hymenophylloides. —E. Abaxial
surface of pinna of A. tripinnatifidus.
hymenophylloides because of the pendulous habit
of the leaves of both species. The broader and longer
leaf blade, larger pinnae, and irregularly shaped
spores of the hybrid (Table 1), however, readily allow
lor its distinction.
Material for chromosomal analyses was collected
from the type locality. Meiotic behavior of chro¬
mosomes was observed following the methods of
Haufler et al. (1985). An exact meiotic chromosome
count could not be obtained because of the limited
410
Novon
Table 1 . Comparison
pinnatifidus.
of Adenophorus x carsonii
with its putative parents, A.
hymenophylloides and A. tri-
Character
A. hymenophylloides
A. x carsonii
A. tripinnatifidus
Rhizome
Short
Short-creeping
Long-creeping
Rhizome scales
Sublinear
Long-lanceolate
Narrowly deltoid
Leaf length (cm)
2-15
12-35
8 50
Leaf habit
Pendulous
Erect when young,
becoming pendulous
Erect, arching
Leaf blade
Sub- to deeply
bipinnatifid
Bipinnatifid
Bi- to tripinnatifid
Pinnae length (mm)
2-10
5-15
10-60
Basal, acroscopic pinnule
Simple
Simple to bifid
Simple to pinnatifid,
with up to 10 lobes
amount of material available and due to irregular
pairing at diakinesis. Several of the chromosomes
appeared to he univalents, and the total number of
meiotic figures was greater than would be expected
in a normally pairing diploid in this genus (i.e., 37
pairs) anil fewer than would he expected in a triploid
or higher polyploid. Adenophorus x carsonii there¬
fore appears to he an infertile diploid.
Preliminary isozyme data (following Ranker et
ah, 1989) were obtained from the limited amount
of material collected (i.e., four individuals; Ranker,
unpublished data). At 10 of the 14 loci scored, the
two parental taxa and their putative hybrid were
invariant for the same allele. At the remaining loci,
hybrid individuals were heterozygous and shared
alleles with at least one parent. At the latter group
of loci, however, parental taxa were highly variable
(Ranker, 1992), and alleles present in the hybrid
were not always detected in the individual parental
plants examined. A more detailed analysis is nec¬
essary, therefore, to attempt to sample more of the
allelic variability present in the potential parental
taxa. Each of the four hybrids sampled possessed a
unique genotype at each of several loci, indicating
that separate hybridization events were responsible
for the origin of each plant.
Acknowledgments. I thank Hampton Carson, Bill
Mull, and Genie frapp for accompanying me in the
field, Chrissen Gemmill, Ann Hambleton, and Kieu
Iran for laboratory assistance, William A. Weber
for assistance with the manuscript, and Karolyn
Harrow for providing the illustrations. This work
was supported by the National Science f oundation
(BSR-9096282), the University of Colorado, and
the University of Hawaii. Permission to collect spec¬
imens was generously granted by the Natural Area
Reserves System Commission, Stale of Hawaii.
Literature Cited
Bishop, L. E. 1974. Revision of the genus Adenophorus
(Grammitidaceae). Brittonia 26: 217-240.
Hauffer, C. H., M. D. Windham, I). M. Britton & S. J.
Robinson. 1985. Triploidy and its evolutionary sig¬
nificance in Cystopteris protrusa. Canad. J. Bot. 63:
1855-1863.
Ranker, U A. 1992. Genetic diversity of endemic Ha¬
waiian epiphytic ferns: Implications for conservation.
Selbyana. 13: (in press).
-, C. H. Hauffer, P. S. Soltis & D. E. Soltis. 1989.
Genetic evidence for allopolyploidy in the neotropical
fern Hemionitis pinnatifida (Adiantaceae) and the
reconstruction of an ancestral genome. Syst. Bot. 14:
439-447.
Additions to Aequatorium and Gynoxys (Asteraceae: Senecioneae) in
Bolivia, Ecuador, and Peru
Harold Robinson and Jose Cuatrecasas
Department of Botany, National Museum of Natural History, Smithsonian Institution, Wash¬
ington, D.C. 20560, U.S.A.
ABSTRACT. Gynoxys carpishensis, G. rima-
chiana, and Senecio tuestae are transferred to Ae-
quatorium, Gynoxys regis is transferred to Para-
gynoxys, and Aequatorium cajamarcense, A.
juninense, A. kingii, A. tovarii , Gynoxys chin-
gualensis, G. jaramilloi, and G. hutchisonii are
described as new.
Specimens submitted for identification over the
years include the following four undescribed species
of Aequatorium B. Nord. and three undescribed
species of Gynoxys Cass, from Bolivia, Ecuador,
and Peru. Two Peruvian species are transferred
from Gynoxys to Aequatorium, one Peruvian spe¬
cies is transferred from Senecio to Aequatorium ,
and an Ecuadorian species is transferred from Gy¬
noxys to Paragynoxys (Cuatrec.) Cuatrec. These
are in addition to results of many separate papers
providing recent descriptions and additions in Ae¬
quatorium (Nordenstam, 1978; Diaz-Piedrahita &
Cuatrecasas, 1990; Jeffrey, 1992) and Gynoxys
(Sagastegui & Dillon, 1985; Dillon & Sagastegui,
1988; Funk & Robinson, 1989; Herrera, A., 1980;
Robinson & Cuatrecasas, 1984; Sagastegui & Tel¬
lez-Alvarado. 1987).
Aequatorium cajamarcense H. Robinson & J.
Cuatrecasas, sp. nov. TYPE: Peru. Cajamarca;
Prov. Cutervo, Distrito San Andres de Cutervo,
Parque Nacional de Cutervo, caserio ‘"Pajon-
al,” camino hacia Jaen (Chorro bianco), 2,600
m, 10 ago. 1987, Diaz X Osores 2585 (ho-
lotype, US; isotype, MO).
A speciebus ceteris hujus generis in foliis ellipticis abax-
ialiter subglabris distinctissimum.
Tree 18 m tall; internodes of young branches
0.8-30.0 cm long, glabrous or glabrescent with
minute hairs near nodes, surface subcarnose, wrin¬
kled when dry. Leaves opposite, petioles 1.0-2.5
cm long; blades subcoriaceous, elliptical, 6 12 cm
long, 2 4 cm wide, base obtuse to shortly acute,
margins remotely, minutely denticulate, apex acute
to shortly and slightly acuminate, upper surface
glabrous except for sparse, minute, granular hairs
along veins, lower surface subglabrous with sparse
granular hairs along veins and over surface, the
hairs with short narrow bases, subglobular, with 1-
3 short points; venation pinnate, with 8-11 veins
on each side. Inflorescence terminal on branches,
broadly corymbose with spreading, densely corym¬
bose branches, ca. 6 cm high and 1 1 cm wide;
peduncles 1-3 mm long, with dense, minute, gran¬
ular pubescence. Heads ca. 6 mm high; calyculus
with an occasional narrowly lanceolate bract; in-
volucral bracts ca. 8, elliptical or oblong, 2.0-2.3
mm long, 1.0 1.2 mm wide, apices rounded, sub-
glabrous outside, with sparse granular pubescence
on median band. Rays ca. 3; corolla yellow, glabrous,
tube ca. 1.5 mm long, limb elliptical, ca. 4 mm
long, 1.5 mm wide. Disk florets ca. 5; corolla yellow,
ca. 5.5 mm long, glabrous, tube ca. 2 mm long,
throat narrowly campanulate, ca. 1 mm long, lobes
ca. 2.5 mm long; anther collar ca. 0.35 mm long,
anther thecae ca. 1.5 mm long, apical appendage
ca. 0.4 mm long, 0.25 mm wide; tips of style branch¬
es rounded, with slightly longer papillae at the apex.
Achenes ca. 1.8 mm long, subglabrous, with small,
scattered, rounded, thin-walled, subsessile glands that
lack evident septa; pappus ca. 4 mm long, with
bristles in 1 crowded series, apices slightly but dis¬
tinctly broadened. Pollen grains 37 40 jitm diam.
The genus Aequatorium was established by Nor-
denstam in 1978 based on two species of Seneci¬
oneae distinguished from Senecio by stellate hairs,
elongate disk corolla lobes, cylindrical anther collars,
and continuous but centrally depressed stigmatic
surfaces on the style branches. The species had some
resemblance to the genus Gynoxys in spite of having
alternate leaves and blunt style tips. Still, some sub¬
sequent additions to the Aequatorium (Diaz-Pied¬
rahita & Cuatrecasas, 1990; Jeffrey, 1992) have
opposite or subopposite leaves and were originally
described in Gynoxys. The new species is one of
the members of the genus with opposite leaves, but
it is most distinct in the very sparse pubescence on
the lower leaf surfaces. The hairs are not truly
stellate, but they are globular with 1 -3 short points.
Only A. rimachiana has pubescence on the abaxial
Novon 2: 411-416. 1992.
412
Novon
leaf surface that is sparse, but that pubescence is
still much denser and more stellate, and the leaves
of that species are alternate with larger blades. The
new species is described as a tree 18 m tall, and as
such may be the largest member of this shrubby
and arborescent genus.
Aequatoriuni earpishense (Cuatrec.) 11. Robin-
son & J. Cuatrecasas, comb. nov. Basionym:
Gvnoxys carpishensis Cuatrec., Brittonia 12:
185. I960.
This species, lrom Carpish Pass in Huanuco, ap¬
pears to be closest to A. stellato-pilosum (Greenm.
ex Greenm. & Cuatrec.) Jeffrey from the Villca-
bamba region of Cuzco. The leaf blades ol A. ear-
pishen.se seem more obviously shallowly dentate with
thicker and more persistent stellate pubescence on
the undersurface.
Aequatorium juninense II. Robinson & ,|. Cua¬
trecasas, sp. nov. TYPE: Peru. Junin: Carpa-
pata, above Uuacapistana, 2,700-3,200 m, 7
June 1929, Killip & Smith 24431 (holotype,
US).
In caulibus sparse et distincte hirsutis et in latninis
foliorum ellipticis et integris distinctissimum.
Shrubs 2 4 m tall, laxly branched; stems slightly
deflected at the upper nodes, strongly ribbed, cov¬
ered by dense yellowish tomentum ol stellate hairs,
sparsely hirsute above with long spreading hairs.
Leaves alternate, petioles 2-3 cm long, surface sim¬
ilar to stems; blades elliptical, 7 14 cm long, 3.0-
6.5 cm wide, base rounded, margins entire, apex
shortly acute, upper surface dark green, with sparse,
evanescent pubescence of stellate hairs, lower sur¬
face with loose tomentum of stellate, interlocking
hairs only partially hiding the leaf surface; venation
pinnate with ca. 10 spreading secondary veins on
each side. Inflorescence terminal on branches,
broadly corymbose, 11-14 cm high and 12-20 cm
wide, with rather spreading branches; peduncles 2
8 mm long, rather densely stellate pubescent. Heads
9 10 mm high; subinvolucral and calyeular bracts
I -2 mm long; involucral bracts ca. 8, 4-6 mm long,
1.5-2.0 mm wide, apices obtuse, glabrous outside.
Ray florets 2 or 3, corolla yellow, glabrous, tube
ca. 5 mm long, limb elliptical, ca. 10 mm long and
2.5 mm wide. Disk florets 7-9; corolla yellow, gla¬
brous, tube ca. 3 mm long, throat narrowly cam-
panulate, ca. 3 mm long, lobes narrowly triangular,
ca. 1.8 mm long; anther collars ca. 0.5 mm long,
thecae ca. 1.7 mm long, apical appendage ca. 0.6
mm long and 0.25 mm wide; style tips rounded,
with short pencil ol hairs. Achenes ca. 3.5 mm long.
glabrous; pappus 5.0-5.5 mm long, apices not
broadened. Pollen grains 35-37 jum diam.
Paratype. Peru. JUNIN: Carpapata, above Huacapis-
tana, 2,700-3,200 m, 7 June 1929, Killip & Smith
24402 (US).
The type and paratype specimens have the same
basic label citing the habitat as edge of forest.
The new species seems closely related to Aequa-
torium repandum (Wedd.) Jeffrey in the general
aspect, the number of florets in the head, and the
comparatively short triangular lobes of the corolla,
shorter than the corolla throat. Aequatorium jun¬
inense differs by the generally more elliptical leaf
blades, and the less dense stellate pubescence on
the undersurfaces not completely hiding the leaf
surface. The younger stems show some erect spread¬
ing hairs similar to those of A. kingii described
below, but the number ol hairs is much less and
they are not as obvious a key character. The pappus
bristles are like those of A. kingii in the lack of
broadened apices.
Aequatorium kingii II. Robinson & J. Cuatre-
casas, sp. nov. TYPE: Bolivia. Cochabamba:
15 km from Colomi, on the road to Tunari,
10,600 ft., 7 Eeb. 1978, King & Bishop 7080
(holotype, US).
In caulibus grosse et dense hirsutis et in laminis foliorum
oblongo-ovatis et in capitulis majoribus distinctissimum.
Frees or large shrubs 3 m tall; moderately branch¬
ing; stems strongly ribbed, sulcate, densely covered
with appressed tomentum of stellate hairs, also
densely hirsute with stiff, spreading, yellowish hairs.
Leaves alternate, petioles 10 25 mm long, surface
similar to stems, with or without large, erect hairs;
blades ovate-oblong, mostly 5-10 cm long and 2.5-
5.5 cm wide, base shallowly but distinctly cordate,
margins shallowly to strongly dentate, apex shortly
acute, upper surface dark green, smooth, veinlets
rarely slightly prominulous, subglabrous, with scat¬
tered, minute, evanescent, stellate hairs, lower sur¬
face with large, yellow, stellate, interlocking hairs,
with or without denser underlayer of minute, whitish
stellate hairs completely hiding surface; veins pin¬
nate with 5-6 spreading secondary veins on each
side which are more crowded and more spreading
near base. Inflorescence terminal on branches,
broadly corymbose with ascending branches, 6-9
cm high, 9-14 cm wide; peduncles 2-12 mm long,
densely covered with grayish, stellate hairs. Heads
7-10 mm high; subinvolucral and few calyeular
bracts linear, 6 9 mm long, ca. 1 mm wide, glabrous
or with few stellate hairs; involucral bracts 8, oblong,
7-10 mm long, 1.5-2.5 mm wide, narrowed above
Volume 2, Number 4
1992
Robinson & Cuatrecasas
Aequatorium and Gynoxys
413
with apices obtuse, glabrous outside. Ray florets 4
or 5; corolla yellow, glabrous, tube 6 mm long, limb
elliptical, ca. 10 mm long and 3 mm wide. Disk
florets 10-14; corolla yellow, glabrous, tube ca. 3.5
mm long, throat narrowly campanulate, 3.5-4.0
mm long, lobes narrowly triangular, ca. 1.8 mm
long, 0.8 mm wide at base; anther collar ca. 0.5
mm long, thecae ca. 1.8 mm long, apical appendage
0.6-0.7 mm long, ca. 0.22 mm wide; tips of style
branches rounded, with stout apical pencil of hairs.
Achenes submature, ca. 2.5 mm long, glabrous;
pappus ca. 7 mm long, in 2 series, apices not broad¬
ened. Pollen grains ca. 37 /um diam.
Paratypes. BOLIVIA. COCHABAMBA: 26 km from Com-
arapa, on road to Cochabamba, 8,500 ft., 5 Feb. 1978,
King & Bishop 7650 (US); Prov. Carrasco, 5 km al NE
de Monte Punco por el camino a Sihuenca, 17°34'S
65°15'W, 2,700 m, 10 Mar. 1988, Solomon & Nee
18067 (MO, US).
Aequatorium kingii seems most similar to A.
fabrisii (Cabrera) Jeffrey of northern Argentina in
the short bases of its leaf blades, ihe ascending
branches of its inflorescences, and the 4 to 5 ray
florets and 10 or more disk florets in the head. The
new species is most distinct in the densely and stiffly
hirsute condition of the stems, a feature seen to
some extent elsewhere in the genus only in A. jun-
inense described above. I he three specimens of the
present species all show leaf blades with slightly but
broadly cordate bases unlike the narrowly cordate
bases sometimes seen in A. repanda (Wedd.) Jef¬
frey, the obtuse to subtruncate bases illustrated for
A. fabrisii (Cabrera, 1978), or the rounded bases
seen in A. juninense. I he type specimen of the new
species lacks the dense layer of smaller hairs on the
lower leaf surface that completely hides the surface,
but such hairs are present in both paratypes.
Aequatorium rimaehianuni (Cuatrec.) 11. Rob¬
inson & J. Cuatrecasas, comb. nov. Basionym:
Gynoxys rimachiana Cuatrec., Phytologia 52:
164. 1982.
Sagastegui & Dillon (1985) noted the presence
of three arborescent species of Gynoxys in the Car-
pish Pass area, including this and G. carpishensis.
The third species mentioned, Gynoxys congestiflora
Sagastegui & Dillon, has strictly opposite leaves,
unbranched, contorted hairs, and more pointed style
tips, and it is not an Aequatorium.
Aequatorium tovarii H. Robinson & J. Cuatre¬
casas, sp. nov. TYPE: Peru. Huancavelica:
Prov. Tayacaja, Arriba de Marcavalle, entre
Huachocolpa y Tintay, 3,300 m, 21 Apr. 1964,
Tovar 4781 (holotype, US).
In laminis foliorum oblanceolatis et in lobis corollae
anguste oblongis distinctissimum.
Shrubs to 2 m high, moderately branched; stems
terete, weakly ribbed, completely covered with dense,
yellowish, appressed tomentum of stellate hairs.
Leaves alternate, petioles 7-15 mm long; blades
oblanceolate, mostly 8-13 cm long and 2.3-4.0 cm
wide, widest near distal third or fourth, base narrowly
rounded, margins subentire, with minute, remote
denticulations, apex obtuse or shortly acute, upper
surface dark green, with scarcely prominulous vein-
lets, subglabrous with few evanescent hairs, with
larger, unbranched hairs on midvein; lower surface
with dense tomentum of stellate hairs; venation pin¬
nate with ca. 8 ascending secondary veins on each
side. Inflorescences terminal on branches, rounded
corymbose, with erect-spreading lower branches from
axils of distal leaves; peduncles 1-7 mm long, dense¬
ly stellate pubescent. Heads 9-10 mm high; calyc-
ular bracts ca. 5, linear, 1-2 mm long; involucral
bracts ca. 8, oblong, ca. 6 mm long and 1.5-2.0
mm wide, apices obtuse, partially obscured outside
by stellate hairs. Ray florets usually 3; corolla yellow,
glabrous, tube ca. 5 mm long, limb elliptical, ca. 6
mm long and 2 mm wide. Disk florets ca. 7; corolla
yellow, 8.0-8.3 mm long, tube 3.5 4.0 mm long,
throat narrowly campanulate, ca. 2 mm long, lobes
narrowly oblong, ca. 2.3 mm long; anther collar ca.
0.4 mm long; thecae ca. 1.7 mm long; apical ap¬
pendage ca. 0.7 mm long and 0.27 mm wide; style
tips rounded, with short apical pencil of hairs. Achenes
submature, ca. 1.8 mm long, glabrous; pappus 6-
7 mm long, with bristles in 2 series, apices slightly
but distinctly broadened. Pollen grains ca. 33 pm
diam.
Paratype. PERU. HUANCAVELICA: Prov. Tayacaja, Aba-
jo de San Antonio, distrito de Surcubamba, 2,200 m, 27
abr. 1963, Tovar 4252 (US).
The specimens are cited from “bosque con pa-
jonal” and “bosque subxerofilo.”
Aequatorium tovarii has the general appearance
of A. repandum from Bolivia and southeasternmost
Peru, but in the former the leaves are distinctly
wider in the distal half, the secondary veins are
more ascending, and the disk corolla lobes are nar¬
rowly oblong and longer than the corolla throat. The
lobes are unlike the narrowly triangular lobes of the
A. repandum group, and are like those of the more
northern species of Aequatorium.
Aequatorium tuestae (Cuatrec.) H. Robinson &
J. Cuatrecasas, comb. nov. Basionym: Senecio
tuestae Cuatrec., Fieldiana 27: 46. 1951.
414
Novon
This species seems somewhat similar to large-
leaved A. carpishense and A. rimachianum, hut it
lacks cordate bases on the leaf blades.
Gynoxys chingualensis H. Robinson & J. Cua-
trecasas, sp. nov. TYPE: Ecuador. Sucumbios:
Paramo Mirador SW of Playon <le San Fran¬
cisco, S of Rio Chingual headwaters, elev.
3,400 3,600 m, king, Peterson & Judziew-
iez 10131 (holotype, US; isotypes, MO, K).
In foliis late oblongo-ovatis subsessilibus supra glabris
distinctissimis.
Shrubs 3 4 m tall, moderately branching; stems
densely brownish velutinous. Leaves opposite, pet¬
ioles 1-2 mm long; leaf blades broadly oblong-ovate,
4.5-7.0 cm long, 3.0-4.3 cm wide, base shallowly
cordate, margins entire with remote, mucronate den¬
tations reflexed against lower surface, apices round¬
ed, upper surface glabrous, with a reticulum ol
prominulous veinlets, lower surface with dense brown
tomentum, with long, unbranched, sinuous hairs and
some T-shaped hairs of similar diameter intermixed;
venation pinnate, with 6 8 secondary veins on each
side, widely forking about halfway to margin. Inflo¬
rescence terminal on branches, broadly corymbose
wi th d ensely corymbose branches, ca. 9 cm high
and 14 cm wide; peduncles 5-10 mm long, densely
brownish tomentose. Meads ca. 12 mm high; below
the heads 2-3 linear, tomentose bracts ca. 8 mm
long; involucral bracts ca. 8, oblong, 8-9 mm long,
1.5 3.0 mm wide, obtuse to shortly acute, densely
tomentose on exposed outer surfaces. Ray florets
ca. 5; corolla yellow, tube 4 mm long, glabrous,
limb ca. 11 mm long and 3 mm wide, pilosulous in
basal sinus, otherwise glabrous. Disk florets ca. 12;
corolla yellow, ca. 10 mm long, glabrous, tube 4.0-
4.5 mm long, throat narrowly campanulate, ca. 3.5
mm long, lobes ca. 2 mm long, 0.8 mm wide at
base; anther collar ca. 0.8 mm long, cylindrical;
thecae ca. 2 mm long; apical appendage ovate-
oblong, 0.8 mm long and 0.3 mm wide; style apex
twice as long as wide to bases of apical hairs. Sub¬
mature achenes ca. 2.5 mm long, glabrous; pappus
ca. 8 mm long, bristles in 2 crowded series, apices
distinctly enlarged with many rows of crowded cells.
The specimen was collected from a paramo with
Espeletia and Calamagrostis.
The subsessile leaves of Gynoxys chingualensis
resemble those of G. hirsuta Wedd. from Depto.
Cundinamarca, Colombia, but the latter has a more
diffuse inflorescence with fewer heads on more nu¬
merous leafy branches, a coarser grayish pubes¬
cence, and smooth upper leaf surfaces with eva¬
nescent hairs. The new species resembles G.
tornenlosissirna Cuatrec. of Amazonas, Peru, in hab¬
it, but the latter has more angled stems, longer
petioles to ca. 1 cm long, and larger leaf blades with
auriculate bases and smooth upper surfaces thinly
velutinous with yellowish hairs.
Gynoxys hutchisonii H. Robinson & ,1. Cuatre-
casas, sp. nov. TYPE: Peru. Piura: above
lluancabamba, road to Piura, 3,000 m, 10
Oct. 1957, P. (.. Hutchison 1609 (holotype,
US).
In foliis parvis et in capitulis eradiatis distincta.
Shrubs 0.5 1.5 m tall, highly branched; stems
terete, scarcely striated, densely pale yellowish to¬
mentose. Leaves opposite, petioles 5-10 mm long;
blades oblong, 1.5 3.0 cm long, 0.8-1.5 cm wide,
base rounded to subtruncate, margins entire, apex
rounded to slightly obtuse, upper surface glabrous
except on midvein, essentially smooth, lower surface
densely tomentose with pale yellowish, unbranched,
sinuous hairs; venation pinnate, with 4 or 5 sec¬
ondary veins on each side sometimes forked near
margin. Inflorescences clustered in pyramidal groups
on leafy branches and branchlets, branches rather
densely to laxly corymbose; peduncles 2-10 mm
long, densely tomentose. Heads 1.0-1.2 cm high;
calyculus bulging without obvious bracts; involucral
bracts ca. 8, oblong, ca. 5 mm long and 1.5 mm
wide, apices shortly acute, densely tomentose on
exposed outer surfaces. Ray florets lacking. Disk
florets 5-12; corolla pale yellow, 7-8 mm long,
glabrous, tube 2.0 2.5 mm long, throat narrowly
campanulate, 3.0-3.5 mm long, lobes ca. 1.8 mm
long, ca. 0.7 mm wide at base; anther collar ca.
0.6 mm long, thecae ca. 1.2 mm long, apical ap¬
pendage ca. 0.6 mm long and 0.25 mm wide; apex
of style twice as long as wide to base of apical hairs.
Achenes 2.5-3.0 mm long, glabrous; pappus in 2
crowded series, 6- 8 mm long, distinctly broadened
at tips. Pollen grains ca. 37 jam diam.
Paratype. PKRU. AMAZONAS: Prov. Chachapoyas: Cer-
ros de Caila-Calla, near km 403 407 of Balsas Leime-
bamba road, 3,400-3,550 in, 18 Aug. 1962, J. J. If ur
duck 1702 (US).
Gynoxys hutchisonii is distinct in the combina¬
tion of the small leaves and the rayless heads. Such
small leaves are not found in other Peruvian species
(Herrera, A., 1980; Sagastegui & Tellez-Alvarado,
1987; Dillon & Sagastegui, 1988), while the many
small-leaved species of Ecuador all have rays. The
rayless condition might occur occasionally in nor¬
mally radiate species of Gynoxys, but a number of
species are characteristically rayless. One group
without rays and with glabrous involucral bracts is
Volume 2, Number 4
1992
Robinson & Cuatrecasas
Aequatorium and Gynoxys
415
represented in Peru hy G. soukupii Cuatrec. Also
rayless is the Peruvian G. longifolia Wedd., with
its unusually long and slender leaves and somewhat
axillary inflorescences.
The two specimens of the new species differ some¬
what in appearance. The paratype is more branched
and has more flexuous stems. The paratype also has
generally longer peduncles and has 5-10 florets in
the heads. The holotype has peduncles under 5 mm
long and mostly 10-12 florets in the heads.
Gynoxys jaramilloi If. Robinson & .1. Cuatre¬
casas, sp. nov. TYPE: Ecuador. Loja: Loma
del Oro, 2,800-3,200 m, without date, Jar-
arnillo, Zak cY 1 alencia 8799 (holotype, US;
isotype, QUA).
In floribus femineis redactis et in pilis crassis contortis
et in pilis pagina abaxiale foliorum valde biformibus dis-
tinctissima.
Shrubs 5 m high, with many short branches; stems
terete with low ribs, bearing numerous, coarse, con¬
torted, retrorse hairs, without underlayer of hairs,
stem surface visible between hairs. Leaves opposite,
petioles 0.5 1.0 cm long, slender, with erect hairs;
blades oblong, mostly 3.0-4.5 cm long, 1.2-2.2 cm
wide, base subtruncate to rounded, margins slightly
sinuous, with minute, remote, mucronate denticu-
lations reflexed against lower surface, apex rounded
to obtuse, upper surface with secondary veins and
some tertiary veinlets slightly prominulous, subglab-
rous, with sparse, evanescent, coarse, contorted hairs,
lower surface densely yellowish tomentose, with
coarse, contorted hairs underlain with fine tomentum
ol small T-shaped hairs; venation pinnate, with 5
6 secondary veins on each side, often forked halfway
to margin. Inflorescence with small groups terminal
on branches and branchlets, pyramidal with cor¬
ymbose branches, 3-4 cm high and 3-5 cm wide;
peduncles 5-13 mm long, densely tomentose with
contorted, brown hairs. Heads ca. 1 cm high; sub-
involucral and calycular bracts few, short, linear,
1.5-2.5 mm long; involucral bracts ca. 8, oblong,
ca. 6 mm long and 1.5-2.0 mm wide, apices obtuse
to shortly acute, densely tomentose outside on ex¬
posed surfaces. Female florets 2-3; corolla yellow,
in form ol small ray with 3 lobes or disciform with
5 lobes, glabrous, tube 4.5-6.0 mm long, limb 1.8-
3.0 mm long, lobes 0.5-1.8 mm long, 0.3 0.4 mm
wide at base; staminodia sometimes present. Disk
florets 10-12; corolla yellow, glabrous, tube ca. 3
mm long, throat narrowly campanulate, ca. 3 mm
long, lobes ca. 1.8 mm long, ca. 0.7 mm wide at
base; anther collar cylindrical, ca. 0.6 mm long,
thecae ca. 1.4 mm long, apical appendage ca. 0.7
mm long and 0.3 mm wide; apex of style 3 times
as long as wide to base of apical hairs. Achenes
immature, ca. 2 mm long, glabrous; pappus ca. 7
mm long, in 2 crowded series, gradually broadened
to tip. Pollen grains ca. 37 /am diam.
The type specimen is said to be from “vegetacion
de bosque andina y paramal con areas de cultivo y
potrero, especies de Weinmannia, Clusia, Podo-
carpus, Diplostephium, Hesperomeles, Clethra."
Gynoxys jaramilloi superficially resembles some
ol the more common members of the genus in south¬
ern Ecuador, such as Gynoxys cuicochensis Cua¬
trec., but the relationship is evidently not close. The
type specimen was first noticed because of the seem¬
ingly rayless heads, some of which are mature enough
to show rays if they were of normal size. A few
lemale florets prove to be present, but they are small
and often radially symmetrical with five lobes. The
one female floret mounted on a microscope slide
had a small raylike limb and small staminodia.
A more interesting distinction of the new species
is in the hairs of the stems and leaves. The stem
hairs are coarse and contorted, and they do not
completely hide the surface of the stem. Similar
unbranched, contorted, but smaller hairs are found
in the denser pubescence on the leaf undersurface,
but those surfaces have a dense underlayer of much
smaller, contorted, T-shaped hairs. Such hairs have
not been noticed in other species of Gynoxys, and
most species obviously have hairs completely cov¬
ering the young stems. Most species of Gynoxys
have not had the hair types of their abaxial leaf
surfaces examined in such detail, but unbranched
and T-shaped hairs of similar size to each other are
intermixed in G. chingualensis described above.
Paragynoxys regis (H. Robins. & Cuatrec.) II.
Robinson & J. Cuatrecasas, comb. nov. Bas-
ionym: Gynoxys regis H. Robins. & Cuatrec.,
Phytologia 56: 370. 1984.
This species is unlike most Gynoxys, and like Par¬
agynoxys in the lack of rays in the heads, the blunt
tips of the style branches, and the long corolla lobes
separated to the base of the throat. The species is
unusual in Paragynoxys by the opposite leaves and
the involucre of 5 8 bracts. Its closest relationship
seems to be with the similarly small-leaved, Peruvian
Gynoxys lopezii Dillon & Sagastegui (1988), which
was transferred to Paragynoxys by Cuatrecasas
(1990: 314).
Literature Cited
Cabrera, A. L. 1978. Flora de la Provincia de Jujuy,
Republica Argentina, parte X — Compositae. Colec-
cion del INTA. 13: 1-726.
Cuatrecasas, J. 1990. Miscellaneous notes on neotrop-
416
Novon
ical flora, XIX. Combinations in Senecioneae, Com-
positae. Phytologia 69: 313-315.
Diaz-Piedrahita, S. & J. Cuatrecasas. 1990. El genero
Aequatorium Nord. (Senecioneae-Asteraceae) en
Colombia. Revista Academia Colombiana Ciencias
Exact. Fisicas y Naturales 17: 659-666.
Dillon, M. O. & A. Sagastegui A. 1988. Additions to
South American Senecioneae (Asteraceae). Brittonia
40: 221-228.
Funk, V. A. & H. Robinson. 1989. A new species of
Gynoxys (Asteraceae: Senecioneae) from northern
Peru. Revista Academia Colombiana Ciencias Exact.
Fisicas y Naturales 17: 243-245.
Herrera, Alarcon de Loja B. 1980. Revision de las
especies peruanas del genero Gynoxys. Bol. Soc.
Peruana Bot. 8(1, 2): 3-74.
Jeffrey, C. 1992. The tribe Senecioneae (Compositae)
in the Mascarene Islands with an annotated world
checklist of the genera of the tribe. Kew Bull. 47:
49-109.
Nordenstam, B. 1978. Taxonomic studies in the tribe
Senecioneae (Compositae). Opera Bot. 44: 1-83.
Robinson, H. & J. Cuatrecasas. 1984. Observations of
the genus Gynoxys in Ecuador (Senecioneae: Aster¬
aceae). Phytologia 56: 368-375.
Sagastegui-A., A. & M. 0. Dillon. 1985. Four new
species of Asteraceae from Peru. Brittonia 37: 6-
13.
-& C. Tellez-Alvarado. 1987. A new species of
Gynoxys (Asteraceae: Senecioneae) from northern
Peru. Brittonia 39: 432-435.
A New Species of Petrea (Verbenaceae) from Brazilian Amazonia
Ricardo M. Rueda
Departamento de Biologia, Universidad Nacional Autonoma de Nicaragua, Leon, Nicaragua;
present address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.
ABSTRACT. During preparation of a revision of the
genus Petrea (Verbenaceae), it has become evident
that six specimens from white sands in Roraima,
Brazil, represent a new species that is herein de¬
scribed and given the name Petrea campinae. The
new species is most closely related to Petrea brev¬
icalyx from the same area.
Petrea campinae Rueda, sp. nov. TYPE: Brazil.
Roraima: along the road from Manaus to Car-
acarai, km 350, left side, 00°06'N, 60°40'W,
12 Feb. 1979 (fl). It . Rodrigues, el at. 10IOO
(holotype, MG; isotypes, COL, INPA). fig¬
ure 1.
Frutex 1.5-5 m altus, ramulis puberulis. Folia breviter
petiolata, laminis subtus foveatis, ellipticis, 5-10 cm lon-
gis, 1.5-6 cm latis, coriaceis, asperatis, apiculatis, venis
secundariis 6 10. Inflorescentiae 10-22 cm longae, 2-
5 cm latae; bracteis 2-4 mm longis, pedicellis 6-10 mm
longis. Flos lobis calycinis 4-8 mm longis rotundatis,
corolla hypocrateriformi, azurea, puberulo-glandulosa, 1.2-
1.5 cm longa, lobis 3-6 mm longis, inaequalibus.
Shrub 1.5-5 m tall; branches tetragonal or sub-
terete, minutely pubescent, pith well developed.
Leaves decussate; petiole 3 8 mm long; blade ellip¬
tic, apiculate or truncate at apex, obtuse at base,
5-10 cm long, 1.5-6 cm wide, coriaceous, asper-
ous, glabrous above, intricately pitted beneath with
abundant short hairs in the pits, the midrib elevated
above, prominently elevated beneath, the secondary
veins brochidodromous, 6 10 on each side forming
an angle of 60° with the midrib, impressed above,
elevated beneath, ascending, with strongly defined
veinlets delimiting the [fits, the margins entire to
slightly undulate. Inflorescences racemiforrn, axil¬
lary or terminal, many-flowered, 10-22 cm long,
2-5 cm wide; bracts setaceous to foliaceous, 2 4
mm long; pedicels 6-10 mm long; rat his, bracts,
and pedicels minutely pubescent. Flowers with calyx
glabrous, the lobes 5, unequal, rounded, 4-8 mm
long; calicinal crest 5-cleft; corolla hypocrateriform,
straight or curved, blue, glandular-pubescent, 1.2-
1.5 cm long, the limb lobed, 5-8 mm wide, the
lobes 5, ovate, 3-6 mm long, unequal; stamens 4,
attached to corolla throat, the filaments stout, about
1.5 mm long, the anthers oblong; ovary globose.
the style included, 2-3 mm long, the stigma capitate;
basal disk about 1 mm long. Fruit not seen.
Paratypes. Brazil. roraima: along BR 174 between
Manaus and Caracarai, 6 km S of equator, in a campina,
00°04'S, 60°40’W, 17 June 1985 (fl), Cordeiro et al.
76 (INPA, SP); Road Manaus-Caracarai, between km
345 and 360, in a campina, 15 Mar. 1984 (fl), J. L. dos
Santos, & Coelho 690 (INPA).
The Cordeiro et al. 76 collection is from a shrub
5 m tall and is anomalous in having one pair of
blades fused on a single petiole. This same specimen
also has very anomalous flowers. Some of the flowers
have 7 sepals and petals and a 7-parted calicinal
crest; there is also a fifth stamen with deformed
anthers. This is the only known case in Petrea L.
where the calyx and corolla are not strictly 5-merous.
Petrea campinae resembles P. brevicalyx Ducke
in having leaves elliptic, intricately pitted beneath,
and calyx with short rounded lobes; it differs by
having unequal calyx lobes, twice as long as those
of the latter species, and it is erect, whereas P.
brevicalyx is a liana.
All the collections of P. campinae come from
sandy campinas in Roraima, from which vegetation
formation the specific epithet is derived. Endemism
in these white sand areas is common, as Prance
(1982) noted for several species of Chrysobalana-
ceae. Another typical campina phenomenon illus¬
trated by this new species is a shrubby habit in a
taxon whose related forest species are lianas. This
also occurs, for example, in the Bignoniaceae in the
same area in the cases of Distictella monophylla
Sandw. and D. laevis (Sandw.) A. Gentry (Gentry,
pers. comm.).
Acknowledgments. Supported by National Sci¬
ence Foundation grant INT-9024094. I thank the
curators of MG and SP for the loan of specimens,
John Myers for the skillful illustration, and Bruce
Holst, Alwyn H. Gentry, and Roy Gereau for com¬
ments on the manuscript.
Literature Cited
Prance, G. T. 1982. Forest refuges: evidence from
woody angiosperms. Pp. 137-156 in G. T. Prance
(editor), Biological Diversification in the Tropics. Co¬
lumbia Univ. Press, New York.
Novon 2: 417-418. 1992.
418
Novon
Figure 1. Petrea campinae Rueda (R". Rodrigues et al. 10100). —A. Habit. —B. Close up of abaxial leaf surface,
showing pits. —C. Flower. —L). Flower, cut longitudinally and laid open.
A Review of the Fern Genus Micro poly podium (Grarnmitidaceae)
Alan R. Smith
University Herbarium, University of California, Berkeley, California 94720, U.S.A.
ABSTRACT. Micropolypodium differs from other
genera of Grarnmitidaceae in having segments or
pinnae each with a single unbranched or 1-forked
vein, erect rhizomes, linear fronds generally less than
1 cm wide, unisoriate segments, and prominent hy-
dathodes. About 30 species are known, most in
tropical America and a few in eastern Asia. New
combinations are made for the known species. A list
of excluded names in Xiphopteris, in which most
of the species have previously been placed, is pro¬
vided.
Recent studies of grammitid ferns (Bishop, 1988,
1989; Smith et ah, 1991; Smith & Moran, 1992;
Parris, 1984, 1990) allude to the patent unnatu¬
ralness of traditional classifications ol the Grarn-
mitidaceae, which have been based in large part on
blade dissection. In an effort to arrive at a more
natural classification ol the family, studies initiated
by Bishop and continued by Smith reveal that the
genus Micro poly podium , established for a few Asian
species, should be expanded to include many tropical
American species heretofore placed variously in
Grammitis (Morton, 1967), Xiphopteris (Copeland,
1952), and formerly Polypodium (Maxon, 1916).
In lieu of a monographic treatment, which is needed
especially for the South American species, and so
that combinations are available lor the forthcoming
Flora Mesoamericana, I here provide a conspectus
ol Micropolypodium and make transfers of those
species generally accepted in modern floristic ac¬
counts for Latin America (Proctor, 1977, 1985,
1989; Mickel & Beitel, 1988; Lellinger, 1989;
Smith, 1981; Stolze, 1981). Following the transfers
to Micropoly podium, 1 list all species that have
been included within Xiphopteris and give their
current taxonomic disposition, if known.
Micropolypodium Hayata, Bot. Mag. Tokyo 42:
341. 1928. TYPE: Micropolypodium pseu-
dotrichomanoides (Hayata) Hayata, based on
Polypodium pseudotrichomanoides Hayata [=
M. okuboi (Yatabe) Hayata]. Figure 1A.
Plants usually epiphytic, rarely on rocks; rhizome
radially symmetrical, suberect or erect, bearing
golden, orangish, or yellow- to orange-brown, often
lustrous, nonclatbrate or weakly clathrate scales that
are entire, irregularly toothed, sparsely glandular,
or setulose on the margin, setulae concolorous or
often darker than scale body, cells of body often
tumid; fronds monomorphic or nearly so, short-stip-
itate, with petioles 0.2-0.5 mm diam., not articulate
to the rhizome; lamina linear, commonly 3-15(-25)
cm long, usually less than l(-l .2) cm wide, shallowly
to usually deeply pinnatifid, pinnatisect, or barely
pinnate, with usually numerous oblong to debate,
decurrent segments (pinnae), these sometimes with
an acroscopic hump (segments gibbous); segments
planar, basiscopic side not folded over sorus; laminar
tissue chartaceous to subcoriaceous; veins simple or
with only an acroscopic fertile vein branch, lacking
a dark sclerenchymatous sheath except at the very
base of the segment (not extending more than
the segment length); hydathodes present, round to
clavate, usually conspicuous adaxially; setae olten
present on axes and lamina, these maroon to red-
brown; hairs simple or often branched, mostly 0.1
0.5 mm long, multicellular with many short cells,
at times glandular, hyaline with darkened cross¬
walls; sori round, superficial, one per pinna, usually
adjacent to midrib at base of costa or on the ac¬
roscopic veinlet, borne along most of the blade length;
paraphyses absent; sporangia glabrous.
The genus comprises about 30 species Irom
southern Mexico to Bolivia and southern Brazil, and
the Antilles; a few species occur in eastern Asia and
Malesia (Japan, China, Sikkim, Philippines, New
Guinea).
Micropolypodium differs from all other neotrop¬
ical genera of Grarnmitidaceae by having segments
(pinnae) with a single unbranched vein or with only
an acroscopic branch (Figs. 1,2A) anil the segments
often gibbous. Additional important (but not unique)
characteristics are the radially symmetric erect rhi¬
zomes (Hayata 1928a, b, 1929), the often golden
brown nonclathrate rhizome scales, linear fronds
usually less than 10 mm wide and often less than
6 mm wide, unisoriate segments (pinnae), prominent
adaxial hydathodes (figs. II), G, H, 2A), and the
long, dark red-brown unbranched setae. I he type
is from eastern Asia, but by far the greatest diversity
in the genus is in the Neotropics, with species most
numerous and diverse in the West Indies and Central
America.
Novon 2: 419-425. 1992.
420
Novon
Figure 1. Representative species of Micropolypodium, habits and details of laminae, rhizome scales, and indument.
— A. M. okuboi ( Tagawa 8303, UC). —B. M. truncicola (Silverstone-Sopkin el nl. 3605, UC). —C. M. setosum
(Hatschbach 44998, UC). — D. M. caucanum, (Liesner 24841, UC). — E. M. setulosum (Stork 2336, UC). —F.
M. hyalinum (Smith & Beliz 2081, UC). —G. M. trichomanoides (II illiams el nl. 41935, UC]). — H. M. nanum
(Moore et al. 9673, UC).
Thus defined. Micropolypodium seems homo¬
geneous, without any readily apparent subgroups. 1
see no close afTinities to any other neotropical or
African genera of Grammitidaceae. The relationship
ol Micropolypodium to paleotropical genera, cur¬
rently under study by Barbara Parris, is not apparent
either. In the Old World, Micropoly podium may
be most similar to Calymmodon, a genus of about
25 mostly Malesian species but extending from Sri
Lanka to Tahiti. Calymmodon agrees with Micro-
polypodium in having ascending, radially symmet¬
ric rhizomes, often golden brown rhizome scales,
usually narrow laminae, nonarticulate stipes, ab¬
sence ol paraphyses, unbranched veins, adaxial by-
Volume 2, Number 4
1992
Smith
Micropolypodium
421
Figure 2. Micropolypodium and Lellingeria spp., details of venation and hydathodes. A. M. basiattenuata ,
cleared lamina with details showing cellular pattern of hydathodes (Underwood 1452, US). — B. Lellingeria harm
(Jenman) A. R. Smith & R. C. Moran, cleared lamina (Stehle 1473, US). —C. Lellingeria schenckii (Hieron.) A.
R. Smith & R. C. Moran, cleared lamina with detail showing cellular pattern of hydathodes (Brade 6583, US). -
D. Lellingeria myosuroides (Sw.) A. R. Smith & R. C. Moran, cleared lamina (Proctor 4363, US).
dathodes, and a single sorus per segment or pinna.
Calymmodon differs in the usually whitish to pale
red-brown setae on the axes and blades, often winged
stipes. 1 -5-branched hyaline hairs lacking darkened
cross-walls on the hair branches, scales with l(-2)
hyaline seta(e) at the tip, sori often confined to the
distal part of frond, thinner laminar tissue, and fertile
segments with basiscopic side folded upwards to form
a pouch for the sorus (see Parris, 1990, tab. 4000).
The similarities between species of Micropoly¬
podium and some species of Lellingeria (Smith et
al., 1991), particularly the L. limula/myosuroides
group, in blade size and dissection are obvious, but
the striking differences in rhizome scales indicate
that the relationship is not a close one. 1 liese scale
differences were noted many years ago by Maxon
(1916: 542), who divided plants of this general form
into two groups, based upon the differences m rhi¬
zome scales. Scales ot Micropolypodium are usually
golden to golden brown and concolorous; occasion¬
ally the lateral walls are slightly darker than the
surface walls, and the scales might be described as
subclathrate. However, they are never strongly
clathrate to the degree found in Lellingeria limula
or Melpomene moniliformis and their allies. Scale
differences in Grammitidaceae have frequently been
noted and used in descriptions and keys, but their
significance in distinguishing large groups of species
has not been realized until recently.
Another difference between the two genera is in
the setae on the stipe and rachis, which are dark
red-brown and simple in Micropolypodium but usu¬
ally whitish or hyaline, often branched, and generally
much shorter in Lellingeria. Narrow-bladed species
of Lellingeria (L. myosuroides group) usually have
dark sclerenchymatous tissue covering the veins (Fig.
2B D). These red-brown cells extend most ol the
length of the vein to the hydathode and are best
seen in cleared fronds. Species of Micropolypodium
generally lack darkened costae, or the sclerenchyma
of the rachis extends less than 0.5 mm as rudi¬
mentary branches at the base oi the segments (Fig.
2A).
Flydathodes in most species of Micropolypodium
are among the most prominent ol any genus in the
family. The cell pattern on the surface of the hy¬
dathodes, as seen in cleared fronds, is strikingly
dissimlar to the epidermal cell pattern, with many
cells rather elongate (Fig. 2A). Although this char¬
acter has not yet been studied in detail, most other
hydathodous grammitid genera (e.g., Lellingeria)
appear to have the cells of the hydathode more
nearly isodiametric, smaller, and less obviously dif¬
ferent from epidermal cells (Fig. 2C).
The setae in Micropolypodium also appear to
differ from those of many other grammitid genera.
A ring of cells, different from the surrounding epi¬
dermal cells, radiates from the base of the seta in
M. basiattenuata and in several other species of
Micropolypodium examined. This specialized cell
pattern at the base of the setae has not been detected
in preliminary surveys of other grammitid genera.
Only a few chromosome counts have been re¬
ported for species belonging to Micropolypodium.
Walker (1985) counted 2n = 74 for M. taenifolium
(as Grammitis taenifolia) from Irinidad. Silva Ar-
422
Novon
aujo (1976) reported 2 n = ca. 36 for M. trichom-
anoides (as Grammitis trie horn an aides) from near
Manaus, Brazil. The voucher (not seen) is probably
misidentified and from the locality may be M. nan-
urn, which is the only species of Mieropolypodium
known from near Manaus. A base number of x —
37 is known in most other genera of Grammitida-
ceae, e.g., Enterosora , Ceradenia, and Melpomene.
In Lellingeria, base numbers of x = 32 and 33 are
so lar known (see Smith et ah, 1991), and these
numbers suggest an additional difference between
Mie ro poly podium and Lellingeria.
American species
Micropolypodiuni uplielnlcpis (C. Morton) A.
R. Smith, comb. nov. Basionym: Grammitis
aphelolepis C. Morton, Contr. U.S. Natl. Herb.
38: 97. 1967. Xiphopteris aphelolepis (C.
Morton) Pichi-Serm. Distribution: Ecuador.
Micro poly podium basiattenuatum (Jenman) A.
R. Smith, comb. nov. Basionym: Polypodium
basiattenuutum Jenman, Bull. Bot. Dept. Ja¬
maica, n.s. 4: 114. 1897. Grammitis basiat-
tenuata (Jenman) Proctor. Xiphopteris bus-
lattenuata (Jenman) Copel. Distribution:
Jamaica, southern Mexico, Guatemala, Hon¬
duras.
Micropolypodiuni blepharideum (Copel.) A. R.
Smith, comb. nov. Basionym: Polypodium ble¬
pharideum Copel., Univ. Calif. Publ. Bot. 19:
304, pi. 64. 1941. Xiphopteris blepharidea
(Copel.) Copel. Distribution: Peru.
Micropolypodiuni caucanum (Hieron.) A. R.
Smith, comb. nov. Basionym: Polypodium cau¬
canum Hieron., Bot. Jahrb. Syst. 34: 503.
1904. Grammitis caucana (Hieron.) C. Mor¬
ton. Xiphopteris caucana (Hieron.) Copel. Dis¬
tribution: Costa Rica to Ecuador and Venezue¬
la. Figure ID.
Micropolypodiuni cookii (L. Underw. & Max-
on) A. R. Smith, comb. nov. Basionym: Po¬
ly podium cookii L. Underw. & Maxon, Contr.
U.S. Natl. Herb. 17: 408. 1914. Grammitis
cookii (L. Underw. & Maxon) F. Seymour.
Xiphopteris cookii (L. Underw. & Maxon) Co-
pel. Distribution: Guatemala to Costa Rica.
Micropolypodiuni grisebachii (L. Underw. ex
C. Chr.) A. R. Smith, comb. nov. Basionym:
Poly podium grisebachii L. Underw. ex C. Chr.,
Index Filic. 531. 1906, based on Polypodium
exiguum Griseb., FI. Br. W. bid. 701. 1864,
non Howard, 1838. Grammitis grisebachii (L
Underw. ex C. Chr.) Proctor. Xiphopteris gri¬
sebachii (L. Underw. ex C. Chr.) Copel. Dis¬
tribution: Jamaica, Lesser Antilles.
Micropolypodiuni hyalinum (Maxon) A. R.
Smith, comb. nov. Basionym: Polypodium hv-
alinurn Maxon, Contr. U.S. Natl. Herb. 17:
406. 1913. Grammitis hyalina (Maxon) F.
Seymour. Xiphopteris hyalina (Maxon) Copel.
Distribution: Costa Rica, Panama, Ecuador.
Figure 1 F.
Micropolypodiuni knowltoniorum (Hodge) A.
R. Smith, comb. nov. Basionym: Polypodium
knowltoniorum Hodge, Amer. Fern .1. 31: 105,
t. 1, figs. 4-6. 1941. Grammitis knowlto-
niorum (Hodge) Proctor. Xiphopteris knowl¬
toniorum (Hodge) Copel. Distribution: Lesser
Antilles.
Micropolypodiuni liesneri (A. R. Smith) A. R.
Smith, comb. nov. Basionym: Grammitis lies¬
neri A. R. Smith, Ann. Missouri Bot. Card. 77:
257. 1990. Distribution: eastern Venezuela.
Micropolypodiuni nanum (Fee) A. R. Smith,
comb. nov. Basionym: Polypodium nanum Fee,
Ceil. Filic. 238. 1852. Grammitis blanchetii
(C. Chr.) A. R. Smith. Polypodium blanchetii
C. Chr. Xiphopteris nana (Fee) Copel. Distri¬
bution: Venezuela, Cuianas, Colombia, Ama¬
zonian Brazil. Figure 111.
Micropolypodiuni ninibatuni (Jenman) A. B.
Smith, comb. nov. Basionym: Poly podium
nimbatum Jenman, J. Bot. 24: 271. 1886.
Grammitis nimbata (Jenman) Proctor. Xiph¬
opteris nimbata (Jenman) Copel. Distribution:
Cuba, Jamaica, Hispaniola; U.S.A. (disjunct in
Macon Co., North Carolina).
Micropolypodiuni perpusilluni (Maxon) A. R.
Smith, comb. nov. Basionym: Polypodium per-
pusillum Maxon, Contr. U.S. Natl. Herb. 17:
409. 1913. Xiphopteris pcrpusilla (Maxon)
Copel. Distribution: Brazil (Minas Gerais).
Micropolypodiuni plicatuni (A. R. Smith) A. R.
Smith, comb. nov. Basionym: Grammitis pti-
cata A. R. Smith, Ann. Missouri Bot. Card.
77: 258. 1990. Distribution: eastern Venezue¬
la.
Micropolypodiuni pulogense (Copel.) A. R.
Smith, comb. nov. Basionym: Polypodium pul-
ogense Copel., Philipp. J. Sci. 6: 148. 1911.
Xiphopteris pulogensis (Copel.) Copel. Distri¬
bution: Philippines.
Micropolypodiuni serricula (Fee) A. R. Smith,
comb. nov. Basionym: Polypodium serricula
Fee, Cen. Fil. 238. 1852. Grammitis serricula
(Fee) Proctor. Xiphopteris serricula (Fee) Co-
pel. Distribution: Lesser Antilles.
Micropolypodiuni setosum (Kaulf.) A. R. Smith,
comb. nov. Basionym: Xiphopteris setosa
Kaulf., Enum. Filic. 275. 1824. Grammitis
setosa (Kaulf.) C. Presl, non Blume, 1828.
Volume 2, Number 4
1992
Smith
Micropolypodium
423
Polypodium micropteris C. Chr. Distribution:
southern Brazil. Figure 1C.
Micropolypodium setulosum (Rosenstock) A.
R. Smith, comb. nov. Basionym: Polypodium
setulosum Rosenstock, Repert. Sp. Nov. Regni
Veg. 10: 277. 1912. Grammitis setulosa (Ro¬
senstock) F. Seymour. Xiphopteris setulosa
(Rosenstock) Copel. Distribution: Costa Rica,
Panama. Figure IE.
Micropoly podium sherringii (Baker) A. R.
Smith, comb. nov. Basionym: Poly podium
sherringii Baker in Jenman, J. Bot. 20: 26.
1882. Grammitis sherringii (Baker) Proctor.
Xiphopteris sherringii (Baker) Copel. Distri¬
bution: Cuba. Jamaica.
Micropolypodium taenifolium (Jenman) A. R.
Smith, comb. nov. Basionym: Polypodium
taenifolium Jenman, Bull. Bot. Dept. Jamaica,
n.s. 4: 114. 1897. Grammitis blepharodes
(Maxon) F. Seymour. Grammitis taenifolia
(Jenman) Proctor. Polypodium blepharodes
Maxon. Xiphopteris blepharodes (Maxon) Co-
pel. Xiphopteris taenifolia (Jenman) Copel.
Distribution: Antilles, northern South America.
Micropoly podium trichomanoides (Sw.) A. R.
Smith, comb. nov. Basionym: Polypodium tri¬
chomanoides Sw., Prodr. 131. 1788. (,ram-
mitis trichomanoides (Sw.) Ching. Xiphopteris
trichomanoides (Sw.) Copel. Distribution: Cuba,
Jamaica, southern Mexico, Guatemala, Vene¬
zuela. Figure 1G.
Micropolypodium truncicola (Klotzsch) A. R.
Smith, comb. nov. Basionym: Polypodium
truncicola Klotzsch, Linnaea 20: 374. 1847.
Grammitis truncicola (Klotzsch) C. Morton.
Poly podium andinum Hook. Xiphopteris
truncicola (Klotzsch) Copel. Distribution: Costa
Rica to Venezuela and Peru. Figure IB.
Micropoly podium williamsii (Maxon) A. R.
Smith, comb. nov. Basionym: Polypodium wil¬
liamsii M axon, Contr. U.S. Natl. Herb. 17:
547, pi. 34. 1916. Grammitis williamsii
(Maxon) Lellinger. Xiphopteris williamsii
(Maxon) Copel. Distribution: Bolivia.
Micropolypodium zurquinum (Copel.) A. R.
Smith, comb. nov. Basionym: Xiphopteris zur-
quina Copel., Amer. Fern J. 42: 99. 1952.
Grammitis zurquina (Copel.) F. Seymour. Dis¬
tribution: Costa Rica.
The following names (listed alphabetically by basionym)
clearly apply to species of Micropolypodium, but until
monographic or floristic work can be undertaken in their
area of occurrence, their status (whether synonyms or
good species) is unclear. They are not likely to supersede
any of the combinations made above.
Polypodium basale Maxon, Amer. Fern J. 52: 110.
1962. Grammitis basalis (Maxon) Lellinger. Distribution:
Ecuador. This species was omitted by Morton (1967).
Polypodium blepharolepis C. Chr., Index Fil. Suppl.
1: 58. 1913. Grammitis blepharolepis (C. Chr.) C. Mor¬
ton. Xiphopteris blepharolepis (C. Chr.) Copel. Distri¬
bution: Ecuador, Peru.
Polypodium buesii Maxon, Contr. Gray Herb. 165:
72. 1947. Grammitis buesii (Maxon) Lellinger. Xiph¬
opteris buesii (Maxon) Copel. Distribution: Peru.
Polypodium daguense Hieron., Bot. Jahrb. Syst. 34:
504. 1904. Grammitis daguensis (Hieron.) C. Morton.
Xiphopteris daguensis (Hieron.) Copel. Distribution: Co¬
lombia.
Polypodium gibbosum Fee, Mem. Foug. 6: 8, pi. 2,
f. 2. 1854. This species was said by Fee to be from
Mexico, probably erroneously.
Polypodium gracillimum Hieron., Hedwigia 48: 250.
1909, non Copel., 1905. = P. blepharolepis C. Chr.
Polypodium pseudotrichomanoides Hayata, Icon. FI.
Formosa 4: 251, f. 176. 1914. Micropolypodium pseu¬
dotrichomanoides (Hayata) Hayata, But. Mag. (Tokyo)
42: 341. 1928. Distribution: Taiwan. This species is prob¬
ably conspecific with Micropolypodium okuboi.
Polypodium pullei Alston, J. Bot. 78: 229. 1940.
Distribution: New Guinea. This is possibly a synonym of
Micro poly podium pulogense (Parris, in litt.).
Polypodium sikkimense Hieron., Hedwigia 44: 97.
1905. Xiphopteris sikkimensis (Hieron.) Copel. Distri¬
bution: Sikkim.
Xiphopteris killipii Copel., Amer. Fern J. 42: 105.
1952. Grammitis killipii (Copel.) Lellinger. Distribution:
Colombia.
Excluded New World Species
The following New World species of Xiphopteris (or
Grammitis) historically have been thought to be closely
allied by Copeland (1952) and others with species here
included in Micro poly podium. Their combinations in
Xiphopteris (or Grammitis, if no combination has been
made in Xiphopteris ), together with their taxonomic dis¬
position, are given in alphabetical order:
Grammitis anamorphosa Proctor = Lellingeria an-
amorphosa (Proctor) A. R. Smith & R. C. Moran
Grammitis nutata (Jenman) Proctor = Lellingeria
nutata (Jenman) A. R. Smith & R. C. Moran
Grammitis pseudomitchellae Lellinger = Lellingeria
pseudomitchellae (Lellinger) A. R. Smith & R. C. Moran
Grammitis ruglessii Proctor = Lellingeria ruglessii
(Proctor) A. R. Smith & R. C. Moran
Xiphopteris allosuroides (Rosenstock) Crabbe = Mel¬
pomene allosuroides (Rosenstock) A. R. Smith & R. C.
Moran
Xiphopteris amphidasyon (Kunze ex Mett.) Alston =
“ Terpsichore”
Xiphopteris apiculata (Kunze ex Klotzsch) Copel. =
Lellingeria apiculata (Kunze ex Klotzsch) A. R. Smith
& R. C. Moran
Xiphopteris aromatica (Maxon) Crabbe = Melpom¬
ene firma (J. Smith) A. R. Smith & R. C. Moran
Xiphopteris auyantepuiensis Vareschi = Cochlidium
serrulatum (Sw.) L. E. Bishop
Xiphopteris capillaris (Desv.) Crabbe = Ceradenia
capillaris (Desv.) L. E. Bishop
Xiphopteris choquetangensis (Rosenstock) Crabbe =
Pecluma choquetangensis (Rosenstock) M. G. Price
Xiphopteris cult rata (Bory ex Willd.) Schelpe =
“ Terpsichore ”
424
Novon
Xiphopteris delitescens (Maxon) Copel. = Lellingeria
delitescens (Maxon) A. R. Smith & R. C. Moran
Xiphopteris extensa Fee = Cochlidium serrulatum
(Sw.) L. E. Bishop
Xiphopteris fabelliformis (Poiret) Schelpe = Mel¬
pomene fiabelliformis ( Poiret) A. R. Smith & R. C. Moran
Xiphopteris hartii (Jenman) Copel. = Lellingeria har-
tii (Jenman) A. R. Smith & R. C. Moran
Xiphopteris heteromorpha (Hook. & Grev.) Crabbe
= “ Terpsichore"
Xiphopteris jamesonii Hook. = Lellingeria myosu-
roides (Sw.) A. R. Smith & R. C. Moran
Xiphopteris lanigera (Desv.) Crabbe = “ Terpsicho¬
re"
Xiphopteris limula (Christ) Pichi-Serm. = Lellingeria
limula (Christ) A. R. Smith & R. C. Moran
Xiphopteris longa (C. Chr.) Alston = “ Terpsichore ”
Xiphopteris luetzelburgii (Rosenstock) Brade = Lel¬
lingeria wittingiana (Fee) A. R. Smith & R. C. Moran?
Xiphopteris mitchellae (Baker ex Hemsley) Copel. =
Lellingeria mitchellae (Baker ex Hemsl.) A. R. Smith &
R. C. Moran
Xiphopteris moniliformis (Lagasca ex Sw.) Crabbe =
Melpomene moniliformis (Lagasca ex Sw.) A. R. Smith
& R. C. Moran
Xiphopteris mortonii Copel., Amer. Fern J. 42: 97.
1952. = Lellingeria
Xiphopteris myosuroides (Sw.) Kaulf. = Lellingeria
myosuroides (Sw.) A. R. Smith & R. C. Moran
Xiphopteris myriophylla (Mett.) Crabbe = “ Terp¬
sichore ”
Xiphopteris organensis (Gardner) Copel. = Lellin¬
geria organensis (Gardner) A. R. Smith & R. C. Moran
Xiphopteris peruviana (Desv.) Crabbe = Melpomene
peruviana (Desv.) A. R. Smith & R. C. Moran
Xiphopteris pilosissima (M. Martens & Galeotti)
Crabbe = Melpomene pilosissima (M. Martens & Gal¬
eotti) A. R. Smith & R. C. Moran
Xiphopteris pozuzoensis (Baker) Crabbe = Ceradenia
pilipes (Hook.) L. E. Bishop
Xiphopteris reitzii Alston = Lellingeria
Xiphopteris schenckii (Hieron.) Copel. = Lellingeria
schenckii (Hieron.) A. R. Smith & R. C. Moran
Xiphopteris serrulata (Sw.) Kaulf. = Cochlidium ser¬
rulatum (Sw.) L. E. Bishop
Xiphopteris shaferi (Maxon) Kramer = Lellingeria
shaferi (Maxon) A. R. Smith & R. C. Moran
Xiphopteris skutchii (Maxon) Copel. = Lellingeria
myosuroides (Sw.) A. R. Smith & R. C. Moran
Xiphopteris sodiroi (Christ & Rosenstock) Crabbe =
Melpomene sodiroi (Christ & Rosenstock) A. R. Smith
& R. C. Moran
Xiphopteris strictissima (Hook.) Vareschi = Lellin¬
geria myosuroides (Sw.) A. R. Smith & L. E. Bishop
Xiphopteris subscabra (Klotzsch) Crabbe = Polypo¬
dium subscabrum Klotzsch (see Stolze, 1991)
Xiphopteris wittingiana (Fee) Brade = Lellingeria
wittingiana (Fee) A. R. Smith & R. C. Moran
Xiphopteris yungensis (Rosenstock) Crabbe = “ Terp¬
sichore"
Facluded Oi.d World species ok Xiphopteris
Xiphopteris aethiopica Pichi-Serm. = Lellingeria ae-
thiopica (Pichi-Serm.) A. R. Smith & R. C. Moran
Xiphopteris albobrunnea (Baker) Schelpe
Xiphopteris alternidens (Cesati) Copel. (see Parris,
1986)
Xiphopteris antipodalis Copel.
Xiphopteris apoensis (Copel.) Copel.
Xiphopteris ascensionensis (Hieron.) Cronk = Lellin¬
geria
Xiphopteris bryophylla (v.A.v.R.) Parris = X. mus-
graviana (see Parris, 1986)
Xiphopteris conjunctisora (Baker) Copel. = Gram-
mitis ? (see Parris, 1983)
Xiphopteris cornigera (Baker) Copel. (see Parris, 1986)
Xiphopteris cucullata (Nees & Blume) Sprengel =
Calymmodon cucullatus (Nees & Blume) C. Presl
Xiphopteris elastica (Bory ex Willd.) Alston = “ Terp¬
sichore "
Xiphopteris exilis Parris (see Parris, 1986)
Xiphopteris glandulosopilosa (Brause) Copel. = X.
musgraviana (see Parris, 1986)
Xiphopteris govidjoaensis (Brause) Copel.
Xiphopteris hecistophylla (Copel.) Copel.
Xiphopteris heterophylla Sprengel = Grammitis ? (see
Parris, 1984)
Xiphopteris hieronymusii (C. Chr.) Holttum. Poly-
podium sertularioides J. Smith ex Hieron. (non Baker,
1876). = X. cornigera
Xiphopteris hildebrandtii (Hieron.) Tard. = Lellin¬
geria hildebrandtii (Hieron.) A. R. Smith & R. C. Moran
Xiphopteris khaoluangensis Tag. & Iwatsuki = X.
cornigera ? (Parris, in litt.)
Xiphopteris murudensis (Copel.) Copel.
Xiphopteris musgraviana (Baker) Parris (see Parris,
1986)
Xiphopteris oosora (Baker) Alston = Lellingeria oos-
ora (Baker) A. R. Smith & R. C. Moran
Xiphopteris orientalis (Desv.) Fourn. = Cochlidium
serrulatum (Sw.) L. E. Bishop
Xiphopteris punctata (Ballard) Alston = Ceradenia
pruinosa (Maxon) L. E. Bishop
Xiphopteris rigescens (Bory ex Willd.) Alston = Mel¬
pomene flabelliformii . (Poiret) A. R. Smith & R. C. Moran
Xiphopteris sajfordii (Maxon) Copel. = Lellingeria
saffordii (Maxon) A. R. Smith & R. C. Moran
Xiphopteris setulifera (v.A.v.R.) Parris = Polypo¬
dium pumHum Brause (non Cockayne, 1909) (see Parris,
1986)
Xiphopteris sparsipilosa (Holttum) Holttum (see Par¬
ris, 1986)
Xiphopteris strangeana Pichi-Serm. = Lellingeria
strangeana (Pichi-Serm.) A. R. Smith & R. C. Moran
Xiphopteris subcoriacea (Copel.) Copel. = Lellingeria
subcoriacea (Copel.) A. R. Smith & R. C. Moran
Xiphopteris subpinnatifida (Blume) Copel. (see Parris,
1986)
Xiphopteris villosissuna (Hook.) Alston = Zygophle-
bia villosissima (Hook.) L. E. Bishop
Xiphopteris zenkeri (Hieron.) Schelpe = Ctenopteris
zenkeri (Hieron.) Tard.
Acknowledgments. I thank Barbara Parris and
Bobbin Moran for helpful comments on this manu¬
script, and Linda Vorobik for executing the illus¬
trations.
Literature Cited
Bishop, L. E. 1988. Ceradenia , a new genus of Gram-
mitidaceae. Amer. Fern J. 78: 1-5.
- . 1989. Zygophlebia, a new genus of Gram-
initidaceae. Amer. Fern J. 79: 103 118.
Volume 2, Number 4
1992
Smith
Micropolypodium
425
Copeland, E. B. 1952. The American species of Xiph-
opteris. Amer. Fern J. 42: 41-52, 93-110.
Hayata, B. 1928a. On the systematic importance of
the stelar system in the Filicales, II. Bot. Mag. Tokyo
42: 301-311.
-. 1928b. On the systematic importance of the
stelar system in the Filicales, III. Bot. Mag. Tokyo
42: 334-348.
-. 1929. Uber die systematische Bedeutung des
stelaren Systemes in den Polypodiaceen. Flora 124:
38-62 + 2 pi.
Lellinger, D. B. 1989. The ferns and fern-allies of Costa
Rica, Panama, and the Choco (Part 1). Pteridologia
2A: 1-364.
Maxon, W. R. 1916. Studies of tropical American
ferns — No. 6. Contr. U.S. Natl. Herb. 17: 541-
608.
Mickel, J. T. & J. M. Beitel. 1988. Pteridophyte flora
of Oaxaca, Mexico. Mem. New York Bot. Card. 46:
1-568.
Morton, C. V. 1967. The genus Grammitis in Ecuador.
Contr. U.S. Natl. Herb. 38: 85-123.
Parris, B. S. 1983. A taxonomic revision of the genus
Grammitis Swartz (Grammitidaceae: Filicales) in New
Guinea. Blumea 29: 13-222.
-. 1984. Another intergeneric hybrid in Gram¬
mitidaceae: Ctenopteris longiceps x Grammitis su-
matrana. Fern Gaz. 12: 337-340.
-. 1986. Grammitidaceae of peninsular Malaysia
and Singapore. Kew Bull. 41: 491-517.
-. 1990. Noteworthy species of Grammitidaceae
from South-east Asia. Hooker’s Icon. PI. 40(4): 1-
129 + i-iv.
Proctor, G. R. 1977. Pteridophyta. Pp. 1414 in R.
A. Howard (editor), Flora of the Lesser Antilles, vol.
2. Arnold Arboretum, Jamaica Plain, Massachusetts.
-. 1985. Ferns of Jamaica. British Museum (Nat¬
ural History), London.
-. 1989. Ferns of Puerto Rico and the Virgin
Islands. Mem. New York Bot. Card. 53: 1-389.
Silva Araujo, I. J. 1976. In: IOPB chromosome number
reports LIII. Taxon 25: 483-500.
Smith, A. R. 1981. Pteridophytes. Pp. 1-370 in D.
E. Breedlove (editor). Flora of Chiapas, Part 2. Cal¬
ifornia Academy of Sciences, San Francisco.
- & R. C. Moran. 1992. Melpomene, a new
genus of Grammitidaceae (Pteridophyta). Novon 2:
426-432.
-, - & L. E. Bishop. 1991. Lellingeria,
a new genus of Grammitidaceae. Amer. Fern J. 81:
76-88.
Stolze, R. G. 1981. Fern and fern allies of Guatemala.
Part 2: Polypodiaceae. Fieldiana, Bot. n.s. 6: 1-522.
-. 1991. The identity of Poly podium subsca-
brum Klotzsch. Amer. Fern J. 81: 139-140.
Walker, T. G. 1985. Cytotaxonomic studies of the ferns
of Trinidad. 2. The cytology and taxonomic impli¬
cations. Bull. Brit. Mus. (Nat. Hist.), Bot. 13: 149—
249.
Melpomene , a New Genus of Grammitidaceae (Pteridophyta)
Alan R. Smith
University Herbarium, University of California, Berkeley, California 94720, U.S.A.
Rabbin C. Moran
Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ABSTRACT. Melpomene, a new genus of Gram¬
mitidaceae, is described, and combinations are made
for the species known to belong to it. Melpomene
can be distinguished from all other Grammitidaceae
by its rhizome scales that are clathrate, basally cor¬
date, and entire except at the apex where provided
with one to ten minute papillae. The genus contains
about 20 species and is primarily neotropical.
This paper is one in a series that have set torth
new generic concepts in neotropical Grammitida¬
ceae. In the series, Grammitis sensu lato has been
divided into smaller monophyletic groups that have
not previously been known or recognized. These
groups, given generic status, are Cochlidium (Bish¬
op, 1978), Ceradenia (Bishop, 1988), Enterosora
(Bishop & Smith, 1992), Grammitis (Bishop, 1977),
Lellingeria (Smith et ah, 1991), and Zygophlebia
(Bishop, 1989). Each group is believed to be mono¬
phyletic because it is defined by several correlating
characteristics. In this paper, we describe another
monophyletic group within the Grammitidaceae:
Melpomene (the name of the Muse of Tragedy).
The distinctness of Melpomene was first recog¬
nized by the late L. Earl Bishop. We have deter¬
mined what species belong to the genus and have
found additional characteristics that define it. There¬
fore, we take sole reponsibility lor the description
of the genus and new combinations.
Melpomene A. R. Smith & R. C. Moran, gen.
nov. TYPE: Polypodium moniliforme Lagasca
ex Sw., Syn. Fil. 33. 1806. = Melpomene
moniliformis (Lagasca ex Sw.) A. R. Smith &
R. G. Moran. Figure 1.
Plantae epiphyticae terrestres vel rupestres; squamae
rhizomatis clathratae denigratae brunneolae vel vulgo ru-
bellae integrae, base cordatae, apice glandulis praeditae;
phyllopodia absentia; folia plerumque pinnatisecta, hy-
dathodis adaxialiter ornata; petioli et rhachides setosi, setis
castaneis; venae non furcatae librae; sori rotundi super-
ficiales.
Epiphytic, epipetric, or terrestrial; rhizome erect
or short- to long-creeping, the scales strongly clath¬
rate, cordate basally and attached at a single point,
usually blackish or reddish, never setulose, with one
to several papillose cells (glandular?) ca. 0.1 mm
long at the apex, these often dislodged and not
present; phyllopodia absent; leaves pinnatifid, pin-
natisect, or rarely 1 -pinnate basally, linear to elliptic,
pubescent and often setose, the hairs 0.1-0.2 mm
long, branched, 2- 8-celled, lax, pale reddish, with
most of the color concentrated at the cross walls,
the setae 0.3-3 mm long, castaneous, multicellular
(the cross walls sometimes difficult to observe), erect
to spreading; hydathodes present; veins free, often
not visible; sori round or (rarely) slightly oblong,
discrete, superficial or rarely slightly sunken, lacking
globose, waxlike paraphyses, but some species with
castaneous receptacular setae.
In his revision of American grammitids (as Cten-
opteris), Copeland (19S5) treated the species of
Melpomene in four of his ten species groups. He
placed most ol the species of Melpomene, however,
under the group of C. moniliformis (Lagasca ex
Sw.) J. Smith (the first of his ten groups). In this
group he also included several species now placed
in Lellingeria and a few species belonging to other
genera. Copeland defined the (,. moniliformis group
very loosely, saying that it consisted of “small, lin¬
ear, coriaceous ferns” that were related to Cten-
opleris moniliformis (= Melpomene moniliformis).
Obviously, Copeland’s characterization of the group
is not sufficient to establish monophyly. Yet Mel¬
pomene does appear to be monophyletic, and we
have found several characteristics of the stem scales
that define it.
All Melpomene species have scales that are clath¬
rate throughout (Fig. IB, 1), I, J, L, M, S). The
scales are entire except at or near the apex, where
one to ten, minute glandular cells occur (Fig. II, J,
O, P). The base of the scales is cordate, sometimes
with the basal auricles overlapping, and attached at
a point (Fig. IB, I), I, L, 0, R). These four char¬
acteristics of the scales distinguish Melpomene from
all other genera in the family.
Lellingeria is the only other genus of neotropical
Novon 2: 426-432. 1992.
Volume 2, Number 4
1992
Smith & Moran
Melpomene
427
Figure 1. Characteristics of Melpomene. A, B. M. sodiroi, Rimbach s.n. (Rosenstock exsic. 13; UC). —A. Habit.
— B. Rhizome scale. C E. M. anfractuosa, Mickel 677] (UC). —C. Habit, with small leaf from root proliferations.
— D. Rhizome scale. —E. Fungal fruiting body from rachis. F, G. M. Jirma, van der Werff & Gudino 11105 (UC).
— F. Habit. —G. Hairs on petiole. H J. M. flabelliformis. Smith el al. 1478 (UC). —H. Habit. —1. Rhizome scale.
—J. Tip of rhizome scale. K-M. M. erecta, Steyermark et al. 100548 (UC). —K. Habit. —L. Rhizome scale. —
M. Detail of rhizome scale. N-P. M. pilosissima, Mickel & Hellwig 3775 (UC). —N. Habit. —0. Rhizome scale.
— P. Tip of rhizome scale. Q S. M. moniliformis, Mickel & Hellwig 4041 (UC). —Q. Habit. —R. Rhizome scale.
—S. Detail of rhizome scale. —T. M. assurgens, habit, van der Werff & Palacios 8960 (UC). —U. M. leptostoma,
habit, Hutchison 440 (UC).
Grammitidaceae that has strongly clathrate stem
scales. Its scales, however, are often setulose mar¬
ginally and/or at the apex and lack glands (or pa¬
pillae) near the apex (Smith et al., 1991). The scales
are also truncate basally, being attached across the
entire length of the base.
In addition to the stem scales, Melpomene has
other characteristics that distinguish it, although
these characteristics are not unique to the genus.
In habit, the leaves are erect, never laxly pendent
as in many American grammitids (the only exception
is M. leptostoma (Fee) A. R. Smith & R. C. Moran,
which may have arching or pendent leaves; Fig.
1U). The laminae are generally thick or coriaceous,
428
Novon
Figure 2. Worldwide distribution of Melpomene. The number on the left is the number of species in the region;
the number in parentheses is the number of those species endemic to the region.
and linear to narrowly elliptic (but M. firma (J.
Smith) A. K. Smith & H. C. Moran may have broadly
elliptic laminae). Dark, castaneous setae are present
in most species, especially along the rachis anti around
the sori. These dark, castaneous setae are usually
absent along the segment margins, with the excep¬
tion of M. pilosissima (M. Martens & Galeotti) A.
R. Smith & R. C. Moran, which has setae along the
margins throughout, and M. anfractuosa (Kunze ex
Klotzsch) A. R. Smith & R. C. Moran, which has
marginal setae near the segment apices. The spo-
rangial capsules are glabrous (never setulose), and
the receptacle lacks glands. All species have con¬
spicuous hydathodes adaxially, which distinguishes
the genus from the anhydathodous Ceradenia, Zyg-
ophlebia, and Knterosora. Melpomene anfractuosa
is unusual in the genus in having plantlets produced
from buds on the roots (Fig. 1C) and by bearing
black club-shaped fruiting bodies of the ascomycete
Acrospermum maxonii Farlow (Fig. IE).
Many species of Melpomene, when dried, emit a
sweet spicy fragrance that does not occur in other
grammitid genera. The fragrance persists for many
decades after the plants are dried. This aroma can
be almost overpowering to some individuals, while
scarcely noticeable by others. Proctor (1985) re¬
ported that specimens of M. firma were still fragrant
after 100 years. Mickel & Beitel (1988: 199) re¬
ported a “decidedly sweet smell on drying” in mem¬
bers of the M. moniliformis group. The species that
have this fragrance often discolor the herbarium
sheet on which they are mounted, turning the paper
brownish. The species that we have noted as having
this fragrance include, but are not restricted to, M.
firma, M. flabellifiormis, M. pilosissima, M. pe¬
ruviana, M. vernicosa, and M. xiphopteroides.
Nothing is known about the chemistry of this fra¬
grance.
Chromosome counts and spores, which are often
helpful in defining fern genera, do not distinguish
Melpomene from other genera of Grammitidaceae.
Only one chromosome count is known in Melpom¬
ene: n = ca. 74 in M. moniliformis from Jamaica
(Walker, 1966). Phis appears to be a tetraploid
number based on 37, which is a common base num¬
ber for grammitid genera. The spores of the Gram¬
mitidaceae are remarkably uniform, and those of
Melpomene photographed (as Grammitis) by 1 ryon
& Lugardon (1991) are similar to those of other
grammitid genera.
The gametophytes of 25 species of Grammiti¬
daceae from Jamaica, including four of Melpomene
{M. anfractuosa, M. firma, M. moniliformis, and
M. pilosissima), were described by Stokey & At¬
kinson (1958). An examination of their work does
not reveal any differences between the gametophytes
of Melpomene and those of other grammitids.
Melpomene is a genus of mid to high elevations,
generally occurring from 1,000 to 4,000 m. In the
New World, it ranges from Mexico to Panama, the
Volume 2, Number 4
1992
Smith & Moran
Melpomene
429
Antilles, and South America from Venezuela to Bo¬
livia, Argentina, and southeastern Brazil. In the Old
World it occurs in Africa, Madagascar, and Reunion
(Fig. 2), but is apparently absent from Malesia,
southeastern Asia, and the Pacific. The genus con¬
tains about 20 species that are badly in need of a
monograph.
The relationships of Melpomene within the Gram-
mitidaceae are not clear. The genus may be most
closely related to the other hydathodous neotropical
genera, particularly Lellingeria (which also has
clathrate scales), or to the species groups that in¬
clude Grammitis taxifolia (L.) Proctor and G. sub-
scabra (Klotzsch) C. Morton, which belong to a yet
unpublished genus Terpsichore, with over 50 New
World species. There is no close affinity to Xiph-
opteris (sensu Copeland, 1956), which we regard
as a mixture of Coehlidium and Micropoly podium.
A few species in the Pacific Basin resemble Mel¬
pomene in having hydathodes and clathrate or sub-
clathrate, entire rhizome scales, e.g., Ctenopteris
lasiostipes (Mett.) Brownlie, C. aff. bleehnoides
(Grev.) W. Wagner & Grether (both from New
Caledonia), and (.. curtisii (Baker) Copel. (Malesia).
These species differ from Melpomene in one or
usually more of the following characteristics: tan¬
gential walls of the rhizome scales slightly darkened
(not clear and transparent as in Melpomene)-, scales
lacking glandlike papillae at the tips, not cordate
basally, sometimes curving inward at the tip (C.
curtisii); branched hairs lacking on the petiole and
rachis; clumped soral setae ( C. lasiostipes); nonaro¬
matic dried leaves. It seems likely that the resem¬
blances in scales are due to convergent evolution
and are not an indication of affinity.
New Combinations
Melpomene allosuroides (Rosenstock) A. R.
Smith & R. C. Moran, comb. nov. Basionym:
Polypodium allosuroides Rosenstock, Meded.
Rijks-Herb. 19: 16. 1913. Ctenopteris allo¬
suroides (Rosenstock) Copel., Philipp. .). Sci.
84: 396. 1955 (1956). Grammitis allosu¬
roides (Rosenstock) Bellinger, Amer. Fern J.
74: 58. 1984. TYPE: Bolivia. Cochabamba?:
Lagodos Valley, 4,000 m, Herzog 2373 (ho-
lotype, S not seen; isotypes, UC, US not seen).
I)is t ribution. Bol i via.
Melpomene anfraetuosa (Kunze ex Klotzsch) A.
R. Smith & R. C. Moran, comb. nov. Basionym:
Polypodium anfractuosum Kunze ex Klotzsch,
Linnaea 20: 375. 1847. Ctenopteris anfrac-
tuosa (Kunze ex Klotzsch) Copel.. Philipp. J.
Sci. 84: 431. 1955 (1956). Grammitis an-
fractuosa (Kunze ex Klotzsch) Proctor, Rho-
dora 63: 35. 1961. TYPE: Venezuela. Merida:
Moritz 330? (isotype, US).
Distribution. Southern Mexico to Guyana and
Peru, Antilles.
Melpomene assurgens (Maxon) A. R. Smith &
R. C. Moran, comb. nov. Basionym: Poly po¬
dium assurgens Maxon, Contr. Gray Herb.
165: 73. 1947. Ctenopteris assurgens (Max¬
on) Copel., Philipp. J. Sci. 84: 386. 1955
(1956). Grammitis assurgens (Maxon) C. Mor¬
ton, Contr. IJ.S. Natl. Herb. 38: 116. 1967.
TYPE: Ecuador. Pichincha: Quito Santo Do¬
mingo road, Haught 3226 (holotype, US not
seen).
Distribution. Ecuador, Colombia, Peru.
Melpomene brevipes (C. Morton) A. R. Smith &
R. C. Moran, comb, et stat. nov. Basionym:
Grammitis sodiroi var. brevipes C. Morton,
Phytologia 22: 79. 1971. TYPE: Ecuador.
Tunguragua: Mt. Tunguragua, Spruce 5270
(holotype, K not seen).
Distribution. Western Ecuador.
Melpomene deltata (Mickel & Beitel) A. R. Smith
& R. C. Moran, comb. nov. Basionym: Gram¬
mitis deltata Mickel & Beitel, Mem. New York
Bot. Card. 46: 198. 1988. TYPE: Mexico.
Oaxaca: Dtto. Mixe, NW slope of Cerro Zem-
poaltepetl, Mickel 4637a (holotype, NY).
Distribution. Southern Mexico, El Salvador.
Melpomene dissimulans (Maxon) A. R. Smith
& R. C. Moran, comb. nov. Basionym: Poly¬
podium dissimulans Maxon, Contr. U.S. Natl.
Herb. 10: 502. 1908. Ctenopteris dissimulans
(Maxon) Copel., Philipp. J. Sci. 84: 415. 1955
(1956). Grammitis dissimulans (Maxon) F.
Seymour, Phytologia 31: 179. 1975. TYPE:
Guatemala. Alta Verapaz: near Coban, von
Tiirckheim (J. D. Smith 834) (holotype, US
not seen).
Distribution. Guatemala.
Melpomene erecla (C. Morton) A. R. Smith &
R. C. Moran, comb. nov. Basionym: Grammitis
erecla C. Morton, Phytologia 22: 72. 1971.
TYPE: Ecuador. Tunguragua: Mt. Tungura-
430
Novon
gua. Spruce 5279A (holotype, K not seen;
isotype, (ill).
Distribution. Venezuela, Ecuador, Peru.
Melpomene firma (J. Smith) A. R. Smith & R.
C. Moran, comh. now Basionym: Polypodium
firmum Klotzsch, Linnaea 20: 378. 1947 (non
Kaulf., 1827). Ctenopterisfirma J. Smith, Hist.
Fil. 184. 1875. Grammitis firma (J. Smith) C.
Morton, Contr. U.S. Natl. Herb. 38: 1 10. 1967.
TYPE: Guyana, Schomburgk 1170 (lectotype,
inferentially chosen by Looser, Revista Univ.
(Santiago) 36(1): 75. 1951, B not seen; iso-
lectotype, K).
Polypodium aromaticum Maxon, Proc. U.S. Natl. Mus.
27: 743. 1904. Ctenopteris aromatica (Maxon)
Copel., Philipp. J. Sci. 84: 418. 1955 (1956). Gram¬
mitis aromatica (Maxon) Proctor, Brit. Fern Gaz.
9: 218. 1965. Xiphopteris aromatica (Maxon)
Crabbe, Brit. Fern Gaz. 9: 318. 1967. TYPE: Ja¬
maica. Blue Mt. Peak, Underwood 1449 (holotype,
NY not seen; fragment US).
Polypodium herzogii Rosenstock, Repert. Spec. Nov.
Regni Veg. 6: 176. 1908. TYPE: Bolivia. Cocha¬
bamba: Incacorral, Herzog 783 (holotype, S not
seen; isotypes, UC, US not seen).
Distribution. Southern Mexico to Honduras, Cos¬
ta Rica to Guyana and Bolivia, Jamaica.
Melpomene flabelliformis (Poiret) A. R. Smith
& R. G. Moran, comb. nov. Basionym: Poly¬
podium fiahclli forme Poiret, Encycl. 5: 519.
1804. Grammitis flabelliformis (Poiret) G.
Morton, Contr. U.S. Natl. Herb. 38: 57. 1967.
Xiphopteris flabelliformis (Poiret) Schelpe, Bol.
Soc. Brot., ser. 2, 41: 217. 1967. TYPE:
Reunion, Gornmerson s.n. (P-JU 1098C).
Polypodium rigescens Bory ex Willd., Sp. PI. ed. 4, 5:
183. 1810. Ctenopteris rigescens (Bory ex Willd.)
J. Smith, Hist. Fil. 184. 1875. Xiphopteris riges¬
cens (Bory ex Willd.) Alston, Bol. Soc. Brot., ser.
2, 30: 27. 1956. Grammitis rigescens (Bory ex
Willd.) Lellinger, Proc. Biol. Soc. Wash. 89: 383.
1985. TYPE: Reunion, Bory s.n. (holotype, B-W
19668; isotype, P-JU 1098C).
Grammitis katasophistes Mickel & Beitel, Mem. New
York Bot. Card. 46: 200. 1988. TYPE: Mexico.
Oaxaca: Dtto. Mixe, NW slope of Cerro Zempoal-
tepetl, Mickel 4837b (holotype, NY).
Distribution. Southern Mexico, Costa Rica to
Venezuela and Peru, southern Brazil, Hispaniola,
Africa, Madagascar, Reunion.
Melpomene leptostoma (Fee) A. R. Smith & R.
C. Moran, comb. nov. Basionym: Polypodium
leptostomum Fee, Mem. Foug. 7: 58. 1857.
Ctenopteris leptostoma (Fee) Copel., Philipp.
.1. Sci. 84: 421. 1955 (1956). Grammitis lep¬
tostoma (Fee) F. Seymour, Phytologia 31:1 79.
1975. TYPE: Mexico. Veracruz: near Orizaba,
Schaffner 210 (holotype, P not found).
Polypodium productum Maxon, Contr. U.S. Natl. Herb.
13: 11. 1909. TYPE: Guatemala. Alta Verapaz:
near Coban, von Tiirckheim II. 1347, pro parte (ho¬
lotype, US).
Distribution. Southern Mexico, Guatemala.
Melpomene melanosticta (Kunze) A. R. Smith
& R. C. Moran, comb. nov. Basionym: Poly¬
podium melanostictum Kunze, Linnaea 9: 44.
1834. Ctenopteris melanosticta (Kunze) Co-
pel., Philipp. J. Sci. 84: 385. 1955 (1956).
Grammitis melanosticta (Kunze) F. Seymour,
Phytologia 31: 179. 1975. TYPE: Peru, Poep-
pig s.n. (holotype, LZ destroyed).
Polypodium calvum Maxon, J. Wash. Acad. Sci. 12:
440. 1922. Ctenopteris calva (Maxon) Copel., Phil¬
ipp. J. Sci. 84: 386. 1955 (1956). TYPE: Cuba.
Oriente: Leon 11131 (holotype, US).
Distribution. Southern Mexico, Guatemala, Cos¬
ta Rica, Venezuela to Bolivia, Brazil, Cuba, Hispan¬
iola.
Melpomene moniliformis (Lagasca ex Sw.) A.
R. Smith & R. C. Moran, comb. nov. Basionym:
Polypodium moniliforme Lagasca ex Sw., Syn.
Fil. 33. 1806. Ctenopteris moniliformis (La¬
gasca ex Sw'.) J. Smith, Hist. Fil. 184. 1875.
Grammitis moniliformis (Lagasca ex Sw.)
Proctor, Brit. Fern Gaz. 9: 219. 1965. Xiph¬
opteris moniliformis (Lagasca ex Sw.) Crabbe,
Brit. Fern Gaz. 9: 319. 1967. TYPE: Peru,
collector not stated [Ruiz & PavonV] (S not
seen).
Distribution. Mexico to Venezuela, Bolivia, and
southern Brazil, Jamaica, Hispaniola.
Melpomene pennellii (Copel.) A. R. Smith & R.
C. Moran, comb. nov. Basionym: Ctenopteris
pennellii Copel., Philipp. J. Sci. 84: 397. 1955
(1956). Grammitis pennellii (Copel.) C. Mor¬
ton, Phytologia 22: 82. 1971. TYPE: Colom¬
bia. Cauca: “San Jose,” San Antonio, Pennell
& Killip 7379 (holotype, IJS).
Distribution. Western Colombia.
Melpomene peruviana (Desv.) A. R. Smith &
R. C. Moran, comb. nov. Basionym: Polypo¬
dium peruvianum Desv., Mem. Soc. Linn. Par-
Volume 2, Number 4
1992
Smith & Moran
Melpomene
431
is 6: 231. 1827. Ctenopteris peruviana (l)esv.)
J. Smith, Hist. Fil. 184. 1875. Grammitis pe¬
ruviana (Desv.) C. Morton, Contr. U.S. Natl.
Herb. 38: 115. 1967. Xiphopteris peruviana
(Desv.) Crabbe, Brit. Fern Gaz. 9: 319. 1967.
TYPE: Peru. Collector not stated (holotype, P).
Polypodium peruvianum var. subgibbosum Hosenstock,
Meded. Rijks-Herh. 19: 16. 1913. TYPE: Bolivia.
Cochabamba?: Valle Lagados, Herzog 2373a (ho¬
lotype, S not seen; isotype, UC).
Distribution. Ecuador to Bolivia, Argentina, and
Brazil.
Melpomene pilosissima (M. Martens & Galeotti)
A. K. Smith & B. C. Moran, comb. nov. Bas-
ionym: Polypodium pilosissimum M. Martens
& Galeotti, Nouv. Mem. Acad. Roy. Sci. Bru¬
xelles 15(5): 39, t. 9, fig. 2. 1842. Ctenopteris
pilosissima (M. Martens & Galeotti) Copel.,
Philipp. .1. Sci. 84: 390. 1955 (1956). Gram¬
mitis pilosissima (M. Martens & Galeotti) (7
Morton, Contr. U.S. Natl. Herb. 38: 1 14. 1967.
Xiphopteris pilosissima (M. Martens & Gal¬
eotti) Crabbe, Brit. Fern Gaz. 9: 319. 1967.
TYPE: Mexico. Oaxaca: Galeotti 6397 (lec-
totype, designated by Smith, FI. Chiapas 2:
124. 1981, BR; isolectotype, K).
Polypodium acrodontium Fee, Crypt. Vase. Bresil 2: 58,
t. 99, fig. 2. 1872. TYPE: Brazil. Glaziou 4409
(holotype, P or RB not seen).
Grammitis zempoaltepetlensis Mickel & Beitel, Mem.
New York Bot. Card. 46: 205. 1988. TYPE: Mex¬
ico. Oaxaca: Dtto. Mixe, NW slope of Cerro Zem-
poaltepetl, trail from Tacoche to Totontepec, Mickel
4638 (holotype, NY).
Distribution. Mexico to Honduras, Costa Rica to
Surinam, Peru and Brazil.
Melpomene pseudonutans (Christ & Rosen-
stock) A. R. Smith & R. C. Moran, comb. nov.
Basionym: Polypodium pseudonutans Christ
& Rosenstock, Repert. Spec. Nov. Regni Veg.
5: 15. 1908. Ctenopteris pseudonutans (Christ
& Rosenstock) Copel., Philipp. J. Sci. 84: 389.
1955 (1956). Grammitis pseudonutans (Christ
& Rosenstock) C. Morton, Contr. U.S. Natl.
Herb. 38: 114. 1967. TYPE: Ecuador. Tun-
guragua: slopes of Mt. Tunguragua, Rimbach
s.n. (holotype, S not seen; isotypes, UC, US
not seen).
Distribution. Colombia, Ecuador, Peru.
Melpomene rosarum (Copel.) A. R. Smith & R.
C. Moran, comb. nov. Basionym: Ctenopteris
rosarum Copel., Philipp. J. Sci. 84: 387. 1955
(1956). Grammitis rosarum (Copel.) C. Mor¬
ton, Contr. U.S. Natl. Herb. 38: 116. 1967.
TYPE: Ecuador. Chimborazo: vicinity of Hui-
gra. Rose 22232 (holotype, US not seen).
Distribution. Colombia, Ecuador.
Melpomene sodiroi (Christ & Rosenstock) A. R.
Smith & R. C. Moran, comb. nov. Basionym:
Polypodium sodiroi Christ & Rosenstock, Re-
pert. Spec. Nov. Regni Veg. 5: 14. 1908.
Grammitis sodiroi (Christ & Rosenstock) C.
Morton, Contr. U.S. Natl. Herb. 38: 1 14. 1967.
Xiphopteris sodiroi (Christ & Rosenstock)
Crabbe, Brit. Fern Gaz. 9: 319. 1967. TYPE:
Ecuador. Tunguragua: base of Mt. Tungura¬
gua, Rimbach 24 (holotype, P? or S?; isotypes
UC, US not seen).
Distribution. Western Colombia, western Ecua¬
dor.
Melpomene vernicosa (Copel.) A. R. Smith &
R. C. Moran, comb. nov. Basionym: Ctenop¬
teris vernicosa Copel., Philipp. J. Sci. 84: 452,
t. 9. 1955 (1956). Grammitis vernicosa (Co-
pel.) C. Morton, Contr. U.S. Natl. Herb. 38:
1 10. 1967. TYPE: Costa Rica. Cartago: Alto
de Estrella, Standley 39140 (lectotype, des¬
ignated by Morton, Contr. U.S. Natl. Herb. 38:
1 10. 1967, US not seen).
Distribution. Costa Rica, Panama, Venezuela,
Colombia, Ecuador.
Melpomene wolfii (Hieron.) A. R. Smith & R.
C. Moran, comb. nov. Basionym: Polypodium
wolfii Hieron., Hedwigia 48: 249. 1909.
Grammitis wolfii (Hieron.) C. Morton, Contr.
U.S. Natl. Herb. 38: 115. 1967. TYPE: Ec¬
uador. Pichincha: near Mindo, at base of Mount
Pichincha, Stiibel 743 (holotype, B not seen).
Distribution. Western Ecuador.
Melpomene xiphopteroides (Liebm.) A. R. Smith
& R. C. Moran, comb. nov. Basionym: Poly¬
podium xiphopteroides Liebm., Kongel.
Danske Vidensk. Selsk. Skr., Naturvidensk. Afd.
ser. 5, 1: 196. 1849. Grammitis xiphopter¬
oides (Liebm.) A. R. Smith, Amer. Fern J. 70:
26. 1980. TYPE: Mexico. Veracruz: “Hac. de
Mirador,” Liebmann [Liebm. PI. Mex. 2548,
FI. Mex. 189] (lectotype, designated by Smith,
FI. Chiapas 2: 125. 1981, C).
432
Novon
Ctenopleris megaloura Copel., Philipp. J. Sci. 84: 391.
1955 (1956). TYPE: Guatemala. Alta Verapaz:
Coban, von Tiirckheim II. 1855 (holotype, US).
Polypodium rigens Maxon, Proc. U.S. Natl. Mus. 27:
741. 1904. Ctenopleris rigens (Maxon) Copel., Phi¬
lipp. J. Sci. 84: 422. 1955 (1956). Grammitis ri¬
gens (Maxon) Proctor, Brit. Fern Gaz. 9: 219. 1965.
TYPE: Jamaica. John Crow Peak, Maxon 134b
(holotype, US).
Distribution. Southern Mexico, Guatemala, Hon¬
duras, Costa Rica, Panama, Greater Antilles, Ven¬
ezuela, Colombia, Ecuador, Peru.
Incertae Seiiis
The following names are not in current use, and
we have not seen their types. Nevertheless, they
apparently belong to Melpomene and have been
inluded here for the benefit of future monographers.
Most will probably prove to be synonyms of the
above names.
Jamesonia adnata Kunze, Farrnkrauter 2: 80, t. 133,
fig. 1. 1851. TYPE: Colombia. Tolima: Paramo de
Tolima, Linden 1006 (holotype, LZ destroyed; is¬
otypes, B not seen, FI not seen).
Polypodium angustissimum Fee, Crypt. Vase. Bresil 2:
55, t. 96, fig. 3. 1873. TYPE: Brazil: Glaziou 5297
(P or RB, not seen).
Polypodium moniliforme var. bogotense Hieron., Hed-
wigia 48: 249. 1909. TYPE: Colombia. Cundina-
marca: Bogota region, Stiibel 431 pro parte (holo¬
type, B not seen).
Polypodium moniliforme var. culebriliense Bosco, Nuo-
vo Giorn. Bot. Ital. N.S., 45: 149. 1938. TYPE:
Ecuador. Chimborazo?: Paramo de Culebrilla, Cres-
pi s.n. (holotype, TO not seen).
Polypodium moniliforme var. major Liebin., Kongel.
Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. ser.
5, 1: 196. 1849. TYPE: Mexico. Veracruz: Pico
de Orizaba, Liebmann [Liebm. PI. Mex. 2533, FI.
Mex. 183] (lectotype, designated by Smith, FI. Chia¬
pas 2: 123. 1981, C).
Polypodium moniliforme var. minus Christ, Bull. Soc.
Bot. Geneve ser. 2, 1: 217. 1909. SYNTYPES:
Costa Rica. Cartago: Crater of Volcan Irazu, Pittier
179 (P not seen, isosyntype US not seen); Brazil.
Rio de Janiero: Ule 3785 (P not seen).
Polypodium pilosissimum var. glabriusculum Mett., Abh.
Senckenberg. Naturf. Ges. 2: 42. 1857. SYN¬
TYPES: Mexico: Leibold 97 (B not seen). Vene¬
zuela. Merida: Moritz 216 (B not seen, US); Distrito
Federal: Funck & Schlim 965 (B not seen).
Polypodium pilosissimum var. hirsutum Mett., Abh.
Senckenberg. Naturf. Ges. 2: 42. 1857. TYPE:
Venezuela. Punch & Schlim 1848 (holotype, B not
seen).
Polypodium suberenatum Hook., Ic. PI. 8: pi. 719. 1848.
TYPE: Ecuador. Pichincha: Jameson 215 (holo¬
type, K not seen).
Polypodium subdicarpon Fee, Crypt. Vase. Bresil 2: 55,
t. 96, fig. 4. 1874. TYPE: Brazil. Glaziou 4410
(holotype, P or RB, not seen).
Acknowledgments. We thank David B. Lellinger
and Barbara S. Parris for helpful reviews ol the
manuscript.
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Pteridophyta. Springer-Verlag, New York.
Walker, T. G. 1966. A cytotaxonomic survey of the
pteridophytes of Jamaica. Trans. Roy. Soc. Edin¬
burgh. 66: 169-237.
Carex david-smithii 1 (Cyperaceae), a New Species from
High Andean Peru
I). /V. Smith f
Department of Botany, Iowa State University, Arnes, Iowa 50011, U.S.A.
A. A. Reznicek
University of Michigan Herbarium, Ann Arbor, Michigan 48109, U.S.A.
Abstract. Carex david-smithii is described as new
from Bolivia. It is most closely related to the Ar¬
gentinean and Chilean C. catamarcensis , but differs
in its dark, almost black, pistillate scales and strongly
outcurved perigynia tapered to the beak. Carex
catamarcensis has stramineous to reddish purple
scales and straight perigynia abruptly contracted
into the beak.
This new species was discovered in the process
of preparing treatments of the monocotyledonous
families included in the flora of the Huascaran Na¬
tional Park and International Biosphere Reserve
(Smith, 1988).
The park includes the bulk of the Cordillera Blan¬
ca in the central Peruvian Andes. It extends from
8°50' to 1 (POO'S latitude, a distance of nearly 160
km, and has an area of 340,000 ha. Elevations
range from 3,240 m to 6,770 m, at the summit of
Nevado Huascaran Sur. The Cordillera Blanca is
the highest tropical mountain range in the world.
The geological structure of the park is complex. The
northern half is a mixture of granitic and sedimen¬
tary rocks, and the southern half is sedimentary.
The rocks of the sedimentary series are often meta¬
morphosed. During several epochs of the Pleisto¬
cene, the Cordillera was extensively glaciated, and
still has many glaciers and icefields. The nature of
the local geology and the effects of glaciation have
resulted in steep and dissected topography.
The climate of the park is a summer rainy type,
dominated by air masses coming from the east. The
annual precipitation is between 700 and 1,100 mm,
as estimated from published data (ONERN, 1972)
and unpublished weather records collected by
ElectroPeru. The mean annual temperatures are 6°
'David Smith, formerly Assistant Curator at the Missouri
Botanical Garden, died tragically in Bolivia Feb. 7, 1991,
after this manuscript was essentially complete. I have
taken the liberty of changing the name of the species to
commemorate him. — A. A. Reznicek.
7°C at 4,000 m. At that elevation and above, freez¬
ing temperatures can occur in all months, and av¬
erage minimum temperatures are below 0°C from
July to December. This general picture is modified
somewhat by the many microclimates created by
the complex topography and geology of the Cordil¬
lera Blanca.
The vegetation of the park is a complex mosaic,
including dwarf woods, shrublands, grasslands,
aquatic and semi-aquatic communities, and extreme¬
ly high Andean types. Of these, grasslands are the
most widespread. Each vegetation type includes sev¬
eral different plant communities.
Carex david-smithii Reznicek, sp. nov. TYPE:
Peru. Ancasb: Huari Prov., Huarascan Na¬
tional Park, upper terrace, Quebrada Pachach-
aca, a lateral valley of Quebrada Rurichinchay,
4,040-4,200 m, 13 June 1986, Smith , Gon¬
zales & Maldonado 12600 (holotype, I SM:
isotypes, CPUN, HUT, ISC, MO, Huascaran
National Park herbarium. Ministry of Agricul¬
ture, Huaraz). Figure 1.
Plantae rhizomatibus brevirepentibus; culmi 50- 1 30
cm alti; vaginae basales brunneae vel dilute purpuras-
centes, glabrae. Folia ca. 15-17, plerumque basalia; lam¬
inae (28-)40-100 cm longae, 6-16 mm latae; vaginae
ca. 8-35 cm longae, glabrae; ligulae ca. 4—15(—40) mm
longae. Inflorescentiae 18-32 cm longae, ramuli primarii
duplo (raro triplo) ramosi; spicae ultimae androgynae ses-
siles, 4.5-8 mm longae, 5-7.5 mm latae; bracteae infimae
laminis 29-60 cm longis, 5.5-10.5 mm latis, plerumque
evaginatis. Perigynia 2.8-3.9 mm longa, 0.9-1.4 mm
lata, patentia, valde extrorsus curvata, plano-convexa,
atrata, glabra. Achenium ca. 1.5 mm longum, ca. 1 mm
latum, juvenile. Styli marcescentes; stigmata 2 interdum
3. Antherae 3, 1.6-2.2 mm longae.
Plants forming extensive clones from stout, short-
creeping rhizomes; roots pale brown, felted with
stramineous root hairs when young; culms 50 130
cm tall, stiffly erect, trigonous, antrorsely scabrous¬
angled on upper portions, with glabrous, brown or
faintly reddish purple tinged bladeless basal sheaths
Novon 2: 433-436. 1992.
434
Novon
Figure 1. Carex david-smithii, drawn from Smith et al. 12600 (ISC), isotype. —A. Habit. —B. Portion of
inflorescence. —C. Sheath and ligule. —D. Pistillate scale. —E. Perigynium, side view. —F. Perigynium, front view.
— G. Perigynium, top view. —H. Achene, front view. —I. Achene, top view. —J. Staminate scale. —K. anther.
Bar equals 2 cm in A, 1 cm in B, C, and 1 mm in D-K.
Volume 2, Number 4
1992
Smith & Reznicek
Carex david-smithii
435
with stramineous veins. Leaves ca. 15-17, mostly
basal; blades (28 -)40-100 cm long, 6-16 mm wide,
plicate, papillose abaxially, glabrous adaxially, the
margins and keel antrorsely scabrous, widest leaves
10-16 mm wide; leaf sheaths ca. 8-35 cm long,
± loosely enveloping culms, glabrous, the lower
faintly purple tinged near base; inner band strami¬
neous or red-streaked near margins, glabrous, faintly
veined, the apex ± deeply concave, not or very
slightly thickened, purple tinged; ligules ca. 4-15
(-40) mm long, obtuse to rounded at apex, the Iree
portion stramineous to purple tinged. Vegetative
shoots unknown. Inflorescences 18-32 cm long,
upper inflorescence branches strongly overlapping,
lowest two inflorescence branches 5.5 15.5 cm dis¬
tant; single at nodes, erect to ascending on short,
stiff, trigonous, ± scabrous-angled peduncles; the
lowest inflorescence branches on peduncles 3-7.8
cm long, the upper sessile; lowermost bracts with
blades 29-60 cm long and 5.5 10.5 mm wide,
sheathless or rarely with a vestigial sheath up to 1.5
cm long, the uppermost bracts much reduced. In¬
florescences highly compounded, with third- or rare¬
ly even fourth-order branching; primary inflores¬
cence branches 6-8, the lowermost 9.5-21 cm long,
ca. 1-2 cm wide, secondary branches ca. 18 40
per primary branch, the lowermost 0.9 2.4 cm long,
ca. 0.8 1.2 cm wide, sessile or rarely with a pe¬
duncle up to 9 mm long. Ultimate branches short,
androgynous, sessile spikes; spikes 4.5-8 mm long,
5-7.5 mm wide, ovate-orbicular with (1 )3—22 per-
igynia and ca. 4 8 staminate flowers; up to ca. 25
per secondary branch. Pistillate scales 2.1-2.8 mm
long, 1.2-1.8 mm wide, ovate to broadly ovate,
acute to acuminate or sometimes with a short awn
up to 0.5 mm long, dark reddish purple to almost
black with slightly paler margins, faintly 1-5-veined.
Staminate scales 2.6 3.3 mm long, 1.2-1.6 mm
wide, elliptic to narrowly obovate, acute to acumi¬
nate, dark reddish purple to almost black with paler
margins, l-3(-5)-veined. Perigynia 2.8-3.9 mm
long, 0.9-1.4 mm wide, spreading and strongly out-
curved, ± planoconvex with lanceolate to narrowly
elliptic faces, dark reddish purple to almost black
distally, lightening to stramineous below, heavily dot¬
ted with reddish tanniferous cells, glabrous, strongly
2-veined on the margins, veinless or more faintly
1 2-veined on the faces, tapering to a bidentate
apex with teeth 0.3 0.4 mm long. Achenes ca. 1.5
mm long, ca. 1 mm wide, lenticular with broadly
obovate faces. Styles withering, ca. 3 mm long,
stigmas 2 or sometimes 3. Anthers 3, 1.6-2.2 mm
long, yellow, dotted with reddish tanniferous cells.
Paratypes. Peru, ancash: Huari Prov., Huascaran
National Park, 2 km below Cahuish Tunnel, 4,400 m,
30 Mar. 1985, Smith & Escalona 10118(C.P\JN, 1IUT,
ISC, MO, USM, Huascaran National Park herbarium);
slopes below and valley of Laguna Ichicpotrero, 4,100
m, 8 May 1986, Smith el al. f24/9(CPUN, HUT, ISC,
MICH, MO, USM, Huascaran National Park herbarium).
Most of the cited specimens of this handsome new
species are immature, and even the holotype does
not have completely fully developed achenes. Thus,
measurements of achenes and, to some extent, per-
igyia are somewhat tentative pending further col¬
lections of mature material. Another new species,
similar to Carex ancashensis but with even longer,
acute ligules, red-dotted inner bands to the sheaths,
apparently more ovate perigynia with shorter beaks
showing no tendency to reflex, and a broader inflo¬
rescence with distant lower branches, is represented
by a specimen from Quebrada Pucaraju, a lateral
valley of Quebrada Rurichinchay, Smith et al.
12698, 15 June 1986 (MO, USM), but the speci¬
men is unfortunately too immature to describe.
This species grows in habitats with saturated soils
or in shallow standing or slowly flowing water in¬
cluding boggy or marshy areas, streamsides, or lake
and pond shores. Specimens have been collected
from 4,040 to 4,400 m in elevation. It has been
found only in Huari Province, on the eastern side
of the Cordillera Blanca, where it has been collected
in the Pucavado (near the Cahuish Tunnel), Car-
huazcancha (in Laguna Ichicpotrero), and Ruri¬
chinchay valleys. All the localities are within the
Huascaran National Park. Flowering occurs in May
and June and fruiting probably from June through
August.
Carex david-smithii is most closely related to C.
catamarcensis Kiik. (including C. latehracteolata
Kiik.) of Argentina and Chile. Both are large plants
with highly compounded inflorescences of very sim¬
ilar structure composed ultimately of small, sessile,
androgynous spikes. Both have stigmas regularly
both two and three in the same inflorescence, and
perigynia heavily dotted with reddish tanniferous
cells. Carex david-smithii differs in having very
dark, almost black pistillate scales, staminate scales,
and distal portions of perigynia. Carex catamar¬
censis has stramineous to reddish purple scales and
perigynia. The perigynia of C. david-smithii are
2.8 3.9 mm long, tapered to the apex, and strongly
outcurved distally, whereas those of C. catamar¬
censis are ca. 2.2-3.5 mm long, abruptly contracted
into a short beak up to 0.6 mm long, and not
outcurved.
Because of its highly compounded inflorescences,
Kiikenthal (1909) placed Carex catamarcensis in
section Fecundae Kiik. (subgenus Carex) close to
C. fecunda Steudel, an arrangement also followed
by Barros (1935). H owever, the uniform, small.
436
Novon
sessile, androgynous spikes and lack of cladopro-
phylls show that this species and C. david-smithii
belong in subgenus f ignea (P. Beauv. ex Lestib. f.)
Peterm. (Reznicek, 1990). As well, in C. fecunda
and its allies, both the first and the larger second-
order branches are on long, flexuous peduncles. In
C. david-smithii and its allies, the first-order branch¬
es of the inflorescence have short, stiff peduncles
and all the other branches are sessile or essentially
so. Fhe sectional affinities of these species within
subgenus Vignea are as yet unclear and await a
sectional revision of the suhgenus.
Acknowledgments. Fieldwork resulting in this pa¬
per was supported by National Geographic Society
Research Grant 3069-85. We thank Susan Rezni¬
cek for the figure.
Literature Cited
Barros, M. 1935. Ciperaceas Argentinas II, generos
Kyllinga , Scirpus , Carex. Anales Mus. Nac. Hist.
Nat. Buenos Aires 38: 133-263.
Kiikenthal, G. 1909. Cyperaceae-Caricoideae. Pp. 1-
824 in A. Engler (editor), Das Pflanzenreich. IV. Vol.
20, Heft 38. Wilhelm Engelmann, Leipzig.
ONERN. 1972. Inventario, evaluacion y uso racional
de los recursos naturales de la costa: Cuencas de los
Rios Santa, Lacrarnarca y Nepena. 3 vols. Oficina
de Evaluacion de Recursos Naturales (ONERN), Lima.
Reznicek, A. A. 1990. Evolution in sedges (Carex,
Cyperaceae). Canad. J. Bot. 68: 1409-1432.
Smith, D. N. 1988. Flora and Vegetation of the Huas-
caran National Park, Ancash, Peru, with Preliminary
Taxonomic Studies for a Manual of the Flora. Doc¬
toral Dissertation, Iowa State University. University
Microfilms, Ann Arbor.
New Combinations in Gnaphalium (Asteraceae: Inuleae)
G. Ledyard Stebbins
Department of Genetics, University of California, Davis, California 95616, U.S.A.
David J. keil
Biological Sciences Department, California Polytechnic State University, San Luis Obispo,
California 93407, U.S.A.
ABSTRACT. Because of extensive morphological in¬
termediacy, plants formerly distinguished at the spe¬
cies level as Gnaphalium beneolens, G. microce-
phalum, G. thermale, and G. wrightii are treated
as intergrading races of the widespread G. canes-
cens. The new combinations, G. canescens subsp.
beneolens, G. canescens subsp. microcephalum , and
G. canescens subsp. thermale are made.
In preparing the treatment of Gnaphalium L. for
the forthcoming flora of California, Thejepson Man¬
ual, we have found that the species currently rec¬
ognized as G. beneolens A. Davidson, G. thermale
E. Nelson, G. microcephalum Nutt., and G. wrigh¬
tii A. Gray intergrade to such a degree with respect
to the characters used in current keys (Ferris. 1960;
Munz, 1959, 1968, 1974) to differentiate them—
decurrent versus nondecurrent leaf bases, nature of
tomentum, character of capitulescence, size and
shape of heads, and acute versus obtuse phyllary
tips—that they cannot consistently be separated
from each other. In addition, G. wrightii of east¬
ernmost California, Arizona, New Mexico, and
northern Mexico is indistinguishable from G. ca¬
nescens DC. of central Mexico. These entities are
therefore all merged into a single polymorphic spe¬
cies, G. canescens. Recognizable geographic ten¬
dencies in variation are maintained as subspecies.
The following new combinations are therefore pro¬
posed:
Gnaphalium canescens DC. subsp. beneolens
(A. Davidson) Stebbins & Keil. comb. nov. Gna¬
phalium beneolens A. Davidson, Bull. So. Calif.
Acad. Sci. 17; 17. 1918.
Gnaphalium canescens DC. subsp. microce¬
phalum (Nutt.) Stebbins & Keil, comb. nov.
Gnaphalium microcephalum Nutt., Trans.
Amer. Phil. Soc. n.s., 7: 404. 1841.
Gnaphalium canescens DC. subsp. thermale (E.
Nelson) Stebbins & Keil. comb. nov. Gna¬
phalium thermale E. Nelson, Bot. Gaz. (Craw-
fordsville) 30: 121. 1900.
Gnaphalium canescens subsp. canescens (in¬
cluding G. wrightii) occurs from the southwestern
United States to central Mexico. In the desert moun¬
tains of eastern California it grades into subspecies
microcephalum, a race chiefly of southern Califor¬
nia. This in turn grades into subspecies beneolens
in the South Coast Ranges of California. Plants of
the Sierra Nevada foothills are mostly assignable to
subspecies beneolens, but above l ,000 m, plants
agree more with subspecies thermale. The latter
ranges eastward to the Rocky Mountains and north¬
ward to British Columbia. Descriptions of these taxa
and their ranges are included in our treatment of
Gnaphalium in The Jepson Manual.
Literature Cited
Ferris, R. S. 1960. Compositae. Pp. 98-613 in L.
Abrams & R. S. Ferris, Illustrated Flora of the Pacific
States, vol 4. Stanford Univ. Press, Stanford.
Munz, P. A. 1959. A California Flora. Univ. California
Press, Berkeley.
-. 1968. Supplement to A California Flora. Univ.
California Press, Berkeley.
-. 1974. A Flora of Southern California. Univ.
California Press, Berkeley.
Novon 2: 437. 1992.
Notes on the Rubiaceae of Peru
Charlotte M. Taylor
Missouri Botanical Garden, P.0. Box 299, St. I.ouis, Missouri 63166-0299, U.S.A.
ABSTRACT. Raritebe axillare, Gonzalagunia mil-
dredae, and G. villosa, all of Peru and (except G.
mildredae) Ecuador, are newly described; Remijia
megistocaula is transferred to Calycophyllum; and
a key is presented to the eight species of Gonza¬
lagunia found in Peru.
These previously undescribed species and the need
for the nomenclatural transfer were discovered dur¬
ing study of specimens recently collected in Peru.
Notes on Calycopuylu m DC.
Calycophyllum megistocaulum (K. Krause) C.
M. Taylor, comb. nov. Basionym: Remijia
megistocaula K. Krause, Bot. Jahrb. Syst. 40.
319. 1908. TYPE: Peru. Huanuco: between
Monzon and Rio Huallaga, W'eberbauer 3687
(holotype, B destroyed; photographs (F negs.
#161, #162), MO).
Calycophyllum acreanum Ducke, Arq. Inst. Biol. Veg.
2: 709. 1935. TYPE: Brazil. Acre: Rio Acre, Ser¬
inga! Iracema, 18 Mar. 1933, Ducke HJBR-24414
(holotype, RB not seen; isotype, P not seen, pho¬
tograph (F neg. # 37200), MO).
Examination of the type photographs shows that
these names represent the same species. The leaves
on the type collection of Calycophyllum acreanum
appear discolored, hut in other respects (size, shape,
and venation pattern) they agree with the photo¬
graph of Remijia megistocaula.
I his species was first described in Remijia DC.,
and was treated in that genus without comment by
Standley (1936), who relied primarily on capsule
dehiscence to distinguish Remijia. However, R.
megistocaula differs from other Remijia species in
its terminal inflorescences, rather than axillary, and
its corollas with shortly funnelform tubes that are
pilose internally and lobes that are convolute in
aestivation, rather than elongated cylindrical tubes
that are glabrous internally and valvate lobes. These
unusual features of R. megistocaula are all char¬
acteristic of Calycophyllum. The capsules with sep-
ticidal dehiscence and valves that split secondarily
from the apex, features that Standley used to char¬
acterize Remijia , are also characteristic of Caly¬
cophyllum. Other features used to characterize Ca¬
lycophyllum are enlarged bracts that completely
enclose developing cymules of flowers, and the pres¬
ence of expanded petaloid calyx lobes that function
as semaphylls (Steyermark, 1974). The enlarged
bracts are found in several species of Calycophyl¬
lum, e.g., C. candidissimum (M. Vahl) DC., C.
multijlorum Griseb., C. spruceanum (Benth.)
Schum., but not in all, e.g., lacking in C. obovatum
(Ducke) Ducke and C. venezuelense Steyerm. Sim¬
ilarly, the expanded calyx lobes are found in some
species, e.g., C. candidissimum, C. obovatum, and
C. venezuelense, but not all, e.g., lacking in C.
multijlorum and C. spruceanum. The size, form,
and texture of the corollas and capsules of Remijia
megistocaula are generally similar to those of Ca¬
lycophyllum. Thus the species seems best placed
in this genus. Ducke commented in his description
of C. acreanum that while he had no doubts about
its generic placement, his new species was somewhat
unusual morphologically within Calycophyllum.
Notes on Raritebe Wernham
The characteristics of Raritebe were described
in detail by Kirkbride (1979). 'These must now be
emended to include axillary as well as terminal in¬
florescences. Kirkbride recognized ordy one variable
species, R. palicoureoides Wernham, although this
may be overly conservative in Central America (cf.
Dwyer, 1980). I he generic name is here treated as
neuter, following Kirkbride (1979); usage by Dwyer
(1980) bas varied.
Raritebe axillare C. M. Taylor, sp. nov. TYPE:
Ecuador. Esmeraldas: Rio Zapallo Grande (trib¬
utary of Rio Cayapa), trail from riverbank op¬
posite to the evangelic mission station, 00°48'N,
78°52'W, 25 Oct. 1982, (imm. fl), Barfod
41055 (holotype, MO; isotype, AAU not seen).
Figure l.
Haec species a congeneris omnibus inflorescentiis ax-
illaribus differt.
Erect, small trees to 3 m tall, strigillose becoming
glabrescent. Leaves opposite; blades elliptic, 16-28
cm long, 7-9.5 cm wide, at apex acute to slightly
acuminate and often falcate, at base acute, char-
Novon 2: 438-442. 1992.
Volume 2, Number 4
1992
Taylor
Rubiaceae of Peru
439
Figure 1. Raritebe axillare C. M. Taylor. —A. Habit. —B. Infructescence. Based on Barfod 41055 (MO).
taceous, on both surfaces glabrous or usually sparse¬
ly to moderately strigillose along midrib and sec¬
ondary veins; secondary veins 11-16 pairs, slightly
to strongly interconnected to form an undulating
submarginal vein, without domatia; petioles 1.5 4
cm long, similar to leaf midrib in pubescence; stip¬
ules persistent on distal 2-3 nodes, chartaceous,
triangular, 3-10 mm long, acute. Inflorescences
axillary, 1-2 per node, open cymose, with peduncles
1- 5 cm long, with thyrses 2-10 cm long, 2-8 cm
wide, with bracts triangular, to 1 mm long, with
flowers ca. 15-30 in cymes of 2-5, with pedicels
2- 5 mm long; branches, bracts, and pedicels green,
strigillose or appressed-pilosulous. Flowers (imma¬
ture) with hypanthium cupuliform to turbinate, stri¬
gillose, ca. 1-2 mm long; calyx limb 1-1.5 mm
long, subtruncate to denticulate with 5 teeth; co¬
rollas salverform, white, externally puberulent to
glabrous, lobes 5, narrowly triangular to linear. In-
fructescences similar to inflorescences; fruit subglo-
bose, smooth, 3-4 mm long, 4-4.5 mm wide; seeds
subglobose, ca. 0.1 mm diam., with reticulate testa.
Distribution. Coastal Ecuador and Amazonian
Peru, in wet forest at 200 250 m elevation.
This species is placed in Haritebe based on its
lack of raphides, paniculate-thyrsoid inflorescences
with cymose branches, flowers with 5-lobed calyx
and corollas, corollas with valvate, reduplicate aes¬
tivation and narrow acute shape in bud, baccate
fruit, and numerous seeds with testa structure similar
to that shown by Garcia Kirkbride (1979: fig. 10)
(Kirkbride, 1979). This new species is sympatric
with and similar to K. palicoureoides, from which
it differs in its axillary rather than terminal inflo¬
rescences. The specific epithet relers to this dis¬
tinction.
Although several specimens of Raritebe pali¬
coureoides show overtopping of old infructescenc.es
by vegetative growth from one axil, the plants de¬
scribed here clearly have axillary inflorescences, as
demonstrated by production of inflorescences si¬
multaneously from both axils of a node. These in¬
florescences are produced both with and proximal
to healthy leaves, and originate from enlarged struc¬
tures that suggest that the production of inflores¬
cences in these axils is indeterminate. Variation is
seen in development of a submarginal vein; similar
variation in all leaf characters is seen in R. pali¬
coureoides. The data accompanying the type col¬
lection indicate that the plant is used locally in the
treatment of “headaches, fever, and body pains”
by patting the body with the plant. Data accom¬
panying the second collection indicate that the plant
is not used medicinally in that area.
Paratypes. Peru, AMAZONAS: Rio Cenepa, vicinity of
Huampami, hill above Quebrada Tujushik, ca. 5 km E
of Chavez Valdivia, 4°30'S, 78°30’W, 24 July 1978 (fr),
Ancuash 1081 (MO).
Notes on Gonzalaguma Rufz Lopez & Pavon
Gonzalagunia has not been treated comprehen¬
sively recently; its last review for Peru was by Stan-
dley (1936). The following key is offered as an
outline of this genus in Peru and as a guide to the
identification of the species newly described here.
Key to Species of Gonzalagi xia in Peru
la. Leaf blades tomentose to floccose on lower surface of lamina, the trichomes generally white.
2a. Leaf blades with sparse pubescence of lower surface of lamina spreading and dull white.
. G. dependens Ruiz Lopez & Pavon
2b. Leaf blades with pubescence of lower surface of lamina appressed and bright white .
. G. chionea Standley ex Steyerm.
lb. Leaf blades glabrous to pilosulous on lower surface of lamina, the trichomes generally colorless to brown.
440
Novon
3a. Leaves subsessile, blades ovate, rounded to somewhat cordate at base .
. G. mildredae Simpson ex C. M. Taylor
3b. Leaves petiolate, blades elliptic to lanceolate, obtuse or cuneate to acute at base.
4a. Stems and lower leaf surfaces spreading-pilosulous, at least along midrib.
5a. Stipules with short base 1-2 mm long, abruptly contracted to a very narrowly triangular
apex 2-10 mm long; corollas with tubes 5-6 mm long; fruit with 4 cocci .
. G. villosa Simpson ex C. M. Taylor
5b. Stipules triangular, uniformly tapered, 10-20 mm long; corollas with tubes 6.5-8 mm long;
fruit with 2 cocci. G. pachystachya Standley
4b. Stems and lower leaf surfaces appressed-strigose to -strigillose.
6a. Corolla tubes 7-8 mm long, lobes 5 6 mm long; pedicels 1-4 mm long .
. G. aflinis Standley ex Steyerm.
6b. Corolla tubes 4-6 mm long, lobes 2 3 mm long; pedicels to 1.5 mm long.
7a. Leaf blades 1.5-5.5 cm wide . G. cornifolia (HBK) Standley
7b. Leaf blades 6-15 cm wide. G. bunchosioides Standley
Gonzalagunia mildredae D. Simpson ex C. M.
Taylor, sp. nov. TYPE: Peru. Huanuco: Prov.
Leoncio Prado, distrito Rupa Rupa, road to
Jacintillo, left bank of Rio Monzon, 20 Mar.
1972 (H), Schunke 5293 (holotype, MO; iso¬
type, F). Figure 2.
Gonzalaguniac sessilifoliae Standley similis, sed ab
ea stipulis longioribus (15-20 mm longis), lobulis calycinis
aequalibus ca. 0.8 mm longis et tubo corollirto longiore
(8-9 mm longo) differt.
Erect shrubs or small trees to 12 m tall, strigillose
to usually strigose throughout. Leaves opposite, sub-
sessile; blades ovate, 6 14 cm long, 2.5-6.5 cm
wide, at apex long-acute to slightly acuminate, at
base rounded to somewhat cordate, chartaceous to
somewhat subcoriaceous, above glabrous to sparsely
strigillose, below strigillose along the veins but gla¬
brous on the remaining surfaces; secondary veins
8-10 pairs, basal 2-4 veins originating from petiole
attachment, without domatia; petioles 0 2 mm long,
similar to leaf midrib in pubescence; stipules per¬
sistent, membranaceous, strigillose to glabrescent,
lanceolate to triangular, 15 20 mm long, acute to
slightly acuminate, sometimes with a thickened mid¬
rib. Inflorescences spiciform, with peduncles 3-6
cm long, with thyrses 20 33 cm long, 1-2 cm wide,
with bracts narrowly triangular to linear, 2-5 mm
long, with flowers ca. 40 60 in sessile cymules of
1 -3, these separated 3-10 mm along axis, with
pedicels 1-3 mm long; axis, peduncle, bracts, and
pedicels green, densely strigillose; flowers with hy-
panthium cupuliform, densely strigillose to pilosu-
lous, 1-1.5 mm long; calyx limb glabrous, 0.8 1
mm long, lobed for ca. l A its length, lobes 4, rounded,
wi th sinuses rounded; corollas salverform, white,
externally densely appressed-pilose to more sparsely
so on lobes, internally pilosulous at apex of tube
with trichomes included, tubes 8 9 mm long, lobes
4, lanceolate, 4-5 mm long, 3-4 mm wide, rounded;
anthers very narrowly oblong, 2.1 2.2 mm long.
Infructescences similar to inflorescences; fruit sub-
globose to oblate, with 4 cocci, 4-5 mm long, 5-6
mm wide when young, becoming white and 8 mm
long, 9 mm wide.
Distribution. Amazonian Peru, in wet forests at
680-1,100 m elevation.
This species is distinguished by its subsessile ovate
leaf blades with the basal 2-4 veins originating from
the petiole attachment and the base rounded to
usually subcordate. In this respect it is similar to
Gonzalagunia sessilifolia Standley of Ecuador,
which can he separated by its broader leaves, 5-9
cm wide, of a markedly more membranaceous tex¬
ture; its shorter stipules, 5-10 mm long, that are
narrowly triangular to subulate; its longer and mark¬
edly unequal calyx lobes, 1-3 mm long on a single
flower; and its smaller corollas, with tubes 2.5-3
mm long and lobes 1-1.5 mm long.
Gonzalagunia mildredae was separated as a new
species and its name chosen by Donald Simpson,
formerly of the Field Museum of Natural History,
who unfortunately did not publish his work. The
name and the type specimen he selected are used
here. The specific epithet honors Mildred Mathias,
an outstanding student of the South American flora.
Paratypes. PERU. HUANUCO: Prov. Leoncio Prado, Tin-
go Maria, 3 Aug. 1964 (fr), Dwyer 6283 (MO); distrito
Rupa Rupa, road from Tingo Maria to Lechuzas Cave,
9°18'S, 75°59'W, 24 Mar. 1982 (fl). King & Ramirez
Rengifo 369 (MO); vicinity of Tingo Maria, trail from
airport bridge to Monzon-Huallaga junction, 21 June
1960 (fl, fr), Mathias & I). Taylor 4063 (F, MO). UCAYALI:
Prov. Coronel Portillo, distrito Padre Abad, Rio Chino W
of Acapulco Restaurant, 6 June 1976 (fl, fr), Seliunke
9164 (F, MO).
Gonzalagunia villosa D. Simpson ex C. M. Tay¬
lor, sp. nov. TYPE: Peru. Ucayali: Prov. Co¬
ronel Portillo, distrito Galleria, Carretera Pu-
callpa Km 90, San Alejandro, 21 Feb. 1972
Volume 2, Number 4
1992
Taylor
Rubiaceae of Peru
441
Figure 2. Gonzalagunia mildredae D. Simpson ex C.
M. Taylor. —A. Habit. —B. Flower. Habit based on
King & Ramirez 369 (MO), flower on Mathias & Taylor
4063 (MO).
(fl, fr), Schunke 5272 (holotype, MO; isotype,
F). Figure 3.
Haec species a parte majore congenerorum indurnento
pilosulo dense etiam a Gonzalagunia pachystachya Stan-
dley corollae tubo 5-6 mm longo et lobulis 3-4 mm longis
instructa differt.
Erect shrubs or small trees to 9 m tall, densely
pilosulous throughout with trichomes spreading and
usually drying brown. Leaves opposite or occasion¬
ally ternate; blades elliptic to lanceolate, 5-28 cm
long, 2-11 cm wide, at apex acute to slightly acu¬
minate, at base acute to cuneate, chartaceous to
membranaceous, above glahrescent to sparsely ap-
pressed-pilosulous or sometimes more densely so on
midrib, below spreading-pilosulous; secondary veins
7-13 pairs, without domatia, with midrib frequently
pale or reddened below; petioles 2-20 mm long,
similar to leaf midrib in pubescence; stipules per¬
sistent, chartaceous, pilosulous, triangular with basal
portion 1—2 mm long and abruptly contracted to a
narrowly triangular to linear tip 2-10 mm long.
Inflorescences spiciform, with peduncles 1.5-3.5
cm long, with thyrses 15 30 cm long, 12 cm wide,
with bracts linear, to 3 mm long, with flowers ca.
60-100 in sessile cymules of 2-5, these separated
3-10 mm along axis, with pedicels 0.5-2.5 mm
long; axis, peduncle, bracts, and pedicels green,
densely strigillose to pilosulous; flowers with hypan-
thium ellipsoid, densely strigillose to pilosulous, 1
1.5 mm long; calyx limb glabrous, 11.5 mm long,
lobed nearly to base, lobes 4, ovate to elliptic, fre¬
quently unequal, rounded, with sinuses acute; co¬
rollas salverform, white, externally moderately to
densely strigillose to often sparsely so on lobes, in¬
Figure 3. Gonzalagunia villosa D. Simpson ex C. M.
Taylor. —A. Habit, smaller leaf showing abaxial surface.
— B. Flower. Based on Schunke 5272 (MO).
ternally pilosulous at apex of tube with trichomes
included, tube 5-6 mm long, lobes 4, triangular, 3
4 mm long, 1-1.2 mm wide, rounded to obtusely
angled, lnfructescences similar to inflorescences; fruit
subglobose to oblate, with 4 cocci, 2-2.5 mm long,
3-4 mm wide, becoming white.
Distribution. Amazonian Ecuador and Peru, in
wet forests at 200-450 m elevation.
Gonzalagunia villosa is distinguished by its dense
pilosulous pubescence, relatively small stipules with
narrowly triangular apices, and relatively short co¬
rollas. Gonzalagunia pachystachya Standley of
Ecuador and Peru is similarly pilosulous, but differs
in its triangular stipules that are not strongly nar¬
rowed at the apex, corollas with tubes 6.5 8 mm
long and lobes 5-6 mm long, and fruit with 2 cocci
and 3-4 mm long by 4-6 mm wide. The specific
epithet refers to the characteristic pubescence.
The flowers of Gonzalagunia villosa are re¬
portedly fragrant. This species shows wide variation
in leaf size. The type specimen has leaves that gen¬
erally represent the smaller end of the range. Similar
variation in vegetative features is seen in many, if
not most, species ol Gonzalagunia , which charac¬
teristically are found in secondary habitats, although
the labels of G. villosa generally attribute it to dense
or mature forest.
Paratypes. ECUADOR. WARD: Jatun Sacha Biological
Station, 8 km E downstream of Puerto Misahualli, on the
Rio Napo, 1°04'S, 77°37'W, 28 Mar. 1990(fl, fr), Bens-
man 80 (MO); Coco, road to Las Areas, spur trail to N,
23 July 1977 (fl), Boeke 2221 (MO); along road SE of
Francisco de Orellano (Coco) on the way to El Auca, 14.6
442
Novon
km past bridge over the Rio Napo, 00°37'S, 76°40'W,
5 Oct. 1980 (fl), Croat 50393 (MO). Peru, amazonas:
SF, of Huampami, trail to Aintami, 17 Dec. 1972 (fl),
Merlin 587 (F, MO); ridge above Quebrada Chikishuk, a
tributary of Huampami, entering from S about 5 km from
confluence with Rio Cenepa, 21 Dec. 1972 (fl), Merlin
053 (F, MO); S of Huampami, trail to house of Theodora,
S of Rio Cenepa, 17 July 1974, Merlin 1083 (F, MO);
Huambisa, valle del Rio Santiago, ca. 65 km al N de
Pingo, Quebrada Caterpiza, 2 3 km atras de la comunidad
de Caterpiza, 19 Sep. 1979 (fr), Huashikat 087 (MO),
11 Feb. 1980 (fl), Huashikat 2039 (MO); Aintami, al
lado del Rio Cenepa, 14 Feb. 1973 (fl, fr), Kayap 357
(F, MO); Huampami, Rio Cenepa, 15 June 1973 (fl, fr),
Kayap 974 (F, MO); Quebrada Satik entsa, 16 July 1974
(fl, fr), Kayap 1111 ( F, MO); al lado del Rio Huampami,
18 July 1974 (fr), Kayap 1220 (F, MO); Rio Cenepa,
vicinity of Huampami, ca. 5 km E of Chavez Valdivia,
10 km en camino de Chigkan entsa, 4°30'S, 78°30'W,
8 Aug. 1978(fr), Kujikat /99(MO); Rio Cenepa, vicinity
of Huampami, ca. 5 km E ot Chavez Valdivia, al lado de
Kachaim, Quebrada Huampami, 4°30'S, 78°30'W, 15
Aug. 1978 (fl), Kujikat 391 (F, MO). SAN MARTIN: Prov.
Lamas, distrito Lamas, N of San Antonio 2-4 km along
Rio Cumbasa, 2 Oct. 4 Nov. 1937 (fl, fr), Melshaw 3529
(MO). UCAYALI: Padre Ahad, Pucallpa, Irezola, Km 79
camino Francisco Basache, 10 Apr. 1989 (fr), h. Chavez
330 (MO).
Acknowledgments. I thank Lennart Andersson
and anonymous reviewers for helpful comments, and
Roy E. Gereau for preparation of the Latin diag¬
noses. 1 am particularly indebted to Amy Pool, who
checked nomenclatural points, and Donald Simpson,
who pointed the way in Gonzalagunia.
Literature Cited
Dwyer, J. I). 1980. Rubiaceae. In: Flora of Panama.
Ann. Missouri But. Card. 67: 1-522.
Garcia Kirkbride, M. C. 1979. Review of neotropical
Isertieae. Brittonia 31: 313-332.
Kirkbride, J. H., Jr. 1979. Raritebe, an overlooked
genus of Central America. Brittonia 31: 299-312.
Standley, P. C. 1936. Rubiaceae. In: Flora of Peru.
Field Mus. Nat. Hist., Bot. Ser. 13(6): 1 261.
Steyermark, J. A. 1974. Rubiaceae. In: Flora de Ven¬
ezuela 9: 1 2070.
A New Species of Puya (Bromeliaceae) from Central Peru
Walter Till
Institut fiir Botanik, Universitat Wien, Bennweg 14, A-1030 Wien, Austria
Abstract. Puya lutheri is described as a new
species of Bromeliaceae: Pitcairnioideae. The di¬
agnostic characteristics distinguishing it from P.fos-
teriana, the closest related species, are given.
While studying specimens of Bromeliaceae from
ISC, I was unable to determine one of Puya with
the available literature (e.g., Smith & Downs, 1974).
The collector wrote “Puya aff. elava-hereulis ,”
while Harry Luther from SEL stated “Puya sp. nov.,
aff. pygmaea L. B. Sm. et P. cryptantha Cuatr.”
I agree with Luther that the present specimen res-
presents a new species, but disagree in its alliance.
After a critical check of many possibilities in the
Flora Neotropica keys, and after comparison with
the descriptions, respectively, I am convinced that
the new species is allied to P. fosteriana L. B. Smith
because ol the overall similarity of both species (see
also Foster, 1984). Using subkey IV of Smith &
Downs’s monograph (1974) and applying the given
characteristics rigorously, the present specimen keys
out as P. trianae Baker from Colombia. But if one
assumes that the “outer bracts conspicuously ser¬
rate,” is instead “entire or very obscurely serrate”
in a closely related species, one reaches P. foster¬
iana L. B. Smith (1950). The serration of bracts is
not a characteristic to infer natural alliances, but
only a tool in an artificial key to separate taxa.
Puya lutheri W. Till, sp. nov. TYPE: Peru. An-
cash: Yungay Province, Huascaran National
Park. Quebrada Huaripampa between Quebra-
da Paria and Moroeocha, 77°33'W, 8°56'S,
alt. 3,930-4,500 m, 13 Jan. 1985, Smith,
Sanchez & Vidaurre 9196 (hclotype, ISC
376409; isotype, MO not seen). Figure 1.
A P. fosteriana L. B. Smith vaginis foliorum angus-
tioribus, laminis foliorum angustioribus subtus adpresso
lepidotis spinis brevioribus, scapo lorigiori, inflorescentia
simplici minore angustioribusque albido-lanuginosa, sepalis
minoribus et petalis angustioribus pallide coeruleis differt.
About 1 m tall (according to Smith); leaves to 50
cm long; sheaths broadly triangular ovate, ca. 5 cm
long and 5 cm wide, pale stramineous and glabrous
on both faces, the margins entire except for the
short spinulose transition with the blades; blades
narrowly triangular, 25-30 mm wide at the base,
evenly tapering to a point, glabrous above, densely
pale appressed-lepidote beneath, the margins laxly
but coarsely antrorse-spinose, spines dark brown,
strictly to slightly curved, 4 6 mm long; scape un¬
known but estimated from the complete height to
be ca. 60 cm long, 15 mm diam. below the inflo¬
rescence; scape bracts unknown; inflorescence ca.
20 cm long, ca. 5 cm diam., dense, simple, whitish
lanuginose; floral bracts covering the flowers except
for the petals, their sheaths ovate, ca. 6 cm long
and 3 cm wide, membranaceous, brownish when
dry, densely white lanuginose, the margins entire,
their blades long-acuminate, laxly short-spinose, and
reflexed, glabrescent; sepals linear-lanceolate, broadly
acute, ca. 27 mm long and 5 mm wide, rigidly
membranaceous, adaxially glabrous, abaxially densely
whitish lanuginose, free, the posterior ones slightly
carinate; petals spathulate, ca. 45 mm long, un-
appendaged, their blades about 9 mm wide, pale
blue (according to Smith). Anthers versatile, 5 mm
long and 1 mm diam., dark brown when dry. Ovary
conical, 10 mm long and 3 mm diam., style slender,
20 mm long, stigma lobes twisted.
Terrestrial in grassland with rare scattered shrubs
or stands of Polylepis sp., “Queshque” bank of
rivulet between two moraines.
Puya lutheri differs from P. fosteriana in its
narrower leaf sheaths and blades (the blades in P.
lutheri being appressed-lepidote beneath), the short¬
er leaf spines, the longer scape, the simple, smaller
and narrower, whitish lanuginose inflorescence, the
smaller sepals, and the narrower, pale blue petals.
Puya lutheri and P. fosteriana both occur at high
elevations (Foster, 1984). The populations of the
species are separated by a distance of about 1,400
km. I his fact should not be over-emphasized, as a
similar disjunction is found in Puya raimondii Harms.
I he species is dedicated to Harry E. Luther from
the Bromeliad Identification Center at the Marie
Selby Botanical Gardens, Sarasota, Florida, who first
recognized it as a novelty.
Acknowledgments. I thank Marshall K. Crosby
(MO) and two anonymous reviewers for critically
reading the manuscript and offering valuable sug¬
gestions.
Novon 2: 443-444. 1992.
444
Novon
■ ^
mi
' ilAU'
tj January 1985
. Vldaurra 9196
Torrostr lal , 1m totul ficlgrtt; I vs to
50cmi Inti 5cm dlo at basoi tls pal«
blua. 'Ouoshquo' Bank of rlvulut bot.can
two moraines.
Bronellacaae - • c s
Puya ff. f
M-. ilTf
Anc.ish. Yungay Prov, Huascaran National
Park* Cuebrada Huarlpanpa between Quo-
brada Parla ind Morococha <77*33’W*
• 56 * S) 3 93 0-4 50 0m. Grassland with r ro
attorod shrubs or stands of Polylopls.
Figure 1. Puya lutheri W. Till, holotype (ISC 376409). Scale bar — 5 cm.
Literature Cited
Foster, M. B. 1984. Puya, the Pineapple’s Andean
ancestor. J. Bromeliad Soc. 34: 147 157.
Smith, L. B. 1950. A new Puya from Bolivia. J. \\ ash-
ington Acad. Sci. 40: 217.
- & R. J. Downs. 1974. Pitcairnioideae (Bro-
meliaceae). In: Flora Neotropica Monograph 14. Haf-
ner Press, New York.
Innovations in the Fern Genus Phanerophlebia
George Yatskievych
Missouri Botanical Garden, I’.O. Box 299, St. Louis, Missouri 63166, U.S.A.
ABSTRACT. Phanerophlebia gastonyi, a new Mex¬
ican species, is described, and P. remotispora is
reduced to varietal status under P. nobilis.
The genus Phanerophlebia (Dryopteridaceae)
comprises eight species of terrestrial ferns ot the
New World tropics and adjacent subtropical regions.
It was last revised taxonomically by Underwood
(1899) and Maxon (1912), whose studies were based
on the relatively few herbarium specimens in exis¬
tence at that time. A recently completed doctoral
dissertation (Yatskievych, 1990) involving biosys-
tematic studies of this genus has not been fully
published, and a taxonomic revision based upon this
research is currently in preparation. Evidence from
several independent experimental data sources can
be correlated with the results of a morphological
study of the wealth of recently collected specimens
of Phanerophlebia to provide a clearer picture of
species limits in the group.
The present note describes a new species and
makes a new combination in Phanerophlebia. These
names are needed for the forthcoming pteridophyte
volume ol Flora Mesoamericana.
Phanerophlebia gastonyi Yatskievych, sp. nov.
TYPE: Mexico. Ca. 13 km NW of Berriozabal,
limestone outcrops in evergreen tropical hard¬
wood forest, elevation 1,000 m. 15 July 1985,
Yatskievych , Gonzalez G. Hanker & C.
Starr H5-IH2 (holotype, MO; isotypes, AR1Z,
CHAPA, MENU, INI), NY, l AMI/.. UC).
Species habitu Phanerophlebiae juglandifoliae (Hum¬
boldt et Bonpland ex Willdenow) J. Smith simillima, sed
differt statura minora; pinnis 1 -3(-4)-jugatis; paleis ad
basem stipitis non imbricatis, bicoloribus, centro lato ni-
grescentibus, margine angusta pallidis; sporis 30-42 /uin
longis; chromosomatum numero n = 41.
Rhizomes to ca. 5 mm diam., short repent. Rhi¬
zome scales 2.5-4.5 mm long, 1.5-2 mm wide,
ovate to elliptic-lanceolate, ciliate, bicolorous with
broad, dark, sclerotic centers and narrow hyaline
margins. Leaves to 45(-55) cm long, the petiole
slightly shorter than to usually longer than lamina.
Petiolar scales usually deciduous with age, most Iy
not overlapping, much like rhizome scales, the
broadest 2 mm wide, the reduced hairlike scales
uncommon among the broader scales. Pinnae I
3(-4) pairs, to 12(-15.5) cm long, ovate to lance-
ovate, sometimes somewhat falcate, the apex acute
to attenuate, the base obliquely cuneate to rounded,
sometimes nearly equilateral, the margins entire to
slightly undulate proximally, spinulose-serrulate only
in distal hall. Gemmiferous buds absent from axils
of distal pinnae. Secondary veins with 1-3 series of
regular marginal anastomoses, 2-4-branched. Sori
in (1)2-4 series between costa and margin, exin-
dusiate. Spores monolete, 30-42 gm long, n = 41.
Illustration. Mickel & Beitel (1988, as P. jug-
landifolia).
Distribution. Moist soil, less commonly on fis¬
sured limestone rock faces, in cloud forests and
montane rainforests; 900-2,200 m. Southern Mex¬
ico (Chiapas, Oaxaca, Veracruz).
This species is named for the eminent American
pteridologist, Gerald J. Gastony, whose advice and
assistance has nurtured my developing interest in
ferns for many years. It is an infrequently collected
taxon and is known from very few sites.
Specimens of P. gastonyi have previously been
identified as P. juglandifolia. The latter species
occurs from southern Mexico to northern Venezuela
and Colombia. The two species are very similar
morphologically, but can be differentiated by die
characters in the following key:
la. Petiolar scales not overlapping, 4-6 mm long,
with broad, dark, sclerotic centers and narrow,
hyaline margins; gemmiferous buds absent;
spores 30-42 /mi long . . Phanerophlebia gastonyi
lb. Petiolar scales usually overlapping, 6-11 mm
long, with dark brown central stripes and lighter
margins of about equal width; gemmiferous buds
often present on rachis in axils of distal pinnae;
spores 41-60 /an long .
. Phanerophlebia juglandifolia
Yatskievych & Gastony (1987) discussed mor¬
phological and allozymic evidence that P. juglan¬
difolia is an allotetraploid with diploid l\ gastonyi
as one of its parental taxa. They speculated that the
diploid P. macrosora (Baker) L. Underwood might
be the other progenitor, but were unable to confirm
this. Yatskievych et al. (1988) used restriction site
analysis of the chloroplast genome to confirm that
P. gastonyi is a maternal parent of P. juglandifolia.
Novon 2: 445-446. 1992.
446
Novon
but also were unable to verify the identity of the
other parental taxon.
Paratypes. MEXICO. CHIAPAS: 13 kin N of Berriozabal,
2 Nov. 1971, Breedlove & Smith 2/652(DS, F, MICH,
NY), oaxaca: trail from San Pedro Nolasco N to the
Llano Verde, 5 Oct. 1969, A lickel & Hellwig 3786B
(NY). VERACRUZ: ca. 12 kin S of Misantla, 3 Jan. 1974,
Conant 726 (GH, MEXU).
Phanerophlebia nob i I is (Schlechtendal &
Chamisso) C. Presl var. remotispora (Four¬
nier) Yatskievych, comb, et stat. nov. Pha¬
nerophlebia remotispora Fournier, Mexic. PI.
1: 100, 1872. Cyrtomium remotisporum
(Fournier) C. Morton, Amer. Fern J. 47: 54,
1957. TYPE: Mexico. Veracruz: Orizaba,
Izhuatlancillo, Aug. 1805-1866, Bourgeon
23 19 (holotype, P; isotypes, BM, GH, k, MICH,
MO, NY, I’, US).
Previous authors (cl. Tryon & Tryon, 1982) uni¬
formly have treated variety remotispora as a distinct
species. These authors emphasized anastomosing
versus free venation patterns as a critical character
in subdivision of Phanerophlebia into two species
groups. Yatskievych et al. (1988) showed that P.
n obi Us and P. remotispora are genetically very
closely related, based on analysis of ehloroplast-DNA
restriction-site variation, and suggested that anas¬
tomosing veins were probably derived twice inde¬
pendently within the genus.
Variety remotispora differs from typical P. no-
bilis only in its venation pattern, with copious mar¬
ginal anastomoses, rather than the free veins found
in variety nobilis. The two varieties are sympatric
in much of Mexico, although they are seldom en¬
countered in mixed populations. Rare specimens ot
intermediate morphology (veins with few, irregular
anastomoses) originate from Michoaean ( Arsene
5961; GH, MO, US) and Veracruz (Fisher s.n
MO), Mexico, and are apparently fully lertile.
Literature Cited
Maxon, W. R. 1912. Notes on the North American
species of Phanerophlebia. Bull. Torrey Bot. Club
39: 23-28.
Mickel, J. T. & J. M. Beitel. 1988. Pteridophyte flora
of Oaxaca, Mexico. Mem. New York Bot. Card. 46:
1-568.
Tryon, R. M. & A. F. Tryon. 1982. Ferns and Allied
Plants, With Special Reference to Tropical America.
Springer-Verlag, New York.
Underwood, L. M. 1899. American ferns—II. The
genus Phanerophlebia. Bull. Torrey Bot. Club 26:
205-215, 2 pi.
Yatskievych, G. 1990. Systematics of the Fern Genus
Cyrtomium in the New World. Ph.D. Dissertation,
Indiana University, Bloomington.
- & G. J. Gastony. 1987. Hybridization and
polyploidy in the fern genus Phanerophlebia. Amer.
J. Bot. 74: 715. [Abstract.]
-, D. B. Stein & G. J. Gastony. 1988. Chloro-
plast DNA evolution and systematics of Phanero¬
phlebia and related fern genera. Proc. Natl. Acad.
U.S.A. 85: 2589-2593.
Volume 2, Number 1, pp. 1-80 of NOVON was published on 18 March 1992.
Volume 2, Number 2, pp. 81-178 of NOVON was published on 9 June 1992.
Volume 2, Number 3, pp. 179-284 of NOVON was published on 13 October 1992.
Volume 2, Number 4, pp. 285-446 of NOVON was published on 22 December 1992.
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